PhytoKeys 59: 1–828 (2016)
doi: 10.3897/phytokeys.59.6261
http://phytokeys.pensoft.net
World checklist of hornworts and liverworts
A peer-reviewed open-access journal
1
CHECKLIST
Launched to accelerate biodiversity research
World checklist of hornworts and liverworts
Lars Söderström1,*, Anders Hagborg2,*, Matt von Konrat2,*,
Sharon Bartholomew-Began3, David Bell4, Laura Briscoe2, Elizabeth Brown5, †,
D. Christine Cargill6, Denise P. Costa7, Barbara J. Crandall-Stotler8,
Endymion D. Cooper9, Gregorio Dauphin10, John J. Engel2, Kathrin Feldberg11,
David Glenny12, S. Robbert Gradstein13, Xiaolan He14, Jochen Heinrichs11,
Jörn Hentschel15, Anna Luiza Ilkiu-Borges16, Tomoyuki Katagiri17,
Nadezhda A. Konstantinova18, Juan Larraín2, David G. Long19, Martin Nebel20,
Tamás Pócs21, Felisa Puche22, Elena Reiner-Drehwald23, Matt A.M. Renner5,
Andrea Sass-Gyarmati21, Alfons Schäfer-Verwimp24, José Gabriel Segarra Moragues25,
Raymond E. Stotler8, †, Phiangphak Sukkharak26, Barbara M. hiers27,
Jaime Uribe28, Jiří Váňa29, Juan Carlos Villarreal30, Martin Wigginton31,
Li Zhang32, Rui-Liang Zhu33
1 Department of Biology, Norwegian University of Science and Technology, N-7491 Trondheim, Norway
2 Department of Science and Education, Field Museum, 1400 South Lake Shore Drive, Chicago, IL 60605–
2496, United States of America 3 Department of Biology, West Chester University, West Chester, PA 19383,
United States of America 4 Department of Botany, University of British Columbia, 6270 University Boulevard,
Vancouver, BC, V6T 1Z4, Canada 5 Royal Botanic Gardens and Domain Trust, Mrs Macquaries Road,
Sydney NSW2000, Australia 6 Centre for Australian National Biodiversity Research, Australian National
Herbarium, GPO Box 1600, Canberra, ACT 2601, Australia 7 Instituto de Pesquisas Jardim Botânico do
Rio de Janeiro, Rua Pacheco Leão 915, 22460-030, Rio de Janeiro, RJ, Brazil 8 Department of Plant Biology,
Southern Illinois University, Carbondale, Illinois 62901-6509, United States of America 9 CMNS-Cell
Biology and Molecular Genetics, 2107 Bioscience Research Building, University of Maryland, College Park,
MD 20742-4451, United States of America 10 Apartado 5-1500, Acosta, Costa Rica 11 Systematic Botany
and Mycology, Ludwig Maximilian University of Munich, Menzinger Str. 67, 80638 Munich, Germany
12 Allan Herbarium, Landcare Research, P O Box 69-040, Lincoln 7608, New Zealand 13 Muséum
National d’Histoire Naturelle, Department Systématique et Evolution, C.P. 39, 57 Rue Cuvier, 75231 Paris
05, France 14 Botany Unit, Finnish Museum of Natural History, University of Helsinki, P.O. Box 7, Helsinki
FI-00014, Finland 15 Department of Systematic Botany with Herbarium Haussknecht and Botanical Garden,
Friedrich Schiller University, Fürstengraben 1, 07737 Jena, Germany 16 Museu Paraense Emilio Goeldi,
Coordenaçao de Botanica, Av. Magalhaes Barata 376, 66040-1 70 Belem, Para, Brazil 17 Department of
Biological Science, Graduate School of Science, Hiroshima University, Kagamiyama 1–3–1, Higashihiroshimashi, Hiroshima 739–8526, Japan 18 N.A. Avrorin Polar-Alpine Botanical Garden–Institute of Kola SC RAS,
184236 Kirovsk-6, Russia 19 Royal Botanic Garden, Edinburgh EH3 5LR, United Kingdom 20 Staatliches
Copyright Lars Söderström et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Museum für Naturkunde Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany 21 Botany Department, Institute
of Biology, Eszterházy Károly College, Eger, Pf. 43, H-3301, Hungary 22 Departamento de Botánica,
Facultad de Ciencias Biológicas, Universitat de València. C/ Dr. Moliner 50, E-46100, Burjassot (Valencia),
Spain 23 Albrecht-von-Haller-Institut für Planzenwissenschaften, Department of Systematics, Biodiversity
and Evolution of Plants, Untere Karspüle 2, 37073 Göttingen, Germany 24 Mittlere Letten 11, 88634
Herdwangen-Schönach, Germany 25 Centro de Investigaciones sobre Desertiicación (CIDE-CSIC-UV-GV),
C/ Carretera de Moncada-Náquera Km. 4.5, E-46113, Moncada (Valencia), Spain 26 Department of Biology,
Faculty of Science, Burapha University, Mueang, 20131 Chonburi, hailand 27 William and Lynda Steere
Herbarium, he New York, Botanical Garden, Bronx, New York 10458-5126, United States of America
28 Instituto de Ciencias Naturales. Universidad Nacional de Colombia. Apartado 7495, Bogotá D.C.,
Colombia 29 Department of Botany, Charles University, Benátská 2, CZ-128 01 Praha 2, Czech Republic
30 Department of Biology, Ludwig-Maximilians-Universität, Menzinger Str. 67, D-80638, München,
Germany 31 36, Big Green, Warmington, Peterborough PE8 6TU, United Kingdom 32 Shenzhen Key
Laboratory of Southern Subtropical Plant Diversity, Fairylake Botanical Garden, 160 Xianhu Rd., Liantang,
Shenzhen 518004, Guangdong, China 33 Department of Biology, School of Life Sciences, East China Normal
University, 3663 Zhong Shan North Road, Shanghai 200062, China
*
Editors
Corresponding author: Lars Söderström (lars.soderstrom@ntnu.no)
Academic editor: Lyubomir Penev | Received 20 June 2015 | Accepted 25 September 2015 | Published 29 January 2016
Citation: Söderström L, Hagborg A, von Konrat M, Bartholomew-Began S, Bell D, Briscoe L, Brown E, Cargill DC,
Costa DP, Crandall-Stotler BJ, Cooper ED, Dauphin G, Engel JJ, Feldberg K, Glenny D, Gradstein SR, He X, Heinrichs
J, Hentschel J, Ilkiu-Borges AL, Katagiri T, Konstantinova NA, Larraín J, Long DG, Nebel M, Pócs T, Felisa Puche F,
Reiner-Drehwald E, Renner MAM, Sass-Gyarmati A, Schäfer-Verwimp A, Moragues JGS, Stotler RE, Sukkharak P,
hiers BM, Uribe J, Váňa J, Villarreal JC, Wigginton M, Zhang L, Zhu R-L (2015) World checklist of hornworts and
liverworts. PhytoKeys 59: 1–828. doi: 10.3897/phytokeys.59.6261
his checklist is dedicated to Elizabeth Brown and Ray Stotler, who sadly left us
during our work with it.
“……Charles Darwin wrote of his desire to provide inancial support ‘for the formation of a perfect
M.S. catalogue of all known plants’ (Darwin 1881, letter 13570). It is a personal embarrassment to
me, and should be chastening to us all, that more than 120 years later we still have not delivered on
that commitment.” (Crane 2004).
Abstract
A working checklist of accepted taxa worldwide is vital in achieving the goal of developing an online
lora of all known plants by 2020 as part of the Global Strategy for Plant Conservation. We here present
the irst-ever worldwide checklist for liverworts (Marchantiophyta) and hornworts (Anthocerotophyta)
that includes 7486 species in 398 genera representing 92 families from the two phyla. he checklist
has far reaching implications and applications, including providing a valuable tool for taxonomists and
systematists, analyzing phytogeographic and diversity patterns, aiding in the assessment of loristic and
World checklist of hornworts and liverworts
3
taxonomic knowledge, and identifying geographical gaps in our understanding of the global liverwort
and hornwort lora. he checklist is derived from a working data set centralizing nomenclature, taxonomy and geography on a global scale. Prior to this efort a lack of centralization has been a major
impediment for the study and analysis of species richness, conservation and systematic research at both
regional and global scales. he success of this checklist, initiated in 2008, has been underpinned by
its community approach involving taxonomic specialists working towards a consensus on taxonomy,
nomenclature and distribution.
Keywords
Marchantiophyta, Anthocerophyta, nomenclature, taxonomy
Introduction
he natural world is changing fast (Balmford et al. 2005). he Global Strategy for
Plant Conservation (GSPC) was adopted by the Conference of Parties (COP) of the
Convention on Biological Diversity (CBD) in April 2002 (http://www.cbd.int/decision/cop/?id=7183; accessed 2014.06.02). he Strategy set out 16 outcome-oriented
targets that were to be achieved by 2010. he GSPC was designed as a framework for
action to halt the loss of plant diversity. Target 1 of the Strategy was to complete “a
widely accessible working list of all known plant species, as a step towards a complete
world Flora” (Lughadha 2004). As early as 1881 Charles Darwin expressed a wish to
have a catalogue of all known plants (Crane 2004). However, over 125 years later, this
wish is not yet fulilled. Without a working checklist, many of the other objectives in
the GSPC cannot be met and botanists around the world cannot communicate about
plants on a global basis (Crane 2004). A working list of known plant species is critical for (I) its underpinning role in the efective implementation of the other targets
through provision of baseline information; (II) increasing the accessibility and use of
accurate botanical name information for research, conservation and sustainable use;
and (III) real-world politics and how taxonomists respond to the decisions of policy
makers (Paton et al. 2008).
Version 1.0 of he Plant List (http://www.theplantlist.org/) was released in December 2010 aimed to be comprehensive for species of vascular plants (lowering
plants, conifers, ferns and their allies) and of bryophytes (mosses, liverworts and hornworts), as a response to the 2010 Target 1 of the GSPC and to a clear global need for
such data. he Plant List is a broad collaboration, coordinated by the Royal Botanic
Gardens, Kew, and the Missouri Botanical Garden (MO), involving diverse partnerships. Paton (2013) noted he Plant List represents a work in progress with future versions planned to improve the quality of names and decrease the number of unresolved
names. Target 1 has since been revised and agreed at the 10th COP of the CBD in Nagoya, Japan, with the goal of developing ‘‘an online lora of all known plants’’ by 2020
(http://www.cbd.int/decision/cop/default.shtml?id=12283; accessed 2014.06.02). Paton (2013) identiied several factors that might assist in the implementation of the
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
revised target, stressing that achieving a working list for plant taxa worldwide was a
vital step to an online lora.
Earlier, Paton et al. (2008) assessed the progress made at that time and discussed
prospects for the completion of Target 1. Paton noted that good progress had been
made in bryophytes (mosses, liverworts and hornworts), ferns and gymnosperms with
widely accessible working lists either complete or almost so for those groups. For bryophytes alone, he tabulated 13,370 accepted species noting that the data was largely
derived from Tropicos. However, although Tropicos is an indispensable reference for
anyone dealing with bryophyte names and the database is very strong for mosses, the
nomenclatural and auxiliary data for liverworts is less complete, especially for larger
genera (von Konrat et al. 2008a). he lack of a central source providing a synthesis of
nomenclatural data and global distributional data was the impetus toward developing
the current checklist of liverworts and hornworts (von Konrat et al. 2008a, 2010d,
Söderström et al. 2012e). Checklists are powerful and important tools that can integrate the almost overwhelmingly scattered information concerning taxonomy, systematics, nomenclature, distribution, and even frequency (Söderström et al. 2008).
Ecological and biological significance of liverworts and hornworts
Liverworts and hornworts (Figure 1) are of critical biological, ecological, and phylogenetic signiicance (e.g. Asakawa 1998, Longton 1992, Hallingbäck and Hodgetts
2000, Gradstein et al. 2001b, Wellman et al. 2003, Qiu et al. 2007). Liverworts are
found on soil, rocks, and trees throughout the world, from coastal Antarctica to the
tundra of the Northern Hemisphere, and from semi-arid areas of Australia to the Amazon rainforests (Hallingbäck and Hodgetts 2000). Although there are xero-tolerant
taxa, the majority of liverworts are found in relatively humid and shaded terrestrial
ecosystems (Gradstein and Costa 2003). Liverworts and hornworts are an important
component of the vegetation in many regions of the world, constituting a major part
of the biodiversity in moist forest, wetlands, mountain, and tundra ecosystems (Hallingbäck and Hodgetts 2000).
Liverworts, hornworts and mosses ofer microhabitats that are critical to the survival
of a tremendous diversity of organisms such as single-celled eukaryotes, protozoa, and
numerous groups of invertebrates (Gerson 1982). heir structural contribution to levels
of diversity might be as signiicant as that of vascular plants, albeit at a smaller scale (von
Konrat et al. 2008a). hese plants are also important environmental and ecological indicators (Rao 1982, Pitcairn et al. 1995, Gradstein et al. 2001b, Giordano et al. 2004). Liverworts, in concert with mosses and hornworts, play a signiicant role in the global carbon
budget (O’Neill 2000) and CO2 exchange (De Lucia et al. 2003), plant succession (Cremer and Mount 1965), net production and phytomass (Frahm 1990), nutrient cycling
(Coxson 1991), and water retention (Pócs 1980a, Gradstein et al. 2001b). hese groups
of land plants also have been used as indicators of past climate change, to validate climate
models, and as potential indicators of global warming (Gignac 2001, Tuba et al. 2011).
World checklist of hornworts and liverworts
5
Figure 1. Habit images of selected lineages of Anthocerotophyta (hornworts) and Marchantiophyta
(liverworts). A Phaeomegaceros squamuliger, photo by J. Hollinger B Haplomitrium mnioides, photo by L.
Zhang C Conocephalum japonicum, photo by D. G. Long D Colura vitiensis, photo by T. Pócs E Pleurozia
gigantea, photo by L. Söderström F Riccardia spegazziniana, photo by Juan Larraín.
Species numbers
here are estimated to exist only 215 hornwort species. By comparison estimates of
liverwort species richness have varied considerably, by as much as 50%, with estimates ranging from 4,500 to 9,000 (e.g. Pearson 1995, Forrest et al. 2006). In the
last decade or so estimates in the range of 5,000 to 6,000 liverwort species have been
widely accepted (e.g. Gradstein and Costa 2003, Gradstein and Ilkiu-Borges 2009,
Heinrichs et al. 2007). von Konrat et al. (2010a) provided a mean estimate of 7,500
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
for the number of liverwort species based on estimating rates of synonymy in a sample
of recently monographed and revised taxa. A standardized global worldwide liverwort
checklist with strong community participation, as presented here, coupled with the
critical need for ongoing monographs and revisions, will aid in arriving at clearer estimates of liverwort and hornwort diversity. Signiicantly, a list of standardized names
reviewed by taxonomic experts will enable more meaningful geographical and biogeographical species comparative studies. he global checklist of accepted liverwort and
hornwort taxa has vast potential, not only in aiding our understanding of liverwort
diversity, patterns, and processes, but also to the broader biological community (von
Konrat et al. 2008a).
Below we provide context to the worldwide checklist, including a brief historical account and current informational resources, structure and layout, systematic and
classiication concepts, and methodology, including a brief overview of the underlying
data set and nomenclatural elements.
Major historical works
Schuster (1966d) has provided a detailed account on the history of hepaticology, up
to the irst half of the 20th century. Earlier, Verdoorn (1934c) also outlined a valuable
historical review of important igures in hepaticology. In the early 19th century, possibly the two most inluential persons who provided an early framework for taxonomic
concepts were W.J. Hooker and G. Raddi. During the middle of the 19th century
C.G. Nees von Esenbeck, C.M. Gottsche and J.B.W. Lindenberg provided vastly signiicant contributions towards developing a classiication framework. Frahm and Eggers (2001) provided a brief biography of these workers. his was later followed by R.
Spruce and the Austrian-born H. Leitgeb in the second half 19th century. Leitgeb in
particular was instrumental in liverwort plant morphology and anatomy including a
six part treatise, “Untersuchungen ueber die Lebermoose”, from 1874 to 1881. Towards the end of 19th century and early 20th century V. Schifner, F. Stephani, and A.
Evans were prominent hepaticologists. In the irst half of the 20th century A. Evans,
H. Buch, F. Verdoorn, and K. Müller (Frib.) were leading inluential hepaticologists.
In the second half of the 20th century, R. Grolle and R.M. Schuster were of extraordinary inluence in working towards contemporary classiication schemes. Many of these
groundbreaking concepts still serve the foundation from which our current classiication scheme is derived. R.M. Schuster’s major contribution to botany and hepaticology
lies in the astounding new diversity of liverworts he added to our knowledge. He described a staggering 463 species, 83 genera and 15 families new to science. It is diicult
to name another contemporary botanist who discovered this much new diversity of a
major clade of land plants (Qiu et al. 2013). R.M. Schuster’s contribution to botany
went beyond hepatics. He was one of the irst botanists who recognized the importance
of Wallace’s Line in plant biogeography, separating Australia of Gondwanaland from
Southeast Asia of Laurasia (Cronquist 1988).
World checklist of hornworts and liverworts
7
Figure 2. Number of novel liverwort species, excluding new combinations, which have been described
over the last 250 years, with an inset of the number described from 2001–2012.
Figure 2 depicts the number of novel liverwort species, excluding new combinations that have been described over the last 250 years. he irst major peak corresponds
to the works of several early 19th century botanists, including Synopsis Hepaticarum
by Gottsche, Lindenberg, and Nees (1844–1847). he three decades leading into, but
prior to the highest peak, between 1860 and 1890, correspond to publications by a
number of prominent bryologists including W. Mitten, J.D. Hooker, T. Taylor, and
V. Schifner. he second and highest peak of almost 1,200 names, in the early 1900s
corresponds largely to the plethora of taxa described by F. Stephani (1898–1924) in
his monumental work Species Hepaticarum. he periods of highest rates of new species described in the 1830s and around 1900 are the same for seed plants (Mutke and
Barthlott 2005). he third peak over the four decades between 1950 and 1980 can
be attributed mainly to the works of R.M. Schuster, H. Inoue, and S. Hattori. H.
Inoue and S. Hattori were inluential on large species-rich genera such as Plagiochila,
Frullania, and Porella. he decline in newly described species since 1970 does not
necessarily translate to the conclusion that taxonomists are closer to discovering all
known species. he almost 200 novel liverwort species that have been described in the
past six years alone still represent a signiicant number, considering the relatively few
liverwort taxonomists and monographers. Moreover, bryological exploration has been
very uneven in many parts of the world. For example, many areas of the Neotropics
still remain without a single bryophyte record (Gradstein et al. 2001a). Paradoxically,
scores of new species are still being discovered and described in relatively well-studied
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
areas such as New Zealand, e.g. over 50 new taxa since 2000 alone (e.g. Engel and
Schuster 2001, Engel and Glenny 2008a, Renner and de Lange 2011, von Konrat et
al. 2012b). Recent attention to cryptic speciation in bryophytes is also revealing novel
liverwort species (e.g. Szweykowski et al. 2005). he combination of collecting in yetto-be explored areas, the continued discovery of species in well-studied regions, and
an increased understanding of the biology of liverworts (including cryptic speciation),
will lead to a signiicant number of newly discovered species in the foreseeable future.
he corollary of this, coupled with increased monographic and revisional work, will be
the increased discovery and the unravelling of synonymy.
Contemporary resources
Data availability and information needs associated with liverworts and hornworts have
been reviewed extensively by von Konrat et al. (2008a, 2010a). A fundamental problem common to all nomenclatural indexing projects is the dispersed nature of the biological literature, some of which may date back over 250 years (Lughadha 2004). Our
own data records, from 1990 to the present alone, indicate there are over 190 periodical titles and non-serials in which liverwort nomenclatural novelties were published.
he most useful and successful web-accessible bryophyte nomenclatural database is
Tropicos (http://www.tropicos.org), which ofers name data with references and type
information as well as links to specimen data of its holdings. Tropicos has therefore become an indispensable reference for anyone dealing with bryophyte names. However,
as stated above, the database is particularly strong for mosses and far less comprehensive for liverworts. Moreover, Tropicos does not prescribe taxonomic disposition of
names, i.e., there is no attempt to adopt a single consistent view on the status of any
particular name (Lughadha 2004, von Konrat et al. 2008a).
A major nomenclatural work is the Index Hepaticarum (http://www.ville-ge.ch/
musinfo/bd/cjb/hepatic/index.php), which includes all efectively published liverwort
epithets spanning 12 volumes with the closing date of 1973. he indices were prepared as a purely nomenclatural resource and did not claim to express any particular
taxonomic concept. Recently, Crosby and Engel (2006) provided an equally valuable
nomenclatural resource and catalog of names at all ranks for liverworts and hornworts
published during 1974 to 2000. Subsequently, indices of the citations for names published for bryophytes have been compiled for the years 2001-2004 (Crosby and Magill
2005) and 2005 (Crosby and Magill 2006). he Early Land Plants Today initiative has
continued this series (von Konrat et al. 2010c, Söderström et al. 2012b, 2014).
he application of molecular phylogenetics continues to generate new insights into
the evolutionary history of liverworts and hornworts. In recent years, inferences made
from these phylogenies have especially revolutionized liverwort classiication (CrandallStotler et al. 2009). Yet, the reconstruction of the phylogenetic history of the Marchantiophyta and Anthocerotophyta remains an ongoing efort and the classiication is luid (von Konrat et al. 2010a). he current checklist largely follows the comprehensive
World checklist of hornworts and liverworts
9
phylogenetic liverwort classiication scheme provided by Crandall- Stotler et al. (2009)
and the hornwort classiication scheme of Stotler and Crandall-Stotler (2005). hus
the checklist largely adheres to their higher classiication namely from class to suborder, and the majority of family and genera arrangements provided therein. However,
there are some notable exceptions, especially in the systematic treatment of liverworts,
which largely either relect publications subsequent to 2009, or are a slight departure
in taxonomic opinion based on earlier works, e.g. the concepts of Saccogynaceae and
Stephaniellaceae depart from Crandall-Stotler et al. (2009). Since 2010 there have
been a number of realignments and novel hypotheses proposed that were generated
from conclusions based on molecular evidence which have inluenced the systematic
treatment presented here. hese include, for example, Adelanthaceae (e.g. Konstantinova and Vilnet 2009, Feldberg et al. 2010b, 2010a, Söderström et al. 2010b, Vilnet
et al. 2010), Anastrophyllaceae (e.g. Söderström et. al 2010b), Gymnomitriaceae (e.g.
Vilnet et al 2007a, 2010, Váňa et al. 2010b, Shaw et al. 2015), Lepidoziaceae (Cooper
2013), Lophocoleaceae (Söderström et al. 2013b), and Scapaniaceae (e.g. Vilnet et al.
2010, Heinrichs et al. 2012a). In all cases, any such departures, new families, or realignments are noted and very briely discussed throughout the checklist. As noted by
Crandall-Stotler et al. (2009) many small families still remain to be investigated using
molecular data, and many of the large families and genera that have been sampled appear to be either polyphyletic or paraphyletic.
he systematic treatment as outlined here also includes the addition of subfamily
and infrageneric classiication; ranks not presented by Crandall-Stotler et al. (2008b,
2009). Although there are arguably a number of ambiguous placements, especially at
the infrageneric rank, and there remain many areas of contention, particularly at the
familial and generic level, the systematic treatment itself serves as an excellent synopsis of contemporary taxonomic opinion, including infrageneric ranks. he infrageneric
treatments will also aid future eforts in providing a taxonomic framework that will be
particularly useful for extremely large genera. In large part, the infrageneric treatments
only deal with those taxa that represent phylogenetic clades with strong support. Care
has been taken to reduce ambiguity and those taxa where infrageneric placement is weak
or without evidence are simply listed under the genus and not placed systematically.
Structure and format of the checklist
he checklist is presented in a taxonomic framework as outlined above with four main
sections. he irst is arranged systematically with taxa ordered alphabetically within
the nearest higher rank. A brief discussion for selected families is provided in the main
body of the text under the respective families. his does not appear uniformly for all
families, but typically for larger more complicated families. Rather, a brief description
is provided mainly where concepts depart from Crandall-Stotler et al. (2009) or if new
information has come to light since that publication. Subfamilies and infrageneric
ranks have been incorporated where appropriate. However, if the subgeneric place-
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
ment is unclear the species are listed under “Incertae sedis”. From the rank of genus and
below, all entries include a three star conidence ranking (described below), accepted
taxon name, authority, abbreviated publication title, page number and date, which is
cross- referenced to the full citation in the reference list, as well as the basionym if appropriate. A typical entry appears as follows:
***
***
Oleolophozia L.Söderstr., De Roo et Hedd., Phytotaxa 3: 50, 2010 (Söderström
et al. 2010b).
Oleolophozia perssonii (H.Buch et S.W.Arnell) L.Söderstr., De Roo et Hedd.,
Phytotaxa 3: 51, 2010 (Söderström et al. 2010b). Bas.: Lophozia perssonii
H.Buch et S.W.Arnell, Bot. Not. 97: 382, 1944 (Buch 1944).
here are comprehensive footnotes throughout the systematic section of the checklist. hese are applied to selected names representing the rank of genus and below only.
Footnotes without any literature citation represent observations from the authors of
the respective family. Footnotes are wide ranging in their content. Generally, footnotes
can be categorized into the following: i) general statements relating to taxonomic opinion, ii) comments on the type specimen, iii) comments about possible conspeciicity,
including citation of conlicting opinions, and iv) comment about species complexes.
his section is followed by a list of taxa in genera that we do not recognize. hose
names represent taxa that have not been studied recently and have not been recognized
in any recent treatment. However, many of those names are old and may gain priority
over some recognized taxa once their identity is determined.
he next section is an alphabetical list of taxa of the ranks of species and below. he
alphabetical list includes the conidence level, the taxon name and authority as well as
a reference to the page of the corresponding entry in the systematic section. hus, the
alphabetical list provides a rapid gateway requiring no prior knowledge of higher taxonomy, but also serves as an index to the corresponding name in the systematic section
that includes more detailed information.
he last section is a reference list where full bibliographic citations are given to all
references for taxa included in the checklist. Titles and references have all been checked
and veriied, except in a few cases where we could not get hold of the publications.
Taxon confidence levels
Signiicantly, each accepted taxon is qualiied using a three level ranking system that
summarizes our knowledge about a taxon. he coding convention we are adopting
largely follows that described by von Konrat et al. (2010d) using one to three stars,
which has been applied to recent regional checklists produced by the Early Land Plants
Today initiative, e.g. Söderström et al. (2010a, 2011a, 2011b, 2012g, 2013e). he
conventions are briely outlined below coupled with samples illustrating how these
conventions are applied in practice. he application of a conidence level to a taxon’s
World checklist of hornworts and liverworts
11
status and whether it represents a genuine “species” that is reached through community
consensus may go towards reining species estimates using an evidence-based approach
(von Konrat et al. 2008a, 2008b). he conidence levels coupled with the detailed
annotations in the form of footnotes may also provide a rapid assessment of taxa and
help aid and drive future research into speciic taxonomic or nomenclatural problems
and issues.
he coding convention:
*
Serious doubts. here are doubts about the value of the taxon. It can be that there
are conlicting views without any substantial evidence in any direction, conlicting
views with substantiating evidence supporting one or both positions, or evidence
pointing towards synonymization but it is premature to do it. We have adhered to
the principle that it is better to keep a taxon with one star (and preferably a note)
than to synonymize it too quickly. Example: Bazzania asymmetrica is conspeciic
with Bazzania macgregorii in Grolle (1968a), but Kitagawa (1979a) kept them
separate. Example: Nardia kamtschatica may be conspeciic with Nardia assamica
(Váňa 1976c), but the type specimen could not be studied.
** Knowledge problem. he taxon is not well known by the person evaluating it. It
may be a newly described species or a species originally not well described and not
restudied recently. Example: Jungermannia erectii Ajit P.Singh et V.Nath was recently described and has not been independently studied by someone with a global
overview. It is therefore diicult to evaluate.
*** Accepted. A good taxon as currently understood based on personal experience or
on taxonomic revisions that have been convincingly performed. Nomenclature
and/or taxonomic position may, however, be questioned. Elements may be excluded from the taxon, but the taxon with the current type will still be accepted.
Example: H. Bischler-Causse revised the genus Marchantia worldwide in a series
of publications. Although she had a broad species concept, she also recognized
infraspeciic taxa. he taxa that she recognized without doubt should be accepted
unless new evidence against it exists. Adopting a narrower species concept so that
many of her subspecies are elevated to species does not change her view of what a
good taxon is.
Methodology
he foundation of the checklist is in the underlying data set from which it is derived. It was briely described by von Konrat et al. (2008a, 2010a). he working
data sets now includes a bibliography of 25,000 publications; approximately 39,000
published liverwort names (including “accepted” taxa, synonyms, invalid and illegitimate names). he data quality and standards were outlined by von Konrat et al.
(2010d). In summary, for authorities and for the citation abbreviations, we follow
the standards set by the on-line version of Authors of Plant Names at the Royal Bota-
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nical Gardens, Kew Website (http://www.ipni.org), with whom we collaborate closely and provide with updated data records. Publications and journal abbreviations
follow Taxonomic Literature: A selective guide to botanical publications and collections
with dates, commentaries and types (Staleu and Cowan 1976–2009) (http://www.
sil.si.edu/digitalcollections/tl- 2/search.cfm) and Botanico Periodicum Huntianum
Lawrence et al. (1968) (http://huntbot.andrew.cmu.edu/HIBD/Departments/Databases.shtml).
here has been an intense systematic efort focusing on data quality. In all but a
few cases nomenclatural data has been veriied against original publications. Söderström and Hagborg have checked and conirmed almost every original publication
(three publications have not been available) for correct author, title and journal/
book citation, date of publication as far as possible, page number for the protologue,
and if the name is validly and legitimately published according to the International
Code of Nomenclature for algae, fungi and plants (Melbourne Code; McNeill et al.
2012). Most signiicantly, the checklist has been community driven and collaborative. Broader participation by taxonomic specialists and regional experts has lead to
the checklist containing high quality data (cf. Söderström et al. 2008). Development
of the checklist has included three international meetings (the irst one in 2009) and
generated 74 published notes on taxonomy and nomenclature under the auspices of
the Early Land Plants Today initiative from 2012 to 2015. he series of meetings
were instrumental in providing a framework for direct interaction with taxonomic
experts, workshops reviewing names, and helping identify potential participants who
were taxonomic experts in speciic taxonomic groups, whether these were individual
genera or entire families.
he notes, published in the journal Phytotaxa, have included 38 authors from
13 countries. hese are the results of several years of revisions of taxonomic groups
by taxonomic experts. Following these revisions and working with participants identifying nomenclatural and taxonomic problems that would impact the worldwide
checklist, Söderström, Hagborg and von Konrat led editorial eforts in compiling
manuscripts with baseline information and nomenclatural or taxonomic issues that
required resolving. he work was then coordinated with the taxonomic experts who
drove and led the process and often identiied further issues. he series of notes
on taxonomy and nomenclature of liverworts and hornworts provided updated nomenclature, corrected invalid and illegitimate names and described new taxa based
on studies (mainly molecular) that did not draw nomenclatural conclusions. hose
notes are all open access and any changes were efective immediately as the Melbourne Code of Botanical Nomenclature allows publishing on internet from 1 Jan.
2012 (Knapp et al. 2011).
his checklist builds on all published taxonomic and nomenclatural papers, that
have come to our attention until June 30, 2015.
World checklist of hornworts and liverworts
13
Summary statistics
We here present the irst-ever worldwide checklist for liverworts (Marchantiophyta)
and hornworts (Anthocerotophyta) that includes 8,078 taxa (species and below) in
7,486 species representing 398 genera, 92 families, 20 orders and 7 classes from the
two phyla. he list includes 3,533 species with three stars, 2,988 species with two stars
and 915 species with one star. he checklist also has extra utility in that it contains
3,106 references in the bibliography that serve as a powerful bibliographic resource for
liverwort and hornwort systematic and taxonomic research.
Concluding remarks
he marked-up publication form of the current checklist by PhytoKeys provides a
virtual instrument with a linked environment both internally (e.g. within an article)
and externally (GBIF, IPNI, Tropicos, Wikispecies, etc.) that will undoubtedly help
accelerate taxonomic research. he published checklist was the irst phase in providing
a worldwide list of accepted names. he next phase is to establish a generally recognized online repository to augment the huge underlying informational auxiliary data
of over 25,000 publications, almost 39,000 published names, and the over 700,000
geographical observations. Several features of Web-based technology make it an essential tool, including 1) a vehicle to facilitate data access, 2) to unify the vastly scattered
data on distributional information and nomenclature, 3) ofering dynamic rather than
static data in a searchable forum. he broader accessibility to the wealth of auxiliary
data will help augment monographic and revisional work for many taxonomic groups,
aid in identifying the need for increased loristic and survey work in many regions
throughout the world, and have broad implications and applications beyond taxonomic research such as conservation science. he current Early Land Plants Today model
has strong participation from taxonomic experts and an online resource will provide an
opportunity to expand stakeholders to include ecologists, conservationists, scientists
from other disciplines and general interest groups. However, such an efort can only be
successful if it comes with sustained funding and infrastructure rather than depending
on an ad hoc commitment by a few individuals, however dedicated.
he project will help augment monographic and revisional work for many taxonomic groups and aid in identifying the need for increased loristic and survey work
in many regions throughout the world. Although there are many challenges ahead to
obtain high quality data, quantifying global liverwort diversity is a tractable, multifaceted and scientiically important goal, and everyone stands to gain by fostering this
endeavour. he success of the project will lie on strong collaboration between institutions and the bryological community in general.
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Taxonomic synopsis
Anthocerotophyta ................................................................................................. 27
Anthocerotopsida de Bary ex Jancz. .................................................................... 27
Anthocerotidae Rosenv. ................................................................................. 27
Anthocerotales Limpr. ............................................................................... 27
Anthocerotaceae Dumort. ...................................................................... 27
Anthoceros L. ................................................................................ 27
Folioceros D.C.Bharadwaj ........................................................... 31
Dendrocerotidae R.J.Duf .............................................................................. 32
Dendrocerotales Hässel .............................................................................. 32
Dendrocerotaceae J.Haseg. .................................................................... 32
Dendroceros Nees ......................................................................... 32
Megaceros Campb. ....................................................................... 27
Nothoceros (R.M.Schust.) J.Haseg............................................... 35
Phaeomegaceros R.J.Duf............................................................. 35
Phymatocerotales R.J.Duf ......................................................................... 36
Phymatocerotaceae R.J.Duf .................................................................. 36
Phymatoceros Stotler.................................................................... 36
Notothylatidae R.J.Duf................................................................................. 36
Notothyladales Hyvönen et Piippo ............................................................ 36
Notothyladaceae Müll.Frib. ex Prosk. .................................................... 36
Notothyladoideae Grolle ................................................................... 37
Notothylas Sull. ex A.Gray ........................................................... 37
Phaeocerotoideae Hässel .................................................................... 38
Mesoceros Piippo.......................................................................... 38
Paraphymatoceros Hässel ............................................................ 38
Phaeoceros Prosk. ........................................................................ 38
Leiosporocerotopsida Stotler et Crand.-Stotl. ................................................. 41
Leiosporocerotales Hässel ........................................................................... 41
Leiosporocerotaceae Hässel ex Ochyra ................................................... 41
Leiosporoceros Hässel................................................................... 41
Marchantiophyta ........................................................................................... 41
Haplomitriopsida Stotler et Crand.-Stotl. ...................................................... 41
Haplomitriidae Stotler et Crand.-Stotl. ...................................................... 41
Calobryales Hamlin ............................................................................... 41
Haplomitriaceae Dědeček .................................................................. 41
Haplomitrium Nees ..................................................................... 41
Treubiidae Stotler et Crand.-Stotl. ............................................................. 42
Treubiales Schljakov .............................................................................. 42
Treubiaceae Verd............................................................................... 42
Apotreubia S.Hatt. et Mizut. ....................................................... 42
Treubia K.I.Goebel...................................................................... 43
World checklist of hornworts and liverworts
15
Jungermanniopsida Stotler et Crand.-Stotl. ........................................................ 43
Jungermanniidae Engl.................................................................................... 43
Jungermanniales H.Klinggr........................................................................ 43
Cephaloziineae Schljakov ....................................................................... 43
Adelanthaceae Grolle ......................................................................... 43
Adelanthoideae K.Feldberg ............................................................ 43
Adelanthus Mitt. ...................................................................... 44
Pseudomarsupidium Herzog ................................................... 44
Wettsteinia Schifn. ................................................................. 45
Jamesonielloideae Inoue ................................................................ 45
Cuspidatula Steph. .................................................................. 45
Denotarisia Grolle ................................................................... 46
Nothostrepta R.M.Schust. ....................................................... 46
Pisanoa Hässel ......................................................................... 46
Protosyzygiella (Inoue) R.M.Schust. ....................................... 46
Syzygiella Spruce...................................................................... 46
Vanaea (Inoue et Gradst.) Inoue et Gradst. ........................... 49
Anastrophyllaceae L.Söderstr. ............................................................ 49
Anastrepta (Lindb.) Schifn. .................................................... 50
Anastrophyllum (Spruce) Steph............................................... 50
Barbilophozia Loeske .............................................................. 51
Biantheridion (Grolle) Konstant. et Vilnet ............................ 52
Chandonanthus Mitt. .............................................................. 52
Crossocalyx Meyl. ..................................................................... 52
Gymnocolea (Dumort.) Dumort. ............................................ 52
Hamatostrepta Váňa et D.G.Long .......................................... 52
Hattoria R.M.Schust. .............................................................. 53
Isopaches H.Buch ..................................................................... 53
Neoorthocaulis L.Söderstr. ...................................................... 53
Orthocaulis H.Buch ................................................................. 53
Plicanthus R.M.Schust. ........................................................... 54
Schizophyllopsis Váňa et L.Söderstr. ....................................... 54
Schljakovia Konstant. et Vilnet ............................................... 54
Schljakovianthus Konstant. et Vilnet ...................................... 55
Sphenolobopsis R.M.Schust. et N.Kitag. ................................. 55
Sphenolobus (Lindb.) Berggr................................................... 55
Tetralophozia (R.M.Schust.) Schljakov .................................. 55
Zantenia (S.Hatt.) Váňa et J.J.Engel....................................... 56
Cephaloziaceae Mig. .......................................................................... 56
Alobielloideae R.M.Schust............................................................. 56
Alobiella (Spruce) Schifn. ....................................................... 56
Alobiellopsis R.M.Schust. ........................................................ 56
Cephalozioideae Müll.Frib. ........................................................... 57
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Cephalozia (Dumort.) Dumort. .............................................. 57
Fuscocephaloziopsis Fulford .................................................... 59
Nowellia Mitt. .......................................................................... 61
Odontoschismatoideae H.Buch ex Grolle ...................................... 62
Odontoschisma (Dumort.) Dumort. ....................................... 62
Schifnerioideae R.M.Schust. ......................................................... 64
Schifneria Steph...................................................................... 64
Trabacelluloideae R.M.Schust. ...................................................... 64
Haesselia Grolle et Gradst. ...................................................... 64
Trabacellula Fulford ............................................................... 64
Cephaloziellaceae Douin.................................................................... 64
Allisoniella E.A.Hodgs. ............................................................ 65
Amphicephalozia R.M.Schust.................................................. 65
Anastrophyllopsis (R.M.Schust.) Váňa et L.Söderstr. ............. 65
Cephalojonesia Grolle.............................................................. 66
Cephalomitrion R.M.Schust. ................................................... 66
Cephaloziella (Spruce) Schifn. ............................................... 66
Cephaloziopsis (Spruce) Schifn. ............................................. 72
Chaetophyllopsis R.M.Schust. ................................................. 72
Cylindrocolea R.M.Schust. ...................................................... 72
Gottschelia Grolle .................................................................... 74
Gymnocoleopsis (R.M.Schust.) R.M.Schust. ........................... 74
Herzogobryum Grolle .............................................................. 74
Kymatocalyx Herzog ................................................................ 74
Lophonardia R.M.Schust. ....................................................... 75
Nothogymnomitrion R.M.Schust............................................. 75
Obtusifolium S.W.Arnell ......................................................... 75
Oleolophozia L.Söderstr. ......................................................... 75
Phycolepidozia R.M.Schust. .................................................... 75
Protolophozia (R.M.Schust.) Schljakov .................................. 75
Lophoziaceae Cavers .......................................................................... 77
Andrewsianthus R.M.Schust. .................................................. 77
Gerhildiella Grolle................................................................... 78
Heterogemma (Jørg.) Konstant. et Vilnet ............................... 78
Lophozia (Dumort.) Dumort. ................................................. 79
Lophoziopsis Konstant. et Vilnet ............................................. 80
Pseudocephaloziella R.M.Schust. ............................................ 81
Trilophozia (R.M.Schust.) Bakalin ......................................... 81
Tritomaria Schifn. ex Loeske ................................................. 82
Scapaniaceae Mig............................................................................... 82
Diplophyllum (Dumort.) Dumort........................................... 82
Douinia (C.E.O.Jensen) H.Buch ............................................ 84
Pseudotritomaria Konstant. et Vilnet ..................................... 84
World checklist of hornworts and liverworts
17
Saccobasis H.Buch ................................................................... 85
Scapania (Dumort.) Dumort. ................................................. 85
Schistochilopsis (N.Kitag.) Konstant. ...................................... 92
Jungermanniineae R.M.Schust. ex Stotler et Crand.-Stotl...................... 93
Acrobolbaceae E.A.Hodgs.................................................................. 93
Enigmella G.A.M.Scott et K.G.Beckm. ................................... 93
Acrobolboideae R.M.Schust. ex Briscoe ......................................... 93
Acrobolbus Nees ....................................................................... 93
Austrolophozioideae R.M.Schust. ex Crand.-Stotl. ........................ 96
Austrolophozia R.M.Schust. .................................................... 96
Goebelobryum Grolle............................................................... 97
Lethocoleoideae Grolle .................................................................. 97
Lethocolea Mitt. ....................................................................... 97
Saccogynidioideae Crand.-Stotl. .................................................... 97
Saccogynidium Grolle .............................................................. 97
Antheliaceae R.M.Schust. .................................................................. 98
Anthelia (Dumort.) Dumort. .................................................. 98
Arnelliaceae Nakai ............................................................................. 99
Arnellia Lindb. ........................................................................ 99
Balantiopsidaceae H.Buch ................................................................. 99
Acroscyphella N.Kitag. et Grolle.............................................. 99
Pseudoisotachis Váňa ............................................................... 99
Balantiopsidoideae J.J.Engel et Váňa ............................................. 99
Balantiopsis Mitt. .................................................................... 99
Isotachidoideae Grolle ................................................................. 101
Isotachis Mitt. ........................................................................ 101
Neesioscyphus Grolle .............................................................. 102
Ruizanthoideae R.M.Schust. ex J.J.Engel et G.L.Merr. ................ 103
Ruizanthus R.M.Schust. ........................................................ 103
Blepharidophyllaceae R.M.Schust. ex J.J.Engel ................................ 103
Blepharidophyllum Ångstr. ................................................... 103
Clandarium (Grolle) R.M.Schust. ........................................ 103
Calypogeiaceae Arnell ...................................................................... 103
Calypogeia Raddi ................................................................... 103
Eocalypogeia (R.M.Schust.) R.M.Schust............................... 107
Metacalypogeia (S.Hatt.) Inoue............................................. 107
Mizutania Furuki et Z.Iwats. ................................................ 107
Mnioloma Herzog .................................................................. 107
Endogemmataceae Konstant. ........................................................... 109
Endogemma Konstant............................................................ 109
Geocalycaceae H.Klinggr. ................................................................ 109
Geocalyx Nees ........................................................................ 109
Gymnomitriaceae H.Klinggr. .......................................................... 109
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Acrolophozia R.M.Schust. ..................................................... 110
Nanomarsupella R.M.Schust. ex A.Hagborg ........................ 110
Paramomitrion R.M.Schust. ................................................. 110
Gymnomitrioideae T.Jensen........................................................ 110
Cryptocoleopsis Amakawa ...................................................... 110
Gymnomitrion Corda ............................................................ 110
Marsupella Dumort. .............................................................. 112
Poeltia Grolle ......................................................................... 114
Prasanthus Lindb. ................................................................. 114
Nardioideae Váňa ........................................................................ 114
Nardia Gray ........................................................................... 114
Gyrothyraceae R.M.Schust. ............................................................. 115
Gyrothyra M.Howe................................................................ 115
Harpanthaceae Arnell ...................................................................... 116
Harpanthus Nees ................................................................... 116
Hygrobiellaceae Konstant. et Vilnet................................................. 116
Hygrobiella Spruce ................................................................ 116
Jackiellaceae R.M.Schust. ................................................................ 116
Jackiella Schifn. .................................................................... 116
Jungermanniaceae Rchb. ................................................................. 117
Delavayelloideae Grolle ............................................................... 117
Delavayella Steph. ................................................................. 117
Liochlaena Nees ..................................................................... 117
Jungermannioideae Dumort. ....................................................... 118
Eremonotus Lindb. et Kaal. ex Pearson ................................ 118
Jungermannia L. .................................................................... 118
Mesoptychioideae R.M.Schust. ................................................... 120
Mesoptychia (Lindb.) A.Evans ............................................... 120
Rivulariella D.H.Wagner ...................................................... 122
Notoscyphaceae Crand.-Stotl. ......................................................... 122
Notoscyphus Mitt. .................................................................. 122
Saccogynaceae Heeg ........................................................................ 122
Saccogyna Dumort. ................................................................ 122
Solenostomataceae Stotler et Crand.-Stotl. ...................................... 123
Aponardia (R.M.Schust.) Váňa ............................................. 123
Arctoscyphus Hässel ............................................................... 123
Cryptocolea R.M.Schust. ....................................................... 123
Diplocolea Amakawa ............................................................. 123
Solenostoma Mitt. .................................................................. 123
Southbyaceae Váňa .......................................................................... 135
Gongylanthus Nees ................................................................ 135
Southbya Spruce .................................................................... 135
Stephaniellaceae R.M.Schust. .......................................................... 136
World checklist of hornworts and liverworts
19
Stephaniella J.B.Jack ............................................................. 136
Stephaniellidium S.Winkl. ex Grolle .................................... 136
Trichotemnomataceae R.M.Schust. ................................................. 136
Trichotemnoma R.M.Schust.................................................. 136
Lophocoleineae Schljakov .................................................................... 137
Blepharostomataceae W.Frey et M.Stech ......................................... 137
Blepharostoma (Dumort.) Dumort. ...................................... 137
Brevianthaceae J.J.Engel et R.M.Schust. .......................................... 137
Brevianthus J.J.Engel et R.M.Schust. ................................... 137
Tetracymbaliella Grolle ......................................................... 137
Chonecoleaceae R.M.Schust. ex Grolle ............................................ 138
Chonecolea Grolle .................................................................. 138
Grolleaceae Solari ex R.M.Schust..................................................... 138
Grollea R.M.Schust. .............................................................. 138
Herbertaceae Müll.Frib. ex Fulford et Hatcher ................................ 138
Herbertus Gray ...................................................................... 138
Triandrophyllum Fulford et Hatcher .................................... 171
Lepicoleaceae R.M.Schust................................................................ 141
Lepicolea Dumort. ................................................................. 142
Vetaforma Fulford et J.Taylor............................................... 142
Lepidoziaceae Limpr. ....................................................................... 142
Meinungeria Frank Müll. ...................................................... 143
Bazzanioideae Rodway................................................................. 143
Acromastigum A.Evans .......................................................... 143
Bazzania Gray ....................................................................... 146
Mastigopelma Mitt................................................................. 163
Drucelloideae R.M.Schust. .......................................................... 163
Drucella E.A.Hodgs. .............................................................. 163
Lembidioideae R.M.Schust.......................................................... 163
Dendrolembidium Herzog..................................................... 163
Hygrolembidium R.M.Schust. ............................................... 164
Isolembidium R.M.Schust. .................................................... 164
Kurzia G.Martens .................................................................. 164
Lembidium Mitt. ................................................................... 168
Megalembidium R.M.Schust. ................................................ 168
Pseudocephalozia R.M.Schust. .............................................. 168
Lepidozioideae Müll.Frib. ........................................................... 169
Ceramanus E.D.Cooper ........................................................ 169
Lepidozia (Dumort.) Dumort. .............................................. 169
Neolepidozia Fulford et J.Taylor........................................... 176
Tricholepidozia (R.M.Schust.) E.D.Cooper ......................... 179
Micropterygioideae Grolle ........................................................... 181
Micropterygium Gottsche ...................................................... 181
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Mytilopsis Spruce ................................................................... 182
Protocephalozioideae R.M.Schust................................................ 182
Protocephalozia (Spruce) K.I.Goebel.................................... 182
Zoopsidoideae R.M.Schust. ......................................................... 183
Amazoopsis J.J.Engel et G.L.Merr. ........................................ 183
Hyalolepidozia S.W.Arnell ex Grolle .................................... 183
Monodactylopsis (R.M.Schust.) R.M.Schust. ........................ 183
Neogrollea E.A.Hodgs............................................................ 183
Odontoseries Fulford ............................................................. 183
Paracromastigum Fulford et J.Taylor ................................... 183
Psiloclada Mitt....................................................................... 185
Pteropsiella Spruce ................................................................ 185
Telaranea Spruce ex Schifn. ................................................. 185
Zoopsidella R.M.Schust. ........................................................ 187
Zoopsis Hook.f. ex Gottsche .................................................. 188
Lophocoleaceae Vanden Berghen..................................................... 189
Bragginsella R.M.Schust........................................................ 189
Chiloscyphus Corda................................................................ 189
Clasmatocolea Spruce ............................................................ 191
Conoscyphus Mitt. .................................................................. 193
Cryptolophocolea L.Söderstr.................................................. 193
Deceptifrons J.J.Engel et Váňa............................................... 196
Evansianthus R.M.Schust. et J.J.Engel ................................. 196
Hepatostolonophora J.J.Engel et R.M.Schust. ...................... 196
Heteroscyphus Schifn. ........................................................... 197
Lamellocolea J.J.Engel ........................................................... 204
Leptophyllopsis R.M.Schust. .................................................. 204
Leptoscyphopsis R.M.Schust. ................................................. 204
Leptoscyphus Mitt. ................................................................. 204
Lophocolea (Dumort.) Dumort. ............................................ 208
Otoscyphus J.J.Engel, Bardat et houvenot ........................... 215
Pachyglossa Herzog et Grolle ................................................ 216
Perdusenia Hässel .................................................................. 216
Pigafettoa C.Massal. .............................................................. 217
Platycaulis R.M.Schust.......................................................... 217
Stolonivector J.J.Engel ........................................................... 217
Xenocephalozia R.M.Schust. ................................................. 217
Mastigophoraceae R.M.Schust......................................................... 217
Dendromastigophora R.M.Schust. ........................................ 217
Mastigophora Nees ................................................................ 217
Plagiochilaceae Müll.Frib................................................................. 218
Acrochila R.M.Schust. ........................................................... 218
Chiastocaulon Carl ................................................................ 219
World checklist of hornworts and liverworts
21
Dinckleria Trevis. .................................................................. 219
Pedinophyllopsis R.M.Schust. et Inoue ................................. 219
Pedinophyllum Lindb. ex Nordst. ......................................... 219
Plagiochila (Dumort.) Dumort. ............................................ 219
Plagiochilidium Herzog......................................................... 252
Plagiochilion S.Hatt............................................................... 252
Pseudolophocolea R.M.Schust. et J.J.Engel ........................... 253
Xenochila R.M.Schust. .......................................................... 253
Pseudolepicoleaceae Fulford et J.Taylor ........................................... 253
Archeophylla R.M.Schust....................................................... 253
Castanoclobos J.J.Engel et Glenny ........................................ 254
Chaetocolea Spruce ................................................................ 254
Herzogiaria Fulford ex Hässel............................................... 254
Isophyllaria E.A.Hodgs. et Allison ........................................ 254
Pseudolepicolea Fulford et J.Taylor ...................................... 254
Temnoma Mitt. ...................................................................... 255
Trichocoleaceae Nakai ..................................................................... 256
Eotrichocolea R.M.Schust...................................................... 256
Leiomitra Lindb. ................................................................... 256
Trichocolea Dumort. ............................................................. 257
Myliineae J.J.Engel et Braggins ex Crand.-Stotl. .................................. 258
Myliaceae Schljakov ......................................................................... 258
Mylia Gray ............................................................................. 258
Perssoniellineae R.M.Schust. ............................................................... 259
Schistochilaceae H.Buch .................................................................. 259
Schistochila Dumort. ............................................................. 259
Porellales Schljakov .................................................................................. 264
Jubulineae Müll.Frib. .......................................................................... 264
Frullaniaceae Lorch.......................................................................... 264
Frullania Raddi ..................................................................... 264
Jubulaceae H.Klinggr....................................................................... 295
Jubula Dumort. ..................................................................... 295
Neohattoria Kamim. .............................................................. 296
Nipponolejeunea S.Hatt. ....................................................... 296
Lejeuneaceae Cavers ........................................................................ 297
Lejeuneoideae ............................................................................. 297
trib. Brachiolejeuneeae ................................................................ 297
subtrib. Brachiolejeuneinae Gradst. ............................................. 297
Acanthocoleus R.M.Schust. .................................................... 297
Blepharolejeunea S.W.Arnell ................................................ 298
Brachiolejeunea (Spruce) Schifn. ......................................... 298
Dicranolejeunea (Spruce) Schifn. ........................................ 299
Lindigianthus Kruijt et Gradst. ............................................. 299
22
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Odontolejeunea (Spruce) Schifn. ......................................... 299
subtrib. Stictolejeuneinae Gradst. ................................................ 299
Neurolejeunea (Spruce) Schifn............................................. 299
Stictolejeunea (Spruce) Schifn. ............................................ 300
trib. Lejeuneeae Dumort. ............................................................ 300
Dactylophorella R.M.Schust.................................................. 300
Metalejeunea Grolle .............................................................. 300
Pictolejeunea Grolle .............................................................. 300
subtrib. Ceratolejeuneinae Gradst. ............................................... 301
Ceratolejeunea (Spruce) J.B.Jack et Steph. ........................... 301
Luteolejeunea Piippo ............................................................. 304
Otigoniolejeunea (Spruce) Schifn. ....................................... 304
subtrib. Cheilolejeuneinae Gradst. ............................................... 304
Aureolejeunea R.M.Schust. ................................................... 304
Cheilolejeunea (Spruce) Steph. ............................................. 305
Cyrtolejeunea A.Evans ........................................................... 316
Cystolejeunea A.Evans............................................................ 316
Omphalanthus Lindenb. et Nees ........................................... 316
subtrib. Cololejeuneinae Gradst. ................................................. 317
Aphanotropis Herzog ............................................................. 317
Calatholejeunea K.I.Goebel .................................................. 317
Cololejeunea (Spruce) Steph. ................................................ 317
Colura (Dumort.) Dumort. ................................................... 340
Diplasiolejeunea (Spruce) Schifn. ........................................ 344
Haplolejeunea Grolle ............................................................. 348
Macrocolura R.M.Schust. ...................................................... 348
Myriocoleopsis Schifn. .......................................................... 348
Nephelolejeunea Grolle .......................................................... 349
Schusterolejeunea Grolle ....................................................... 349
Siphonolejeunea Herzog ........................................................ 350
Tuyamaella S.Hatt................................................................. 350
subtrib. Cyclolejeuneinae Gradst. ................................................ 350
Bromeliophila R.M.Schust. ................................................... 350
Cyclolejeunea A.Evans ........................................................... 351
Prionolejeunea (Spruce) Schifn............................................ 351
subtrib. Drepanolejeuneinae Gradst. ........................................... 353
Capillolejeunea S.W.Arnell ................................................... 353
Drepanolejeunea (Spruce) Steph........................................... 353
Vitalianthus R.M.Schust. et Giancotti .................................. 361
subtrib. Echinolejeuneinae Gradst. .............................................. 361
Anoplolejeunea (Spruce) Schifn. .......................................... 361
Echinolejeunea R.M.Schust. .................................................. 362
Kymatolejeunea Grolle .......................................................... 362
World checklist of hornworts and liverworts
23
subtrib. Leiolejeuneinae Schäf.-Verw. et Heinrichs ...................... 362
Leiolejeunea A.Evans ............................................................. 362
subtrib. Lejeuneinae Gradst. ........................................................ 362
Harpalejeunea (Spruce) Schifn. ........................................... 362
Hattoriolejeunea Mizut. ........................................................ 364
Lejeunea Lib. ......................................................................... 364
Microlejeunea (Spruce) Steph. .............................................. 385
Taxilejeunea (Spruce) Steph. ................................................ 388
subtrib. Lepidolejeuneinae Gradst. .............................................. 391
Lepidolejeunea R.M.Schust. .................................................. 391
Otolejeunea Grolle et Tixier .................................................. 392
Rectolejeunea A.Evans ........................................................... 393
subtrib. Leptolejeuneinae Heinrichs et Schäf.-Verw. .................... 394
Leptolejeunea (Spruce) Steph. ............................................... 394
subtrib. Pycnolejeuneinae Heinrichs et Schäf.-Verw. .................... 396
Pycnolejeunea (Spruce) Schiffn. ........................................... 396
subtrib. Xylolejeuneinae Henrichs et Schäf.-Verw. ....................... 398
Xylolejeunea Xiao L.He et Grolle .......................................... 398
trib. Symbiezidieae Gradst. .......................................................... 398
Symbiezidium Trevis. ............................................................ 398
Ptychanthoideae Mizut. ............................................................... 399
Acrolejeunea (Spruce) Schifn. .............................................. 399
Archilejeunea (Spruce) Steph. ............................................... 401
Bryopteris (Nees) Lindenb..................................................... 403
Caudalejeunea Schifn........................................................... 403
Cephalantholejeunea R.M.Schust. ........................................ 405
Cephalolejeunea Mizut. ......................................................... 405
Frullanoides Raddi ................................................................ 405
Fulfordianthus Gradst. .......................................................... 405
Lopholejeunea (Spruce) Steph. .............................................. 406
Marchesinia Gray .................................................................. 408
Mastigolejeunea (Spruce) Steph. ........................................... 409
Phaeolejeunea Mizut. ............................................................ 410
Ptychanthus Nees ................................................................... 411
Schifneriolejeunea Verd........................................................ 411
Spruceanthus Verd. ................................................................ 412
hysananthus Lindenb. ......................................................... 413
Tuzibeanthus S.Hatt.............................................................. 414
Verdoornianthus Gradst. ....................................................... 414
Porellineae R.M.Schust. ....................................................................... 415
Goebeliellaceae Verd........................................................................ 415
Goebeliella Steph. .................................................................. 415
Lepidolaenaceae Nakai .................................................................... 415
24
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Gackstroemia Trevis. ............................................................. 415
Lepidogyna R.M.Schust. ........................................................ 416
Lepidolaena Dumort. ............................................................ 416
Porellaceae Cavers nom. conserv........................................................ 417
Ascidiota C.Massal. ................................................................ 417
Porella L................................................................................. 417
Radulineae R.M.Schust. ...................................................................... 424
Radulaceae Müll.Frib. ..................................................................... 424
Radula Dumort. .................................................................... 425
Ptilidiales Schljakov ................................................................................. 434
Herzogianthaceae Stotler et Crand.-Stotl. ........................................ 434
Herzogianthus R.M.Schust. ................................................... 434
Neotrichocoleaceae Inoue ................................................................ 434
Neotrichocolea S.Hatt. ........................................................... 434
Trichocoleopsis S.Okamura ................................................... 434
Ptilidiaceae H.Klinggr. .................................................................... 434
Ptilidium Nees ....................................................................... 434
Metzgeriidae Barthol.-Began ........................................................................ 435
Metzgeriales Chalaud ............................................................................... 435
Aneuraceae H.Klinggr. .................................................................... 435
Aneura Dumort. .................................................................... 435
Lobatiriccardia (Mizut. et S.Hatt.) Furuki........................... 437
Riccardia Gray ....................................................................... 438
Verdoornia R.M.Schust. ........................................................ 456
Metzgeriaceae H.Klinggr. ................................................................ 456
Metzgeria Raddi..................................................................... 457
Steereella Kuwah. .................................................................. 461
Vandiemenia Hewson ........................................................... 461
Pleuroziales Schljakov .............................................................................. 462
Pleuroziaceae Müll.Frib. .................................................................. 462
Pleurozia Dumort. ................................................................ 462
Pelliidae He-Nygrén .................................................................................... 463
Fossombroniales Schljakov ....................................................................... 463
Calyculariineae He-Nygrén.................................................................. 463
Calyculariaceae He-Nygrén ............................................................. 463
Calycularia Mitt. ................................................................... 463
Fossombroniineae R.M.Schust. ex Stotler et Crand.-Stotl. ................... 463
Allisoniaceae Schljakov .................................................................... 463
Allisonia Herzog .................................................................... 463
Fossombroniaceae Hazsl. nom. conserv. ............................................ 463
Fossombronia Raddi .............................................................. 463
Petalophyllaceae Stotler et Crand.-Stotl. .......................................... 468
Petalophyllum Nees et Gottsche ............................................ 468
World checklist of hornworts and liverworts
25
Sewardiella Kashyap ............................................................. 468
Makinoiineae He-Nygrén .................................................................... 468
Makinoaceae Nakai ......................................................................... 468
Makinoa Miyake.................................................................... 468
Pallaviciniales W.Frey et M.Stech ............................................................ 468
Pallaviciniineae R.M.Schust. ................................................................ 468
Hymenophytaceae R.M.Schust. ....................................................... 468
Hymenophyton Dumort......................................................... 468
Moerckiaceae K.I.Goebel ex Stotler et Crand.-Stotl. ........................ 469
Hattorianthus R.M.Schust. et Inoue..................................... 469
Moerckia Gottsche ................................................................. 469
Pallaviciniaceae Mig......................................................................... 469
Pallavicinioideae Mig. ex Grolle .................................................. 469
Jensenia Lindb. ...................................................................... 469
Pallavicinia Gray ................................................................... 470
Podomitrium Mitt. ................................................................ 471
Symphyogynoideae R.M.Schust. ex Grolle .................................. 471
Greeneothallus Hässel............................................................ 471
Seppeltia Grolle ..................................................................... 471
Symphyogyna Nees et Mont. .................................................. 471
Symphyogynopsis Grolle......................................................... 473
Xenothallus R.M.Schust. ....................................................... 473
Sandeothallaceae R.M.Schust. ......................................................... 474
Sandeothallus R.M.Schust..................................................... 474
Phyllothalliineae R.M.Schust. .............................................................. 474
Phyllothalliaceae E.A.Hodgs. ex T.Katag. ........................................ 474
Phyllothallia E.A.Hodgs. ....................................................... 474
Pelliales He-Nygrén ................................................................................. 474
Noterocladaceae W.Frey et M.Stech ................................................ 474
Noteroclada Taylor ex Hook.f. et Wilson ............................. 474
Pelliaceae H.Klinggr. ....................................................................... 475
Pellia Raddi ........................................................................... 475
Marchantiopsida Cronquist.............................................................................. 476
Blasiidae He-Nygrén .................................................................................... 476
Blasiales Stotler et Crand.-Stotl. ............................................................... 476
Blasiaceae H.Klinggr........................................................................ 476
Blasia L. ................................................................................. 476
Cavicularia Steph. ................................................................. 476
Marchantiidae Engl...................................................................................... 476
Lunulariales H.Klinggr. ........................................................................... 476
Lunulariaceae H.Klinggr.................................................................. 476
Lunularia Adans. ................................................................... 476
Marchantiales Limpr. ............................................................................... 476
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Aytoniaceae Cavers .......................................................................... 476
Asterella P.Beauv. .................................................................. 477
Cryptomitrium Austin ex Underw. ....................................... 481
Mannia Corda ....................................................................... 481
Plagiochasma Lehm. .............................................................. 482
Reboulia Raddi ...................................................................... 483
Cleveaceae Cavers ............................................................................ 484
Athalamia Falc. ...................................................................... 484
Clevea Lindb. ......................................................................... 484
Peltolepis Lindb. .................................................................... 484
Sauteria Nees ......................................................................... 485
Conocephalaceae Müll.Frib. ex Grolle ............................................. 485
Conocephalum Hill ................................................................ 485
Corsiniaceae Engl. ........................................................................... 486
Corsinioideae Schifn....................................................................... 486
Corsinia Raddi ....................................................................... 486
Cronisioideae R.M.Schust. .............................................................. 486
Cronisia Berk. ........................................................................ 486
Cyathodiaceae Stotler et Crand.-Stotl. ............................................. 486
Cyathodium Kunze ................................................................ 486
Dumortieraceae D.G.Long .............................................................. 487
Dumortiera Nees ................................................................... 487
Exormothecaceae Müll.Frib. ex Grolle ............................................. 487
Aitchisoniella Kashyap........................................................... 487
Exormotheca Mitt. ................................................................. 488
Stephensoniella Kashyap ....................................................... 488
Marchantiaceae Lindl. ..................................................................... 488
Bucegioideae R.M.Schust. ........................................................... 488
Bucegia Radian ...................................................................... 488
Marchantioideae Schifn. ............................................................. 489
Marchantia L. ........................................................................ 489
Preissia Corda ........................................................................ 492
Monocleaceae A.B.Frank ................................................................. 492
Monoclea Hook. .................................................................... 492
Monosoleniaceae Inoue ................................................................... 492
Monosolenium Grif. .............................................................. 492
Oxymitraceae Müll.Frib. ex Grolle .................................................. 493
Oxymitra Bisch. ex Lindenb.................................................. 493
Ricciaceae Rchb. .............................................................................. 493
Riccia L. ................................................................................. 493
Ricciocarpos Corda ................................................................ 503
Targioniaceae Dumort..................................................................... 503
World checklist of hornworts and liverworts
27
Targionia L. ........................................................................... 503
Wiesnerellaceae Inoue...................................................................... 504
Wiesnerella Schifn. ............................................................... 504
Neohodgsoniales D.G.Long ..................................................................... 504
Neohodgsoniaceae D.G.Long .......................................................... 504
Neohodgsonia Perss................................................................ 504
Sphaerocarpales Cavers ............................................................................ 504
Monocarpaceae D.J.Carr ex Schelpe ................................................ 504
Monocarpus D.J.Carr ............................................................ 504
Riellaceae Engl................................................................................. 504
Austroriella Cargill et J.Milne ............................................... 504
Riella Mont. ........................................................................... 505
Sphaerocarpaceae Heeg.................................................................... 506
Geothallus Campb. ................................................................ 506
Sphaerocarpos Boehm. .......................................................... 506
Taxonomic list
ANTHOCEROTOPHYTA
ANTHOCEROTOPSIDA de Bary ex Jancz.
Anthocerotidae Rosenv.
Anthocerotales Limpr.
***
Anthocerotaceae Dumort.
by J.C. Villarreal and D.C. Cargill
Notes on nomenclature and taxonomy can also be found in Cargill et al. (2013b) and
Villarreal et al. (2015)
*** Anthoceros L., Sp. Pl. 1: 1139, 1753 (Linnaeus 1753). 1
*** Anthoceros adscendens Lehm. et Lindenb., Nov. Stirp. Pug. 4: 24, 1832 (Lehmann
1832).
*** Anthoceros agrestis Paton, J. Bryol. 10 (3): 257, 1979 (Paton 1979a), nom. conserv. 2
*** Anthoceros alpinus Steph., Sp. Hepat. (Stephani) 6: 425, 1923 (Stephani 1923).
1
2
Anthoceros includes Aspiromitus, but some taxa have neither been transferred nor synonymized. hey
are listed in the “Names in genera not currently accepted” section below.
Anthoceros agrestis is conserved against the earlier Anthoceros nagasakiensis Steph. 1916. Anthoceros
cristatus Steph. 1916 is also an earlier synonym and a proposal to conserve Anthoceros agrestis against
it is in progress.
28
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4
5
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Anthoceros angustifolius Gottsche, Lindenb. et Nees, Syn. Hepat. 4: 582, 1846
(Gottsche et al. 1846). 3
Anthoceros angustus Steph., Sp. Hepat. (Stephani) 5: 1001, 1916 (Stephani 1916b).
Anthoceros bharadwajii Udar et A.K.Asthana, Proc. Indian Natl. Sci. Acad., B 51
(4): 484, 1985 (Udar and Asthana 1985b).
Anthoceros buettneri Steph., Sp. Hepat. (Stephani) 5: 997, 1916 (Stephani 1916b).
Anthoceros capricornii Cargill et G.A.M.Scott, J. Hattori Bot. Lab. 82: 55, 1997
(Cargill and Scott 1997).
Anthoceros caucasicus Steph., Izv. Kavkazsk. Muz. 8: 87, 1914 (Voronov’ 1914).
Anthoceros cavernosus Steph., Sp. Hepat. (Stephani) 5: 998, 1916 (Stephani
1916b).
Anthoceros chambensis Kashyap, J. Bombay Nat. Hist. Soc. 25 (2): 281, 1917
(Kashyap 1917).
Anthoceros chungii Khanna, J. Indian Bot. Soc. 17 (5/6): 316, 1938 (Khanna 1938).
Anthoceros crispatus Grif., Not. pl. asiat. 2: 349, 1849 (Griith 1849).
Anthoceros dimorphus Sim, Trans. Roy. Soc. South Africa 15 (1): 114, 1926 (Sim
1926).
Anthoceros erectus Kashyap, New Phytol. 14 (1): 9, 1915 (Kashyap 1915).
Anthoceros expansus (Steph.) J.C.Villarreal et Cargill, Phytotaxa 208 (1): 92, 2015
(Villarreal et al. 2015). Bas.: Aspiromitus expansus Steph., Sp. Hepat. (Stephani) 5:
961, 1916 (Stephani 1916b).
Anthoceros ferdinandi-muelleri Steph., Sp. Hepat. (Stephani) 5: 1007, 1916
(Stephani 1916b). 4
Anthoceros fragilis Steph., Sp. Hepat. (Stephani) 5: 1006, 1916 (Stephani 1916b).
Anthoceros fusiformis Austin, Bull. Torrey Bot. Club 6 (4): 28, 1875 [1876] (Austin
1875a).
Anthoceros fusiformis var. taiwanensis J.Haseg., Acta Phytotax. Geobot. 44 (2): 100,
1993 (Hasegawa 1993b).
Anthoceros gasongorii Gola, Mem. Reale Accad. Sci. Torino (ser. 2) 65 (1): 10,
1916 (Gola 1916).
Anthoceros granulatus Gottsche, Mexik. Leverm.: 275, 1863 (Gottsche 1863).
Anthoceros harrisanus (Steph.) Parihar, Univ. Allahabad Stud., Bot. 1961-2: 31,
1962 (Parihar 1962). Bas.: Aspiromitus harrisanus Steph., Sp. Hepat. (Stephani) 5:
965, 1916 (Stephani 1916b).
Anthoceros helmsii Steph., Hedwigia 32 (3): 142, 1893 (Stephani 1893b). 5
Anthoceros jamesonii Taylor ex Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 171,
1855 (Mitten 1855).
Anthoceros angustifolius (type from Europe) has neither been recognized in any recent treatment nor
synonymized. he name may have priority once its identity is determined.
Anthoceros ferdinandi-muelleri is similar to Anthoceros punctatus, but Anthoceros still needs to be revised for Australia.
Anthoceros helmsii was treated as conspeciic with Anthoceros muscoides by Campbell (1982), but it is
in need of a revision.
World checklist of hornworts and liverworts
*
29
***
Anthoceros javanicoides H.A.Mill., Phytologia 47 (4): 319, 1981 (Miller 1981).
Nom. nov. pro Anthoceros javanicus Steph., Sp. Hepat. (Stephani) 5: 988, 1916
(Stephani 1916b), nom. illeg. 6
Anthoceros jungermannioides Schwein., Spec. Fl. Amer. Crypt.: 25, 1821 (Schweinitz
1821).
Anthoceros kajumas (K.I.Goebel) Prosk., Bull. Torrey Bot. Club 78 (4): 347, 1951
(Proskauer 1951a). Bas.: Aspiromitus kajumas K.I.Goebel, Ann. Jard. Bot. Buitenzorg 39: 69, 1928 (Goebel 1928).
Anthoceros koshii Khanna, J. Indian Bot. Soc. 15 (4): 237, 1936 (Khanna 1936).
Anthoceros lamellatus Steph., Sp. Hepat. (Stephani) 5: 1000, 1916 (Stephani
1916b).
Anthoceros laminifer Steph., J. Linn. Soc., Bot. 29 (201): 266, 1892 (Stephani 1892b).
Anthoceros macounii M.Howe, Bull. Torrey Bot. Club 25 (1): 19, 1898 (Howe
1898a).
Anthoceros macrosporus Steph., Sp. Hepat. (Stephani) 5: 1005, 1916 (Stephani 1916b).
Anthoceros maritimus Steph., Sp. Hepat. (Stephani) 5: 984, 1916 (Stephani 1916b).
Anthoceros megasporus Meijer, J. Hattori Bot. Lab. 18: 6, 1957 (Meijer 1957).
Anthoceros muscoides Colenso, Trans. & Proc. New Zealand Inst. 16: 361, 1884
(Colenso 1884).
Anthoceros myriandroecius Steph., Wiss. Ergebn. Deut. Zentr.-Afr. Exped. (190708), Bot. 2: 134, 1911 (Stephani 1911a).
Anthoceros natalensis Steph., Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr.
88: 732, 1913 (Stephani 1913b).
Anthoceros neesii Prosk., Leberm. Eur. 2 (9): 1312, 1958 (Proskauer 1958). Nom.
nov. pro Anthoceros punctatus α* monocarpus Nees, Naturgesch. Eur. Leberm. 4:
339, 1838 (Nees 1838a).
Anthoceros niger Steph., Sp. Hepat. (Stephani) 5: 1005, 1916 (Stephani 1916b).
Anthoceros orizabensis (Steph.) Hässel, Candollea 45 (1): 211, 1990 (Hässel
1990b). Bas.: Aspiromitus orizabensis Steph., Sp. Hepat. (Stephani) 5: 965, 1916
(Stephani 1916b).
Anthoceros pandei Udar et A.K.Asthana, J. Indian Bot. Soc. 64: 305, 1985 (Udar
and Asthana 1985a).
Anthoceros patagonicus Hässel, Candollea 45 (1): 207, 1990 (Hässel 1990b).
Anthoceros patagonicus subsp. gremmenii J.C.Villarreal, J.J.Engel et Váňa, Mem.
New York Bot. Gard. 105: 32, 2013 (Váňa and Engel 2013).
Anthoceros peruvianus Steph., Sp. Hepat. (Stephani) 5: 999, 1916 (Stephani 1916b).
Anthoceros pinnatus Steph., Bol. Soc. Brot. 4: 154 [182], 1885 [1886] (Stephani
1885c).
Anthoceros punctatus L., Sp. Pl. 1: 1139, 1753 (Linnaeus 1753). 7
6
7
Anthoceros javanicoides belongs to the Anthoceros punctatus species complex (Söderström et al. 2010a).
Anthoceros punctatus is a species complex in need of a revision.
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8
9
10
11
12
13
14
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Anthoceros pusillus Colenso, Trans. & Proc. New Zealand Inst. 18: 255, 1886
(Colenso 1886b).
Anthoceros rosulans J.Haseg., J. Hattori Bot. Lab. 60: 379, 1986 (Hasegawa 1986b).
Anthoceros sambesianus Steph., Sp. Hepat. (Stephani) 5: 996, 1916 (Stephani 1916b).
Anthoceros scariosus Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 230, 1869 (Austin 1869).
Anthoceros schroederi Steph., 52 (5): 306, 1912 (Stephani 1912a).
Anthoceros serratus Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9):
90, 1911 (Stephani 1911b).
Anthoceros simulans M.Howe, Proc. Calif. Acad. Sci. (ser. 4) 21 (17): 204, 1934
(Howe 1934).
Anthoceros spongiosus Steph., Sp. Hepat. (Stephani) 5: 1003, 1916 (Stephani
1916b).
Anthoceros subtilis Steph., Sp. Hepat. (Stephani) 5: 1003, 1916 (Stephani 1916b).
Anthoceros telaganus Steph., Sp. Hepat. (Stephani) 5: 1005, 1916 (Stephani 1916b).
Anthoceros tristanianus J.C.Villarreal, J.J.Engel et Váňa, Mem. New York Bot.
Gard. 105: 33, 2013 (Váňa and Engel 2013).
Anthoceros tuberculatus Lehm. et Lindenb., Nov. Stirp. Pug. 4: 25, 1832 (Lehmann
1832).
Anthoceros venosus Lindenb. et Gottsche, Syn. Hepat. 4: 584, 1846 (Gottsche et al. 1846).
Excluded from the genus
Anthoceros aethyopicus Gola, Ann. Bot. (Rome) 13 (1): 73, 1914 (Gola 1914a). 8
Anthoceros brunnthaleri Steph., Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr. 88: 732, 1913 (Stephani 1913b). 9
Anthoceros loribundus Steph., Sp. Hepat. (Stephani) 5: 977, 1916 (Stephani
1916b). 10
Anthoceros mildbraedii Steph., Sp. Hepat. (Stephani) 6: 428, 1923 (Stephani 1923). 11
Anthoceros parvifrons Steph., 52 (5): 307, 1912 (Stephani 1912a). 12
Anthoceros pseudocostus Steph., Sp. Hepat. (Stephani) 5: 977, 1916 (Stephani 1916b). 13
Anthoceros rossoi Gola, Mem. Reale Accad. Sci. Torino (ser. 2) 65 (1): 10, 1916
(Gola 1916). 14
Anthoceros aethyopicus is probably a Phaeoceros species (Wigginton 2009).
Anthoceros brunnthaleri was annotated by Hasegawa in 1993 as probably conspeciic with Phymatoceros
bulbiculosus but the synonymy seems not to have been published.
Anthoceros loribundus type specimen in G was annotated by Hasegawa in 1993 as conspeciic with
Anthoceros parvifrons which is a Phaeoceros species.
Anthoceros mildbraedii is probably conspeciic with Phaeoceros carolinianus (Wigginton and Grolle 1996).
Anthoceros parvifrons is annotated as a Phaeoceros species by Hasegawa in 1993 but the new combination
has never been published.
Anthoceros pseudocostus is a Phaeoceros species of uncertain status (Grolle 1995).
Anthoceros rossoi is a Phaeoceros species of uncertain status (Wigginton 2009).
World checklist of hornworts and liverworts
31
*** Folioceros D.C.Bharadwaj, Geophytology 1 (1): 9, 1971 (Bharadwaj 1971).
*** Folioceros amboinensis (Schifn.) Piippo, Acta Bot. Fenn. 148: 36, 1993 (Piippo
1993b). Bas.: Anthoceros amboinensis Schifn., Leberm., Forschungsr. Gazelle 4
(4): 45, 1890 (Schifner 1890).
** Folioceros apiahynus (Steph.) Hässel, Candollea 45 (1): 215, 1990 (Hässel 1990b).
Bas.: Anthoceros apiahynus Steph., Sp. Hepat. (Stephani) 5: 999, 1916 (Stephani
1916b).
** Folioceros argillaceus (Steph.) J.C.Villarreal et Cargill, Phytotaxa 208 (1): 93, 2015
(Villarreal et al. 2015). Bas.: Aspiromitus argillaceus Steph., Sp. Hepat. (Stephani)
5: 970, 1916 (Stephani 1916b).
*** Folioceros assamicus D.C.Bharadwaj, Geophytology 1 (1): 10, 1971 (Bharadwaj 1971).
** Folioceros dilatatus (Steph.) J.C.Villarreal et Cargill, Phytotaxa 208 (1): 93, 2015
(Villarreal et al. 2015). Bas.: Anthoceros dilatatus Steph., Bot. Jahrb. Syst. 8 (2): 95,
1886 (Stephani 1886d).
*** Folioceros dixitianus (Mahab.) D.C.Bharadwaj, Geophytology 5 (2): 227, 1975
(Bharadwaj 1975). Bas.: Aspiromitus dixitianus Mahab., Curr. Sci. 10 (12): 532,
1941 (Mahabale 1941).
*** Folioceros fuciformis (Mont.) D.C.Bharadwaj, Geophytology 5 (2): 227, 1975
(Bharadwaj 1975). Bas.: Anthoceros fuciformis Mont., Ann. Sci. Nat. Bot. (sér. 2)
20: 296, 1844 (Montagne 1844b).
*** Folioceros glandulosus (Lehm. et Lindenb.) D.C.Bharadwaj, Geophytology 5 (2):
227, 1975 (Bharadwaj 1975). Bas.: Anthoceros glandulosus Lehm. et Lindenb.,
Nov. Stirp. Pug. 4: 26, 1832 (Lehmann 1832).
*** Folioceros incurvus (Steph.) D.C.Bharadwaj, Geophytology 5 (2): 227, 1975
(Bharadwaj 1975). Bas.: Anthoceros incurvus Steph., Hedwigia 32 (3): 143, 1893
(Stephani 1893b).
*** Folioceros indicus D.C.Bharadwaj, Geophytology 8 (1): 114, 1978 (Bharadwaj 1978).
*** Folioceros kashyapii S.C.Srivast. et A.K.Asthana, Bryologist 92 (2): 219, 1989
(Srivastava and Asthana 1989).
*** Folioceros mangaloreus (Steph.) D.C.Bharadwaj, Geophytology 5 (2): 227, 1975
(Bharadwaj 1975). Bas.: Aspiromitus mangaloreus Steph., Sp. Hepat. (Stephani) 5:
967, 1916 (Stephani 1916b).
*** Folioceros paliformis D.K.Singh, Bull. Bot. Surv. India 29: 176, 1987 [1989] (Singh
1987b).
*** Folioceros physocladus D.C.Bharadwaj, Geophytology 8 (1): 115, 1978 (Bharadwaj
1978). Based on: Anthoceros physocladus Schifn. ex Pandé, Proc. Indian Sci. Congr.
Assoc. 47 (2): 98, 1960 (Pandé 1960), nom. inval.
*** Folioceros pinnilobus (Steph.) D.C.Bharadwaj, Geophytology 5 (2): 227, 1975
(Bharadwaj 1975). Bas.: Anthoceros pinnilobus Steph., Akad. Wiss. Wien, Math.Naturwiss. Kl., Denkschr. 81: 299, 1907 (Stephani 1907a).
*** Folioceros satpurensis D.C.Bharadwaj et K.P.Srivast., Geophytology 8 (1): 112,
1978 (Bharadwaj 1978). Based on: Anthoceros satpurensis K.P.Srivast., Proc. Indian Sci. Congr. Assoc. 47 (3): 337, 1960 (Srivastava 1960), nom. inval.
32
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Folioceros udarii A.K.Asthana et S.C.Srivast., Cryptog. Bryol. Lichénol. 7 (2): 151,
1986 (Asthana and Srivastava 1986).
*** Folioceros verruculosus (J.Haseg.) R.L.Zhu et M.J.Lai, Ann. Bot. Fenn. 48 (5): 383,
2011 (Wang et al. 2011). Bas.: Anthoceros verruculosus J.Haseg., Acta Phytotax.
Geobot. 44 (2): 103, 1993 (Hasegawa 1993b).
Dendrocerotidae R.J.Duff, J.C.Villarreal, Cargill et Renzaglia
Dendrocerotales Hässel
***
Dendrocerotaceae J.Haseg.
by J.C. Villarreal and D.C. Cargill
*** Dendroceros Nees, Syn. Hepat. 4: 579, 1846 (Gottsche et al. 1846).
** subg. Apoceros R.M.Schust., Phytologia 63 (3): 200, 1987 (Schuster 1987b).
*** Dendroceros cavernosus J.Haseg., J. Hattori Bot. Lab. 47: 306, 1980 (Hasegawa 1980).
*** Dendroceros diicilis Steph., Sp. Hepat. (Stephani) 5: 1009, 1917 (Stephani
1917b).
*** Dendroceros muelleri Steph., Hedwigia 28 (2): 133, 1889 (Stephani 1889a).
*** Dendroceros ogeramnangus Piippo, Acta Bot. Fenn. 148: 40, 1993 (Piippo 1993b).
*** Dendroceros pedunculatus Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 14, 1909
(Stephani 1909c).
** Dendroceros subdiicilis S.Hatt., Bot. Mag. (Tokyo) 64 (755/756): 119, 1951
(Hattori 1951c).
** subg. Dendroceros
*** Dendroceros acutilobus Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 18, 1909
(Stephani 1909c).
*** Dendroceros borbonicus Steph., Bull. Soc. Roy. Bot. Belgique, Compt. Rend. 32
(2): 31, 1893 [1894] (Stephani 1893e).
*** Dendroceros crispus (Sw.) Nees, Syn. Hepat. 4: 581, 1846 (Gottsche et al. 1846).
Bas.: Anthoceros crispus Sw., Prodr. (Swartz): 146, 1788 (Swartz 1788).
*** Dendroceros foliicola J.Haseg., J. Hattori Bot. Lab. 47: 296, 1980 (Hasegawa 1980).
*** Dendroceros japonicus Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 15, 1909
(Stephani 1909c).
*** Dendroceros javanicus (Nees) Nees, Syn. Hepat. 4: 582, 1846 (Gottsche et al. 1846).
Bas.: Anthoceros javanicus Nees, Enum. Pl. Crypt. Javae: 1, 1830 (Nees 1830).
*** Dendroceros subplanus Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 20, 1909
(Stephani 1909c).
*** Dendroceros tubercularis S.Hatt., Bot. Mag. (Tokyo) 58 (685): 6, 1944 (Hattori 1944b).
World checklist of hornworts and liverworts
33
*** Dendroceros validus Steph., Sp. Hepat. (Stephani) 5: 1016, 1917 (Stephani 1917b).
** Dendroceros vesconianus Gottsche, J. Bot. (Morot) 12: 150, 1898 (Bescherelle 1898).
** Dendroceros wattsianus Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 17, 1909
(Stephani 1909c).
*
***
**
**
**
**
***
*
**
***
**
**
*
*
***
**
**
**
Incertae sedis
Dendroceros adglutinatus (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn.
Hepat. 4: 580, 1846 (Gottsche et al. 1846). Bas.: Monoclea adglutinata Hook.f. et
Taylor, London J. Bot. 4: 96, 1845 (Hooker and Taylor 1845). 15
Dendroceros africanus Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 18, 1909
(Stephani 1909c).
Dendroceros allionii Steph., Sp. Hepat. (Stephani) 5: 1014, 1917 (Stephani 1917b).
Dendroceros australis Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 17, 1909 (Stephani 1909c).
Dendroceros breutelii Nees, Syn. Hepat. 4: 581, 1846 (Gottsche et al. 1846).
Dendroceros breutelii var. surinamensis Lindenb. et Gottsche, Linnaea 24 (6): 639,
1851 [1852] (Lindenberg and Gottsche 1851a).
Dendroceros cichoraceus (Mont.) Gottsche, Bot. Zeitung (Berlin) Beil. 16: 16, 1858
(Gottsche 1858). Bas.: Anthoceros cichoraceus Mont., Ann. Sci. Nat. Bot. (sér. 3) 4:
355, 1845 (Montagne 1845b).
Dendroceros crassicostatus Steph., Sp. Hepat. (Stephani) 5: 1015, 1917 (Stephani 1917b).
Dendroceros crassinervis (Nees) Gottsche, Bot. Zeitung (Berlin) Beil. 16: 18, 1858
(Gottsche 1858). Bas.: Anthoceros crassinervis Nees, Syn. Hepat. 4: 589, 1846
(Gottsche et al. 1846).
Dendroceros crispatus (Hook.) Nees, Syn. Hepat. 4: 579, 1846 (Gottsche et al.
1846). Bas.: Monoclea crispata Hook., Bot. Misc. 1: 117, 1830 (Hooker 1830).
Dendroceros crispatus var. simplicior Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 574, 1885 (Spruce 1885).
Dendroceros cucullatus Steph., Sp. Hepat. (Stephani) 6: 429, 1923 (Stephani 1923). 16
Dendroceros exalatus Steph., Sitzungsber. Naturf. Ges. Leipzig 36: 14, 1909
(Stephani 1909c). 17
Dendroceros gracilis Steph., Sp. Hepat. (Stephani) 5: 1015, 1917 (Stephani 1917b).
Dendroceros granulatus Mitt., Fl. vit.: 419, 1871 [1873] (Mitten 1871).
Dendroceros herasii M.Infante, J. Bryol. 32 (4): 285, 2010 (Infante 2010).
Dendroceros humboldtensis Hürl., Bauhinia 1 (3): 254, 1960 (Hürlimann 1960).
Dendroceros paivae C.A.Garcia, Sérgio et J.C.Villarreal, Cryptog. Bryol. 33 (1): 5,
2012 (Garcia et al. 2012).
15 Dendroceros adglutinatus is conspeciic with Dendroceros crispus in Proskauer (1960), but later authors
accept it.
16 Dendroceros cucullatus was treated as conspeciic with Dendroceros diicilis by Hasegawa (1980), but
re-instated by Chantanaorrapint et al. (2014).
17 Dendroceros exalatus was reported from the Moluccas in Stephani (1909c), but Hasegawa (1980)
states that this is a Brazilian species. Dendroceros has not been recently revised for Brazil.
34
*
*
**
***
**
**
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Dendroceros rarus Steph., Sp. Hepat. (Stephani) 5: 1014, 1917 (Stephani 1917b).
Dendroceros reticulus Herzog, Memoranda Soc. Fauna Fl. Fennica 26: 37, 1950
[1951] (Herzog 1950b). 18
Dendroceros rigidus Steph., Sp. Hepat. (Stephani) 5: 1017, 1917 (Stephani 1917b).
Dendroceros seramensis J.Haseg., Acta Phytotax. Geobot. 37 (1/3): 10, 1986
(Hasegawa 1986a).
Dendroceros subtropicus C.J.Wild, Trans. Nat. Hist. Soc. Queensland 1: 49, 1893
(Wild 1893).
Dendroceros tahitensis Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 30 (5): 138,
1873 (Ångström 1873).
*** Megaceros Campb., Ann. Bot. (Oxford) 21 (4): 484, 1907 (Campbell 1907).
** Megaceros aneuriformis Steph., Sp. Hepat. (Stephani) 5: 949, 1916 (Stephani 1916b).
*** Megaceros austronesophilus Cargill et Seppelt, Austral. Syst. Bot. 26 (5): 372, 2013
(Cargill et al. 2013a).
** Megaceros ciliatus K.I.Goebel, Ann. Jard. Bot. Buitenzorg 39: 74, 1928 (Goebel 1928).
*** Megaceros denticulatus (Lehm.) Steph., Sp. Hepat. (Stephani) 5: 956, 1916
(Stephani 1916b). Bas.: Anthoceros denticulatus Lehm., Nov. Stirp. Pug. 10: 25,
1857 (Lehmann 1857).
*** Megaceros lagellaris (Mitt.) Steph., Sp. Hepat. (Stephani) 5: 951, 1916 (Stephani
1916b). Bas.: Anthoceros lagellaris Mitt., Fl. vit.: 419, 1871 [1873] (Mitten 1871).
*** Megaceros gracilis (Reichardt) Steph., Sp. Hepat. (Stephani) 5: 955, 1916 (Stephani 1916b). Bas.: Anthoceros gracilis Reichardt, Verh. K.K. Zool.-Bot. Ges. Wien
16: 957, 1866 (Reichardt 1866).
** Megaceros leptohymenius (Hook.f. et Taylor) Steph., Sp. Hepat. (Stephani) 5: 955,
1916 (Stephani 1916b). Bas.: Monoclea leptohymenia Hook.f. et Taylor, London J.
Bot. 3: 575, 1844 (Hooker and Taylor 1844d).
*** Megaceros pellucidus (Colenso) E.A.Hodgs., J. Roy. Soc. New Zealand 2 (1): 115,
1972 (Hodgson 1972). Bas.: Anthoceros pellucidus Colenso, Trans. & Proc. New
Zealand Inst. 17: 263, 1885 (Colenso 1885).
** Megaceros tjibodensis Campb., Ann. Bot. (Oxford) 21 (4): 484, 1907 (Campbell 1907).
*
*
Excluded from the genus 19
Megaceros lavens (Spruce) Campb., Ann. Bot. (Oxford) 21 (4): 483, 1907 (Campbell 1907). Bas.: Anthoceros lavens Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 575, 1885 (Spruce 1885).
Megaceros jamesonii (Taylor) Steph., Biblioth. Bot. 87 (2): 268, 1916 (Stephani 1916a).
Bas.: Dendroceros jamesonii Taylor, London J. Bot. 7: 285, 1848 (Taylor 1848b).
18 Dendroceros reticulus was mentioned by Hasegawa (1980). He did not synonymise it with Dendroceros
javanicus, he just stated that Herzog thought it similar to Dendroceros elegans which is conspeciic with
Dendroceros javanicus.
19 Megaceros lavens and Megaceros jamesonii are probably conspeciic with Nothoceros vincentianus
(Villarreal et al. 2010a).
World checklist of hornworts and liverworts
35
*** Nothoceros (R.M.Schust.) J.Haseg., J. Hattori Bot. Lab. 76: 32, 1994 (Hasegawa
1994b). Bas.: Megaceros subg. Nothoceros R.M.Schust., Phytologia 63 (3): 200,
1987 (Schuster 1987b).
*** Nothoceros aenigmaticus J.C.Villarreal et K.D.McFarland, Bryologist 113 (1): 109,
2010 (Villarreal et al. 2010b). Based on: Megaceros aenigmaticus R.M.Schust.,
Hepat. Anthocerotae N. Amer. 6: 830, 1992 (Schuster 1992d), nom. inval.
*** Nothoceros canaliculatus (Pagán) J.C.Villarreal, Hässel et N.Salazar, Bryologist 110
(2): 283, 2007 (Villarreal et al. 2007). Bas.: Dendroceros canaliculatus Pagán, Bryologist 45 (4): 111, 1942 (Pagán 1942a).
*** Nothoceros endiviifolius (Mont.) J.Haseg. ex J.C.Villarreal, Hässel et N.Salazar,
Bryologist 110 (2): 283, 2007 (Villarreal et al. 2007). Bas.: Anthoceros endiviifolius
Mont., Voy. Pole Sud, Bot. 1: 211, 1845 (Montagne 1845c).
** Nothoceros fuegiensis (Steph.) J.C.Villarreal, Bryologist 113 (1): 109, 2010 (Villarreal et al. 2010b). Bas.: Megaceros fuegiensis Steph., Kungl. Svenska Vetensk.-Akad.
Handl. (n.ser.) 46 (9): 91, 1911 (Stephani 1911b).
*** Nothoceros giganteus (Lehm. et Lindenb.) J.Haseg. ex J.C.Villarreal, Hässel et
N.Salazar, Bryologist 110 (2): 283, 2007 (Villarreal et al. 2007). Bas.: Anthoceros
giganteus Lehm. et Lindenb., Nov. Stirp. Pug. 4: 25, 1832 (Lehmann 1832).
*** Nothoceros minarum (Nees) J.C.Villarreal, Molec. Phylogen. Evol. 78: 34, 2014
(Villarreal and Renner 2014). Bas.: Anthoceros minarum Nees, Naturgesch. Eur.
Leberm. 4: 340, 1838 (Nees 1838a).
** Nothoceros renzagliensis J.C.Villarreal, L.V.Campos et Uribe, Syst. Bot. 37 (1): 32,
2012 (Villarreal et al. 2012).
*** Nothoceros schizophyllus (Steph.) J.C.Villarreal, Molec. Phylogen. Evol. 78: 34,
2014 (Villarreal and Renner 2014). Bas.: Megaceros schizophyllus Steph., Sp.
Hepat. (Stephani) 5: 949, 1916 (Stephani 1916b).
*** Nothoceros superbus J.C.Villarreal, Hässel et N.Salazar, Bryologist 110 (2): 280,
2007 (Villarreal et al. 2007).
*** Nothoceros vincentianus (Lehm. et Lindenb.) J.C.Villarreal, Bryologist 113 (1):
111, 2010 (Villarreal et al. 2010b). Bas.: Anthoceros vincentianus Lehm. et Lindenb., Nov. Stirp. Pug. 6: 16, 1834 (Lehmann 1834).
** Phaeomegaceros R.J.Duf, J.C.Villarreal, Cargill et Renzaglia, Bryologist 110
(2): 241, 2007 (Duf et al. 2007).
*** Phaeomegaceros coriaceus (Steph.) R.J.Duf, J.C.Villarreal, Cargill et Renzaglia,
Bryologist 110 (2): 241, 2007 (Duf et al. 2007). Bas.: Anthoceros coriaceus Steph.,
Sp. Hepat. (Stephani) 5: 991, 1916 (Stephani 1916b).
*** Phaeomegaceros imbriatus (Gottsche) R.J.Duf, J.C.Villarreal, Cargill et Renzaglia, Bryologist 110 (2): 241, 2007 (Duf et al. 2007). Bas.: Anthoceros imbriatus
Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 187, 1864 (Gottsche 1864).
*** Phaeomegaceros foveatus (J.Haseg.) J.C.Villarreal, Nova Hedwigia 91 (3/4): 352,
2010 (Villarreal et al. 2010a). Bas.: Phaeoceros foveatus J.Haseg., Bryol. Res. 7 (12):
374, 2001 (Hasegawa 2001).
36
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Phaeomegaceros hirticalyx (Steph.) R.J.Duf, J.C.Villarreal, Cargill et Renzaglia,
Bryologist 110 (2): 241, 2007 (Duf et al. 2007). Bas.: Aspiromitus hirticalyx Steph., Sp. Hepat. (Stephani) 5: 966, 1916 (Stephani 1916b).
*** Phaeomegaceros plicatus (Mitt.) J.C.Villarreal, J.J.Engel et Váňa, Mem. New York
Bot. Gard. 105: 85, 2013 (Váňa and Engel 2013). Bas.: Anthoceros plicatus Mitt.,
Rep. Challenger, Bot. 1 (2): 178, 1884 (Mitten 1884a).
*** Phaeomegaceros skottsbergii (Steph.) R.J.Duf, J.C.Villarreal, Cargill et Renzaglia,
Bryologist 110 (2): 241, 2007 (Duf et al. 2007). Bas.: Anthoceros skottsbergii Steph.,
Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9): 90, 1911 (Stephani 1911b).
*** Phaeomegaceros squamuliger (Spruce) J.C.Villarreal, Nova Hedwigia 91 (3/4): 351,
2010 (Villarreal et al. 2010a). Bas.: Anthoceros squamuliger Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 576, 1885 (Spruce 1885).
** Phaeomegaceros squamuliger subsp. hasselii J.C.Villarreal, Cargill et Goinet, Nova
Hedwigia 91 (3/4): 352, 2010 (Villarreal et al. 2010a).
Phymatocerotales R.J.Duff, J.C.Villarreal, Cargill et Renzaglia
***
Phymatocerotaceae R.J.Duff, J.C.Villarreal, Cargill et Renzaglia
by J.C. Villarreal and D.C. Cargill
*** Phymatoceros Stotler, W.T.Doyle et Crand.-Stotl., Phytologia 87 (2): 113,
2005 (Stotler et al. 2005).
*** Phymatoceros bulbiculosus (Brot.) Stotler, W.T.Doyle et Crand.-Stotl., Phytologia
87 (2): 114, 2005 (Stotler et al. 2005). Bas.: Anthoceros bulbiculosus Brot., Fl. lusit.
2: 430, 1804 [1805] (Brotero 1804).
*** Phymatoceros phymatodes (M.Howe) R.J.Duf, J.C.Villarreal, Cargill et Renzaglia,
Bryologist 110 (2): 240, 2007 (Duf et al. 2007). Bas.: Anthoceros phymatodes
M.Howe, Bull. Torrey Bot. Club 25 (1): 12, 1898 (Howe 1898a).
Notothylatidae R.J.Duff, J.C.Villarreal, Cargill et Renzaglia
Notothyladales Hyvönen et Piippo
***
Notothyladaceae Müll.Frib. ex Prosk.
by J.C. Villarreal and D.C. Cargill
World checklist of hornworts and liverworts
**
37
Notothyladoideae Grolle
*** Notothylas Sull. ex A.Gray, Amer. J. Sci. Arts (ser. 2) 1 (1): 74, 1846 (Gray 1846).
** subg. Notothylas
*** Notothylas anaporata Udar et D.K.Singh, Rev. Bryol. Lichénol. 45 (2): 202, 1979
(Udar and Singh 1979b).
*** Notothylas breutelii (Gottsche) Gottsche, Bot. Zeitung (Berlin) Beil. 16: 21, 1858
(Gottsche 1858). Bas.: Anthoceros breutelii Gottsche, Syn. Hepat. 4: 583, 1846
(Gottsche et al. 1846).
*** Notothylas depressispora J.Haseg., Acta Phytotax. Geobot. 30 (1/3): 26, 1979
(Hasegawa 1979).
*** Notothylas dissecta Steph., Sp. Hepat. (Stephani) 5: 1020, 1917 (Stephani 1917b).
*** Notothylas himalayensis Udar et D.K.Singh, J. Bryol. 11 (3): 451, 1981 (Udar and
Singh 1981a).
*** Notothylas indica Kashyap, Proc. Lahore Philos. Soc. 4: 54, 1925 (Kashyap and
Dutt 1925).
*** Notothylas javanica (Sande Lac.) Gottsche, Bot. Zeitung (Berlin) Beil. 16: 20,
1858 (Gottsche 1858). Bas.: Blasia javanica Sande Lac., Syn. hepat. jav.: 94, 1856
[1857] (Sande Lacoste 1856b).
*** Notothylas orbicularis (Schwein.) Sull., Amer. J. Sci. Arts (ser. 2) 1 (1): 75, 1846
(Gray 1846). Bas.: Targionia orbicularis Schwein., Spec. Fl. Amer. Crypt.: 23,
1821 (Schweinitz 1821).
*** Notothylas pandei Udar et V.Chandra, Geophytology 7 (2): 142, 1977 (Udar and
Chandra 1977).
** subg. Notothyloides A.K.Asthana et S.C.Srivast., Bryophyt. Biblioth. 42: 106,
1991 (Asthana and Srivastava 1991).
*** Notothylas khasiana Udar et D.K.Singh, J. Indian Bot. Soc. 60: 112, 1981 (Udar
and Singh 1981b).
*** Notothylas pleidereri Udar et D.K.Singh, Lindbergia 5 (1): 28, 1979 (Udar and
Singh 1979a).
Incertae sedis
*** Notothylas decurva (Mitt.) Steph., Cat. Afr. Pl. (Hiern) 2 (2): 320, 1901 (Stephani
1901d). Bas.: Anthoceros decurvus Mitt., Trans. Linn. Soc. London 23 (1): 58,
1860 (Mitten 1860a).
*** Notothylas labellata Steph., Cat. Afr. Pl. (Hiern) 2 (2): 320, 1901 (Stephani 1901d).
*** Notothylas galapagensis M.Howe, Proc. Calif. Acad. Sci. (ser. 4) 21 (17): 203, 1934
(Howe 1934).
** Notothylas irregularis Chantanaorr., Acta Bot. Hung. 56 (3/4): 270, 2014 (Chantanaorrapint 2014).
38
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Notothylas kashyapii D.K.Singh, Indian J. Forest. 23 (4): 386, 2000 (Singh and
Semwal 2000).
*** Notothylas nepalensis D.K.Singh, J. Bombay Nat. Hist. Soc. 84 (3): 650, 1987
[1988] (Singh 1987a).
*** Notothylas temperata J.Haseg., Acta Phytotax. Geobot. 30 (1/3): 20, 1979 (Hasegawa 1979).
*** Notothylas udarii D.K.Singh et Semwal, Phytotaxonomy 1: 35, 2001 (Singh and
Semwal 2001).
* Notothylas verdoornii Khanna, Rev. Bryol. Lichénol. 6: 118, 1933 (Khanna 1933).
*** Notothylas vitalii Udar et D.K.Singh, Misc. Bryol. Lichenol. 8 (9): 173, 1980
(Udar and Singh 1980).
** Notothylas yunnanensis T.Peng et R.L.Zhu, Phytotaxa 156 (3): 157, 2014 (Peng
and Zhu 2014).
**
Phaeocerotoideae Hässel
*
*
*
Mesoceros Piippo, Acta Bot. Fenn. 148: 30, 1993 (Piippo 1993b). 20
Mesoceros mesophoros Piippo, Acta Bot. Fenn. 148: 30, 1993 (Piippo 1993b).
Mesoceros porcatus Piippo, Haussknechtia, Beih. 9: 279, 1999 (Piippo 1999).
** Paraphymatoceros Hässel, Phytologia 88 (2): 208, 2006 (Hässel 2006a).
*** Paraphymatoceros diadematus Hässel, Phytologia 88 (2): 209, 2006 (Hässel 2006a).
*** Paraphymatoceros hallii (Austin) Hässel, Phytologia 88 (2): 209, 2006 (Hässel
2006a). Bas.: Anthoceros hallii Austin, Bull. Torrey Bot. Club 6 (4): 26, 1875
[1876] (Austin 1875a).
*** Paraphymatoceros pearsonii (M.Howe) J.C.Villarreal et Cargill, Phytotaxa 208 (1):
94, 2015 (Villarreal et al. 2015). Bas.: Anthoceros pearsonii M.Howe, Bull. Torrey
Bot. Club 25 (1): 8, 1898 (Howe 1898a).
*** Paraphymatoceros proskaueri (Stotler, Crand.-Stotl. et W.T.Doyle) J.C.Villarreal
et Cargill, Phytotaxa 208 (1): 94, 2015 (Villarreal et al. 2015). Bas.: Phaeoceros
proskaueri Stotler, Crand.-Stotl. et W.T.Doyle, Fieldiana, Bot. (n.ser.) 47: 232,
2008 (Crandall-Stotler et al. 2008a).
*** Phaeoceros Prosk., Bull. Torrey Bot. Club 78 (4): 346, 1951 (Proskauer 1951a).
** Phaeoceros austroandinus Hässel, Candollea 44 (2): 721, 1989 (Hässel 1989a).
** Phaeoceros bolusii (Sim) S.W.Arnell, Hepat. South Africa: 401, 1963 (Arnell
1963b). Bas.: Anthoceros bolusii Sim, Trans. Roy. Soc. South Africa 15 (1): 114,
1926 (Sim 1926).
20 Mesoceros was described from a collection that contains a mixture of both Anthoceros and Phaeoceros
material.
World checklist of hornworts and liverworts
39
** Phaeoceros brevicapsulus (Steph.) Hässel, Candollea 44 (2): 725, 1989 (Hässel
1989a). Bas.: Anthoceros brevicapsulus Steph., Sp. Hepat. (Stephani) 5: 981, 1916
(Stephani 1916b).
*** Phaeoceros carolinianus (Michx.) Prosk., Bull. Torrey Bot. Club 78 (4): 347, 1951
(Proskauer 1951a). Bas.: Anthoceros carolinianus Michx., Fl. bor.-amer. (Michaux)
2: 280, 1803 (Michaux 1803).
*** Phaeoceros delicatus E.O.Campb. et Outred, New Zealand J. Bot. 33 (3): 285,
1995 (Campbell and Outred 1995).
*** Phaeoceros dendroceroides (Steph.) Hässel, Beih. Nova Hedwigia 134: 487, 2009
(Hässel and Rubies 2009). Bas.: Anthoceros dendroceroides Steph., Sp. Hepat. (Stephani) 5: 984, 1916 (Stephani 1916b). 21
*** Phaeoceros engelii Cargill et Fuhrer, Fieldiana, Bot. (n.ser.) 47: 248, 2008 (Cargill
and Fuhrer 2008).
** Phaeoceros erectus Udar et D.K.Singh, Geophytology 11 (2): 257, 1981 (Udar and
Singh 1981c). Nom. nov. pro Anthoceros erectus Steph., Sp. Hepat. (Stephani) 5:
984, 1916 (Stephani 1916b), nom. illeg.
*** Phaeoceros evanidus (Steph.) Cargill et Fuhrer, Fieldiana, Bot. (n.ser.) 47: 245,
2008 (Cargill and Fuhrer 2008). Bas.: Anthoceros evanidus Steph., Sp. Hepat.
(Stephani) 5: 990, 1916 (Stephani 1916b).
*** Phaeoceros exiguus (Steph.) J.Haseg., J. Hattori Bot. Lab. 60: 387, 1986 (Hasegawa 1986b). Bas.: Anthoceros exiguus Steph., Sp. Hepat. (Stephani) 5: 988, 1916
(Stephani 1916b).
*** Phaeoceros lexivalvis (Gottsche et Nees) Hässel, Candollea 44 (2): 728, 1989 (Hässel 1989a). Bas.: Anthoceros lexivalvis Gottsche et Nees, Syn. Hepat. 4: 586, 1846
(Gottsche et al. 1846).
*** Phaeoceros fulvisporus (Steph.) J.Haseg., Trop. Bryol. 8: 52, 1993 (Hasegawa
1993a). Bas.: Anthoceros fulvisporus Steph., 52 (5): 306, 1912 (Stephani 1912a).
*** Phaeoceros gemmifer (Horik.) J.Haseg., J. Hattori Bot. Lab. 57: 252, 1984 (Hasegawa 1984). Bas.: Anthoceros gemmifer Horik., Sci. Rep. Tôhoku Imp. Univ., Ser. 4,
Biol. 4 (2): 426, 1929 (Horikawa 1929a).
** Phaeoceros gualaquizanus (Steph.) Gradst., J. Hattori Bot. Lab. 45: 123, 1979
(Gradstein and Hekking 1979). Bas.: Anthoceros gualaquizanus Steph., Sp. Hepat.
(Stephani) 5: 979, 1916 (Stephani 1916b).
*** Phaeoceros himalayensis (Kashyap) Prosk. ex Bapna et G.G.Vyas, J. Hattori Bot.
Lab. 25: 88, 1962 (Bapna and Vyas 1962). Bas.: Anthoceros himalayensis Kashyap,
New Phytol. 14 (1): 8, 1915 (Kashyap 1915).
*** Phaeoceros huebschmannii Hässel, Veröf. Geobot. Inst. ETH Stiftung Rübel
Zürich 91: 301, 1986 (Hässel 1986b).
*** Phaeoceros inlatus (Steph.) Cargill et Fuhrer, Fieldiana, Bot. (n.ser.) 47: 246, 2008
(Cargill and Fuhrer 2008). Bas.: Anthoceros inlatus Steph., Sp. Hepat. (Stephani)
5: 990, 1916 (Stephani 1916b).
21 Phaeoceros dendroceroides may belong to Phaeomegaceros
40
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Phaeoceros kashyapii A.K.Asthana et S.C.Srivast., Bryophyt. Biblioth. 42: 129,
1991 (Asthana and Srivastava 1991).
*** Phaeoceros laevis (L.) Prosk., Bull. Torrey Bot. Club 78 (4): 347, 1951 (Proskauer
1951a). Bas.: Anthoceros laevis L., Sp. Pl. 1: 1139, 1753 (Linnaeus 1753).
** Phaeoceros maranguensis (Steph.) Bapna, Univ. Udaipur Res. Stud. 3: 138, 1966
(Bapna 1966). Bas.: Aspiromitus maranguensis Steph., Sp. Hepat. (Stephani) 5:
960, 1916 (Stephani 1916b).
* Phaeoceros microsporus (Steph.) Hässel, Candollea 44 (2): 721, 1989 (Hässel
1989a). Bas.: Aspiromitus microsporus Steph., Sp. Hepat. (Stephani) 5: 963, 1916
(Stephani 1916b). 22
*** Phaeoceros minutus (Mitt.) S.W.Arnell, Hepat. South Africa: 403, 1963 (Arnell 1963b).
Bas.: Anthoceros minutus Mitt., J. Linn. Soc., Bot. 16 (91): 195, 1877 (Mitten 1877).
*** Phaeoceros mohrii (Austin) Hässel, Candollea 44 (2): 721, 1989 (Hässel 1989a). Bas.:
Anthoceros mohrii Austin, Bull. Torrey Bot. Club 6 (52): 304, 1879 (Austin 1879).
*** Phaeoceros oreganus (Austin) Hässel, Candollea 44 (2): 718, 1989 (Hässel 1989a).
Bas.: Anthoceros oreganus Austin, Bull. Torrey Bot. Club 6 (4): 26, 1875 [1876]
(Austin 1875a).
*** Phaeoceros parvulus (Schifn.) J.Haseg., J. Hattori Bot. Lab. 57: 248, 1984 (Hasegawa 1984). Bas.: Anthoceros parvulus Schifn., Österr. Bot. Z. 49 (11): 391, 1899
(Schifner 1899c).
*** Phaeoceros perpusillus Chantanaorr., Acta Bot. Hung. 51 (1/2): 30, 2009 (Chantanaorrapint 2009).
** Phaeoceros pichinchensis (Spruce) Hässel, Candollea 44 (2): 725, 1989 (Hässel
1989a). Bas.: Anthoceros pichinchensis Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 577, 1885 (Spruce 1885).
** Phaeoceros propagulifer (Steph.) Prosk., Bull. Torrey Bot. Club 78 (4): 347, 1951
(Proskauer 1951a). Bas.: Anthoceros propagulifer Steph., Sp. Hepat. (Stephani) 5:
1001, 1916 (Stephani 1916b).
* Phaeoceros striatisporus J.Haseg., J. Hattori Bot. Lab. 75, 268 (Hasegawa 1994a).23
*** Phaeoceros tenuis (Spruce) Hässel, Veröf. Geobot. Inst. ETH Stiftung Rübel
Zürich 91: 303, 1986 (Hässel 1986b). Bas.: Anthoceros tenuis Spruce, Bull. Soc.
Bot. France (Congr. Bot.) 36: cxcvi, 1889 [1890] (Spruce 1889).
** Phaeoceros tigrinus (Gola) J.C.Villarreal, Phytotaxa 208 (1): 95, 2015 (Villarreal et al.
2015). Bas.: Anthoceros tigrinus Gola, Ann. Bot. (Rome) 13 (1): 72, 1914 (Gola 1914a).
*** Phaeoceros tuberosus (Taylor) Prosk., J. Indian Bot. Soc. 42A: 185, 1964 (Proskauer
1964). Bas.: Anthoceros tuberosus Taylor, London J. Bot. 5: 412, 1846 (Taylor 1846b).
*** Phaeoceros udarii A.K.Asthana et V.Nath, Proc. Natl. Acad. Sci. India, B 63 (4):
461, 1993 (Asthana and Nath 1993).
** Phaeoceros wrightii (Steph.) Hässel, Candollea 44 (2): 722, 1989 (Hässel 1989a). Bas.:
Anthoceros wrightii Steph., Sp. Hepat. (Stephani) 5: 999, 1916 (Stephani 1916b).
22 Phaeoceros microsporus may be conspeciic with Phaeoceros mohrii in Schuster (1992d), but he did not
study any material.
23 Phaeoceros striatisporus was nested within Phaeoceros in Li et al. (2011). Morphologically it was supposed
to difer by its unique spores which, however, turns out to be fungal infections (Villarreal & Renner 2012).
World checklist of hornworts and liverworts
41
LEIOSPOROCEROTOPSIDA Stotler et Crand.-Stotl.
Leiosporocerotales Hässel
***
Leiosporocerotaceae Hässel ex Ochyra
by J.C. Villarreal and D.C. Cargill
*** Leiosporoceros Hässel, J. Bryol. 14 (2): 255, 1986 (Hässel 1986a).
*** Leiosporoceros dussii (Steph.) Hässel, J. Bryol. 14 (2): 255, 1986 (Hässel 1986a).
Bas.: Anthoceros dussii Steph., Hedwigia 32 (3): 142, 1893 (Stephani 1893b).
MARCHANTIOPHYTA
HAPLOMITRIOPSIDA Stotler et Crand.-Stotl.
Haplomitriidae Stotler et Crand.-Stotl.
Calobryales Hamlin
***
Haplomitriaceae Dědeček
by S. Bartholomew-Began
he treatment of Haplomitriaceae follows Bartholomew-Began (1991). Morphogenetic and molecular studies support the infrageneric ranks of Haplomitrium (CrandallStotler et al. 2009, Forrest et al. 2006).
*** Haplomitrium Nees, Naturgesch. Eur. Leberm. 1: 109, 1833 (Nees 1833c) nom.
conserv.
*** subg. Calobryum (Nees) R.M.Schust., Nova Hedwigia 13 (1/2): 40, 1967
(Schuster 1967c). Bas.: Calobryum Nees, Syn. Hepat. 4: 507, 1846 (Gottsche et
al. 1846).
*** Haplomitrium blumei (Nees) R.M.Schust., J. Hattori Bot. Lab. 26: 225, 1963
(Schuster 1963b). Bas.: Monoclea blumei Nees, Enum. Pl. Crypt. Javae: 2, 1830
(Nees 1830).
*** Haplomitrium mnioides (Lindb.) R.M.Schust., J. Hattori Bot. Lab. 26: 225, 1963
(Schuster 1963b). Bas.: Rhopalanthus mnioides Lindb., Not. Sällsk. Fauna Fl.
Fenn. Förh. 13: 391, 1874 (Lindberg 1874a).
*** subg. Haplomitrium
** sect. Archibryum (R.M.Schust.) J.J.Engel, Ann. Missouri Bot. Gard. 68 (4):
675, 1981 (Engel 1981). Bas.: Haplomitrium subg. Archibryum R.M.Schust.,
Nova Hedwigia 13 (1/2): 28, 1967 (Schuster 1967c).
42
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Haplomitrium gibbsiae (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 225, 1963
(Schuster 1963b). Bas.: Calobryum gibbsiae Steph., Sp. Hepat. (Stephani) 6: 76,
1917 (Stephani 1917a).
*** Haplomitrium intermedium Berrie, Proc. Linn. Soc. New South Wales (ser. 2) 87
(399): 191, 1963 (Berrie 1963).
** sect. Haplomitrium
*** Haplomitrium hookeri (Lyell ex Sm.) Nees, Naturgesch. Eur. Leberm. 1: 111, 1833
(Nees 1833c). Bas.: Jungermannia hookeri Lyell ex Sm., Engl. Bot. 35: tab. 2555,
1814 (Smith and Sowerby 1814).
*** Haplomitrium hookeri var. minutum (E.O.Campb.) Barthol.-Began, Bryophyt.
Biblioth. 41: 230, 1991 (Bartholomew-Began 1991). Bas.: Steereomitrium
minutum E.O.Campb., Mem. New York Bot. Gard. 45: 569, 1987 (Campbell
1987).
*** Haplomitrium monoicum J.J.Engel, Ann. Missouri Bot. Gard. 68 (4): 668, 1981
[1982] (Engel 1981).
*** Haplomitrium ovalifolium R.M.Schust., Bryologist 74 (2): 136, 1971 (Schuster 1971d).
Treubiidae Stotler et Crand.-Stotl.
Treubiales Schljakov
***
Treubiaceae Verd.
by R. Stotler and B.J. Crandall-Stotler
Stech et al. (2002) showed a clear molecular distinction between Treubia and Apotreubia, which is in accordance with their morphological diferences and supports their
recognition as separate genera.
*** Apotreubia S.Hatt. et Mizut., Bryologist 69 (4): 491, 1966 [1967] (Hattori et al.
1966).
*** Apotreubia hortoniae Konstant., Phytotaxa 76 (3): 33, 2013 (Konstantinova et al.
2013b). Based on: Apotreubia hortoniae R.M.Schust. et Konstant., J. Hattori Bot.
Lab. 78: 55, 1995 (Schuster and Konstantinova 1995), nom. inval.
*** Apotreubia nana (S.Hatt. et Inoue) S.Hatt. et Mizut., Bryologist 69 (4): 492, 1966
[1967] (Hattori et al. 1966). Bas.: Treubia nana S.Hatt. et Inoue, J. Hattori Bot.
Lab. 11: 99, 1954 (Hattori and Inoue 1954).
** Apotreubia pusilla (R.M.Schust.) Grolle, Acta Bot. Fenn. 125: 63, 1984 (Grolle
and Piippo 1984). Bas.: Treubia pusilla R.M.Schust., Nova Hedwigia 15: 515,
1968 (Schuster 1968b).
World checklist of hornworts and liverworts
43
*** Apotreubia yunnanensis Higuchi, Cryptog. Bryol. Lichénol. 19 (4): 321, 1998 (Higuchi 1998).
*** Treubia K.I.Goebel, Ann. Jard. Bot. Buitenzorg 9 (1): 1, 1890 [1891] (Goebel
1890) nom. conserv.
*** Treubia insignis K.I.Goebel, Ann. Jard. Bot. Buitenzorg 9 (1): 1, 1890 [1891]
(Goebel 1890).
** Treubia insignis subsp. bracteata (Steph.) R.M.Schust. et G.A.M.Scott, J. Hattori
Bot. Lab. 32: 241, 1969 (Schuster and Scott 1969). Bas.: Treubia bracteata Steph.,
Bot. Jahrb. Syst. 23 (1/2, 3): 302, 1896 (Stephani 1896a).
** Treubia insignis subsp. caledonica R.M.Schust. et G.A.M.Scott, J. Hattori Bot.
Lab. 32: 243, 1969 (Schuster and Scott 1969).
** Treubia insignis subsp. vitiensis R.M.Schust. et G.A.M.Scott, J. Hattori Bot. Lab.
32: 242, 1969 (Schuster and Scott 1969).
*** Treubia lacunosa (Colenso) Prosk., Bryologist 58 (3): 199, 1955 (Proskauer 1955).
Bas.: Noteroclada lacunosa Colenso, Trans. & Proc. New Zealand Inst. 18: 248,
1886 (Colenso 1886b).
** Treubia lacunosoides T.Pfeif., W.Frey et M.Stech, Nova Hedwigia 75 (1/2): 249,
2002 (Pfeifer et al. 2002).
*** Treubia pygmaea R.M.Schust., Phytologia 56 (7): 460, 1985 (Schuster 1985c).
*** Treubia scapanioides R.M.Schust., J. Hattori Bot. Lab. 32: 246, 1969 (Schuster
and Scott 1969).
*** Treubia tahitensis (Nadeaud) Besch., J. Bot. (Morot) 12: 147, 1898 (Bescherelle
1898). Bas.: Gottschea tahitensis Nadeaud, Énum. Pl. Tahiti: 7, 1873 (Nadeaud
1873).
*** Treubia tasmanica R.M.Schust. et G.A.M.Scott, J. Hattori Bot. Lab. 32: 248,
1969 (Schuster and Scott 1969).
JUNGERMANNIOPSIDA Stotler et Crand.-Stotl.
Jungermanniidae Engl.
Jungermanniales H.Klinggr.
Cephaloziineae Schljakov
***
Adelanthaceae Grolle
by K. Feldberg, J. Váňa and J. Heinrichs
he subfamily Jamesonielloideae was excluded from Lophoziaceae/Scapaniaceae by De
Roo et al. (2007) and this status was conirmed by e.g. Vilnet et al. (2010). he family
was described and deined by Feldberg et al. (2010a). Some taxonomic and nomenclatural notes can also be found in Feldberg et al. (2010b, 2011), Váňa et al. (2014d).
44
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Adelanthoideae K.Feldberg, Heinrichs et Váňa
*** Adelanthus Mitt., J. Proc. Linn. Soc., Bot. 7 (28): 243, 1864 (Mitten 1864b)
nom. conserv.
** sect. Adelanthus
*** Adelanthus falcatus (Hook.) Mitt., J. Proc. Linn. Soc., Bot. 7 (28): 243, 1864
(Mitten 1864b). Bas.: Jungermannia falcata Hook., Musci Exot. 1: tab. 89, 1818
(Hooker 1818).
*** Adelanthus occlusus (Hook.f. et Taylor) Carrington, Trans. Bot. Soc. Edinburgh
10: 381, 1870 (Carrington 1870). Bas.: Jungermannia occlusa Hook.f. et Taylor,
London J. Bot. 3: 369, 1844 (Hooker and Taylor 1844a).
** sect. Calyptrocolea (R.M.Schust.) Grolle, J. Hattori Bot. Lab. 35: 331, 1972
(Grolle 1972c). Bas.: Calyptrocolea R.M.Schust., Rev. Bryol. Lichénol. 34 (3/4):
685, 1966 (Schuster 1966a).
*** Adelanthus aureomarginatus R.M.Schust., Phytologia 39 (4): 250, 1978 (Schuster
1978a).
*** Adelanthus gemmiparus (R.M.Schust.) E.A.Hodgs., Trans. Roy. Soc. New Zealand, Biol. Sci. 11 (18): 241, 1970 (Hodgson 1970). Bas.: Calyptrocolea gemmipara R.M.Schust., Rev. Bryol. Lichénol. 34 (3/4): 695, 1966 [1967] (Schuster
1966a).
** Adelanthus lingulatus J.J.Engel et Váňa, Mem. New York Bot. Gard. 105: 26,
2013 (Váňa and Engel 2013).
*** Adelanthus tenuis J.J.Engel et Grolle, J. Hattori Bot. Lab. 35: 333, 1972 (Grolle
1972c).
** sect. Lindenbergiani Grolle, J. Hattori Bot. Lab. 35: 331, 1972 (Grolle
1972c).
*** Adelanthus carabayensis (Mont.) Grolle, J. Hattori Bot. Lab. 35: 348, 1972 (Grolle
1972c). Bas.: Plagiochila carabayensis Mont., Ann. Sci. Nat. Bot. (sér. 4) 5: 348,
1856 (Montagne 1856c).
*** Adelanthus integerrimus Grolle, J. Hattori Bot. Lab. 35: 340, 1972 (Grolle
1972c).
*** Adelanthus lindenbergianus (Lehm.) Mitt., J. Proc. Linn. Soc., Bot. 7 (28): 244,
1864 (Mitten 1864b). Bas.: Jungermannia lindenbergiana Lehm., Linnaea 4: 367,
1829 (Lehmann 1829).
** sect. Pittieri Grolle, J. Hattori Bot. Lab. 35: 331, 1972 (Grolle 1972c).
*** Adelanthus pittieri (Steph.) Grolle, J. Hattori Bot. Lab. 35: 337, 1972 (Grolle
1972c). Bas.: Tylimanthus pittieri Steph., Sp. Hepat. (Stephani) 6: 250, 1922
(Stephani 1922).
*** Adelanthus squarrosus Grolle, J. Hattori Bot. Lab. 67: 243, 1989 (Grolle 1989d).
World checklist of hornworts and liverworts
45
*** Pseudomarsupidium Herzog, Svensk Bot. Tidskr. 47 (1): 42, 1953 (Herzog
1953b).
*** Pseudomarsupidium aureocinctum (R.M.Schust.) J.J.Engel, Novon 17 (3): 312,
2007 (Engel 2007). Bas.: Adelanthus decipiens subsp. aureocinctus R.M.Schust.,
Phytologia 39 (4): 250, 1978 (Schuster 1978a).
*** Pseudomarsupidium borneensis (Grolle) Váňa, L.Söderstr., A.Hagborg et von Konrat, Phytotaxa 65: 60, 2012 (Váňa et al. 2012f). Bas.: Adelanthus borneensis Grolle,
J. Hattori Bot. Lab. 35: 362, 1972 (Grolle 1972c).
*** Pseudomarsupidium decipiens (Hook.) Grolle, Trans. Brit. Bryol. Soc. 4 (3): 443,
1963 (Grolle 1963a). Bas.: Jungermannia decipiens Hook., Brit. Jungermann.: tab.
50, 1813 (Hooker 1813).
*** Pseudomarsupidium piliferum (Steph.) Herzog ex Grolle, Trans. Brit. Bryol. Soc.
4 (3): 443, 1963 (Grolle 1963a). Bas.: Marsupidium piliferum Steph., Bull. Herb.
Boissier (sér. 2) 8 (8): 602 (386), 1908 (Stephani 1908e).
*** Wettsteinia Schifn., Ann. Jard. Bot. Buitenzorg, suppl. 2: 44, 1898 (Schifner
1898c).
*** Wettsteinia densiretis (Herzog) Grolle, J. Hattori Bot. Lab. 28: 99, 1965 (Grolle
1965f). Bas.: Tylimanthus densiretis Herzog, Nat. Hist. Juan Fernandez (Botany) 2
(5): 712, 1942 (Herzog 1942a).
*** Wettsteinia inversa (Sande Lac.) Schifn., Ann. Jard. Bot. Buitenzorg, suppl. 2:
45, 1898 (Schifner 1898c). Bas.: Plagiochila inversa Sande Lac., Ann. Mus. Bot.
Lugduno-Batavi 1: 289, 1864 (Sande Lacoste 1864).
*** Wettsteinia rotundifolia (Horik.) Grolle, J. Hattori Bot. Lab. 28: 100, 1965 (Grolle
1965f). Bas.: Adelanthus rotundifolius Horik., J. Sci. Hiroshima Univ., Ser. B, Div.
2, Bot. 2: 181, 1934 (Horikawa 1934).
*** Wettsteinia schusteriana Grolle, J. Hattori Bot. Lab. 28: 99, 1965 (Grolle 1965f).
**
Jamesonielloideae Inoue
*** Cuspidatula Steph., Bull. Herb. Boissier (sér. 2) 1 (10): 1141 (124), 1901
(Stephani 1901c).
*** Cuspidatula contracta (Reinw., Blume et Nees) Steph., Bull. Herb. Boissier (sér. 2)
1 (11): 1141 (124), 1901 (Stephani 1901c). Bas.: Jungermannia contracta Reinw.,
Blume et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1):
233, 1824 [1825] (Reinwardt et al. 1824a).
*** Cuspidatula laccida (Steph.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 142, 2010 (Feldberg et al. 2010b). Bas.: Anastrophyllum laccidum
Steph., Sp. Hepat. (Stephani) 6: 105, 1917 (Stephani 1917a).
*** Cuspidatula lexicaulis (Nees) Váňa et L.Söderstr., Phytotaxa 76 (3): 35, 2013
(Váňa et al. 2013h). Bas.: Jungermannia lexicaulis Nees, Linnaea 6 (4): 604, 1831
(Nees 1831).
46
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Cuspidatula kirkii (Steph.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 142, 2010 (Feldberg et al. 2010b). Bas.: Jamesoniella kirkii Steph.,
Hedwigia 34 (2): 47, 1895 (Stephani 1895c).
*** Cuspidatula monodon (Taylor) Steph., Bull. Herb. Boissier (sér. 2) 1 (11): 1143
(126), 1901 (Stephani 1901c). Bas.: Jungermannia monodon Taylor, Nov. Stirp.
Pug. 8: 7, 1844 (Lehmann 1844).
*** Cuspidatula orbicularis (Grolle) Váňa et L.Söderstr., Phytotaxa 76 (3): 36, 2013
(Váňa et al. 2013h). Bas.: Jamesoniella orbicularis Grolle, Feddes Repert. 82 (1):
42, 1971 (Grolle 1971b).
*** Cuspidatula robusta (Austin) Váňa et L.Söderstr., Phytotaxa 76 (3): 36, 2013 (Váňa
et al. 2013h). Bas.: Jungermannia robusta Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 219, 1869 (Austin 1869).
** Denotarisia Grolle, Feddes Repert. 82 (1): 6, 1971 (Grolle 1971b).
*** Denotarisia linguifolia (De Not.) Grolle, Feddes Repert. 82 (1): 6, 1971 (Grolle 1971b).
Bas.: Plagiochila linguifolia De Not., Epat. Borneo: 13, 1874 (De Notaris 1874).
** Nothostrepta R.M.Schust., Phytologia 45 (5): 420, 1980 (Schuster 1980b).
*** Nothostrepta biida (Steph.) R.M.Schust., Phytologia 45 (5): 420, 1980 (Schuster
1980b). Bas.: Plagiochila biida Steph., Annuario Reale Ist. Bot. Roma 2: 86, 1885
[1886] (Stephani 1885d).
*** Nothostrepta longissima (Steph.) R.M.Schust., Phytologia 45 (5): 420, 1980
(Schuster 1980b). Bas.: Anastrophyllum longissimum Steph., Bih. Kongl. Svenska
Vetensk.-Akad. Handl. 26 (III, 17): 13, 1901 (Stephani 1901b).
** Pisanoa Hässel, Lindbergia 14 (3): 179, 1988 [1989] (Hässel 1988).
*** Pisanoa chilensis Hässel, Lindbergia 14 (3): 179, 1988 [1989] (Hässel 1988).
** Protosyzygiella (Inoue) R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 334,
1980 (Schuster 1980c). Bas.: Syzygiella subg. Protosyzygiella Inoue, J. Hattori Bot.
Lab. 29: 180, 1966 (Inoue 1966c).
*** Protosyzygiella pseudoconnexa (Herzog) R.M.Schust., Hepat. Anthocerotae N.
Amer. 4: 334, 1980 (Schuster 1980c). Bas.: Plagiochila pseudoconnexa Herzog,
Rev. Bryol. Lichénol. 21 (3/4): 259, 1952 [1953] (Herzog 1952d).
*** Syzygiella Spruce, J. Bot. 14: 234, 1876 (Spruce 1876a). 24
*** subg. Anomalae (Inoue) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 143, 2010 (Feldberg et al. 2010b). Bas.: Syzygiella sect. Anomalae
Inoue, J. Hattori Bot. Lab. 29: 183, 1966 (Inoue 1966c).
24 Syzygiella includes Jamesoniella (cf. Feldberg et al. 2010a), but one taxon has neither been transferred
nor synonymized. It is listed in the “Names in genera not currently accepted” section below.
World checklist of hornworts and liverworts
47
*** Syzygiella anomala (Lindenb. et Gottsche) Steph., Bull. Herb. Boissier (sér. 2)
2 (5): 471 (190), 1902 (Stephani 1902f). Bas.: Plagiochila anomala Lindenb. et
Gottsche, Syn. Hepat. 5: 646, 1847 (Gottsche et al. 1847).
*** Syzygiella bilobata Inoue, J. Hattori Bot. Lab. 29: 186, 1966 (Inoue 1966c).
** Syzygiella ciliata Gradst. et A.R.Benitez, Nova Hedwigia 99 (1/2): 115, 2014
(Gradstein and Benitez 2014).
*** Syzygiella concreta (Gottsche) Spruce, J. Bot. 14: 234, 1876 (Spruce 1876a). Bas.:
Jungermannia concreta Gottsche, Mexik. Leverm.: 82, 1863 (Gottsche 1863).
*** Syzygiella manca (Mont.) Steph., Hedwigia 31 (1): 14, 1892 (Jack and Stephani
1892). Bas.: Chiloscyphus mancus Mont., Syll. Gen. Sp. Crypt.: 63, 1856 (Montagne 1856b).
*** Syzygiella pectiniformis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 501, 1885
(Spruce 1885).
*** Syzygiella tonduzana Steph., Sp. Hepat. (Stephani) 6: 118, 1917 (Stephani 1917a).
*** Syzygiella trigonifolia (Steph.) Herzog, Hedwigia 74 (2): 87, 1934 (Herzog 1934a).
Bas.: Jamesoniella trigonifolia Steph., Biblioth. Bot. 87 (2): 185, 1916 (Stephani 1916a).
*** subg. Cryptochila (R.M.Schust.) K.Feldberg, Váňa, Hentschel et Heinrichs,
Cryptog. Bryol. 31 (2): 143, 2010 (Feldberg et al. 2010b). Bas.: Cryptochila
R.M.Schust., J. Hattori Bot. Lab. 26: 284, 1963 (Schuster 1963b).
*** Syzygiella acinacifolia (Hook.f. et Taylor) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31 (2): 143, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia acinacifolia Hook.f. et Taylor, London J. Bot. 3: 367, 1844 (Hooker and
Taylor 1844a).
*** Syzygiella nigrescens (Steph.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31 (2): 143, 2010 (Feldberg et al. 2010b). Bas.: Jamesoniella nigrescens
Steph., Hedwigia 34 (2): 48, 1895 (Stephani 1895c).
*** Syzygiella paludosa (Steph.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 143, 2010 (Feldberg et al. 2010b). Bas.: Jamesoniella paludosa Steph.,
Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26 (III, 17): 11, 1901 (Stephani 1901b).
*** Syzygiella pseudocclusa (E.A.Hodgs.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31 (2): 143, 2010 (Feldberg et al. 2010b). Bas.: Jamesoniella pseudocclusa
E.A.Hodgs., Trans. Roy. Soc. New Zealand 85 (4): 583, 1958 (Hodgson 1958).
*** Syzygiella sonderi (Gottsche) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31 (2): 143, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia sonderi
Gottsche, Linnaea 28 (5): 550, 1856 [1857] (Gottsche 1856).
*** Syzygiella spegazziniana (Spruce ex C.Massal.) K.Feldberg, Váňa, Hentschel et
Heinrichs, Cryptog. Bryol. 31 (2): 144, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia spegazziniana Spruce ex C.Massal., Nuovo Giorn. Bot. Ital. 17 (3): 216,
1885 (Massalongo 1885).
*** subg. Pseudoplagiochila Inoue, J. Hattori Bot. Lab. 29: 182, 1966 (Inoue
1966c).
*** Syzygiella ovalifolia Inoue, J. Hattori Bot. Lab. 29: 191, 1966 (Inoue 1966c).
48
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*
Syzygiella securifolia (Nees) Inoue, J. Hattori Bot. Lab. 46: 232, 1979 (Inoue
1979a). Bas.: Plagiochila securifolia Nees, Sp. Hepat. (Lindenberg) 2-4: 58, 1840
(Lindenberg 1840). 25
*** Syzygiella subintegerrima (Reinw., Blume et Nees) Spruce, J. Linn. Soc., Bot.
30 (210): 362, 1895 (Gepp 1895b). Bas.: Jungermannia subintegerrima Reinw.,
Blume et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1):
238, 1824 [1825] (Reinwardt et al. 1824a).
*** Syzygiella tasmanica (Hook.f. et Taylor) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31 (2): 144, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia tasmanica Hook.f. et Taylor, London J. Bot. 5: 274, 1846 (Taylor 1846a).
*** subg. Roivainenia (Perss.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31 (2): 144, 2010 (Feldberg et al. 2010b). Bas.: Roivainenia Perss.,
Nova Hedwigia 3 (1): 43, 1961 (Persson and Grolle 1961).
*** Syzygiella jacquinotii (Mont.) Hentschel, K.Feldberg, Váňa et Heinrichs, Cryptog.
Bryol. 31 (2): 144, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia jacquinotii
Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 250, 1843 (Montagne 1843).
*** subg. Syzygiella
*** Syzygiella autumnalis (DC.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31 (2): 144, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia autumnalis DC., Fl.
Franç. (DC. & Lamarck), 5 (6): 202, 1815 (De Candolle and Lamarck 1815).
*** Syzygiella campanulata Herzog, Rev. Bryol. Lichénol. 11 (1): 9, 1938 [1939] (Herzog
1938a).
*** Syzygiella colorata (Lehm.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 144, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia colorata
Lehm., Linnaea 4: 366, 1829 (Lehmann 1829).
** Syzygiella colorata var. collenchymata J.J.Engel et Váňa, Mem. New York Bot.
Gard. 105: 102, 2013 (Váňa and Engel 2013).
*** Syzygiella contigua Steph., Bull. Herb. Boissier (sér. 2) 2 (5): 470 (189), 1902
(Stephani 1902f). Nom. nov. pro Jungermannia contigua Gottsche, Ann. Sci. Nat.
Bot. (sér. 5) 1: 118, 1864 (Gottsche 1864), nom. illeg.
*** Syzygiella elongella (Taylor) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 144, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia elongella
Taylor, London J. Bot. 5: 274, 1846 (Taylor 1846a).
*** Syzygiella macrocalyx (Mont.) Spruce, J. Bot. 14: 234, 1876 (Spruce 1876a). Bas.:
Jungermannia macrocalyx Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 248, 1843 (Montagne 1843).
*** Syzygiella nipponica (S.Hatt.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31 (2): 145, 2010 (Feldberg et al. 2010b). Bas.: Jamesoniella nipponica
S.Hatt., J. Jap. Bot. 19 (11): 350, 1943 (Hattori 1943b).
25 Syzygiella securifolia is a sister taxon to Syzygiella subintegerrima (Feldberg et al. 2010b) and it is debatable if they are diferent enough to warrant separation on the species level. Morphologically they are
very similar (Söderström et al. 2010a).
World checklist of hornworts and liverworts
49
** Syzygiella oenops (Lindenb. et Gottsche) K.Feldberg, Váňa, Hentschel et Heinrichs,
Cryptog. Bryol. 31 (2): 145, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia
oenops Lindenb. et Gottsche, Syn. Hepat. 5: 673, 1847 (Gottsche et al. 1847).
*** Syzygiella perfoliata (Sw.) Spruce, J. Bot. 14: 234, 1876 (Spruce 1876a). Bas.: Jungermannia perfoliata Sw., Prodr. (Swartz): 143, 1788 (Swartz 1788).
*** Syzygiella purpurascens (Steph.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31 (2): 145, 2010 (Feldberg et al. 2010b). Bas.: Jamesoniella purpurascens Steph., Bull. Soc. Roy. Bot. Belgique, Compt. Rend. 30 (2): 200, 1891
[1892] (Stephani 1891b).
*** Syzygiella rubricaulis (Nees) Steph., Bull. Herb. Boissier (sér. 2) 2 (5): 468 (187),
1902 (Stephani 1902f). Bas.: Jungermannia rubricaulis Nees, Fl. Bras. (Martius) 1
(1): 344, 1833 (Nees 1833a).
*** Syzygiella setulosa Steph., Bull. Herb. Boissier (sér. 2) 2 (5): 469 (188), 1902
(Stephani 1902f).
*** Syzygiella teres (Carrington et Pearson) Váňa, Phytotaxa 76 (3): 35, 2013 (Váňa et
al. 2013h). Bas.: Jungermannia teres Carrington et Pearson, Pap. & Proc. Roy. Soc.
Tasmania 1887: 9, 1888 (Carrington and Pearson 1888b).
*** Syzygiella undata (Mont.) K.Feldberg, Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 145, 2010 (Feldberg et al. 2010b). Bas.: Jungermannia undata
Mont., Ann. Sci. Nat. Bot. (sér. 4) 14: 183, 1860 (Montagne 1860).
Incertae sedis
*** Syzygiella eatonii (Austin) Inoue, J. Jap. Bot. 37 (12): 359, 1962 (Inoue 1962a).
Bas.: Plagiochila eatonii Austin, Trans. Connecticut Acad. Arts 8 (15): 257, 1891
(Evans 1891).
*** Syzygiella uleana Steph., Hedwigia 44 (4): 224, 1905 (Stephani 1905a).
** Vanaea (Inoue et Gradst.) Inoue et Gradst., Trop. Bryol. 1: 33, 1989 (Gradstein
and Florschütz-de Waard 1989). Bas.: Anastrophyllum subg. Vanaea Inoue et Gradst.,
Bull. Natl. Sci. Mus. Tokyo, B 14 (3): 88, 1988 (Inoue and Gradstein 1988).
*** Vanaea plagiochiloides (Inoue et Gradst.) Inoue et Gradst., Trop. Bryol. 1: 33,
1989 (Gradstein and Florschütz-de Waard 1989). Bas.: Anastrophyllum plagiochiloides Inoue et Gradst., Bull. Natl. Sci. Mus. Tokyo, B 14 (3): 88, 1988 (Inoue
and Gradstein 1988).
***
Anastrophyllaceae L.Söderstr., De Roo et Hedd.
by J. Váňa and L. Söderström
Anastrophyllaceae was described by Söderström et. al (2010b) from elements usually
included in Lophoziaceae. Further taxonomic and nomenclatural notes can be found
in Váňa et al. (2013k, 2013a). he complex of Chandonanthus/Plicanthus/Tetralophozia
should be checked using molecular methods before generic and speciic status of most
50
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
of the species can be conirmed. he placement of Hattoria and Zantenia in the family is
still provisional. he placement of Isopaches is also unclear, de Roo et al. (2007) placed it
in the family, but the study of Vilnet et al. (2010) showed that it can not be placed there.
*** Anastrepta (Lindb.) Schifn., Hepat. (Engl.-Prantl): 85, 1893 (Schifner 1893b).
Bas.: Jungermannia sect. Anastrepta Lindb., Kongl. Svenska Vetensk.-Akad. Handl.
(n.ser.) 23 (5): 40, 1889 (Lindberg and Arnell 1889).
*** Anastrepta orcadensis (Hook.) Schifn., Hepat. (Engl.-Prantl): 85, 1893 (Schifner
1893b). Bas.: Jungermannia orcadensis Hook., Brit. Jungermann.: tab. 71, 1815
(Hooker 1815).
*** Anastrophyllum (Spruce) Steph., Hedwigia 32 (3): 139, 1893 (Stephani 1893b).
Bas.: Jungermannia subg. Anastrophyllum Spruce, J. Bot. 14: 235, 1876 (Spruce
1876a).
*** Anastrophyllum alpinum Steph., Sp. Hepat. (Stephani) 6: 103, 1917 (Stephani 1917a).
*** Anastrophyllum assimile (Mitt.) Steph., Hedwigia 32 (3): 140, 1893 (Stephani
1893b). Bas.: Jungermannia assimilis Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 93,
1860 [1861] (Mitten 1860c).
*** Anastrophyllum auritum (Lehm.) Steph., Bull. Herb. Boissier (sér. 2) 1 (11): 1137
(120), 1901 (Stephani 1901c). Bas.: Jungermannia aurita Lehm., Linnaea 4: 368,
1829 (Lehmann 1829).
*** Anastrophyllum ciliatum Steph., Hedwigia 32 (3): 139, 1893 (Stephani 1893b).
*** Anastrophyllum donnianum (Hook.) Steph., Hedwigia 32 (3): 140, 1893 (Stephani
1893b). Bas.: Jungermannia donniana Hook., Brit. Jungermann.: tab. 39, 1813
(Hooker 1813).
*** Anastrophyllum ellipticum Inoue, Bull. Natl. Sci. Mus. Tokyo, B 4 (1): 13, 1978
(Inoue 1978b). 26
** Anastrophyllum esenbeckii (Mont.) Steph., Hedwigia 32 (3): 140, 1893 (Stephani
1893b). Bas.: Jungermannia esenbeckii Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 247,
1843 (Montagne 1843).
*** Anastrophyllum issum Steph., Bull. Herb. Boissier 5 (10): 845, 1897 (Stephani 1897c).
*** Anastrophyllum joergensenii Schifn., Hedwigia 49 (4): 396, 1910 (Schifner 1910b).
* Anastrophyllum lignicola D.B.Schill et D.G.Long, Ann. Bot. Fenn. 39 (2): 130,
2002 (Schill and Long 2002). 27
*** Anastrophyllum michauxii (F.Weber) H.Buch, Memoranda Soc. Fauna Fl. Fennica
8: 289, 1932 [1933] (Buch 1932). Bas.: Jungermannia michauxii F.Weber, Hist.
Musc. Hepat. Prodr.: 76, 1815 (Weber 1815).
26 Anastrophyllum ellipticum is only tentatively placed in the genus following Mamontov & Vilnet
(2013).
27 Anastrophyllum lignicola is possibly conspeciic with Anastrophyllum ellipticum and its generic placement is uncertain.
World checklist of hornworts and liverworts
51
*** Anastrophyllum nigrescens (Mitt.) Steph., Hedwigia 32 (3): 140, 1893 (Stephani
1893b). Bas.: Jungermannia nigrescens Mitt., Hooker’s J. Bot. Kew Gard. Misc. 3:
358, 1851 (Mitten 1851).
*** Anastrophyllum obtusum Herzog, Trans. Brit. Bryol. Soc. 1 (4): 285, 1950 (Herzog
1950a).
*** Anastrophyllum piligerum (Nees) Steph., Hedwigia 32 (3): 140, 1893 (Stephani
1893b). Bas.: Jungermannia piligera Nees, Nova Acta Phys.-Med. Acad. Caes.
Leop.-Carol. Nat. Cur. 12 (1): 414, 1824 [1825] (Reinwardt et al. 1824b).
*** Anastrophyllum squarrosum Herzog, Ann. Bryol. 5: 72, 1932 (Herzog 1932b).
*** Anastrophyllum stellatum R.M.Schust., Phytologia 39 (4): 243, 1978 (Schuster
1978a).
*** Anastrophyllum tubulosum (Nees) Grolle, J. Hattori Bot. Lab. 28: 101, 1965
(Grolle 1965e). Bas.: Jungermannia tubulosa Nees, Enum. Pl. Crypt. Javae: 32,
1830 (Nees 1830).
*** Barbilophozia Loeske, Verh. Bot. Vereins Prov. Brandenburg 49 (1): 37, 1908
(Loeske 1908).
** subg. Barbilophozia
*** Barbilophozia barbata (Schmidel ex Schreb.) Loeske, Verh. Bot. Vereins Prov.
Brandenburg 49 (1): 37, 1908 (Loeske 1908). Bas.: Jungermannia barbata Schmidel ex Schreb., Spic. Fl. Lips.: 107, 1771 (Schreber 1771).
*** Barbilophozia hatcheri (A.Evans) Loeske, Verh. Bot. Vereins Prov. Brandenburg
49 (1): 37, 1908 (Loeske 1908). Bas.: Jungermannia hatcheri A.Evans, Bull. Torrey
Bot. Club 25 (8): 417, 1898 (Evans 1898).
*** Barbilophozia lycopodioides (Wallr.) Loeske, Verh. Bot. Vereins Prov. Brandenburg
49 (1): 37, 1908 (Loeske 1908). Bas.: Jungermannia lycopodioides Wallr., Comp. l.
Germ. 2 (III): 76, 1831 (Bluf and Fingerhuth 1831).
*** Barbilophozia rubescens (R.M.Schust. et Damsh.) Kartt. et L.Söderstr., Ann. Bot. Fenn.
29 (2): 120, 1992 (Söderström et al. 1992). Bas.: Lophozia rubescens R.M.Schust. et
Damsh., Phytologia 63 (5): 325, 1987 (Schuster and Damsholt 1987).
** subg. Sudeticae (Schljakov) L.Söderstr., De Roo et Hedd., Phytotaxa 3: 50,
2010 (Söderström et al. 2010b). Bas.: Lophozia sect. Sudeticae Schljakov, Pečen.
Mchi Sev. SSSR 3: 113, 1980 (Shliakov 1980a).
*** Barbilophozia sudetica (Nees ex Huebener) L.Söderstr., De Roo et Hedd., Phytotaxa 3: 50, 2010 (Söderström et al. 2010b). Bas.: Jungermannia sudetica Nees ex
Huebener, Hepaticol. germ.: 142, 1834 (Hübener 1834). 28
28 Barbilophozia sudetica was transferred from Lophozia by Söderström et al. (2010a), but transferred to
the new genus Pseudolophozia Konstant. and Vilnet (sister to Barbilophozia) by Konstantinova and
Vilnet (2009).
52
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Biantheridion (Grolle) Konstant. et Vilnet, Arctoa 18: 67, 2009 [2010] (Konstantinova and Vilnet 2009). Bas.: Jamesoniella sect. Biantheridion Grolle, Trans.
Brit. Bryol. Soc. 4 (4): 662, 1964 (Grolle 1964i).
*** Biantheridion undulifolium (Nees) Konstant. et Vilnet, Arctoa 18: 67, 2009 [2010]
(Konstantinova and Vilnet 2009). Bas.: Jungermannia schraderi β undulifolia Nees,
Naturgesch. Eur. Leberm. 1: 306, 1833 (Nees 1833c).
*** Chandonanthus Mitt., Handb. N. Zeal. l. 2: 753, 1867 (Hooker 1867).
*** Chandonanthus squarrosus (Menzies) Mitt., Handb. N. Zeal. l. 2: 753, 1867
(Hooker 1867). Bas.: Jungermannia squarrosa Menzies, Musci Exot. 1: tab. 78,
1818 (Hooker 1818).
*** Crossocalyx Meyl., Bull. Soc. Vaud. Sci. Nat. 60 (249): 266, 1939 (Meylan 1939).
*** Crossocalyx hellerianus (Nees ex Lindenb.) Meyl., Bull. Soc. Vaud. Sci. Nat. 60
(249): 266, 1939 (Meylan 1939). Bas.: Jungermannia helleriana Nees ex Lindenb.,
Syn. hepat. eur: 64, 1829 (Lindenberg 1829).
*** Crossocalyx tenuis (Harry Williams) Schljakov, Novosti Sist. Nizš. Rast. 15: 246,
1978 (Shliakov 1978). Bas.: Anastrophyllum tenue Harry Williams, Bryologist 71
(1): 34, 1968 (Williams 1968).
*** Gymnocolea (Dumort.) Dumort., Recueil Observ. Jungerm.: 17, 1835 (Dumortier 1835). Bas.: Jungermannia sect. Gymnocolea Dumort., Syll. Jungerm. Europ.:
52, 1831 (Dumortier 1831).
*** Gymnocolea borealis (Frisvoll et Moen) R.M.Schust., Lindbergia 12 (1): 7, 1986
(Schuster 1986). Bas.: Lophozia borealis Frisvoll et Moen, Lindbergia 6 (2): 138,
1980 [1981] (Frisvoll and Moen 1980).
** Gymnocolea fascinifera Potemkin, Arctoa 2: 76, 1993 (Potemkin 1993).
*** Gymnocolea inlata (Huds.) Dumort., Recueil Observ. Jungerm.: 17, 1835 (Dumortier 1835). Bas.: Jungermannia inlata Huds., Fl. Angl. (Hudson), ed. 2: 511,
1778 (Hudson 1778).
* Gymnocolea inlata subsp. acutiloba (Schifn.) R.M.Schust. et Damsh. ex L.Söderstr. et Váňa, Lindbergia 27 (1): 43, 2002 (Söderström et al. 2002). Bas.: Lophozia
acutiloba Schifn., Hedwigia 48 (3): 187, 1909 (Schifner 1909a). 29
*** Hamatostrepta Váňa et D.G.Long, Fieldiana, Bot. (n.ser.) 47: 134, 2008 (Váňa
and Long 2008).
*** Hamatostrepta concinna Váňa et D.G.Long, Fieldiana, Bot. (n.ser.) 47: 134, 2008
(Váňa and Long 2008).
29 Gymnocolea inlata subsp. acutiloba is a problematic taxon that sometimes has been treated as conspeciic
with Gymnocolea inlata, sometimes as a separate species.
World checklist of hornworts and liverworts
53
** Hattoria R.M.Schust., Rev. Bryol. Lichénol. 30 (1/2): 69, 1961 (Schuster 1961a).
*** Hattoria yakushimensis (Horik.) R.M.Schust., Rev. Bryol. Lichénol. 30 (1/2): 70,
1961 (Schuster 1961a). Bas.: Anastrophyllum yakushimense Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 149, 1934 (Horikawa 1934).
*** Isopaches H.Buch, Memoranda Soc. Fauna Fl. Fennica 8: 287, 1932 [1933]
(Buch 1932).
*** Isopaches alboviridis (R.M.Schust.) Schljakov, Novosti Sist. Nizš. Rast. 16: 205,
1979 (Shliakov 1979). Bas.: Lophozia alboviridis R.M.Schust., Hepat. Anthocerotae N. Amer. 2: 487, 1969 (Schuster 1969b).
*** Isopaches bicrenatus (Schmidel ex Hofm.) H.Buch, Memoranda Soc. Fauna Fl.
Fennica 8: 288, 1932 [1933] (Buch 1932). Bas.: Jungermannia bicrenata Schmidel
ex Hofm., Deutschl. Fl., heil 2 (Hofm.): 11 (addenda), 1795 [1796] (Hofmann 1795).
*** Isopaches decolorans (Limpr.) H.Buch, Memoranda Soc. Fauna Fl. Fennica 8:
288, 1932 [1933] (Buch 1932). Bas.: Jungermannia decolorans Limpr., Jahresber.
Schles. Ges. Vaterl. Cult. 57: 316, 1879 [1880] (Limpricht 1879).
*** Isopaches pumicicola (Berggr.) Bakalin, Arctoa 17: 162, 2008 [2009] (Bakalin
2008b). Bas.: Lophozia pumicicola Berggr., New Zealand Hepat.: 21, 1898
(Berggren 1898).
*** Neoorthocaulis L.Söderstr., De Roo et Hedd., Phytotaxa 3: 49, 2010 (Söderström et al. 2010b).
*** Neoorthocaulis attenuatus (Mart.) L.Söderstr., De Roo et Hedd., Phytotaxa 3: 49,
2010 (Söderström et al. 2010b). Bas.: Jungermannia quinquedentata δ attenuata
Mart., Fl. crypt. erlang.: 177, 1817 (Martius 1817).
*** Neoorthocaulis binsteadii (Kaal.) L.Söderstr., De Roo et Hedd., Phytotaxa 3: 49,
2010 (Söderström et al. 2010b). Bas.: Jungermannia binsteadii Kaal., Skr. Vidensk.Selsk. Christiana, Math.-Naturvidensk. Kl. 1898 (9): 9, 1898 (Kaalaas 1898).
*** Neoorthocaulis loerkei (F.Weber et D.Mohr) L.Söderstr., De Roo et Hedd., Phytotaxa 3: 50, 2010 (Söderström et al. 2010b). Bas.: Jungermannia loerkei F.Weber
et D.Mohr, Bot. Taschenb. (Weber): 410, 1807 (Weber and Mohr 1807).
** Neoorthocaulis hyperboreus (R.M.Schust.) L.Söderstr., De Roo et Hedd., Phytotaxa 3: 50, 2010 (Söderström et al. 2010b). Bas.: Lophozia loerkei var. hyperborea
R.M.Schust., Bull. Natl. Mus. Canada 164: 21, 1959 (Schuster et al. 1959).
*** Orthocaulis H.Buch, Memoranda Soc. Fauna Fl. Fennica 8: 293, 1932 [1933]
(Buch 1932).
*** Orthocaulis atlanticus (Kaal.) H.Buch, Memoranda Soc. Fauna Fl. Fennica 8: 294,
1932 [1933] (Buch 1932). Bas.: Jungermannia atlantica Kaal., Skr. Vidensk.-Selsk.
Christiana, Math.-Naturvidensk. Kl. 1898 (9): 11, 1898 (Kaalaas 1898).
* Orthocaulis cavifolius H.Buch et S.W.Arnell, Memoranda Soc. Fauna Fl. Fennica
26: 71, 1951 (Buch 1951).
54
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Plicanthus R.M.Schust., Nova Hedwigia 74 (3/4): 484, 2002 (Schuster 2002a).
*** Plicanthus birmensis (Steph.) R.M.Schust., Beih. Nova Hedwigia 119: 223, 2002
(Schuster 2002b). Bas.: Chandonanthus birmensis Steph., Bull. Soc. Roy. Bot. Belgique 38 (1): 43, 1899 (Stephani 1899h). 30
** Plicanthus diicilis (Steph.) L.Söderstr. et Váňa, Phytotaxa 81 (1): 30, 2013 (Váňa
et al. 2013a). Bas.: Chandonanthus diicilis Steph., J. & Proc. Roy. Soc. New
South Wales 48 (1/2): 101, 1914 (Stephani and Watts 1914).
** Plicanthus giganteus (Steph.) R.M.Schust., Nova Hedwigia 74 (3/4): 485, 2002
(Schuster 2002a). Bas.: Chandonanthus giganteus Steph., Wiss. Ergebn. Deut.
Zentr.-Afr. Exped. (1907-08), Bot. 2: 124, 1911 (Stephani 1911a). 31
*** Plicanthus hirtellus (F.Weber) R.M.Schust., Nova Hedwigia 74 (3/4): 492, 2002
(Schuster 2002a). Bas.: Jungermannia hirtella F.Weber, Hist. Musc. Hepat. Prodr.:
50, 1815 (Weber 1815).
*** Schizophyllopsis Váňa et L.Söderstr., Phytotaxa 152 (1): 48, 2013 (Váňa et al.
2013f). Nom. nov. pro Anastrophyllum subg. Schizophyllum R.M.Schust., Hepat.
Anthocerotae N. Amer. 2: 739, 1969 (Schuster 1969b). 32
** Schizophyllopsis aristata (Herzog ex N.Kitag.) Váňa et L.Söderstr., Phytotaxa 152
(1): 48, 2013 (Váňa et al. 2013f). Bas.: Anastrophyllum bidens var. aristatum Herzog ex N.Kitag., J. Hattori Bot. Lab. 33: 216, 1970 (Kitagawa 1970).
*** Schizophyllopsis bidens (Reinw., Blume et Nees) Váňa et L.Söderstr., Phytotaxa
152 (1): 48, 2013 (Váňa et al. 2013f). Bas.: Jungermannia bidens Reinw., Blume
et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 208,
1824 [1825] (Reinwardt et al. 1824a).
** Schizophyllopsis lanciloba (Steph.) Váňa et L.Söderstr., Phytotaxa 152 (1): 48, 2013
(Váňa et al. 2013f). Bas.: Anastrophyllum lancilobum Steph., Sp. Hepat. (Stephani)
6: 107, 1917 (Stephani 1917a).
*** Schizophyllopsis papillosa (J.J.Engel et Braggins) Váňa et L.Söderstr., Phytotaxa 152
(1): 48, 2013 (Váňa et al. 2013f). Bas.: Anastrophyllum papillosum J.J.Engel et
Braggins, J. Bryol. 20 (2): 381, 1998 (Engel and Braggins 1998).
*** Schizophyllopsis sphenoloboides (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 152
(1): 49, 2013 (Váňa et al. 2013f). Bas.: Anastrophyllum sphenoloboides R.M.Schust.,
Hepat. Anthocerotae N. Amer. 2: 741, 1969 (Schuster 1969b).
*** Schljakovia Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010] (Konstantinova and
Vilnet 2009).
30 Plicanthus birmensis seems to be closely related to Tetralophozia (Vilnet et al. 2010), but Grolle (1995)
considered it a possibly depauperate form of Plicanthus hirtellus. However, it may be a good species,
possibly of the genus Tetralophozia.
31 Plicanthus giganteus is often treated as a subspecies of Plicanthus hirtellus (e.g. Pócs and Lye 1999).
32 Schizophyllopsis was originally published as Schizophyllum (R.M.Schust.) Váňa et L.Söderstr. (Váňa et al.
2013k), but that is a later homonym of Schizophyllum Fr. (1815; Fungus) and Nutt. (1841; Asteraceae).
World checklist of hornworts and liverworts
55
*** Schljakovia kunzeana (Huebener) Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010]
(Konstantinova and Vilnet 2009). Bas.: Jungermannia kunzeana Huebener, Hepaticol. germ.: 115, 1834 (Hübener 1834).
*** Schljakovianthus Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010] (Konstantinova and Vilnet 2009).
*** Schljakovianthus quadrilobus (Lindb.) Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010]
(Konstantinova and Vilnet 2009). Bas.: Jungermannia quadriloba Lindb., Kongl.
Svenska Vetensk.-Akad. Handl. (n.ser.) 23 (5): 55, 1889 (Lindberg and Arnell 1889).
*** Sphenolobopsis R.M.Schust. et N.Kitag., Nova Hedwigia 22: 152, 1971 [1972]
(Schuster 1971b).
*** Sphenolobopsis pearsonii (Spruce) R.M.Schust., Nova Hedwigia 22: 153, 1971
[1972] (Schuster 1971b). Bas.: Jungermannia pearsonii Spruce, J. Bot. 19: 33,
1881 (Spruce 1881b).
*** Sphenolobus (Lindb.) Berggr., New Zealand Hepat.: 22, 1898 (Berggren 1898).
Bas.: Jungermannia sect. Sphenolobus Lindb., Not. Sällsk. Fauna Fl. Fenn. Förh.
13: 369, 1874 (Lindberg 1874a).
*** Sphenolobus austroamericanus (Váňa) Váňa, Phytotaxa 81 (1): 30, 2013 (Váňa et
al. 2013a). Bas.: Anastrophyllum austroamericanum Váňa, J. Hattori Bot. Lab. 48:
225, 1980 (Váňa 1980).
*** Sphenolobus minutus (Schreb. ex D.Crantz) Berggr., New Zealand Hepat.: 22,
1898 (Berggren 1898). Bas.: Jungermannia minuta Schreb. ex D.Crantz, Forts.
Hist. Grönland: 285, 1770 (Crantz 1770; non vidi).
*** Sphenolobus saxicola (Schrad.) Steph., Bull. Herb. Boissier (sér. 2) 2 (2): 168 (160),
1902 (Stephani 1902d). Bas.: Jungermannia saxicola Schrad., Syst. Samml. Crypt.
Gew. 2: 4, 1797 (Schrader 1797).
** Tetralophozia (R.M.Schust.) Schljakov, Novosti Sist. Nizš. Rast. 13: 227, 1976
(Shliakov 1976). Bas.: Chandonanthus subg. Tetralophozia R.M.Schust., J. Hattori
Bot. Lab. 23: 206, 1960 [1961] (Schuster 1960a).
*** Tetralophozia cavallii (Gola) Váňa, Trop. Bryol. 8: 102, 1993 (Váňa 1993). Bas.:
Blepharostomum cavallii Gola, Ann. Bot. (Rome) 6 (2): 274, 1907 (Gola 1907).
*** Tetralophozia iliformis (Steph.) Urmi, J. Bryol. 12 (3): 394, 1983 (Urmi 1983).
Bas.: Chandonanthus iliformis Steph., Sp. Hepat. (Stephani) 3: 644, 1909 (Stephani
1909a).
** Tetralophozia pilifera (Steph.) R.M.Schust., Nova Hedwigia 74 (3/4): 482, 2002
(Schuster 2002a). Bas.: Chandonanthus pilifer Steph., Sp. Hepat. (Stephani) 3:
644, 1909 (Stephani 1909a).
*** Tetralophozia setiformis (Ehrh.) Schljakov, Novosti Sist. Nizš. Rast. 13: 228, 1976
(Shliakov 1976). Bas.: Jungermannia setiformis Ehrh., Hannover. Mag. 22 (8):
142, 1784 (Ehrhart 1784).
56
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Zantenia (S.Hatt.) Váňa et J.J.Engel, Mem. New York Bot. Gard. 105: 29, 2013
(Váňa and Engel 2013). Bas.: Anastrophyllum subg. Zantenia S.Hatt., Bot. Mag.
(Tokyo) 79 (937): 342, 1966 (Hattori 1966a).
*** Zantenia borneensis (Herzog) Váňa et J.J.Engel, Mem. New York Bot. Gard. 105:
29, 2013 (Váňa and Engel 2013). Bas.: Anastrophyllum borneense Herzog, Trans.
Brit. Bryol. Soc. 1 (4): 282, 1950 (Herzog 1950a).
*** Zantenia denticulata (Grolle) Váňa et J.J.Engel, Mem. New York Bot. Gard. 105:
30, 2013 (Váňa and Engel 2013). Bas.: Anastrophyllum denticulatum Grolle, Nova
Hedwigia 16: 148, 1968 (Grolle 1968d).
*** Zantenia karstenii (Schifn.) Váňa et J.J.Engel, Mem. New York Bot. Gard. 105:
30, 2013 (Váňa and Engel 2013). Bas.: Anastrophyllum karstenii Schifn., Nova Acta
Acad. Caes. Leop.-Carol. German. Nat. Cur. 60 (2): 268, 1893 (Schifner 1893a).
*** Zantenia prionophylla (S.Hatt.) Váňa et J.J.Engel, Mem. New York Bot. Gard.
105: 30, 2013 (Váňa and Engel 2013). Bas.: Anastrophyllum prionophyllum S.Hatt.,
Bot. Mag. (Tokyo) 79 (937): 342, 1966 (Hattori 1966a).
***
Cephaloziaceae Mig.
by J. Váňa with contributions by S.R. Gradstein (Odontoschisma)
Cephaloziaceae was recently studied by Vilnet et al. (2012) and Feldberg et al. (2013).
Some taxonomic and nomenclatural notes were published by Váňa et al. (2013g,
2013i) and Gradstein et al. (2014b).
**
Alobielloideae R.M.Schust.
** Alobiella (Spruce) Schifn., Hepat. (Engl.-Prantl): 98, 1893 (Schifner 1893b).
Bas.: Cephalozia subg. Alobiella Spruce, Cephalozia: 28, 1882 (Spruce 1882).
*** Alobiella husnotii (Spruce) Schifn., Hepat. (Engl.-Prantl): 98, 1893 (Schifner
1893b). Bas.: Cephalozia husnotii Spruce, Cephalozia: 30, 1882 (Spruce 1882).
** Alobiellopsis R.M.Schust., Nova Hedwigia 10 (1/2): 25, 1965 (Schuster 1965b).
*** Alobiellopsis acroscypha (Spruce) R.M.Schust., Nova Hedwigia 10 (1/2): 25, 1965
(Schuster 1965b). Bas.: Cephalozia acroscypha Spruce, Cephalozia: 30, 1882
(Spruce 1882).
*** Alobiellopsis dominicensis (Spruce) Fulford, Mem. New York Bot. Gard. 11 (3):
350, 1968 (Fulford 1968). Bas.: Alobiella dominicensis Spruce, J. Linn. Soc., Bot.
30 (210): 355, 1895 (Gepp 1895b).
*** Alobiellopsis heteromorpha (Lehm.) R.M.Schust., Bull. Natl. Sci. Mus. Tokyo
(n.ser.) 12 (3): 682, 1969 (Schuster 1969a). Bas.: Jungermannia heteromorpha
Lehm., Linnaea 4: 362, 1829 (Lehmann 1829).
World checklist of hornworts and liverworts
57
*** Alobiellopsis parvifolia (Steph.) R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.)
12 (3): 679, 1969 (Schuster 1969a). Bas.: Alobiella parvifolia Steph., Bull. Herb.
Boissier (sér. 2) 8 (8): 568 (352), 1908 (Stephani 1908e).
*** Alobiellopsis pillansii (Sim) R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 12
(3): 683, 1969 (Schuster 1969a). Bas.: Cephalozia pillansii Sim, Trans. Roy. Soc.
South Africa 15 (1): 87, 1926 (Sim 1926).
**
Cephalozioideae Müll.Frib.
*** Cephalozia (Dumort.) Dumort., Recueil Observ. Jungerm.: 18, 1835 (Dumortier 1835). Bas.: Jungermannia sect. Cephalozia Dumort., Syll. Jungerm. Europ.:
60, 1831 (Dumortier 1831).
*** Cephalozia acuminata (Herzog) Váňa, Phytotaxa 112 (1): 11, 2013 (Váňa et al.
2013g). Bas.: Hygrobiella acuminata Herzog, Trans. Brit. Bryol. Soc. 1 (4): 292,
1950 (Herzog 1950a).
** Cephalozia acutiloba (Inoue) Váňa, Phytotaxa 112 (1): 11, 2013 (Váňa et al.
2013g). Bas.: Metahygrobiella acutiloba Inoue, Bull. Natl. Sci. Mus. Tokyo (n.ser.)
10 (2): 155, 1967 (Inoue 1967c).
*** Cephalozia albula Steph., Hedwigia 32 (5): 318, 1893 (Stephani 1893d). Nom.
nov. pro Jungermannia albula Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 93, 1860
[1861] (Mitten 1860c), nom. illeg.
*** Cephalozia ambigua C.Massal., Malpighia 21 (7/8): 310, 1907 (Massalongo 1907).
*** Cephalozia austrigena R.M.Schust. ex J.J.Engel, Novon 17 (3): 312, 2007 (Engel
2007). Based on: Cephalozia bicuspidata subsp. austrigena R.M.Schust., Hepat.
Anthocerotae N. Amer. 3: 712, 1974 (Schuster 1974), nom. inval.
*** Cephalozia badia (Gottsche) Steph., Bull. Herb. Boissier (sér. 2) 8 (7): 483 (313),
1908 (Stephani 1908f). Bas.: Jungermannia badia Gottsche, Int. Polarforsch.,
Deutsch. Exped. 2: 452, 1890 (Gottsche 1890).
*** Cephalozia bicuspidata (L.) Dumort., Recueil Observ. Jungerm.: 18, 1835 (Dumortier 1835). Bas.: Jungermannia bicuspidata L., Sp. Pl. 1: 1132, 1753 (Linnaeus 1753).
* Cephalozia bicuspidata subsp. lammersiana (Huebener) R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 730, 1974 (Schuster 1974). Bas.: Jungermannia lammersiana Huebener, Flora 15 (20): 306, 1832 (Hübener 1832).
*** Cephalozia chilensis (J.J.Engel et R.M.Schust.) R.M.Schust., Beih. Nova Hedwigia 119: 29, 2002 (Schuster 2002b). Bas.: Metahygrobiella chilensis J.J.Engel et
R.M.Schust., Brittonia 40 (2): 203, 1988 (Engel and Schuster 1988).
*** Cephalozia conchata (Grolle et Váňa) Váňa, Syst. Bot. 40 (1): 38, 2015 (Shaw et
al. 2015). Bas.: Jungermannia conchata Grolle et Váňa, Fragm. Florist. Geobot. 37
(1): 3, 1992 (Grolle and Váňa 1992).
*** Cephalozia crossii Spruce, Cephalozia: 46, 1882 (Spruce 1882).
*** Cephalozia darjeelingensis Udar et D.Kumar, Geophytology 6 (1): 36, 1976 (Udar
and Kumar 1976).
58
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Cephalozia drucei (R.M.Schust.) Váňa, Phytotaxa 112 (1): 11, 2013 (Váňa et al.
2013g). Bas.: Metahygrobiella drucei R.M.Schust., J. Hattori Bot. Lab. 26: 273,
1963 (Schuster 1963b).
*** Cephalozia fuegiensis Váňa, Phytotaxa 112 (1): 12, 2013 (Váňa et al. 2013g). Nom. nov.
pro Hygrobiella dusenii Steph., Sp. Hepat. (Stephani) 6: 444, 1924 (Stephani 1924).
*** Cephalozia hamatiloba Steph., Bull. Herb. Boissier (sér. 2) 8 (6): 427 (303), 1908
(Stephani 1908g).
*** Cephalozia hamatiloba subsp. siamensis (N.Kitag.) Váňa, Acta Bot. Fenn. 177: 16,
2004 (Koponen et al. 2004). Bas.: Cephalozia siamensis N.Kitag., J. Hattori Bot.
Lab. 32: 293, 1969 (Kitagawa 1969c).
*** Cephalozia lacinulata (J.B.Jack ex Gottsche et Rabenh.) Spruce, Cephalozia: 45,
1882 (Spruce 1882). Bas.: Jungermannia lacinulata J.B.Jack ex Gottsche et Rabenh., Hepat. Eur., Leberm. 62-64: no. 624, 1877 (Gottsche and Rabenhorst
1877).
*** Cephalozia lucens (A.Evans) Steph., Bull. Herb. Boissier (sér. 2) 8 (7): 496 (326),
1908 (Stephani 1908f). Bas.: Jungermannia lucens A.Evans, Trans. Connecticut
Acad. Arts 8 (15): 258, 1891 (Evans 1891).
*** Cephalozia macgregorii (Steph.) Váňa, Phytotaxa 112 (1): 12, 2013 (Váňa et al. 2013g).
Bas.: Hygrobiella macgregorii Steph., Hedwigia 34 (2): 45, 1895 (Stephani 1895c).
*** Cephalozia macounii (Austin) Austin, Hepat. bor.-amer.: 14, 1873 (Austin 1873).
Bas.: Jungermannia macounii Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 222,
1869 (Austin 1869).
*** Cephalozia maxima Steph., Sp. Hepat. (Stephani) 6: 441, 1924 (Stephani 1924).
*** Cephalozia mollusca (De Not.) Váňa, Phytotaxa 112 (1): 12, 2013 (Váňa et al. 2013g).
Bas.: Jungermannia mollusca De Not., Epat. Borneo: 16, 1874 (De Notaris 1874).
* Cephalozia neesiana Steph., Bull. Herb. Boissier (sér. 2) 8 (6): 429 (305), 1908
(Stephani 1908g). 33
*** Cephalozia nishimurae (N.Kitag.) Váňa, Phytotaxa 112 (1): 12, 2013 (Váňa et al.
2013g). Bas.: Hygrobiella nishimurae N.Kitag., Misc. Bryol. Lichenol. 9 (4): 69,
1982 (Kitagawa 1982).
** Cephalozia pachygyna R.M.Schust. ex J.J.Engel, Novon 17 (3): 313, 2007 (Engel 2007).
** Cephalozia physocaula (Hook.f. et Taylor) Steph., Bull. Herb. Boissier (sér. 2) 8
(7): 485 (315), 1908 (Stephani 1908f). Bas.: Jungermannia physocaula Hook.f. et
Taylor, London J. Bot. 3: 455, 1844 (Hooker and Taylor 1844b).
** Cephalozia schusteriana J.J.Engel, Novon 17 (3): 313, 2007 (Engel 2007).
*** Cephalozia stolonacea (Herzog) Váňa, Phytotaxa 112 (1): 12, 2013 (Váňa et al.
2013g). Bas.: Hygrobiella stolonacea Herzog, Trans. Brit. Bryol. Soc. 1 (4): 293,
1950 (Herzog 1950a).
*** Cephalozia tubulata (Hook.f. et Taylor) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 417, 1877 (Trevisan 1877). Bas.: Jungermannia tubulata Hook.f. et Taylor, London J. Bot. 3: 463, 1844 (Hooker and Taylor 1844b).
33 Cephalozia neesiana may be conspeciic with Cephalozia hamatiloba (Váňa 1988).
World checklist of hornworts and liverworts
*
*
*
*
*
59
Excluded from the genus
Cephalozia hians Steph., Sp. Hepat. (Stephani) 6: 441, 1924 (Stephani 1924). 34
Cephalozia indica Udar et D.Kumar, Geophytology 6 (1): 42, 1976 (Udar and
Kumar 1976). 35
Cephalozia kodaikanalensis G.Asthana et Saumya Srivast., Geophytology 43 (1):
63, 2013 (Asthana and Srivastava 2013). 36
Cephalozia parvifolia Arnell, Rev. Bryol. 25 (1): 1, 1898 (Arnell 1898). 37
Cephalozia tricuspidata (Nees) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 417, 1877 (Trevisan 1877). Bas.: Jungermannia tricuspidata
Nees, Enum. Pl. Crypt. Javae: 31, 1830 (Nees 1830). 38
*** Fuscocephaloziopsis Fulford, Mem. New York Bot. Gard. 11 (3): 353, 1968
(Fulford 1968).
** Fuscocephaloziopsis ainis (Lindb. ex Steph.) Váňa et L.Söderstr., Phytotaxa 112
(1): 9, 2013 (Váňa et al. 2013g). Bas.: Cephalozia ainis Lindb. ex Steph., Bull.
Herb. Boissier (sér. 2) 8 (4): 277 (291), 1908 (Stephani 1908j).
*** Fuscocephaloziopsis africana (Váňa) Váňa et L.Söderstr., Phytotaxa 112 (1): 9,
2013 (Váňa et al. 2013g). Bas.: Cephalozia africana Váňa, Beih. Nova Hedwigia
90: 187, 1988 (Váňa 1988).
*** Fuscocephaloziopsis albescens (Hook.) Váňa et L.Söderstr., Phytotaxa 112 (1): 9,
2013 (Váňa et al. 2013g). Bas.: Jungermannia albescens Hook., Brit. Jungermann.:
tab. 72, 1815 (Hooker 1815).
* Fuscocephaloziopsis albescens var. islandica (Nees) Váňa et L.Söderstr., Phytotaxa
112 (1): 9, 2013 (Váňa et al. 2013g). Bas.: Jungermannia islandica Nees, Naturgesch. Eur. Leberm. 2: 29, 1836 (Nees 1836).
*** Fuscocephaloziopsis baldwinii (C.M.Cooke) Váňa et L.Söderstr., Phytotaxa 112
(1): 9, 2013 (Váňa et al. 2013g). Bas.: Cephalozia baldwinii C.M.Cooke, Trans.
Connecticut Acad. Arts 12 (1): 35, 1904 (Cooke 1904).
** Fuscocephaloziopsis biloba (Herzog) Fulford, Mem. New York Bot. Gard. 11 (3):
355, 1968 (Fulford 1968). Bas.: Alobiella biloba Herzog, Feddes Repert. Spec.
Nov. Regni Veg. 57 (1/2): 166, 1955 (Herzog 1955).
*** Fuscocephaloziopsis catenulata (Huebener) Váňa et L.Söderstr., Phytotaxa 112 (1):
9, 2013 (Váňa et al. 2013g). Bas.: Jungermannia catenulata Huebener, Hepaticol.
germ.: 169, 1834 (Hübener 1834).
*** Fuscocephaloziopsis catenulata subsp. nipponica (S.Hatt.) Váňa et L.Söderstr., Phytotaxa 112 (1): 9, 2013 (Váňa et al. 2013g). Bas.: Cephalozia nipponica S.Hatt.,
Bull. Tokyo Sci. Mus. 11: 74, 1944 (Hattori 1944d).
34
35
36
37
38
Cephalozia hians is probably an Isotachis or Lophocolea species.
Cephalozia indica is a Lophocolea species and probably conspeciic with “Cephalozia” kodaikanalensis.
Cephalozia kodaikanalensis is a Lophocolea species, possibly conspeciic with “Cephalozia” indica.
Cephalozia parvifolia is a Cephaloziella species although the type specimen has not been found.
Cephalozia tricuspidata is a Lepidozia species (Schifner 1898b).
60
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Fuscocephaloziopsis connivens (Dicks.) Váňa et L.Söderstr., Phytotaxa 112 (1): 9,
2013 (Váňa et al. 2013g). Bas.: Jungermannia connivens Dicks., Fasc. Pl. Crypt.
Brit. 4: 19, 1801 (Dickson 1801).
*** Fuscocephaloziopsis connivens subsp. issa (Steph.) Váňa et L.Söderstr., Phytotaxa
112 (1): 9, 2013 (Váňa et al. 2013g). Bas.: Cephalozia issa Steph., Hedwigia 30
(5): 204, 1891 (Stephani 1891a).
*** Fuscocephaloziopsis connivens subsp. sandwicensis (Mont.) Váňa et L.Söderstr.,
Phytotaxa 112 (1): 9, 2013 (Váňa et al. 2013g). Bas.: Jungermannia sandwicensis
Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 249, 1843 (Montagne 1843).
*** Fuscocephaloziopsis crassifolia (Lindenb. et Gottsche) Váňa et L.Söderstr., Phytotaxa 112 (1): 10, 2013 (Váňa et al. 2013g). Bas.: Jungermannia crassifolia Lindenb.
et Gottsche, Syn. Hepat. 5: 685, 1847 (Gottsche et al. 1847).
*** Fuscocephaloziopsis gollanii (Steph.) Váňa et L.Söderstr., Phytotaxa 112 (1): 10,
2013 (Váňa et al. 2013g). Bas.: Cephalozia gollanii Steph., Bull. Herb. Boissier
(sér. 2) 8 (6): 428 (304), 1908 (Stephani 1908g).
*** Fuscocephaloziopsis leucantha (Spruce) Váňa et L.Söderstr., Phytotaxa 112 (1): 10,
2013 (Váňa et al. 2013g). Bas.: Cephalozia leucantha Spruce, Cephalozia: 68, 1882
(Spruce 1882).
*** Fuscocephaloziopsis loitlesbergeri (Schifn.) Váňa et L.Söderstr., Phytotaxa 112 (1):
10, 2013 (Váňa et al. 2013g). Bas.: Cephalozia loitlesbergeri Schifn., Österr. Bot.
Z. 62 (1): 10, 1912 (Schifner 1912b).
*** Fuscocephaloziopsis lunulifolia (Dumort.) Váňa et L.Söderstr., Phytotaxa 112 (1):
10, 2013 (Váňa et al. 2013g). Bas.: Jungermannia lunulifolia Dumort., Syll. Jungerm. Europ.: 61, 1831 (Dumortier 1831). 39
*** Fuscocephaloziopsis macrostachya (Kaal.) Váňa et L.Söderstr., Phytotaxa 112 (1):
10, 2013 (Váňa et al. 2013g). Bas.: Cephalozia macrostachya Kaal., Rev. Bryol. 29
(1): 8, 1902 (Kaalaas 1902).
*** Fuscocephaloziopsis macrostachya subsp. australis (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 112 (1): 10, 2013 (Váňa et al. 2013g). Bas.: Cephalozia macrostachya subsp. australis R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 754, 1974
(Schuster 1974).
*** Fuscocephaloziopsis macrostachya subsp. macrostachya var. spinilora (Schifn.) Váňa
et L.Söderstr., Phytotaxa 112 (1): 10, 2013 (Váňa et al. 2013g). Bas.: Cephalozia
spinilora Schifn., Hedwigia 54 (6): 323, 1914 (Schifner 1914a).
*** Fuscocephaloziopsis monticola (J.D.Godfrey) Váňa et L.Söderstr., Phytotaxa 112
(1): 11, 2013 (Váňa et al. 2013g). Bas.: Schoieldia monticola J.D.Godfrey, Bryologist 79 (3): 315, 1976 (Godfrey 1976).
*** Fuscocephaloziopsis pachycaulis (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 112
(1): 11, 2013 (Váňa et al. 2013g). Bas.: Cephalozia pachycaulis R.M.Schust., Bryologist 96 (4): 623, 1993 (Schuster 1993b).
39 Fuscocephaloziopsis lunulifolia is a species complex also including Fuscocephaloziopsis ainis and Fuscocephaloziopsis schusteri.
World checklist of hornworts and liverworts
61
*** Fuscocephaloziopsis pleniceps (Austin) Váňa et L.Söderstr., Phytotaxa 112 (1): 11,
2013 (Váňa et al. 2013g). Bas.: Jungermannia pleniceps Austin, Proc. Acad. Nat.
Sci. Philadelphia 21: 222, 1869 (Austin 1869).
** Fuscocephaloziopsis pleniceps var. caroliniana (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 112 (1): 11, 2013 (Váňa et al. 2013g). Bas.: Cephalozia pleniceps var. caroliniana R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 780, 1974 (Schuster 1974).
*** Fuscocephaloziopsis pulvinata (Steph.) Fulford, Mem. New York Bot. Gard. 11 (3):
355, 1968 (Fulford 1968). Bas.: Alobiella pulvinata Steph., Bull. Herb. Boissier
(sér. 2) 8 (8): 572 (356), 1908 (Stephani 1908e).
* Fuscocephaloziopsis schusteri (Sushil K.Singh et D.K.Singh) Váňa, Phytotaxa 183
(4): 291, 2014 (Váňa et al. 2014a). Bas.: Cephalozia schusteri Sushil K.Singh et
D.K.Singh, Lindbergia 32 (1): 1, 2007 (Singh and Singh 2007a).
*** Fuscocephaloziopsis subintegra Gradst. et Váňa, Cryptog. Bryol. 25 (3): 274, 2004
(Parolly et al. 2004).
*** Fuscocephaloziopsis zoopsioides (Horik.) Váňa et L.Söderstr., Phytotaxa 112 (1): 11,
2013 (Váňa et al. 2013g). Bas.: Cephalozia zoopsioides Horik., J. Sci. Hiroshima
Univ., Ser. B, Div. 2, Bot. 2: 178, 1934 (Horikawa 1934).
*** Nowellia Mitt., Nat. hist. Azores: 321, 1870 (Mitten 1870).
** sect. Acronowellia R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 816, 1974
(Schuster 1974).
*** Nowellia reedii H.Rob., Bryologist 73 (1): 150, 1970 (Robinson 1970).
*** Nowellia yunckeri Fulford, Mem. New York Bot. Gard. 11 (3): 329, 1968 (Fulford
1968).
** sect. Metanowellia (Grolle) R.M.Schust., Hepat. Anthocerotae N. Amer. 3:
816, 1974 (Schuster 1974). Bas.: Nowellia subg. Metanowellia Grolle, J. Hattori
Bot. Lab. 31: 33, 1968 (Grolle 1968b).
*** Nowellia borneensis (De Not.) Schifn., Hepat. (Engl.-Prantl): 98, 1893 (Schifner
1893b). Bas.: Jungermannia curvifolia var. borneensis De Not., Epat. Borneo: 19,
1874 (De Notaris 1874).
*** Nowellia dominicensis Steph., Sp. Hepat. (Stephani) 6: 443, 1924 (Stephani 1924).
*** Nowellia evansii Grolle, J. Hattori Bot. Lab. 31: 33, 1968 (Grolle 1968b).
*** Nowellia langii Pearson, J. Linn. Soc., Bot. 46 (305): 25, 1922 (Pearson 1922b).
*** Nowellia pusilla Grolle, J. Hattori Bot. Lab. 31: 37, 1968 (Grolle 1968b).
*** Nowellia wrightii (Gottsche ex Spruce) Steph. ex Duss, Enum. musc. Antilles
franç., Hép.: 21, 1903 (Duss 1903). Bas.: Cephalozia wrightii Gottsche ex Spruce,
J. Linn. Soc., Bot. 30 (210): 354, 1895 (Gepp 1895b).
62
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** sect. Nowellia
*** Nowellia aciliata (P.C.Chen et P.C.Wu) Mizut., Hikobia 11: 469, 1994 (Mizutani 1994). Bas.: Nowellia curvifolia var. aciliata P.C.Chen et P.C.Wu, Obs. l.
Hwangs.: 6, 1965 (Chen and Wu 1965).
*** Nowellia curvifolia (Dicks.) Mitt., Nat. hist. Azores: 321, 1870 (Mitten 1870). Bas.:
Jungermannia curvifolia Dicks., Fasc. Pl. Crypt. Brit. 2: 15, 1790 (Dickson 1790).
**
Odontoschismatoideae H.Buch ex Grolle
*** Odontoschisma (Dumort.) Dumort., Recueil Observ. Jungerm.: 19, 1835 (Dumortier 1835). Bas.: Pleuroschisma sect. Odontoschisma Dumort., Syll. Jungerm.
Europ.: 68, 1831 (Dumortier 1831).
** sect. Cladopodiella (H.Buch) Gradst., S.C.Aranda et Vanderp., Taxon 63 (5):
1017, 2014 (Aranda et al. 2014). Bas.: Cladopodiella H.Buch, Memoranda Soc.
Fauna Fl. Fennica 1: 89, 1927 (Buch 1927).
*** Odontoschisma francisci (Hook.) L.Söderstr. et Váňa, Phytotaxa 112 (1): 12, 2013
(Váňa et al. 2013g). Bas.: Jungermannia francisci Hook., Brit. Jungermann.: tab.
49, 1813 (Hooker 1813).
** sect. Denudata R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 833, 1974
(Schuster 1974).
*** Odontoschisma brasiliense Steph., Bull. Herb. Boissier (sér. 2) 8 (8): 585 (369),
1908 (Stephani 1908e).
** Odontoschisma cleeii Gradst., S.C.Aranda et Vanderp., Taxon 63 (5): 1017, 2014
(Aranda et al. 2014).
*** Odontoschisma denudatum (Mart.) Dumort., Recueil Observ. Jungerm.: 19, 1835
(Dumortier 1835). Bas.: Jungermannia scalaris var. denudata Mart., Fl. crypt. erlang.: 183, 1817 (Martius 1817).
** Odontoschisma denudatum subsp. naviculare (Steph.) Gradst., S.C.Aranda et Vanderp., Taxon 63 (5): 1019, 2014 (Aranda et al. 2014). Bas.: Jamesoniella navicularis Steph., Sp. Hepat. (Stephani) 6: 101, 1917 (Stephani 1917a).
** Odontoschisma denudatum subsp. sandvicense (Ångstr.) Gradst., S.C.Aranda et
Vanderp., Taxon 63 (5): 1019, 2014 (Aranda et al. 2014). Bas.: Sphagnoecetis
sandvicensis Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 29 (4): 22, 1872 (Ångström 1872).
*** Odontoschisma elongatum (Lindb.) A.Evans, Rhodora 14 (157): 13, 1912 (Evans
1912b). Bas.: Odontoschisma denudatum f. elongatum Lindb., Helsingf. Dagbl.
1874 (45, 16 Feb): 2, 1874 (Lindberg 1874b).
*** Odontoschisma engelii Gradst. et Burghardt, Fieldiana, Bot. (n.ser.) 47: 194, 2008
(Gradstein and Burghardt 2008).
World checklist of hornworts and liverworts
63
*** Odontoschisma longilorum (Taylor) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser.
3), C. Sci. Mat. 4 (13): 419, 1877 (Trevisan 1877). Bas.: Sphagnoecetis longilora
Taylor, London J. Bot. 5: 281, 1846 (Taylor 1846a).
*** Odontoschisma macounii (Austin) Underw., Bull. Illinois State Lab. Nat. Hist. 2
(1): 92, 1884 (Underwood 1884). Bas.: Sphagnoecetis macounii Austin, Bull. Torrey Bot. Club 3 (3): 13, 1872 (Austin 1872).
*** Odontoschisma portoricense (Hampe et Gottsche) Steph., Hedwigia 27 (11/12):
296, 1888 (Stephani 1888c). Bas.: Sphagnoecetis portoricensis Hampe et Gottsche,
Linnaea 25 (3): 343, 1852 [1853] (Hampe and Gottsche 1852).
** Odontoschisma pseudogrosseverrucosum Gradst., S.C.Aranda et Vanderp., Taxon 63
(5): 1019, 2014 (Aranda et al. 2014).
*** Odontoschisma purpuratum Herzog, Trans. Brit. Bryol. Soc. 1 (4): 297, 1950
(Herzog 1950a).
*** Odontoschisma soratamum Fulford, Mem. New York Bot. Gard. 11 (3): 338, 1968
(Fulford 1968).
*** Odontoschisma variabile (Lindenb. et Gottsche) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci. Mat. 4 (13): 419, 1877 (Trevisan 1877). Bas.: Sphagnoecetis
variabilis Lindenb. et Gottsche, Syn. Hepat. 5: 688, 1847 (Gottsche et al. 1847).
** Odontoschisma zhui Gradst., S.C.Aranda et Vanderp., Taxon 63 (5): 1020, 2014
(Aranda et al. 2014).
** sect. Iwatsukia (N.Kitag.) Gradst., S.C.Aranda et Vanderp., Phytotaxa 162 (4):
232, 2014 (Gradstein et al. 2014b). Bas.: Iwatsukia N.Kitag., J. Hattori Bot. Lab.
27: 178, 1964 (Kitagawa 1964).
*** Odontoschisma biidum (Fulford) Gradst., S.C.Aranda et Vanderp., Phytotaxa 162
(4): 232, 2014 (Gradstein et al. 2014b). Bas.: Cladomastigum biidum Fulford,
Acta Bot. Venez. 2 (5/8): 80, 1967 (Fulford 1967).
*** Odontoschisma jishibae (Steph.) L.Söderstr. et Váňa, Phytotaxa 112 (1): 13, 2013
(Váňa et al. 2013g). Bas.: Cephalozia jishibae Steph., Sp. Hepat. (Stephani) 6: 437,
1924 (Stephani 1924).
*** Odontoschisma spinosum (Fulford) Gradst., S.C.Aranda et Vanderp., Phytotaxa
162 (4): 232, 2014 (Gradstein et al. 2014b). Bas.: Cladomastigum spinosum Fulford, Mem. New York Bot. Gard. 23: 840, 1972 (Fulford 1972).
** sect. Neesia Gradst., S.C.Aranda et Vanderp., Taxon 63 (5): 1017, 2014 (Aranda et al. 2014).
*** Odontoschisma luitans (Nees) L.Söderstr. et Váňa, Phytotaxa 112 (1): 12, 2013 (Váňa
et al. 2013g). Bas.: Jungermannia luitans Nees, Flora 6 (2): 30, 1823 (Link 1823).
** sect. Odontoschisma
*** Odontoschisma grosseverrucosum Steph., Bull. Herb. Boissier (sér. 2) 8 (8): 593
(377), 1908 (Stephani 1908e).
64
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Odontoschisma sphagni (Dicks.) Dumort., Recueil Observ. Jungerm.: 19, 1835
(Dumortier 1835). Bas.: Jungermannia sphagni Dicks., Fasc. Pl. Crypt. Brit. 1: 6,
1785 (Dickson 1785).
** Odontoschisma steyermarkii Gradst. et Ilk.-Borg., Nova Hedwigia 100 (1/2): 38,
2015 [2014 online] (Gradstein and Ilkiu-Borges 2015).
*
**
Incertae sedis
Odontoschisma obcordatum (Spruce) Steph., Bull. Herb. Boissier (sér. 2) 8 (8): 586
(370), 1908 (Stephani 1908e). Bas.: Cephalozia obcordata Spruce, Cephalozia: 61,
1882 (Spruce 1882). 40
Schiffnerioideae R.M.Schust.
** Schifneria Steph., Österr. Bot. Z. 44 (1): 1, 1894 (Stephani 1894c).
*** Schifneria hyalina Steph., Österr. Bot. Z. 44 (1): 1, 1894 (Stephani 1894c).
**
Trabacelluloideae R.M.Schust.
** Haesselia Grolle et Gradst., J. Hattori Bot. Lab. 64: 327, 1988 (Grolle and Gradstein 1988).
*** Haesselia acuminata Gradst., Trop. Bryol. 1: 30, 1989 (Gradstein and Florschützde Waard 1989).
*** Haesselia roraimensis Grolle et Gradst., J. Hattori Bot. Lab. 64: 327, 1988 (Grolle
and Gradstein 1988).
** Trabacellula Fulford, Acta Bot. Venez. 2 (5/8): 86, 1967 (Fulford 1967).
*** Trabacellula tumidula Fulford, Acta Bot. Venez. 2 (5/8): 86, 1967 (Fulford 1967).
***
Cephaloziellaceae Douin
by J. Váňa
he treatment of Cephaloziellaceae follows what was outlined in Váňa et al. (2013e).
Some nomenclatural and taxonomic notes can also be found in Söderström et al. (2013a)
and Váňa et al. (2013b, 2013l). he placement of Phycolepidozia follows Gradstein et al.
(2014a).
40 Odontoschisma obcordatum is to be excluded from Odontoschisma but its identity remains unclear
as the type is not found. Based on the original description this species might belong to Alobiellopsis
(Gradstein & Ilkiu-Borges 2014: 81).
World checklist of hornworts and liverworts
65
*** Allisoniella E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot. 3 (4): 80, 1965
(Hodgson 1965).
** sect. Allisoniella
*** Allisoniella nigra (Rodway) R.M.Schust., Nova Hedwigia 22: 137, 1971 [1972]
(Schuster 1971b). Bas.: Sphenolobus niger Rodway, Tasm. Bryoph.: 33, 1917
(Rodway 1917b).
** Allisoniella nigra var. acutiloba J.J.Engel, Novon 17 (3): 313, 2007 (Engel 2007).
** Allisoniella nigra subsp. novaezelandiae R.M.Schust., Nova Hedwigia 22: 143,
1971 [1972] (Schuster 1971b).
*** Allisoniella recurva R.M.Schust., Nova Hedwigia 22: 146, 1971 [1972] (Schuster
1971b).
*** Allisoniella subbipartita (C.Massal.) R.M.Schust. et J.J.Engel, Nova Hedwigia
22: 147, 1971 [1972] (Schuster 1971b). Bas.: Cephalozia subbipartita C.Massal.,
Nuovo Giorn. Bot. Ital. 17 (3): 235, 1885 (Massalongo 1885).
*** Allisoniella tasmanica R.M.Schust., Nova Hedwigia 22: 145, 1971 [1972] (Schuster
1971b).
** sect. Protomarsupella (R.M.Schust.) R.M.Schust., Nova Hedwigia 22: 150,
1971 (Schuster 1971b). Bas.: Protomarsupella R.M.Schust., Rev. Bryol. Lichénol.
34 (1/2): 264, 1966 (Schuster 1966b).
*** Allisoniella scottii (R.M.Schust.) R.M.Schust., Nova Hedwigia 22: 151, 1971
[1972] (Schuster 1971b). Bas.: Protomarsupella scottii R.M.Schust., Rev. Bryol.
Lichénol. 34 (1/2): 267, 1966 (Schuster 1966b).
*** Amphicephalozia R.M.Schust., Nova Hedwigia 22: 131, 1971 [1972] (Schuster
1971b).
*** Amphicephalozia africana Váňa et M.Wigginton, J. Bryol. 30 (1): 55, 2008 (Váňa
and Wigginton 2008).
*** Amphicephalozia amplexicaulis R.M.Schust., Nova Hedwigia 22: 133, 1971 [1972]
(Schuster 1971b).
*** Amphicephalozia geisslerae Pócs et Váňa, Polish Bot. J. 46 (2): 145, 2001 (Pócs and
Váňa 2001).
*** Anastrophyllopsis (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 81 (1): 15,
2013 (Váňa et al. 2013k). Bas.: Anastrophyllum sect. Anastrophyllopsis R.M.Schust.,
Beih. Nova Hedwigia 119: 310, 2002 (Schuster 2002b).
*** Anastrophyllopsis involutifolia (Mont. ex Gottsche, Lindenb. et Nees) Váňa et L.Söderstr.,
Phytotaxa 81 (1): 15, 2013 (Váňa et al. 2013k). Bas.: Jungermannia involutifolia Mont.
ex Gottsche, Lindenb. et Nees, Syn. Hepat. 1: 81, 1844 (Gottsche et al. 1844).
*** Anastrophyllopsis revoluta (Steph.) Váňa et L.Söderstr., Phytotaxa 81 (1): 16, 2013
(Váňa et al. 2013k). Bas.: Anastrophyllum revolutum Steph., Hedwigia 32 (3): 139,
1893 (Stephani 1893b).
66
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Anastrophyllopsis subcomplicata (Lehm. et Lindenb.) Váňa et L.Söderstr., Phytotaxa 81 (1): 16, 2013 (Váňa et al. 2013k). Bas.: Jungermannia subcomplicata Lehm.
et Lindenb., Nov. Stirp. Pug. 7: 4, 1838 (Lehmann 1838).
** Cephalojonesia Grolle, Rev. Bryol. Lichénol. 37 (4): 763, 1970 [1971] (Grolle
and Vanden Berghen 1970).
*** Cephalojonesia incuba Grolle et Vanden Berghen, Rev. Bryol. Lichénol. 37 (4):
764, 1970 [1971] (Grolle and Vanden Berghen 1970).
*** Cephalojonesia incuba subsp. mexicana Burghardt, Gradst. et Váňa, J. Hattori Bot.
Lab. 100: 35, 2006 (Burghardt et al. 2006).
** Cephalomitrion R.M.Schust., Nova Hedwigia 61 (3/4): 550, 1995 (Schuster 1995b).
*** Cephalomitrion aterrimum (Steph.) R.M.Schust., Nova Hedwigia 61 (3/4): 554,
1995 (Schuster 1995b). Bas.: Cephalozia aterrima Steph., Bull. Herb. Boissier (sér.
2) 8 (7): 501 (331), 1908 (Stephani 1908f).
*** Cephaloziella (Spruce) Schifn., Hepat. (Engl.-Prantl): 98, 1893 (Schifner
1893b) nom. conserv. Bas.: Cephalozia subg. Cephaloziella Spruce, Cephalozia:
62, 1882 (Spruce 1882). 41
** subg. Cephaloziella
*** Cephaloziella aenigmatica R.M.Schust., Nova Hedwigia 63 (1/2): 20, 1996 (Schuster
1996d).
*** Cephaloziella anthelioides S.W.Arnell, Bot. Not. 105: 322, 1952 (Arnell 1952a).
*** Cephaloziella arctogena (R.M.Schust.) Konstant., Arctoa 3: 126, 1994 (Konstantinova and Vasil’ev 1994). Bas.: Cephaloziella rubella var. arctogena R.M.Schust.,
Hepat. Anthocerotae N. Amer. 4: 125, 1980 (Schuster 1980c).
** Cephaloziella arenaria (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 280, 1963
(Schuster 1963b). Bas.: Cephalozia arenaria Steph., Bull. Herb. Boissier (sér. 2) 8
(7): 504 (334), 1908 (Stephani 1908f).
*** Cephaloziella aspericaulis Jørg., Bergens Mus. Skr. (n.ser.) 16: 197, 1934 (Jørgensen
1934).
*** Cephaloziella baumgartneri Schifn., Verh. K.K. Zool.-Bot. Ges. Wien 56 (3): 273,
1906 (Schifner 1906c).
** Cephaloziella breviperianthia C.Gao, Fl. Hepat. Chin. Boreali-Orient.: 208, 1981
(Gao and Chang 1981).
** Cephaloziella brinkmannii Douin, Mém. Soc. Bot. France 29: 75, 1920 (Douin 1920).
*** Cephaloziella capensis (Sim) S.W.Arnell, Bot. Not. 105: 326, 1952 (Arnell 1952a). Bas.:
Cephalozia capensis Sim, Trans. Roy. Soc. South Africa 15 (1): 87, 1926 (Sim 1926).
41 Cephaloziella is a large genus and the subgeneric division is unclear. he traditional subdivisions used
here do not necessary agree with recent phylogenetic studies.
World checklist of hornworts and liverworts
67
*** Cephaloziella capillaris (Steph.) Douin, Mém. Soc. Bot. France 29: 59, 1920
(Douin 1920). Bas.: Cephalozia capillaris Steph., Bull. Herb. Boissier (sér. 2) 8
(7): 496 (326), 1908 (Stephani 1908f).
*** Cephaloziella crassigyna (R.M.Schust.) R.M.Schust., Nova Hedwigia 61 (3/4): 556,
1995 (Schuster 1995b). Bas.: Cephaloziella aterrima var. crassigyna R.M.Schust.,
Nova Hedwigia 22: 211, 1971 [1972] (Schuster 1971b).
*** Cephaloziella crispata N.Kitag., J. Hattori Bot. Lab. 32: 301, 1969 (Kitagawa 1969c).
*** Cephaloziella densifolia R.M.Schust., Nova Hedwigia 22: 199, 1971 [1972]
(Schuster 1971b).
** Cephaloziella densifolia var. dubia R.M.Schust., Nova Hedwigia 22: 202, 1971
[1972] (Schuster 1971b).
*** Cephaloziella divaricata (Sm.) Schifn., Hepat. (Engl.-Prantl): 99, 1893 (Schifner
1893b). Bas.: Jungermannia divaricata Sm., Engl. Bot. 10: tab. 719, 1800 (Smith
and Sowerby 1800).
** Cephaloziella divaricata var. scabra (M.Howe) Haynes, Bryologist 12 (4): 68, 1909
(Haynes 1909). Bas.: Cephalozia divaricata var. scabra M.Howe, Mem. Torrey
Bot. Club 7: 129, 1899 (Howe 1899). 42
** Cephaloziella dusenii Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26 (III,
6): 49, 1900 (Stephani 1900b).
*** Cephaloziella elachista (J.B.Jack ex Gottsche et Rabenh.) Schifn., Sitzungsber.
deutsch. naturwiss.-med. Vereins Böhmen “Lotos” Prag 48: 336, 1900 (Schifner 1900d). Bas.: Jungermannia elachista J.B.Jack ex Gottsche et Rabenh., Hepat.
Eur., Leberm. 58-59: no. 574, 1873 (Gottsche and Rabenhorst 1873a).
** Cephaloziella elachista var. spinophylla (C.Gao) C.Gao, Fl. Bryoph. Sin. 9: 181,
2003 (Gao 2003). Bas.: Cephaloziella spinophylla C.Gao, Fl. Hepat. Chin. BorealiOrient.: 208, 1981 (Gao and Chang 1981).
** Cephaloziella elegans (Heeg) Schifn., Sitzungsber. deutsch. naturwiss.-med. Vereins
Böhmen “Lotos” Prag 48: 336, 1900 (Schifner 1900d). Bas.: Cephalozia elegans
Heeg, Rev. Bryol. 20 (5): 82, 1893 (Heeg 1893).
*** Cephaloziella exigua R.M.Schust., Nova Hedwigia 63 (1/2): 49, 1996 (Schuster 1996d).
*** Cephaloziella exililora (Taylor) Douin, Mém. Soc. Bot. France 29: 72, 1920
(Douin 1920). Bas.: Jungermannia exililora Taylor, London J. Bot. 5: 279, 1846
(Taylor 1846a).
** Cephaloziella fragillima (Spruce) Fulford, Mem. New York Bot. Gard. 11 (4): 409,
1976 (Fulford 1976). Bas.: Cephalozia fragillima Spruce, Mem. Torrey Bot. Club
1 (3): 131, 1890 (Spruce 1890).
*** Cephaloziella garsidei S.W.Arnell, Rev. Bryol. Lichénol. 23 (1/2): 173, 1954 (Arnell
1954a).
*** Cephaloziella grandiretis (R.M.Schust.) R.M.Schust., Nova Hedwigia 63 (1/2): 35,
1996 (Schuster 1996d). Bas.: Cephaloziella byssacea subsp. grandiretis R.M.Schust.,
Nova Hedwigia 22: 195, 1971 [1972] (Schuster 1971b).
42 he correct name for the variety was discussed by Söderström et al. (2012d).
68
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Cephaloziella grimsulana (J.B.Jack ex Gottsche et Rabenh.) Lacout., Hépat. France: 52,
1905 (Lacouture 1905). Bas.: Jungermannia grimsulana J.B.Jack ex Gottsche et Rabenh., Hepat. Eur., Leberm. 53–55: no. 526, 1872 (Gottsche and Rabenhorst 1872).
*** Cephaloziella hampeana (Nees) Schifn. ex Loeske, Moosl. Harz.: 92, 1903 (Loeske 1903). Bas.: Jungermannia hampeana Nees, Naturgesch. Eur. Leberm. 3: 560,
1838 (Nees 1838b).
** Cephaloziella hebridensis Steph., Hedwigia 32 (5): 316, 1893 (Stephani 1893d).
*** Cephaloziella herzogiana (Pandé et K.P.Srivast.) Udar et D.Kumar, Geophytology
6 (1): 45, 1976 (Udar and Kumar 1976). Bas.: Cephalozia herzogiana Pandé et
K.P.Srivast., Feddes Repert. Spec. Nov. Regni Veg. 58: 75, 1955 (Pandé and Srivastava 1955).
** Cephaloziella heteroica (C.M.Cooke) Douin, Mém. Soc. Bot. France 29: 76, 1920
(Douin 1920). Bas.: Cephalozia heteroica C.M.Cooke, Trans. Connecticut Acad.
Arts 12 (1): 38, 1904 (Cooke 1904).
** Cephaloziella hyalina Douin, Mém. Soc. Bot. France 29: 77, 1920 (Douin 1920).
* Cephaloziella hyalina var. rappii (Douin) R.M.Schust., Hepat. Anthocerotae N.
Amer. 4: 56, 1980 (Schuster 1980c). Bas.: Cephaloziella rappii Douin, Mém. Soc.
Bot. France 29: 77, 1920 (Douin 1920).
*** Cephaloziella inaequalis R.M.Schust., Nova Hedwigia 22: 186, 1971 [1972]
(Schuster 1971b).
*** Cephaloziella invisa R.M.Schust., Nova Hedwigia 63 (1/2): 30, 1996 (Schuster
1996d).
** Cephaloziella kilohanensis (C.M.Cooke) Douin, Mém. Soc. Bot. France 29: 85,
1920 (Douin 1920). Bas.: Cephalozia kilohanensis C.M.Cooke, Trans. Connecticut Acad. Arts 12 (1): 37, 1904 (Cooke 1904).
*** Cephaloziella longii Váňa, Folia Geobot. Phytotax. 27 (2): 193, 1992 (Váňa 1992).
*** Cephaloziella lycopodioides (Sim) S.W.Arnell, Bot. Not. 105: 321, 1952 (Arnell
1952a). Bas.: Cephalozia lycopodioides Sim, Trans. Roy. Soc. South Africa 15 (1):
85, 1926 (Sim 1926).
** Cephaloziella mammillifera R.M.Schust. et Damsh., Phytologia 63 (5): 327, 1987
(Schuster and Damsholt 1987).
*** Cephaloziella massalongi (Spruce) Müll.Frib., Lebermoose 2 (17): 191, 1913 (Müller
1913a). Bas.: Cephalozia massalongi Spruce, Cephalozia: 71, 1882 (Spruce 1882).
*** Cephaloziella muelleriana R.M.Schust., Nova Hedwigia 63 (1/2): 24, 1996 (Schuster
1996d).
*** Cephaloziella natalensis (Sim) S.W.Arnell, Hepat. South Africa: 338, 1963 (Arnell
1963b). Bas.: Cephalozia natalensis Sim, Trans. Roy. Soc. South Africa 15 (1): 86,
1926 (Sim 1926).
** Cephaloziella nicholsonii Douin, Rev. Bryol. 40 (6): 81, 1913 (Douin 1913b).
*** Cephaloziella nothogena R.M.Schust., Nova Hedwigia 63 (1/2): 37, 1996 (Schuster
1996d).
** Cephaloziella obtusilobula R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 108,
1980 (Schuster 1980c).
World checklist of hornworts and liverworts
69
** Cephaloziella patulifolia (Steph.) Douin, Mém. Soc. Bot. France 29: 70, 1920
(Douin 1920). Bas.: Cephalozia patulifolia Steph., Bull. Herb. Boissier (sér. 2) 8
(7): 509 (339), 1908 (Stephani 1908f).
*** Cephaloziella phyllacantha (C.Massal. et Carestia) Müll.Frib., Lebermoose 2 (17):
194, 1913 (Müller 1913a). Bas.: Anthelia phyllacantha C.Massal. et Carestia, Nuovo Giorn. Bot. Ital. 12 (4): 340, 1880 (Massalongo and Carestia 1880).
*** Cephaloziella polystratosa (R.M.Schust. et Damsh.) Konstant., Bot. Zhurn. (Moscow & Leningrad) 85 (10): 127, 2000 (Konstantinova 2000). Bas.: Cephaloziella
byssacea var. polystratosa R.M.Schust. et Damsh., Phytologia 63 (5): 327, 1987
(Schuster and Damsholt 1987).
*** Cephaloziella pseudocrassigyna R.M.Schust. ex J.J.Engel, Novon 17 (3): 313, 2007
(Engel 2007).
*** Cephaloziella pulcherrima R.M.Schust., Nova Hedwigia 22: 203, 1971 [1972]
(Schuster 1971b).
** Cephaloziella pulcherrima subsp. sphagnicola R.M.Schust., Nova Hedwigia 22:
205, 1971 [1972] (Schuster 1971b).
** Cephaloziella pungens Steph. ex Fulford, Mem. New York Bot. Gard. 11 (4): 410,
1976 (Fulford 1976).
*** Cephaloziella rubella (Nees) Warnst., Krypt.-Fl. Brandenburg, Leber- & Torfmoose: 231, 1902 (Warnstorf 1902). Bas.: Jungermannia rubella Nees, Naturgesch. Eur. Leberm. 2: 236, 1836 (Nees 1836).
*** Cephaloziella schelpei S.W.Arnell, Bot. Not. 110 (1): 18, 1957 (Arnell 1957a).
*** Cephaloziella spegazziniana (C.Massal.) Douin, Mém. Soc. Bot. France 29: 69,
1920 (Douin 1920). Bas.: Cephalozia spegazziniana C.Massal., Nuovo Giorn. Bot.
Ital. 17 (3): 234, 1885 (Massalongo 1885).
*** Cephaloziella spinicaulis Douin, Mém. Soc. Bot. France 29: 62, 1920 (Douin 1920).
*** Cephaloziella spinigera (Lindb.) Jørg., Bergens Mus. Skr. (n.ser.) 16: 189, 1934
(Jørgensen 1934). Bas.: Cephalozia spinigera Lindb., Musci Scand.: 4, 1879 (Lindberg 1879).
*** Cephaloziella stellulifera (Taylor ex Carrington et Pearson) Croz., Rev. Bryol. 30 (2):
31, 1903 (Crozals 1903a). Bas.: Jungermannia stellulifera Taylor ex Carrington et
Pearson, Hepat. Brit. Exsicc. Fasc. I: no. 32, 1878 (Carrington and Pearson 1878).
*** Cephaloziella stephanii Schifn. ex Douin, Mém. Soc. Bot. France 29: 85, 1920
(Douin 1920).
** Cephaloziella stolonifera R.M.Schust., Phytologia 39 (6): 426, 1978 (Schuster 1978b).
** Cephaloziella subtilis (Lindenb. et Gottsche) Steph., Hedwigia 32 (5): 318, 1893
(Stephani 1893d). Bas.: Jungermannia subtilis Lindenb. et Gottsche, Syn. Hepat.
5: 685, 1847 (Gottsche et al. 1847).
*** Cephaloziella sumatrana Schifn. ex Douin, Mém. Soc. Bot. France 29: 59, 1920
(Douin 1920).
*** Cephaloziella tabularis S.W.Arnell, Bot. Not. 105: 318, 1952 (Arnell 1952a).
*** Cephaloziella transvaalensis S.W.Arnell, Bot. Not. 110 (1): 19, 1957 (Arnell 1957a).
*** Cephaloziella triplicata S.W.Arnell, Bot. Not. 115: 203, 1962 (Arnell 1962a).
70
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Cephaloziella umtaliensis S.W.Arnell, Bot. Not. 110 (1): 20, 1957 (Arnell 1957a).
*** Cephaloziella uncinata R.M.Schust., Meddel. Grønland 199 (1): 316, 1974
(Schuster and Damsholt 1974).
** Cephaloziella uncinata var. brevigyna R.M.Schust. et Damsh., Phytologia 63 (5):
327, 1987 (Schuster and Damsholt 1987).
** Cephaloziella uncinata var. sphagnicola R.M.Schust., Meddel. Grønland 199 (1):
323, 1974 (Schuster and Damsholt 1974).
*** Cephaloziella vaginans Steph., Wiss. Ergebn. Deut. Zentr.-Afr. Exped. (1907-08),
Bot. 2: 119, 1911 (Stephani 1911a).
*** Cephaloziella varians (Gottsche) Steph., Wiss. Ergebn. Schwed. Südpolar-Exped.
[1901–1903] 4 (1): 4, 1905 (Stephani 1905e). Bas.: Jungermannia varians
Gottsche, Int. Polarforsch., Deutsch. Exped. 2: 452, 1890 (Gottsche 1890).
*** Cephaloziella verrucosa Steph., Hedwigia 32 (5): 318, 1893 (Stephani 1893d).
* Cephaloziella villaumei (Steph.) Váňa, Phytotaxa 112 (1): 3, 2013 (Váňa et al.
2013e). Bas.: Cephalozia villaumei Steph., Sp. Hepat. (Stephani) 6: 437, 1924
(Stephani 1924). 43
** Cephaloziella violacea Schljakov, Novosti Sist. Nizš. Rast. 15: 242, 1978 (Shliakov 1978).
** Cephaloziella welwitschii (Steph.) Douin, Mém. Soc. Bot. France 29: 58, 1920
(Douin 1920). Bas.: Cephalozia welwitschii Steph., Bull. Herb. Boissier (sér. 2) 8
(6): 432 (308), 1908 (Stephani 1908g).
** subg. Dichiton (Mont.) Müll.Frib., Lebermoose 2 (27): 787, 1916 (Müller
1916). Bas.: Dichiton Mont., Syll. Gen. Sp. Crypt.: 52, 1856 (Montagne 1856b).
*** Cephaloziella calyculata (Durieu et Mont.) Müll.Frib., Lebermoose 2 (27): 787,
1916 (Müller 1916). Bas.: Jungermannia calyculata Durieu et Mont., Ann. Sci.
Nat. Bot. (sér. 3) 11: 34, 1849 (Montagne 1849).
*** Cephaloziella integerrima (Lindb.) Warnst., Krypt.-Fl. Brandenburg, Leber- &
Torfmoose: 232, 1902 (Warnstorf 1902). Bas.: Cephalozia integerrima Lindb.,
Acta Soc. Sci. Fenn. 10: 502, 1875 (Lindberg 1875).
** subg. Evansia (Douin et Schifn.) Müll.Frib., Lebermoose 2 (27): 787, 1916
(Müller 1916). Bas.: Evansia Douin et Schifn., Rev. Bryol. 40 (5): 66, 1913
(Douin 1913a).
*** Cephaloziella antillana (Besch. et Spruce) Fulford, Mem. New York Bot. Gard. 11
(4): 405, 1976 (Fulford 1976). Bas.: Cephalozia antillana Besch. et Spruce, Bull. Soc.
Bot. France (Congr. Bot.) 36: clxxxiii, 1889 [1890] (Bescherelle and Spruce 1889).
*** Cephaloziella dentata (Raddi) Steph., Bull. Herb. Boissier 5 (2): 78, 1897 (Stephani
1897b). Bas.: Jungermannia dentata Raddi, Jungermanniogr. Etrusca: 21, 1818
(Raddi 1818a).
*** Cephaloziella hirta (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 280, 1963
(Schuster 1963b). Bas.: Cephalozia hirta Steph., Bull. Herb. Boissier (sér. 2) 8 (8):
561 (345), 1908 (Stephani 1908e).
43 Cephaloziella villaumei is possibly conspeciic with Cephaloziella anthelioides.
World checklist of hornworts and liverworts
71
** Cephaloziella squarrosula (Trevis.) Steph., Hedwigia 32 (5): 318, 1893 (Stephani
1893d). Bas.: Cephalozia squarrosula Trevis., Mem. Reale Ist. Lombardo Sci. (Ser.
3), C. Sci. Mat. 4 (13): 417, 1877 (Trevisan 1877).
*** Cephaloziella subspinosa R.M.Schust., Nova Hedwigia 22: 210, 1971 [1972]
(Schuster 1971b).
** subg. Prionolobus (Spruce) Müll.Frib., Lebermoose 2 (16): 110, 1912 (Müller
1912). Bas.: Cephalozia subg. Prionolobus Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 508, 1885 (Spruce 1885).
*** Cephaloziella acanthophora (S.Hatt.) Horik., Hikobia 1 (2): 79, 1951 (Horikawa
1951c). Bas.: Prionolobus acanthophorus S.Hatt., Bull. Tokyo Sci. Mus. 11: 29,
1944 (Hattori 1944d).
*** Cephaloziella biokoensis Váňa et Frank Müll., Trop. Bryol. 24: 1, 2003 (Váňa and
Müller 2003).
*** Cephaloziella granatensis (J.B.Jack ex Steph.) Fulford, Mem. New York Bot. Gard.
11 (4): 411, 1976 (Fulford 1976). Bas.: Cephalozia granatensis J.B.Jack ex Steph.,
Bull. Herb. Boissier (sér. 2) 8 (7): 500 (330), 1908 (Stephani 1908f).
*** Cephaloziella grisea R.M.Schust., Phytologia 39 (6): 425, 1978 (Schuster 1978b).
* Cephaloziella meghalayensis Udar et Ad.Kumar, Lindbergia 8 (1): 34, 1982 (Udar
and Kumar 1982d). 44
*** Cephaloziella microphylla (Steph.) Douin, Mém. Soc. Bot. France 29: 59, 1920
(Douin 1920). Bas.: Cephalozia microphylla Steph., Bull. Herb. Boissier (sér. 2) 8
(7): 513 (343), 1908 (Stephani 1908f).
*** Cephaloziella tenuissima (Lehm.) Steph., Hedwigia 32 (5): 318, 1893 (Stephani 1893d).
Bas.: Jungermannia tenuissima Lehm., Linnaea 4: 367, 1829 (Lehmann 1829).
*** Cephaloziella turneri (Hook.) Müll.Frib., Lebermoose 2 (17): 202, 1913 (Müller
1913a). Bas.: Jungermannia turneri Hook., Brit. Jungermann.: tab. 29, 1812
(Hooker 1812).
Incertae sedis
** Cephaloziella dentifolia Udar et Ad.Kumar, Lindbergia 8 (1): 30, 1982 (Udar and
Kumar 1982d).
** Cephaloziella ilum (Trevis.) Steph., Hedwigia 32 (5): 318, 1893 (Stephani 1893d).
Bas.: Cephalozia ilum Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 417, 1877 (Trevisan 1877).
* Cephaloziella lexuosa C.Gao et K.C.Chang, Bull. Bot. Res., Harbin 4 (3): 88,
1984 (Chang and Gao 1984). 45
** Cephaloziella intricata Schifn. ex Douin, Mém. Soc. Bot. France 29: 59, 1920
(Douin 1920).
44 Cephaloziella meghalayensis is probably conspeciic with Cephaloziella acanthophora or Cephaloziella
microphylla.
45 Cephaloziella lexuosa may be a form of Plicanthus birmensis or a juvenile form of Tetralophozia iliformis.
72
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Cephaloziella levieri Schifn. ex Douin, Mém. Soc. Bot. France 29: 80, 1920
(Douin 1920).
*** Cephaloziella pellucida R.M.Schust., Nova Hedwigia 63 (1/2): 57, 1996 (Schuster
1996d). 46
* Cephaloziella pygmaea (Spruce) Váňa, Phytotaxa 183 (4): 290, 2014 (Váňa et al.
2014a). Bas.: Cephalozia pygmaea Spruce, Cephalozia: 69, 1882 (Spruce 1882). 47
* Cephaloziella secundifolia Pearson, Ann. Bryol. 4: 106, 1931 (Pearson 1931b). 48
** Cephaloziella sinensis Douin, Rev. Bryol. 41 (1): 8, 1914 (Douin 1914).
** Cephaloziopsis (Spruce) Schifn., Hepat. (Engl.-Prantl): 85, 1893 (Schifner
1893b). Bas.: Jungermannia sect. Cephaloziopsis Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 511, 1885 (Spruce 1885).
*** Cephaloziopsis intertexta (Gottsche) R.M.Schust., Nova Hedwigia 22: 183, 1971
[1972] (Schuster 1971b). Bas.: Jungermannia intertexta Gottsche, Syn. Hepat. 1:
107, 1844 (Gottsche et al. 1844).
** Chaetophyllopsis R.M.Schust., J. Hattori Bot. Lab. 23: 69, 1960 [1961] (Schuster
1960b).
*** Chaetophyllopsis whiteleggei (Carrington et Pearson) R.M.Schust. ex Hamlin, Rec.
Domin. Mus. 7: 255, 1972 (Hamlin 1972). Bas.: Jungermannia whiteleggei Carrington et Pearson, Proc. Linn. Soc. New South Wales (ser. 2) 2 (4): 1051, 1888
(Carrington and Pearson 1888a).
** Cylindrocolea R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 12 (3): 666,
1969 (Schuster 1969a).
** subg. Cylindrocolea
*** Cylindrocolea kiaeri (Austin) Váňa, Phytotaxa 112 (1): 2, 2013 (Váňa et al. 2013e). Bas.:
Jungermannia kiaeri Austin, Bull. Torrey Bot. Club 6 (3): 18, 1875 (Austin 1875b).
*** Cylindrocolea sanctae-helenae M.Wigginton, Polish Bot. J. 58 (1): 107, 2013 (Wigginton 2013).
** sect. Cylindrocolea
*** Cylindrocolea andersonii R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 33, 1980
(Schuster 1980c).
*** Cylindrocolea brasiliensis D.P.Costa, N.D.Santos et Váňa, Bryologist 111 (4): 667,
2008 (Costa et al. 2008).
*** Cylindrocolea chevalieri (Steph.) R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.)
12 (3): 666, 1969 (Schuster 1969a). Bas.: Alobiella chevalieri Steph., Bull. Herb.
Boissier (sér. 2) 8 (8): 567 (351), 1908 (Stephani 1908e).
46 Cephaloziella pellucida is probably not a Cephaloziella, but the type specimen has not been found.
47 Cephaloziella pygmaea is possibly conspeciic with Cephaloziella granatensis (Váňa et al. 2014a).
48 Cephaloziella secundifolia could not be studied by Fulford (1976) and she did not know what it is.
World checklist of hornworts and liverworts
73
*** Cylindrocolea gittinsii (E.W.Jones) R.M.Schust., Nova Hedwigia 22: 172, 1971
[1972] (Schuster 1971b). Bas.: Cephaloziella gittinsii E.W.Jones, Trans. Brit.
Bryol. Soc. 3 (3): 437, 1958 (Jones 1958).
*** Cylindrocolea madagascariensis (Steph.) R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 12 (3): 666, 1969 (Schuster 1969a). Bas.: Lophozia madagascariensis
Steph., Sp. Hepat. (Stephani) 6: 112, 1917 (Stephani 1917a).
*** Cylindrocolea nigerica (E.W.Jones) R.M.Schust., Nova Hedwigia 22: 172, 1971
[1972] (Schuster 1971b). Bas.: Cephaloziella nigerica E.W.Jones, Trans. Brit. Bryol. Soc. 3 (3): 435, 1958 (Jones 1958).
*** Cylindrocolea novae-caledoniae (Grolle) R.M.Schust., Nova Hedwigia 22: 161,
1971 [1972] (Schuster 1971b). Bas.: Cephaloziella novae-caledoniae Grolle, Rev.
Bryol. Lichénol. 29 (3/4): 208, 1960 [1961] (Grolle 1960a).
*** Cylindrocolea planifolia (Steph.) R.M.Schust., Nova Hedwigia 22: 164, 1971
[1972] (Schuster 1971b). Bas.: Cephaloziella planifolia Steph., Hedwigia 32 (5):
317, 1893 (Stephani 1893d).
*** Cylindrocolea sprucei R.M.Schust., Nova Hedwigia 22: 163, 1971 [1972] (Schuster
1971b).
*** Cylindrocolea ugandica (E.W.Jones) R.M.Schust., Nova Hedwigia 22: 171, 1971
[1972] (Schuster 1971b). Bas.: Cephaloziella ugandica E.W.Jones, Trans. Brit.
Bryol. Soc. 3 (3): 433, 1958 (Jones 1958).
** sect. Platycaulis R.M.Schust., Nova Hedwigia 22: 173, 1971 (Schuster
1971b).
*** Cylindrocolea recurvifolia (Steph.) Inoue, J. Jap. Bot. 47 (11): 348, 1972 (Inoue
1972a). Bas.: Cephalozia recurvifolia Steph., Bull. Herb. Boissier (sér. 2) 8 (7): 497
(327), 1908 (Stephani 1908f).
*** Cylindrocolea tagawae (N.Kitag.) R.M.Schust., Nova Hedwigia 22: 174, 1971
[1972] (Schuster 1971b). Bas.: Cephaloziella tagawae N.Kitag., J. Hattori Bot.
Lab. 32: 303, 1969 (Kitagawa 1969c).
** subg. Cylindroscyphus (Douin) R.M.Schust., Hepat. Anthocerotae N. Amer.
4: 20, 1980 (Schuster 1980c). Bas.: Cephaloziella subg. Cylindroscyphus Douin,
Mém. Soc. Bot. France 29: 56, 1920 (Douin 1920).
*** Cylindrocolea abyssinica (Gola) Váňa, Phytotaxa 112 (1): 2, 2013 (Váňa et al.
2013e). Bas.: Cephaloziella abyssinica Gola, Ann. Bot. (Rome) 13 (1): 68, 1914
(Gola 1914a).
** Cylindrocolea obtusifolia Fulford, Mem. New York Bot. Gard. 11 (4): 401, 1976
(Fulford 1976).
** Cylindrocolea reticulata Udar et Ad.Kumar, Lindbergia 8 (3): 181, 1982 [1983]
(Udar and Kumar 1982c).
*** Cylindrocolea rhizantha (Mont.) R.M.Schust., Nova Hedwigia 22: 175, 1971
[1972] (Schuster 1971b). Bas.: Jungermannia rhizantha Mont., Hist. Phys. Cuba,
Bot., Pl. Cell.: 454, 1842 (Montagne 1842a).
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Gottschelia Grolle, J. Hattori Bot. Lab. 31: 13, 1968 (Grolle 1968c).
*** Gottschelia maxima (Steph.) Grolle, J. Bryol. 25 (1): 6, 2003 (Grolle et al. 2003). Bas.:
Tylimanthus maximus Steph., Sp. Hepat. (Stephani) 6: 249, 1922 (Stephani 1922).
*** Gottschelia schizopleura (Spruce) Grolle, J. Hattori Bot. Lab. 31: 16, 1968 (Grolle
1968c). Bas.: Jungermannia schizopleura Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 517, 1885 (Spruce 1885).
** Gymnocoleopsis (R.M.Schust.) R.M.Schust., Phytologia 39 (4): 243, 1978
(Schuster 1978a). Bas.: Gymnocolea subg. Gymnocoleopsis R.M.Schust., Bryologist
70 (1): 111, 1967 (Schuster 1967b).
*** Gymnocoleopsis capensis (S.W.Arnell) R.M.Schust., J. Hattori Bot. Lab. 78: 123,
1995 (Schuster 1995c). Bas.: Lophozia capensis S.W.Arnell, Svensk Bot. Tidskr. 47
(1): 112, 1953 (Arnell 1953a).
*** Gymnocoleopsis cylindriformis (Mitt.) R.M.Schust., J. Hattori Bot. Lab. 78: 126,
1995 (Schuster 1995c). Bas.: Jungermannia cylindriformis Mitt., J. Linn. Soc., Bot.
15 (84): 196, 1876 (Mitten 1876b).
*** Herzogobryum Grolle, Rev. Bryol. Lichénol. 32 (3/4): 160, 1963 [1964] (Grolle
1963d). Nom. nov. pro Chondrophyllum Herzog, Rev. Bryol. Lichénol. 21 (1/2):
46, 1952 (Herzog 1952f).
*** Herzogobryum atrocapillum (Hook.f. et Taylor) Grolle, Österr. Bot. Z. 113 (2):
228, 1966 (Grolle 1966j). Bas.: Gymnomitrion atrocapillum Hook.f. et Taylor,
London J. Bot. 5: 258, 1846 (Taylor 1846a).
*** Herzogobryum iliforme R.M.Schust., Phytologia 45 (5): 422, 1980 (Schuster 1980b).
*** Herzogobryum molle Grolle, Österr. Bot. Z. 113 (2): 226, 1966 (Grolle 1966j).
*** Herzogobryum vermiculare (Schifn.) Grolle, J. Hattori Bot. Lab. 28: 103, 1965
(Grolle 1965e). Bas.: Gymnomitrion vermiculare Schifn., Leberm., Forschungsr.
Gazelle 4 (4): 2, 1890 (Schifner 1890).
** Kymatocalyx Herzog, Memoranda Soc. Fauna Fl. Fennica 25: 56, 1950 (Herzog
1950c).
*** Kymatocalyx africanus Váňa et M.Wigginton, Haussknechtia, Beih. 9: 158, 1999
(Gradstein and Váňa 1999).
*** Kymatocalyx dominicensis (Spruce) Váňa, Österr. Bot. Z. 118 (5): 575, 1970 (Váňa
1970b). Bas.: Jungermannia dominicensis Spruce, J. Linn. Soc., Bot. 30 (210): 363,
1895 (Gepp 1895b).
*** Kymatocalyx madagascariensis (Steph.) Gradst. et Váňa, Haussknechtia, Beih. 9:
164, 1999 (Gradstein and Váňa 1999). Bas.: Acrobolbus madagascariensis Steph.,
Bull. Herb. Boissier (sér. 2) 2 (5): 460 (179), 1902 (Stephani 1902f).
*** Kymatocalyx rhizomaticus (Herzog) Gradst. et Váňa, Haussknechtia, Beih. 9: 166,
1999 (Gradstein and Váňa 1999). Bas.: Stenorrhipis rhizomatica Herzog, Trans.
Brit. Bryol. Soc. 1 (4): 290, 1950 (Herzog 1950a).
World checklist of hornworts and liverworts
75
*** Lophonardia R.M.Schust., Phytologia 39 (4): 244, 1978 (Schuster 1978a).
*** Lophonardia jamesonii (Mont.) L.Söderstr. et Váňa, Phytotaxa 81 (1): 19, 2013
(Söderström et al. 2013a). Bas.: Jungermannia jamesonii Mont., Syll. Gen. Sp.
Crypt.: 60, 1856 (Montagne 1856b).
*** Lophonardia laxifolia (Mont.) L.Söderstr. et Váňa, Phytotaxa 81 (1): 20, 2013
(Söderström et al. 2013a). Bas.: Sarcocyphos laxifolius Mont., Ann. Sci. Nat. Bot.
(sér. 3) 4: 346, 1845 (Montagne 1845b).
*** Lophonardia tristaniana (S.W.Arnell) L.Söderstr. et Váňa, Phytotaxa 81 (1): 20,
2013 (Söderström et al. 2013a). Bas.: Jungermannia tristaniana S.W.Arnell, Results Norweg. Sci. Exped. Tristan da Cunha 42: 12, 1958 (Arnell 1958b).
*** Nothogymnomitrion R.M.Schust., J. Hattori Bot. Lab. 80: 43, 1996 (Schuster 1996a).
*** Nothogymnomitrion erosum (Carrington et Pearson) R.M.Schust., J. Hattori Bot.
Lab. 80: 44, 1996 (Schuster 1996a). Bas.: Cesius erosus Carrington et Pearson, Pap.
& Proc. Roy. Soc. Tasmania 1887: 8, 1888 (Carrington and Pearson 1888b).
*** Obtusifolium S.W.Arnell, Ill. Moss Fl. Fennosc. Hep.: 309, 1956 (Arnell 1956b).
*** Obtusifolium obtusum (Lindb.) S.W.Arnell, Ill. Moss Fl. Fennosc. Hep.: 133, 1956
(Arnell 1956b). Bas.: Jungermannia obtusa Lindb., Musci Scand.: 7, 1879 (Lindberg 1879).
*** Oleolophozia L.Söderstr., De Roo et Hedd., Phytotaxa 3: 50, 2010 (Söderström
et al. 2010b).
*** Oleolophozia perssonii (H.Buch et S.W.Arnell) L.Söderstr., De Roo et Hedd., Phytotaxa 3: 51, 2010 (Söderström et al. 2010b). Bas.: Lophozia perssonii H.Buch et
S.W.Arnell, Bot. Not. 97: 382, 1944 (Buch 1944).
** Phycolepidozia R.M.Schust., Bull. Torrey Bot. Club 93 (6): 438, 1966 [1967]
(Schuster 1966f).
** subg. Metaphycolepidozia Gradst., J.-P.Frahm et U.Schwarz, Taxon 63 (3):
506, 2014 (Gradstein et al. 2014a).
** Phycolepidozia indica Gradst., J.-P.Frahm et U.Schwarz, Taxon 63 (3): 499, 2014
(Gradstein et al. 2014a).
** subg. Phycolepidozia
*** Phycolepidozia exigua R.M.Schust., Bull. Torrey Bot. Club 93 (6): 440, 1966
[1967] (Schuster 1966f).
** Protolophozia (R.M.Schust.) Schljakov, Novosti Sist. Nizš. Rast. 16: 204, 1979
(Shliakov 1979). Bas.: Lophozia subg. Protolophozia R.M.Schust., Nova Hedwigia
15: 472, 1968 (Schuster 1968b).
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Protolophozia androgyna R.M.Schust. ex Váňa et L.Söderstr., Phytotaxa 76 (3): 51,
2013 (Váňa et al. 2013l). Based on: Lophozia androgyna R.M.Schust., Beih. Nova
Hedwigia 119: 266, 2002 (Schuster 2002b), nom. inval.
** Protolophozia autoica (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 76 (3): 51,
2013 (Váňa et al. 2013l). Bas.: Lophozia autoica R.M.Schust., Nova Hedwigia 15:
479, 1968 (Schuster 1968b).
** Protolophozia crispata (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 76 (3): 51,
2013 (Váňa et al. 2013l). Bas.: Lophozia crispata R.M.Schust., Nova Hedwigia 15:
474, 1968 (Schuster 1968b).
*** Protolophozia druceae (Grolle et E.A.Hodgs.) Váňa et L.Söderstr., Phytotaxa 76
(3): 51, 2013 (Váňa et al. 2013l). Bas.: Lophozia druceae Grolle et E.A.Hodgs., J.
Roy. Soc. New Zealand 2 (1): 112, 1972 (Hodgson 1972).
*** Protolophozia elongata (Steph.) Schljakov, Novosti Sist. Nizš. Rast. 16: 204, 1979
(Shliakov 1979). Bas.: Lophozia elongata Steph., Bull. Herb. Boissier (sér. 2) 2 (1):
41 (141), 1902 (Stephani 1902c).
*** Protolophozia herzogiana (E.A.Hodgs. et Grolle) Váňa et L.Söderstr., Phytotaxa 76
(3): 51, 2013 (Váňa et al. 2013l). Bas.: Lophozia herzogiana E.A.Hodgs. et Grolle,
Rev. Bryol. Lichénol. 31 (3/4): 152, 1962 [1963] (Grolle 1962c).
*** Protolophozia lancistipa (Grolle) Váňa et L.Söderstr., Phytotaxa 76 (3): 51, 2013
(Váňa et al. 2013l). Bas.: Andrewsianthus lancistipus Grolle, Marion Prince Edw.
Is: 230, 1971 (Grolle 1971d).
*** Protolophozia leucorhiza (Mitt.) Váňa et L.Söderstr., Phytotaxa 76 (3): 51, 2013
(Váňa et al. 2013l). Bas.: Jungermannia leucorhiza Mitt., J. Linn. Soc., Bot. 15
(82): 68, 1876 (Mitten 1876a).
*** Protolophozia longilora (Herzog) L.Söderstr. et Váňa, Phytotaxa 76 (3): 51, 2013
(Váňa et al. 2013l). Bas.: Orthocaulis longilorus Herzog, Rev. Bryol. Lichénol. 23
(1/2): 32, 1954 (Herzog 1954).
** Protolophozia monoica (E.A.Hodgs.) Váňa et L.Söderstr., Phytotaxa 76 (3): 52,
2013 (Váňa et al. 2013l). Bas.: Metahygrobiella monoica E.A.Hodgs., Trans. Roy.
Soc. New Zealand, Bot. 3 (4): 76, 1965 (Hodgson 1965).
*** Protolophozia multicuspidata (Hook.f. et Taylor) Váňa et L.Söderstr., Phytotaxa
76 (3): 52, 2013 (Váňa et al. 2013l). Bas.: Jungermannia multicuspidata Hook.f. et
Taylor, London J. Bot. 3: 375, 1844 (Hooker and Taylor 1844a).
** Protolophozia nivicola (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 76 (3): 52,
2013 (Váňa et al. 2013l). Bas.: Lophozia nivicola R.M.Schust., Nova Hedwigia 15:
477, 1968 (Schuster 1968b).
*** Protolophozia perssoniana (H.A.Mill.) Váňa et L.Söderstr., Phytotaxa 76 (3): 52,
2013 (Váňa et al. 2013l). Bas.: Lophozia perssoniana H.A.Mill., Ark. Bot. (n.ser.)
5 (2): 508, 1963 (Miller 1963).
** Protolophozia subalpina (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 76 (3): 52,
2013 (Váňa et al. 2013l). Bas.: Lophozia autoica var. subalpina R.M.Schust., Nova
Hedwigia 15: 482, 1968 (Schuster 1968b).
World checklist of hornworts and liverworts
77
*** Protolophozia tasmanica (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 76 (3): 52,
2013 (Váňa et al. 2013l). Bas.: Lophozia tasmanica R.M.Schust., Nova Hedwigia
15: 484, 1968 (Schuster 1968b).
*** Protolophozia verruculosa (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 76 (3): 52,
2013 (Váňa et al. 2013l). Bas.: Lophozia verruculosa R.M.Schust., Phytologia 39
(4): 242, 1978 (Schuster 1978a).
***
Lophoziaceae Cavers
by L. Söderström and J. Váňa
Lophoziaceae was broadly deined by de Roo et al. (2007) excluding Anastrophyllaceae and
Jamesonielloideae and also moving several traditionally included genera to other families.
Vilnet et al. (2010) further reined the family. Nomenclatural and taxonomic notes can also
be found in Vilnet et al. (2007b, 2008), Söderström et al. (2013c) and Váňa et al. (2013m).
he placement of Gerhildiella and Pseudocephaloziella in the family is provisional.
*** Andrewsianthus R.M.Schust., Rev. Bryol. Lichénol. 30 (1/2): 66, 1961 (Schuster
1961a).
** subg. Andrewsianthus
*** Andrewsianthus aberrans (Nees et Mont.) Grolle, Trans. Brit. Bryol. Soc. 4 (3):
440, 1963 (Grolle 1963a). Bas.: Jungermannia aberrans Nees et Mont., Ann. Sci.
Nat. Bot. (sér. 2) 19: 250, 1843 (Montagne 1843).
*** Andrewsianthus bidens (Mitt. ex Steph.) R.M.Schust., Nova Hedwigia 15: 492,
1968 (Schuster 1968b). Bas.: Lophozia bidens Mitt. ex Steph., Bull. Herb. Boissier
(sér. 2) 2 (1): 41 (141), 1902 (Stephani 1902c).
*** Andrewsianthus bilobus (Mitt.) Grolle, Trans. Brit. Bryol. Soc. 4 (3): 437, 1963
(Grolle 1963a). Bas.: Gymnanthe biloba Mitt., J. Proc. Linn. Soc., Bot. 7 (27):
166, 1863 (Mitten 1863).
*** Andrewsianthus carinatus Grolle, Marion Prince Edw. Is: 229, 1971 (Grolle 1971d).
*** Andrewsianthus cavifolius Grolle et Váňa, J. Hattori Bot. Lab. 38: 640, 1974 (Váňa
1974b).
*** Andrewsianthus chimbuensis R.M.Schust., Nova Hedwigia 15: 491, 1968 (Schuster
1968b).
*** Andrewsianthus kinabaluensis N.Kitag., J. Hattori Bot. Lab. 33: 205, 1970 (Kitagawa
1970).
*** Andrewsianthus koponenii Váňa et Piippo, Ann. Bot. Fenn. 26 (3): 284, 1989
(Váňa and Piippo 1989).
*** Andrewsianthus mizutanii N.Kitag., J. Hattori Bot. Lab. 32: 307, 1969 (Kitagawa
1969a).
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Andrewsianthus papillosus N.Kitag., J. Hattori Bot. Lab. 33: 207, 1970 (Kitagawa
1970).
*** Andrewsianthus perigonialis (Hook.f. et Taylor) R.M.Schust., Beih. Nova Hedwigia 119: 336, 2002 (Schuster 2002b). Bas.: Jungermannia perigonialis Hook.f. et
Taylor, London J. Bot. 3: 368, 1844 (Hooker and Taylor 1844a).
*** Andrewsianthus puniceus (Nees) R.M.Schust. ex Grolle, Trans. Brit. Bryol. Soc.
4 (3): 439, 1963 (Grolle 1963a). Bas.: Jungermannia punicea Nees, Enum. Pl.
Crypt. Javae: 32, 1830 (Nees 1830).
*** Andrewsianthus recurvifolius (Nees) R.M.Schust., Hepat. Anthocerotae N. Amer.
2: 710, 1969 (Schuster 1969b). Bas.: Jungermannia recurvifolia Nees, Enum. Pl.
Crypt. Javae: 32, 1830 (Nees 1830).
*** Andrewsianthus sundaicus (Schifn.) R.M.Schust., Hepat. Anthocerotae N. Amer.
2: 710, 1969 (Schuster 1969b). Bas.: Anastrophyllum sundaicum Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 67: 202, 1898 (Schifner 1898a).
*** Andrewsianthus zantenii Váňa, J. Hattori Bot. Lab. 38: 645, 1974 (Váňa 1974b).
** subg. Cephalolobus (R.M.Schust.) R.M.Schust., Beih. Nova Hedwigia 119:
326, 2002 (Schuster 2002b). Bas.: Cephalolobus R.M.Schust., Rev. Bryol. Lichénol. 34 (1/2): 244, 1966 (Schuster 1966b).
*** Andrewsianthus hodgsoniae (R.M.Schust.) R.M.Schust. ex J.J.Engel, Novon 17 (3):
311, 2007 (Engel 2007). Bas.: Cephalolobus hodgsoniae R.M.Schust., Rev. Bryol.
Lichénol. 34 (1/2): 254, 1966 (Schuster 1966b).
*** Andrewsianthus marionensis (S.W.Arnell) Grolle, Marion Prince Edw. Is: 232,
1971 (Grolle 1971d). Bas.: Lophozia marionensis S.W.Arnell, Svensk Bot. Tidskr.
47 (3): 421, 1953 (Arnell 1953c).
*** Andrewsianthus scabrellus (C.Massal.) R.M.Schust. ex J.J.Engel, Novon 17 (3):
311, 2007 (Engel 2007). Bas.: Cephalozia scabrella C.Massal., Nuovo Giorn. Bot.
Ital. 17 (3): 233, 1885 (Massalongo 1885).
** Andrewsianthus sphenoloboides (R.M.Schust.) R.M.Schust. ex J.J.Engel, Novon 17
(3): 311, 2007 (Engel 2007). Bas.: Cephalolobus sphenoloboides R.M.Schust., Rev.
Bryol. Lichénol. 34 (1/2): 251, 1966 (Schuster 1966b).
** Gerhildiella Grolle, Rev. Bryol. Lichénol. 34 (1/2): 187, 1966 (Grolle 1966a).
*** Gerhildiella rossneriana Grolle, Rev. Bryol. Lichénol. 34 (1/2): 187, 1966 (Grolle
1966a).
*** Heterogemma (Jørg.) Konstant. et Vilnet, Arctoa 18: 67, 2009 [2010] (Konstantinova and Vilnet 2009). Bas.: Lophozia sect. Heterogemma Jørg., Bergens Mus.
Skr. (n.ser.) 16: 146, 1934 (Jørgensen 1934).
*** Heterogemma capitata (Hook.) Konstant. et Vilnet, Arctoa 18: 67, 2009 [2010]
(Konstantinova and Vilnet 2009). Bas.: Jungermannia capitata Hook., Brit. Jungermann.: tab. 80, 1816 (Hooker 1816a).
World checklist of hornworts and liverworts
79
*** Heterogemma laxa (Lindb.) Konstant. et Vilnet, Arctoa 18: 67, 2009 [2010]
(Konstantinova and Vilnet 2009). Bas.: Jungermannia laxa Lindb., Acta Soc. Sci.
Fenn. 10: 529, 1875 (Lindberg 1875).
*** Heterogemma patagonica (Herzog et Grolle) L.Söderstr. et Váňa, Phytotaxa 97 (2):
29, 2013 (Söderström et al. 2013c). Bas.: Lophozia patagonica Herzog et Grolle,
Rev. Bryol. Lichénol. 28 (3/4): 343, 1959 [1960] (Grolle 1959a).
*** Lophozia (Dumort.) Dumort., Recueil Observ. Jungerm.: 17, 1835 (Dumortier
1835). Bas.: Jungermannia sect. Lophozia Dumort., Syll. Jungerm. Europ.: 53,
1831 (Dumortier 1831). 49
*** Lophozia ascendens (Warnst.) R.M.Schust., Bryologist 55 (3): 180, 1952 (Schuster
1952). Bas.: Sphenolobus ascendens Warnst., Hedwigia 57 (1/2): 63, 1916 (Warnstorf 1916).
* Lophozia austrosibirica Bakalin, Ann. Bot. Fenn. 40 (1): 49, 2003 (Bakalin 2003).
** Lophozia ciliata Damsh., L.Söderstr. et H.Weibull, Lindbergia 25 (1): 3, 2000
(Söderström et al. 2000).
*** Lophozia guttulata (Lindb. et Arnell) A.Evans, Proc. Wash. Acad. Sci. 2: 302, 1900
(Evans 1900c). Bas.: Jungermannia guttulata Lindb. et Arnell, Kongl. Svenska Vetensk.-Akad. Handl. (n.ser.) 23 (5): 51, 1889 (Lindberg and Arnell 1889).
* Lophozia jamaicensis (Nees) Steph., Bull. Herb. Boissier (sér. 2) 1 (2): 128 (155),
1901 (Stephani 1901h). Bas.: Jungermannia jamaicensis Nees, Syn. Hepat. 1: 105,
1844 (Gottsche et al. 1844). 50
** Lophozia lacerata N.Kitag., Hikobia 3 (3): 172, 1963 (Kitagawa 1963a).
** Lophozia lantratoviae Bakalin, Ann. Bot. Fenn. 40 (1): 47, 2003 (Bakalin 2003).
** Lophozia murmanica Kaal., Rep. Second Norweg. Arctic Exped. 11: 34, 1906
(Bryhn 1906).
** Lophozia paciica Bakalin, Bryologist 114 (2): 302, 2011 (Bakalin 2011).
* Lophozia pallida (Steph.) Grolle, J. Jap. Bot. 39 (6): 174, 1964 (Grolle 1964a).
Bas.: Anastrophyllum pallidum Steph., Bull. Herb. Boissier (sér. 2) 1 (10): 1131
(114), 1901 (Stephani 1901c). 51
** Lophozia savicziae Schljakov, Novosti Sist. Nizš. Rast. 10: 299, 1973 (Shliakov 1973).
* Lophozia schusterana Schljakov, Novosti Sist. Nizš. Rast. 12: 320, 1975 (Shliakov 1975).
*** Lophozia silvicola H.Buch, Beret. 18 Skand. Naturforskarmøde: 228, 1929 (Buch
1929).
*** Lophozia silvicoloides N.Kitag., J. Hattori Bot. Lab. 28: 276, 1965 (Kitagawa 1965).
49 Lophozia has many nomenclatural and taxonomic problems. his treatment follows Söderström et al.
(2013c).
50 Lophozia jamaicensis is probably not a Lophozia, but the type specimen is missing in Nees’s herbarium
(Söderström et al. 2010a).
51 Lophozia pallida may be conspeciic with Lophozia guttulata (Bakalin 2003).
80
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*
Lophozia subapiculata R.M.Schust. et Damsh., Meddel. Grønland 199 (1): 104,
1974 (Schuster and Damsholt 1974). 52
** Lophozia udarii S.Srivast., S.C.Srivast. et K.K.Rawat, Nelumbo 55: 130, 2013
(Srivastava et al. 2013). 53
*** Lophozia ventricosa (Dicks.) Dumort., Recueil Observ. Jungerm.: 17, 1835 (Dumortier 1835). Bas.: Jungermannia ventricosa Dicks., Fasc. Pl. Crypt. Brit. 2: 14,
1790 (Dickson 1790). 54
* Lophozia wenzelii (Nees) Steph., Bull. Herb. Boissier (sér. 2) 2 (1): 35 (135), 1902
(Stephani 1902c). Bas.: Jungermannia wenzelii Nees, Naturgesch. Eur. Leberm. 2:
58, 1836 (Nees 1836). 55
*
Excluded from the genus
Lophozia serpens (Dumort.) Dumort., Recueil Observ. Jungerm.: 17, 1835 (Dumortier 1835). Bas.: Jungermannia serpens Dumort., Syll. Jungerm. Europ.: 56,
1831 (Dumortier 1831). 56
** Lophoziopsis Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010] (Konstantinova
and Vilnet 2009).
*** Lophoziopsis excisa (Dicks.) Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010] (Konstantinova and Vilnet 2009). Bas.: Jungermannia excisa Dicks., Fasc. Pl. Crypt.
Brit. 3: 11, 1793 (Dickson 1793).
** Lophoziopsis excisa var. elegans (R.M.Schust.) Konstant. et Vilnet, Arctoa 18: 66,
2009 [2010] (Konstantinova and Vilnet 2009). Bas.: Lophozia excisa var. elegans
R.M.Schust., Hepat. Anthocerotae N. Amer. 2: 522, 1969 (Schuster 1969b).
** Lophoziopsis excisa var. infuscata (R.M.Schust. et Damsh.) Konstant. et Vilnet,
Arctoa 18: 66, 2009 [2010] (Konstantinova and Vilnet 2009). Bas.: Lophozia excisa var. infuscata R.M.Schust. et Damsh., Meddel. Grønland 199 (1): 94, 1974
(Schuster and Damsholt 1974).
** Lophoziopsis excisa var. succulenta (R.M.Schust. et Damsh.) Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010] (Konstantinova and Vilnet 2009). Bas.: Lophozia excisa var.
52 Lophozia subapiculata may be a form of Lophozia guttulata, but Bakalin (2005) treated it as conspeciic with Lophozia ventricosa.
53 Lophozia udarii has recently been described using the generic concept of Lophozia in the wide sense
(as e.g. Schuster 1969b). We do not know where to refer the species. It is possibly a Schistochilopsis
species close to Schistochilopsis incisa.
54 Lophozia ventricosa is very variable and many segregates at various subspeciic levels have been described. he species complex also includes at least Lophozia austrosibirica, Lophozia schusteriana and
Lophozia wenzelii.
55 Lophozia wenzelii is closely related to Lophozia ventricosa according to De Roo et al. (2007) and
Vilnet et al. (2010) based on molecular and morphological evidence. It is variable and many varieties
are described. he variability and relationships to related species needs further study.
56 Lophozia serpens is probably conspeciic with Schistochilopsis incisa.
World checklist of hornworts and liverworts
***
**
***
**
***
**
*
**
81
succulenta R.M.Schust. et Damsh., Meddel. Grønland 199 (1): 96, 1974 (Schuster
and Damsholt 1974).
Lophoziopsis longidens (Lindb.) Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010]
(Konstantinova and Vilnet 2009). Bas.: Jungermannia longidens Lindb., Helsingf.
Dagbl. 1876 (323, 26 Nov.): 2, 1876 (Lindberg 1876a).
Lophoziopsis longidens subsp. arctica (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa
97 (2): 28, 2013 (Söderström et al. 2013c). Bas.: Lophozia longidens subsp. arctica
R.M.Schust., Hepat. Anthocerotae N. Amer. 2: 539, 1969 (Schuster 1969b). 57
Lophoziopsis pellucida (R.M.Schust.) Konstant. et Vilnet, Arctoa 18: 67, 2009
[2010] (Konstantinova and Vilnet 2009). Bas.: Lophozia pellucida R.M.Schust.,
Canad. J. Bot. 39 (4): 978, 1961 (Schuster 1961b).
Lophoziopsis pellucida var. minor (R.M.Schust.) L.Söderstr. et Váňa, Phytotaxa
97 (2): 28, 2013 (Söderström et al. 2013c). Bas.: Lophozia pellucida var. minor
R.M.Schust., Canad. J. Bot. 39 (4): 984, 1961 (Schuster 1961b).
Lophoziopsis polaris (R.M.Schust.) Konstant. et Vilnet, Arctoa 18: 67, 2009
[2010] (Konstantinova and Vilnet 2009). Bas.: Lophozia alpestris subsp. polaris
R.M.Schust., Hepat. Anthocerotae N. Amer. 2: 614, 1969 (Schuster 1969b).
Lophoziopsis polaris var. sphagnorum (R.M.Schust.) Konstant. et Vilnet, Arctoa
18: 67, 2009 [2010] (Konstantinova and Vilnet 2009). Bas.: Lophozia alpestris f.
sphagnorum R.M.Schust., Hepat. Anthocerotae N. Amer. 2: 619, 1969 (Schuster
1969b).
Lophoziopsis propagulifera (Gottsche) Konstant. et Vilnet, Arctoa 18: 67, 2009
[2010] (Konstantinova and Vilnet 2009). Bas.: Jungermannia propagulifera
Gottsche, Int. Polarforsch., Deutsch. Exped. 2: 451, 1890 (Gottsche 1890). 58
Lophoziopsis rubrigemma (R.M.Schust.) Konstant. et Vilnet, Arctoa 18: 67, 2009
[2010] (Konstantinova and Vilnet 2009). Bas.: Lophozia rubrigemma R.M.Schust.,
Hepat. Anthocerotae N. Amer. 2: 621, 1969 (Schuster 1969b).
** Pseudocephaloziella R.M.Schust., Phytologia 39 (4): 243, 1978 (Schuster 1978a).
*** Pseudocephaloziella epiphytica R.M.Schust., Phytologia 39 (4): 243, 1978 (Schuster
1978a).
*** Trilophozia (R.M.Schust.) Bakalin, Monogr. Lophozia: 34, 2005 (Bakalin
2005). Bas.: Tritomaria subg. Trilophozia R.M.Schust., Amer. Midl. Naturalist 49
(2): 382, 1953 (Schuster 1953).
*** Trilophozia quinquedentata (Huds.) Bakalin, Monogr. Lophozia: 34, 2005 (Bakalin 2005). Bas.: Jungermannia quinquedentata Huds., Fl. Angl. (Hudson): 433,
1762 (Hudson 1762).
57 Lophoziopsis longidens var. arctica was treated as a form of Lophoziopsis pellucida by Bakalin (2005).
58 Lophoziopsis propagulifera belongs to the Lophoziopsis excisa species complex. It is probably only a
modiication (cf. discussion by Váňa and Engel 2013).
82
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Trilophozia quinquedentata var. asymmetrica (Horik.) L.Söderstr. et Váňa, Phytotaxa
97 (2): 32, 2013 (Söderström et al. 2013c). Bas.: Lophozia asymmetrica Horik., J. Sci.
Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 153, 1934 (Horikawa 1934).
*** Tritomaria Schifn. ex Loeske, Hedwigia 49 (1/2): 13, 1909 (Loeske 1909).
Based on: Tritomaria Schifn., Ber. Naturwiss.-Med. Vereins Innsbruck 31
[Beilage]: 12, 1908 (Schifner 1908a).
*** Tritomaria exsecta (Schmidel) Schifn. ex Loeske, Hedwigia 49 (1/2): 13, 1909
(Loeske 1909). Bas.: Jungermannia exsecta Schmidel, Syst. Samml. Crypt. Gew. 2:
5, 1797 (Schrader 1797), nom. conserv.
** Tritomaria exsecta subsp. novaezelandiae J.J.Engel, Bryologist 109 (1): 61, 2006
(Engel 2006b).
*** Tritomaria exsectiformis (Breidl.) Schifn. ex Loeske, Hedwigia 49 (1/2): 13, 1909
(Loeske 1909). Bas.: Jungermannia exsectiformis Breidl., Mitt. Naturwiss. Vereins
Steiermark 30: 321, 1894 (Breidler 1894).
** Tritomaria exsectiformis subsp. arctica R.M.Schust., Hepat. Anthocerotae N.
Amer. 2: 661, 1969 (Schuster 1969b).
** Tritomaria exsectiformis subsp. camerunensis S.W.Arnell ex Váňa, Phytotaxa 167
(2): 216, 2014 (Váňa et al. 2014b). Based on: Tritomaria camerunensis S.W.Arnell,
Svensk Bot. Tidskr. 52 (1): 64, 1958 (Arnell 1958a), nom. inval.
*** Tritomaria ferruginea (Grolle) Váňa, Phytotaxa 81 (1): 24, 2013 (Váňa et al.
2013m). Bas.: Andrewsianthus ferrugineus Grolle, Khumbu Himal 1 (4): 275,
1966 (Grolle 1966k).
* Tritomaria koreana Bakalin, S.S.Choi et B.Y.Sun, Arctoa 18: 163, 2009 [2010]
(Bakalin et al. 2009a).
* Tritomaria mexicana Bakalin, Arctoa 17: 162, 2008 [2009] (Bakalin 2008b).
*** Tritomaria scitula (Taylor) Jørg., Bergens Mus. Aarbok 1919/20 (7): 3, 1922 (Jørgensen 1922). Bas.: Jungermannia scitula Taylor, London J. Bot. 5: 274, 1846
(Taylor 1846a).
***
Scapaniaceae Mig.
by J. Váňa
Scapaniaceae, in its classical concept, was recognized as monophyletic and nested
within Lophoziaceae s. lat. (cf. Davis 2004, Schill et al. 2004, Yatsentyuk et al. 2004,
He-Nygrén et al. 2006). More recent molecular studies classiied Scapaniaceae as a
sister clade to Lophoziaceae, and the genus Schistochilopsis, a segregate of the genus
Lophozia s. lat., more closely related to Scapaniaceae than to Lophoziaceae (cf. de Roo
et al. 2007, Vilnet et al. 2007). he genus Scapania was recently studied by morphological methods by Potemkin (2002), by molecular methods by Vilnet et al. (2007)
and especially Heinrichs et al. (2012a). Some taxonomic and nomenclatural notes
World checklist of hornworts and liverworts
83
can be found in Váňa et al. (2012a, 2013j, 2015) and Konstantinova et al. (2013a).
he placement of Pseudotritomaria, Saccobasis and Schistochilopsis in the family follows
Vilnet et al. (2010).
*** Diplophyllum (Dumort.) Dumort., Recueil Observ. Jungerm.: 15, 1835 (Dumortier 1835) nom. conserv. Bas.: Jungermannia sect. Diplophyllum Dumort.,
Syll. Jungerm. Europ.: 44, 1831 (Dumortier 1831).
*** subg. Austrodiplophyllum R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 11
(1): 18, 1968 (Schuster 1968a).
** Diplophyllum recurvifolium C.Massal., Atti Reale Ist. Veneto Sci. Lett. Arti 87 (2):
221, 1928 (Massalongo 1928).
*** Diplophyllum squarrosum Steph., Sp. Hepat. (Stephani) 4: 116, 1910 (Stephani
1910b).
*** Diplophyllum verrucosum R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 11 (1):
19, 1968 (Schuster 1968a).
*** subg. Diplophyllum
*** sect. Diplophyllum
*** Diplophyllum albicans (L.) Dumort., Recueil Observ. Jungerm.: 16, 1835 (Dumortier
1835). Bas.: Jungermannia albicans L., Sp. Pl. 1: 1133, 1753 (Linnaeus 1753).
*** sect. Protodiplophyllum (R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 76
(3): 29, 2013 (Váňa et al. 2013j). Bas.: Diplophyllum subg. Protodiplophyllum
R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 192, 1974 (Schuster 1974).
*** Diplophyllum africanum S.W.Arnell, Ark. Bot. (n.ser.) 3 (16): 527, 1956 (Arnell
1956e).
*** Diplophyllum andicola R.M.Schust., Phytologia 39 (4): 248, 1978 (Schuster
1978a).
*** Diplophyllum andrewsii A.Evans, Bryologist 25 (2): 28, 1922 (Evans 1922).
* Diplophyllum androgynum J.J.Engel et G.L.Merr., J. Hattori Bot. Lab. 84: 277,
1998 (Engel and Smith Merrill 1998).
* Diplophyllum angustifolium J.J.Engel et G.L.Merr., J. Hattori Bot. Lab. 84: 262,
1998 (Engel and Smith Merrill 1998).
*** Diplophyllum apiculatum (A.Evans) Steph., Sp. Hepat. (Stephani) 4: 110, 1910
(Stephani 1910b). Bas.: Diplophylleia apiculata A.Evans, Bot. Gaz. 34 (5): 372,
1902 (Evans 1902b).
** Diplophyllum dioicum R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 11 (1):
20, 1968 (Schuster 1968a).
** Diplophyllum exiguum Steph., Sp. Hepat. (Stephani) 6: 500, 1924 (Stephani 1924).
* Diplophyllum gemmiparum J.J.Engel et G.L.Merr., J. Hattori Bot. Lab. 84: 255,
1998 (Engel and Smith Merrill 1998).
84
*
***
*
***
***
**
***
***
**
***
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Diplophyllum incrassatum J.J.Engel et G.L.Merr., J. Hattori Bot. Lab. 84: 265,
1998 (Engel and Smith Merrill 1998).
Diplophyllum nanum Herzog, Memoranda Soc. Fauna Fl. Fennica 26: 48, 1950
[1951] (Herzog 1950b).
Diplophyllum novum J.J.Engel et G.L.Merr., J. Hattori Bot. Lab. 84: 274, 1998
(Engel and Smith Merrill 1998).
Diplophyllum obtusatum (R.M.Schust.) R.M.Schust., Hepat. Anthocerotae N.
Amer. 3: 215, 1974 (Schuster 1974). Bas.: Diplophyllum apiculatum var. obtusatum R.M.Schust., Amer. Midl. Naturalist 49 (2): 432, 1953 (Schuster 1953).
Diplophyllum obtusifolium (Hook.) Dumort., Recueil Observ. Jungerm.: 16, 1835
(Dumortier 1835). Bas.: Jungermannia obtusifolia Hook., Brit. Jungermann.: tab.
26, 1812 (Hooker 1812). 59
Diplophyllum obtusifolium subsp. domesticum (Gottsche) Váňa, Phytotaxa 76 (3):
29, 2013 (Váňa et al. 2013j). Bas.: Jungermannia domestica Gottsche, Linnaea 28
(5): 548, 1856 [1857] (Gottsche 1856).
Diplophyllum serrulatum (Müll.Frib.) Steph., Sp. Hepat. (Stephani) 4: 112, 1910
(Stephani 1910b). Bas.: Diplophylleia serrulata Müll.Frib., Bull. Herb. Boissier
(sér. 2) 3 (1): 34, 1903 (Müller 1903).
Diplophyllum taxifolium (Wahlenb.) Dumort., Recueil Observ. Jungerm.: 16,
1835 (Dumortier 1835). Bas.: Jungermannia taxifolia Wahlenb., Fl. Lapp. (Wahlenberg): 389, 1812 (Wahlenberg 1812).
Diplophyllum taxifolium var. mucronatum R.M.Schust., Hepat. Anthocerotae N.
Amer. 3: 203, 1974 (Schuster 1974).
Diplophyllum trollii Grolle, Khumbu Himal 1 (4): 273, 1966 (Grolle 1966k).
*** Douinia (C.E.O.Jensen) H.Buch, Scapan. N.-Eur. Sib.: 13, 1928 (Buch 1928).
Bas.: Diplophylleia subg. Douinia C.E.O.Jensen, Danmarks mosser: 145, 1915
(Jensen 1915).
*** Douinia imbricata (M.Howe) Konstant. et Vilnet, Phytotaxa 76 (3): 31, 2013
(Konstantinova et al. 2013a). Bas.: Scapania imbricata M.Howe, Bull. New York
Bot. Gard. 2 (6): 104, 1901 (Howe 1901b).
*** Douinia ovata (Dicks.) H.Buch, Scapan. N.-Eur. Sib.: 14, 1928 (Buch 1928). Bas.:
Jungermannia ovata Dicks., Fasc. Pl. Crypt. Brit. 3: 11, 1793 (Dickson 1793).
*** Douinia plicata (Lindb.) Konstant. et Vilnet, Phytotaxa 76 (3): 31, 2013 (Konstantinova et al. 2013a). Bas.: Diplophyllum plicatum Lindb., Acta Soc. Sci. Fenn.
10: 235, 1872 [1873] (Lindberg 1872b).
** Pseudotritomaria Konstant. et Vilnet, Arctoa 18: 66, 2009 [2010] (Konstantinova and Vilnet 2009).
59 Diplophyllum obtusifolium is a species complex also including Diplophyllum androgynum, Diplophyllum
angustifolium, Diplophyllum gemmiparum, Diplophyllum incrassatum and Diplophyllum novum.
World checklist of hornworts and liverworts
85
*** Pseudotritomaria heterophylla (R.M.Schust.) Konstant. et Vilnet, Arctoa 18: 66,
2009 [2010] (Konstantinova and Vilnet 2009). Bas.: Tritomaria heterophylla
R.M.Schust., Canad. J. Bot. 36 (2): 272, 1958 (Schuster 1958b).
** Saccobasis H.Buch, Memoranda Soc. Fauna Fl. Fennica 8: 291, 1932 [1933]
(Buch 1932).
*** Saccobasis polita (Nees) H.Buch, Memoranda Soc. Fauna Fl. Fennica 8: 292, 1932
[1933] (Buch 1932). Bas.: Jungermannia polita Nees, Naturgesch. Eur. Leberm. 2:
145, 1836 (Nees 1836).
** Saccobasis polymorpha (R.M.Schust.) Schljakov, Novosti Sist. Nizš. Rast. 16: 205,
1979 (Shliakov 1979). Bas.: Tritomaria polita subsp. polymorpha R.M.Schust.,
Hepat. Anthocerotae N. Amer. 2: 700, 1969 (Schuster 1969b).
*** Scapania (Dumort.) Dumort., Recueil Observ. Jungerm.: 14, 1835 (Dumortier
1835) nom. conserv. Bas.: Radula sect. Scapania Dumort., Syll. Jungerm. Europ.:
38, 1831 (Dumortier 1831).
** subg. Ascapania Grolle, Khumbu Himal 1 (4): 268, 1966 (Grolle 1966k).
*** Scapania contorta Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 101, 1860 [1861] (Mitten
1860c).
** subg. Gracilidae (H.Buch) Váňa, Hentschel, Joch.Müll. et Heinrichs, PhytoKeys 10: 15, 2012 (Váňa et al. 2012a). Bas.: Scapania sect. Gracilidae H.Buch,
Scapan. N.-Eur. Sib.: 106, 1928 (Buch 1928).
*** Scapania ampliata Steph., Bull. Herb. Boissier 5 (2): 106, 1897 (Stephani 1897b).
** Scapania ampliata subsp. queenslandica M.L.Hicks, J. Hattori Bot. Lab. 69: 130,
1991 (Hicks 1991).
*** Scapania bolanderi Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 218, 1869 (Austin
1869).
*** Scapania gracilis Lindb., Morgonbladet (Helsinki) 1873 (286, 9 Dec): 2, 1873
(Lindberg 1873a).
** Scapania macroparaphyllia T.Cao, C.Gao et J.Sun, Acta Phytotax. Sin. 42 (2):
180, 2004 (Cao et al. 2004).
*** Scapania maxima Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 223,
1934 (Horikawa 1934).
*** Scapania nipponica (Amakawa et S.Hatt.) Amakawa, J. Hattori Bot. Lab. 30:
319, 1967 (Amakawa 1967a). Bas.: Scapania bolanderi var. nipponica Amakawa et
S.Hatt., J. Hattori Bot. Lab. 14: 83, 1955 (Amakawa and Hattori 1955).
** Scapania paraphyllia T.Cao, C.Gao, J.Sun et B.R.Zuo, Acta Phytotax. Sin. 45 (3):
311, 2007 (Zuo et al. 2007b).
*** Scapania subnimbosa Steph., Sp. Hepat. (Stephani) 4: 150, 1910 (Stephani 1910b).
86
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** subg. Macroscapania R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 248, 1974
(Schuster 1974).
* Scapania geppii Steph., Hedwigia 44 (1): 14, 1904 (Stephani 1904i).
*** Scapania portoricensis Hampe et Gottsche, Linnaea 25 (3): 342, 1852 [1853]
(Hampe and Gottsche 1852).
** Scapania portoricensis var. boliviensis (Steph.) Herzog, Ann. Bryol. 1: 110, 1928
(Herzog 1928). Bas.: Scapania boliviensis Steph., Biblioth. Bot. 87 (2): 231, 1916
(Stephani 1916a).
** Scapania portoricensis var. organensis (Herzog) Herzog, Ann. Bryol. 1: 110, 1928
(Herzog 1928). Bas.: Scapania organensis Herzog, Repert. Spec. Nov. Regni Veg.
21 (1/7): 27, 1925 (Herzog 1925a).
** Scapania portoricensis var. roraimensis Warnst., Hedwigia 63 (2): 109, 1921 (Warnstorf
1921).
** subg. Plicaticalyx Müll.Frib., Bull. Herb. Boissier (sér. 2) 3 (1): 36, 1903 (Müller
1903).
** sect. Planifoliae (Müll.Frib.) Potemkin, J. Hattori Bot. Lab. 85: 56, 1998
(Potemkin 1998). Bas.: Scapania Gruppe Planifoliae Müll.Frib., Nova Acta Acad.
Caes. Leop.-Carol. German. Nat. Cur. 83: 286, 1905 (Müller 1905).
*** Scapania davidii Potemkin, Ann. Bot. Fenn. 38 (2): 83, 2001 (Potemkin 2001).
** Scapania ferruginaeoides T.Cao, C.Gao et J.Sun, Guihaia 24 (1): 23, 2004 (Sun et
al. 2004).
** Scapania gaochii X.Fu ex T.Cao, Phytotaxa 97 (2): 26, 2013 (Cao et al. 2013).
Based on: Scapania gaochii X.Fu ex T.Cao, Acta Bot. Yunnan. 25 (5): 541, 2003
(Cao et al. 2003), nom. inval.
*** Scapania harae Amakawa, J. Hattori Bot. Lab. 27: 5, 1964 (Amakawa 1964a).
*** Scapania nimbosa Taylor, Nov. Stirp. Pug. 8: 6, 1844 (Lehmann 1844).
*** Scapania ornithopodioides (With.) Waddell, Moss Exch. Club Cat. Hepat.: 4, 1897
(Waddell 1897). Bas.: Jungermannia ornithopodioides With., Bot. arr. veg. Gr. Brit.
2: 695, 1776 (Withering 1776).
*** Scapania rotundifolia W.E.Nicholson, Symb. Sin. 5: 31, 1930 (Nicholson et al. 1930).
*** Scapania secunda Steph., Mém. Soc. Nat. Sci. Nat. Math. Cherbourg 29: 226,
1894 (Stephani 1894b).
*** Scapania zhukovae Potemkin, Arctoa 9: 121, 2000 (Potemkin 2000b).
** sect. Plicaticalyx (Müll.Frib.) Potemkin, Ann. Bot. Fenn. 39 (4): 326, 2002
(Potemkin 2002). Bas.: Scapania subg. Plicaticalyx Müll.Frib., Bull. Herb. Boissier
(sér. 2) 3 (1): 36, 1903 (Müller 1903).
*** Scapania ciliatospinosa Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 222,
1934 (Horikawa 1934).
World checklist of hornworts and liverworts
87
*** Scapania ferruginea (Lehm. et Lindenb.) Lehm. et Lindenb., Syn. Hepat. 1: 72,
1844 (Gottsche et al. 1844). Bas.: Jungermannia ferruginea Lehm. et Lindenb.,
Nov. Stirp. Pug. 4: 20, 1832 (Lehmann 1832).
*** Scapania hians Steph. ex Müll.Frib., Nova Acta Acad. Caes. Leop.-Carol. German.
Nat. Cur. 83: 223, 1905 (Müller 1905).
** Scapania hians subsp. salishensis J.D.Godfrey et G.Godfrey, Bryologist 81 (3): 362,
1978 (Godfrey and Godfrey 1978).
*** Scapania orientalis Steph. ex Müll.Frib., Bull. Herb. Boissier (sér. 2) 1 (6): 606,
1901 (Müller 1901a).
*** Scapania pseudocontorta Potemkin, Arctoa 9: 115, 2000 (Potemkin 2000b).
*** Scapania sinikkae Potemkin, Ann. Bot. Fenn. 38 (2): 85, 2001 (Potemkin 2001).
*** Scapania spiniloba Potemkin, Arctoa 9: 117, 2000 (Potemkin 2000b).
** subg. Pseudomacrodiplophyllum Váňa, Hentschel, Joch.Müll. et Heinrichs,
PhytoKeys 10: 15, 2012 (Váňa et al. 2012a).
*** Scapania microdonta (Mitt.) Müll.Frib., Nova Acta Acad. Caes. Leop.-Carol.
German. Nat. Cur. 83: 262, 1905 (Müller 1905). Bas.: Martinellius microdontus
Mitt., Trans. Linn. Soc. London, Bot. 3 (3): 196, 1891 (Mitten 1891).
*** subg. Scapania
** sect. Aequilobae (Müll.Frib.) H.Buch, Scapan. N.-Eur. Sib.: 110, 1928 (Buch
1928). Bas.: Scapania Gruppe Aequilobae Müll.Frib., Nova Acta Acad. Caes.
Leop.-Carol. German. Nat. Cur. 83: 219, 1905 (Müller 1905).
*** Scapania aequiloba (Schwägr.) Dumort., Recueil Observ. Jungerm.: 14, 1835
(Dumortier 1835). Bas.: Jungermannia aequiloba Schwägr., Hist. Musc. Hepat.
Prodr.: 24, 1814 (Schwägrichen 1814).
*** Scapania aspera M.Bernet et Bernet, Cat. hép. Suisse: 42, 1888 (Bernet 1888).
** sect. Americanae Váňa, Hentschel, Joch.Müll. et Heinrichs, PhytoKeys 10: 15,
2012 (Váňa et al. 2012a).
*** Scapania americana Müll.Frib., Bull. Herb. Boissier (sér. 2) 3 (1): 44, 1903 (Müller
1903).
** sect. Apiculatae H.Buch, Scapan. N.-Eur. Sib.: 53, 1928 (Buch 1928).
*** Scapania apiculata Spruce, Hep. Pyr. Exsic.: no. 15, 1847 (Spruce 1847).
*** Scapania carinthiaca J.B.Jack ex Lindb., Rev. Bryol. 7 (4): 77, 1880 (Lindberg
1880a).
*** Scapania carinthiaca var. massalongi Müll.Frib., Bull. Herb. Boissier (sér. 2) 1 (6):
598, 1901 (Müller 1901a).
*** Scapania umbrosa (Schrad.) Dumort., Recueil Observ. Jungerm.: 14, 1835 (Dumortier 1835). Bas.: Jungermannia umbrosa Schrad., Syst. Samml. Crypt. Gew. 2:
5, 1797 (Schrader 1797).
88
**
***
***
***
***
***
***
***
***
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
sect. Ciliatae Grolle, Khumbu Himal 1 (4): 272, 1966 (Grolle 1966k).
Scapania bhutanensis Amakawa, Fl. E. Himalaya 2: 230, 1971 (Hattori 1971a).
Scapania ciliata Sande Lac., Prolus. l. jap.: 209, 1867 (Sande Lacoste 1867).
Scapania ciliata subsp. hawaiica (Müll.Frib.) Potemkin, Ann. Bot. Fenn. 39 (4):
321, 2002 (Potemkin 2002). Bas.: Scapania hawaiica Müll.Frib., Nova Acta Acad.
Caes. Leop.-Carol. German. Nat. Cur. 83: 160, 1905 (Müller 1905).
Scapania hirosakiensis Steph. ex Müll.Frib., Nova Acta Acad. Caes. Leop.-Carol.
German. Nat. Cur. 83: 120, 1905 (Müller 1905).
Scapania hollandiae W.S.Hong, Bryologist 83 (1): 56, 1980 (Hong 1980).
Scapania koponenii Potemkin, Ann. Bot. Fenn. 37 (1): 41, 2000 (Potemkin
2000c).
Scapania lepida Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 101, 1860 [1861] (Mitten
1860c).
Scapania sandei Schifn. ex Müll.Frib., Bull. Herb. Boissier (sér. 2) 1 (6): 612,
1901 (Müller 1901a).
** sect. Compactae (Müll.Frib.) H.Buch, Scapan. N.-Eur. Sib.: 101, 1928 (Buch
1928). Bas.: Scapania Gruppe Compactae Müll.Frib., Nova Acta Acad. Caes.
Leop.-Carol. German. Nat. Cur. 83: 53, 1905 (Müller 1905).
*** Scapania compacta (Roth) Dumort., Recueil Observ. Jungerm.: 14, 1835 (Dumortier 1835). Bas.: Jungermannia compacta Roth, Tent. Fl. Germ. 3: 375, 1800
(Roth 1800).
*** Scapania kaurinii Ryan, Bot. Not. 42: 210, 1889 (Ryan 1889).
*** Scapania spitsbergensis (Lindb.) Müll.Frib., Bull. Herb. Boissier (sér. 2) 1 (6): 607,
1901 (Müller 1901a). Bas.: Martinellius spitsbergensis Lindb., Kongl. Svenska Vetensk.-Akad. Handl. (n.ser.) 23 (5): 31, 1889 (Lindberg and Arnell 1889).
** sect. Curtae (Müll.Frib.) H.Buch, Scapan. N.-Eur. Sib.: 55, 1928 (Buch 1928).
Bas.: Scapania Gruppe Curtae Müll.Frib., Nova Acta Acad. Caes. Leop.-Carol.
German. Nat. Cur. 83: 245, 1905 (Müller 1905).
*** Scapania curta (Mart.) Dumort., Recueil Observ. Jungerm.: 14, 1835 (Dumortier
1835). Bas.: Jungermannia curta Mart., Fl. crypt. erlang.: 148, 1817 (Martius 1817).
** Scapania curta var. grandiretis R.M.Schust., Hepat. Anthocerotae N. Amer. 3:
393, 1974 (Schuster 1974).
** Scapania curta var. isoloba R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 396,
1974 (Schuster 1974).
*** Scapania diplophylloides Amakawa et S.Hatt., J. Hattori Bot. Lab. 9: 59, 1953
(Amakawa and Hattori 1953).
*** Scapania esterhuyseniae S.W.Arnell, Bot. Not. 110 (1): 26, 1957 (Arnell 1957a).
*** Scapania fulfordiae W.S.Hong, Bryologist 83 (1): 46, 1980 (Hong 1980).
*** Scapania gamundiae R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 11 (1): 14,
1968 (Schuster 1968a).
World checklist of hornworts and liverworts
89
*** Scapania helvetica Gottsche, Hepat. Eur., Leberm. 42-44: no. 426, 1868 (Gottsche
and Rabenhorst 1868).
*** Scapania irrigua (Nees) Nees, Syn. Hepat. 1: 67, 1844 (Gottsche et al. 1844). Bas.:
Jungermannia irrigua Nees, Naturgesch. Eur. Leberm. 1: 193, 1833 (Nees 1833c).
** Scapania irrigua subsp. rufescens (Loeske) R.M.Schust., Hepat. Anthocerotae N.
Amer. 3: 471, 1974 (Schuster 1974). Bas.: Scapania irrigua f. rufescens Loeske,
Moosl. Harz.: 71, 1903 (Loeske 1903).
*** Scapania lingulata H.Buch, Meddel. Soc. Fauna Fl. Fenn. 42: 92, 1916 (Buch 1916).
** Scapania lingulata var. microphylla (Warnst.) R.M.Schust., Hepat. Anthocerotae
N. Amer. 3: 415, 1974 (Schuster 1974). Bas.: Scapania microphylla Warnst., Hedwigia 63 (2): 75, 1921 (Warnstorf 1921).
** Scapania magadanica S.S.Choi, Bakalin et B.Y.Sun, Bot. Paciica 1: 46, 2012
(Choi et al. 2012).
*** Scapania mucronata H.Buch, Meddel. Soc. Fauna Fl. Fenn. 42: 91, 1916 (Buch 1916).
*** Scapania obcordata (Berggr.) S.W.Arnell, Ark. Bot. (n.ser.) 4 (6): 117, 1959 (Arnell and Mårtensson 1959). Bas.: Sarcocyphos obcordatus Berggr., Kongl. Svenska
Vetensk.-Akad. Handl. (n.ser.) 13 (7): 96, 1875 (Berggren 1875).
** Scapania parvifolia Warnst., Hedwigia 63 (2): 78, 1921 (Warnstorf 1921).
*** Scapania praetervisa Meyl., Jahresber. Naturf. Ges. Graubündens (n.f.) 64: 364,
1926 (Meylan 1926).
*** Scapania scandica (Arnell et H.Buch) Macvicar, Stud. handb. Brit. hepat. (ed. 2):
394, 1926 (Macvicar 1926). Bas.: Martinellius scandicus Arnell et H.Buch, Bot.
Not. 74: 1, 1921 (Arnell and Buch 1921).
** Scapania scandica var. argutedentata H.Buch, Scapan. N.-Eur. Sib.: 75, 1928
(Buch 1928).
** Scapania scandica var. dimorpha R.M.Schust., Hepat. Anthocerotae N. Amer. 3:
453, 1974 (Schuster 1974).
** Scapania scandica var. grandiretis (Schljakov) Schljakov, Pečen. Mchi Sev. SSSR
4: 152, 1981 (Shliakov 1981). Bas.: Scapania parvifolia var. grandiretis Schljakov,
Novosti Sist. Nizš. Rast. 8: 332, 1971 (Shliakov 1971).
*** Scapania uliginosa (Lindenb.) Dumort., Recueil Observ. Jungerm.: 14, 1835 (Dumortier 1835). Bas.: Jungermannia undulata var. uliginosa Lindenb., Syn. hepat.
eur: 58, 1829 (Lindenberg 1829).
*** Scapania valdonii Váňa, Bedn.-Ochyra et Cykowska, Nova Hedwigia 89 (1/2):
126, 2009 (Váňa et al. 2009).
*** Scapania zemliae S.W.Arnell, Svensk Bot. Tidskr. 41: 215, 1947 (Arnell 1947).
** sect. Cuspiduligerae H.Buch, Scapan. N.-Eur. Sib.: 125, 1928 (Buch 1928).
*** Scapania cuspiduligera (Nees) Müll.Frib., Lebermoose 2 (22): 472, 1915 (Müller
1915a). Bas.: Jungermannia cuspiduligera Nees, Naturgesch. Eur. Leberm. 1: 180,
1833 (Nees 1833c).
** Scapania cuspiduligera var. diplophyllopsis R.M.Schust., Hepat. Anthocerotae N.
Amer. 3: 361, 1974 (Schuster 1974).
90
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** sect. Grolleoscapania Potemkin, Ann. Bot. Fenn. 39 (4): 329, 2002 (Potemkin 2002).
*** Scapania karl-muelleri Grolle, Khumbu Himal 1 (4): 270, 1966 (Grolle 1966k).
** sect. Hyperboreae Váňa, Hentschel, Joch.Müll. et Heinrichs, PhytoKeys 10:
16, 2012 (Váňa et al. 2012a).
*** Scapania hyperborea Jørg., Forh. Vidensk.-Selsk. Kristiania 1894 (8): 56, 1894
(Jørgensen 1894).
*** Scapania paludicola Loeske et Müll.Frib., Lebermoose 2 (21): 425, 1915 (Müller
1915b).
** Scapania paludicola var. viridigemma R.M.Schust., Bull. Natl. Mus. Canada 122:
20, 1950 [1951] (Schuster 1950).
*** Scapania tundrae (Arnell) H.Buch, Scapan. N.-Eur. Sib.: 99, 1928 (Buch 1928).
Bas.: Martinellius tundrae Arnell, Bot. Not. 74: 289, 1921 (Arnell 1921).
** sect. Kaalaasia (H.Buch) Jørg., Bergens Mus. Skr. (n.ser.) 16: 210, 1934 (Jørgensen 1934). Bas.: Scapania subg. Kaalaasia H.Buch, Scapan. N.-Eur. Sib.: 47,
1928 (Buch 1928).
*** Scapania calcicola (Arnell et J.Perss.) Ingham, Naturalist (Hull) 564: 11, 1904
(Ingham 1904). Bas.: Martinellius calcicola Arnell et J.Perss., Rev. Bryol. 30 (6):
97, 1903 (Arnell 1903).
*** Scapania gymnostomophila Kaal., Bot. Not. 49: 21, 1896 (Kaalaas 1896).
*** Scapania ligulifolia R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 306, 1974
(Schuster 1974).
*** Scapania pseudocalcicola R.M.Schust., Phytologia 63 (5): 327, 1987 (Schuster and
Damsholt 1987).
Scapania sect. Muelleria Potemkin, Ann. Bot. Fenn. 39 (4): 320, 2002 (Potemkin
2002).
*** Scapania himalayica Müll.Frib. ex Herzog, Ann. Bryol. 12: 81, 1939 (Herzog 1939b).
*** Scapania schljakovii Potemkin, Ann. Bot. Fenn. 38 (2): 87, 2001 (Potemkin 2001).
*
** sect. Nemorosae (Müll.Frib.) H.Buch, Scapan. N.-Eur. Sib.: 152, 1928 (Buch
1928). Bas.: Scapania Gruppe Nemorosae Müll.Frib., Nova Acta Acad. Caes.
Leop.-Carol. German. Nat. Cur. 83: 155, 1905 (Müller 1905).
*** Scapania brevicaulis Taylor, London J. Bot. 5: 272, 1846 (Taylor 1846a).
*** Scapania crassiretis Bryhn, Rev. Bryol. 19 (1): 7, 1892 (Bryhn 1892).
* Scapania degenii Schifn. ex Müll.Frib., Lebermoose 2 (22): 497, 1915 (Müller 1915a). 60
* Scapania glaucoviridis Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 221,
1934 (Horikawa 1934). 61
60 Scapania degenii is conspeciic with Scapania brevicaulis in Potemkin (1998), but recognized by
Konstantinova et al. (2009).
61 Scapania glaucoviridis was treated as conspeciic with Scapania parvitexta by Potemkin (2002), but
accepted by Zuo et al. (2007a).
World checklist of hornworts and liverworts
91
** Scapania grossidens Steph. ex Müll.Frib., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 83: 146, 1905 (Müller 1905).
*** Scapania hedbergii S.W.Arnell, Ark. Bot. (n.ser.) 3 (16): 556, 1956 (Arnell 1956e).
*** Scapania integerrima Steph., Sp. Hepat. (Stephani) 4: 148, 1910 (Stephani 1910b).
*** Scapania matveyevae Potemkin, Arctoa 9: 97, 2000 (Potemkin 2000a).
*** Scapania nemorea (L.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 160, 1963 [1964]
(Grolle 1963d). Bas.: Jungermannia nemorea L., Syst. Nat., ed. 10., 2: 1337, 1759
(Linnaeus 1759).
** Scapania parvidens Steph., Hedwigia 44 (1): 15, 1904 (Stephani 1904i). 62
*** Scapania parvitexta Steph., Bull. Herb. Boissier 5 (2): 107, 1897 (Stephani 1897b).
*** Scapania rigida Nees, Syn. Hepat. 1: 69, 1844 (Gottsche et al. 1844).
** sect. Scapania
*** Scapania gigantea Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 1: 15, 1931
(Horikawa 1931b).
** Scapania grandiloba Steph., Sp. Hepat. (Stephani) 6: 502, 1924 (Stephani 1924).
*** Scapania komagadakensis Amakawa, J. Hattori Bot. Lab. 31: 96, 1968 (Amakawa
1968a).
*** Scapania obscura (Arnell et C.E.O.Jensen) Schifn., Österr. Bot. Z. 58 (10): 377,
1908 (Schifner 1908b). Bas.: Martinellius obscurus Arnell et C.E.O.Jensen, Moose
Sarekgeb.: 91, 1907 (Arnell and Jensen 1907).
** Scapania paludosa (Müll.Frib.) Müll.Frib., Mitt. Bad. Bot. Vereins 4 (182/183):
287, 1902 (Müller 1902). Bas.: Scapania undulata var. paludosa Müll.Frib., Beih.
Bot. Centralbl. 10 (4/5): 220, 1901 (Müller 1901b).
*** Scapania ruidula Warnst., Hedwigia 63 (2): 94, 1921 (Warnstorf 1921).
*** Scapania serrulata R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 539, 1974
(Schuster 1974).
*** Scapania subalpina (Nees ex Lindenb.) Dumort., Recueil Observ. Jungerm.: 14,
1835 (Dumortier 1835). Bas.: Jungermannia subalpina Nees ex Lindenb., Syn.
hepat. eur: 55, 1829 (Lindenberg 1829).
** Scapania subalpina var. haynesiae Frye et L.Clark, Univ. Wash. Publ. Biol. 6 (4):
638, 1946 (Frye and Clark 1946).
** Scapania subalpina var. muddiae C.D.Bird et W.S.Hong, Bryologist 83 (1): 51,
1980 (Hong 1980).
*** Scapania undulata (L.) Dumort., Recueil Observ. Jungerm.: 14, 1835 (Dumortier
1835). Bas.: Jungermannia undulata L., Sp. Pl. 1: 1132, 1753 (Linnaeus 1753).
** sect. Scapaniella (H.Buch) Potemkin, J. Hattori Bot. Lab. 85: 43, 1998 (Potemkin
1998). Bas.: Scapaniella H.Buch, Scapan. N.-Eur. Sib.: 33, 1928 (Buch 1928).
62 Scapania parvidens was treated as conspeciic with Scapania parvitexta by Potemkin (2002), but accepted
by Zuo et al. (2007a) and Choi et al. (2012).
92
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Scapania glaucocephala (Taylor) Austin, Bull. Torrey Bot. Club 6 (16): 85, 1876
(Austin 1876c). Bas.: Jungermannia glaucocephala Taylor, London J. Bot. 5: 277,
1846 (Taylor 1846a).
*** Scapania glaucocephala var. saxicola (R.M.Schust.) Potemkin, Bryologist 102 (1):
36, 1999 (Potemkin 1999). Bas.: Scapania saxicola R.M.Schust., Amer. Midl.
Naturalist 49 (2): 448, 1953 (Schuster 1953).
* Scapania scapanioides (C.Massal.) Grolle, Feddes Repert. 87 (3/4): 235, 1976
(Grolle 1976a). Bas.: Jungermannia scapanioides C.Massal., Hepaticol. ven.: 64,
1879 (Massalongo 1879). 63
** sect. Simmonsiae (R.M.Schust.) Váňa, Hentschel, Joch.Müll. et Heinrichs,
PhytoKeys 10: 16, 2012 (Váňa et al. 2012a). Bas.: Scapania subsect. Simmonsiae
R.M.Schust., Hepat. Anthocerotae N. Amer. 3: 612, 1974 (Schuster 1974).
*** Scapania simmonsii Bryhn et Kaal., Rep. Second Norweg. Arctic Exped. 11: 51,
1906 (Bryhn 1906).
** sect. Sphaeriferae Konstant. et Potemkin, Ann. Bot. Fenn. 31 (2): 125, 1994
(Konstantinova and Potemkin 1994).
*** Scapania sphaerifera H.Buch et Tuom., Memoranda Soc. Fauna Fl. Fennica 11:
227, 1936 (Buch and Tuomikoski 1936).
** sect. Stephaniae Potemkin, J. Hattori Bot. Lab. 85: 57, 1998 (Potemkin 1998).
Based on: Scapania sect. Stephaniae Amakawa et S.Hatt., J. Hattori Bot. Lab. 12:
94, 1954 (Amakawa and Hattori 1954). nom. inval.
*** Scapania griithii Schifn., Österr. Bot. Z. 49 (6): 204, 1899 (Schifner 1899a).
*** Scapania javanica Gottsche, Natuurk. Tijdschr. Ned.-Indië 4: 575, 1853
(Gottsche 1853).
* Scapania javanica var. scabra Schifn., Arch. Hydrobiol., suppl. 21 (3/4): 397,
1955 (Schifner 1955).
*** Scapania ligulata Steph., Hedwigia 44 (1): 14, 1904 (Stephani 1904i).
** Scapania ligulata subsp. stephanii (Müll.Frib.) Potemkin, Piippo et T.J.Kop., Ann. Bot.
Fenn. 41 (6): 423, 2004 (Potemkin et al. 2004). Bas.: Scapania stephanii Müll.Frib.,
Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 83: 273, 1905 (Müller 1905).
** sect. Verrucosae Potemkin, J. Hattori Bot. Lab. 85: 54, 1998 (Potemkin 1998).
*** Scapania udarii S.C.Srivast. et A.Srivast., J. Indian Bot. Soc. 72: 237, 1993 (Srivastava and Srivastava 1993).
*** Scapania verrucosa Heeg, Rev. Bryol. 20 (5): 81, 1893 (Heeg 1893).
*** Schistochilopsis (N.Kitag.) Konstant., Arctoa 3: 125, 1994 (Konstantinova and
Vasil’ev 1994). Bas.: Lophozia subg. Schistochilopsis N.Kitag., J. Hattori Bot. Lab.
28: 289, 1965 (Kitagawa 1965).
63 Scapania scapanioides is possibly conspeciic with Scapania glaucocephala (Potemkin 2002).
World checklist of hornworts and liverworts
93
*** Schistochilopsis cornuta (Steph.) Konstant., Arctoa 3: 125, 1994 (Konstantinova
and Vasil’ev 1994). Bas.: Schistochila cornuta Steph., Sp. Hepat. (Stephani) 4: 84,
1909 (Stephani 1909d).
*** Schistochilopsis grandiretis (Lindb. ex Kaal.) Konstant., Arctoa 3: 125, 1994 (Konstantinova and Vasil’ev 1994). Bas.: Jungermannia grandiretis Lindb. ex Kaal., Nyt
Mag. Naturvidensk. 33 (4/5): 322, 1893 (Kaalaas 1893b).
** Schistochilopsis hyperarctica Konstant. et L.Söderstr., Phytotaxa 162 (4): 240,
2014 (Konstantinova et al. 2014b). Based on: Lophozia hyperarctica R.M.Schust.,
Canad. J. Bot. 39 (4): 967, 1961 (Schuster 1961b), nom. inval.
*** Schistochilopsis incisa (Schrad.) Konstant., Arctoa 3: 125, 1994 (Konstantinova
and Vasil’ev 1994). Bas.: Jungermannia incisa Schrad., Syst. Samml. Crypt. Gew.
2: 5, 1797 (Schrader 1797).
* Schistochilopsis nakanishii (Inoue) Konstant., Arctoa 3: 125, 1994 (Konstantinova
and Vasil’ev 1994). Bas.: Lophozia nakanishii Inoue, Bull. Natl. Sci. Mus. Tokyo
(n.ser.) 9 (1): 37, 1966 (Inoue 1966a). 64
** Schistochilopsis opacifolia (Culm. ex Meyl.) Konstant., Arctoa 3: 125, 1994 (Konstantinova and Vasil’ev 1994). Bas.: Lophozia opacifolia Culm. ex Meyl., Beitr.
Kryptogamenl. Schweiz 6 (4): 174, 1924 (Meylan 1924). 65
*** Schistochilopsis setosa (Mitt.) Konstant., Arctoa 3: 125, 1994 (Konstantinova and
Vasil’ev 1994). Bas.: Jungermannia setosa Mitt., J. Proc. Linn. Soc., Bot. 5 (18):
92, 1860 [1861] (Mitten 1860c).
Jungermanniineae R.M.Schust. ex Stotler et Crand.-Stotl.
***
Acrobolbaceae E.A.Hodgs.
by L. Briscoe and J.J. Engel
** Enigmella G.A.M.Scott et K.G.Beckm., J. Bryol. 17 (2): 297, 1992 (Beckmann
and Scott 1992).
** Enigmella thallina G.A.M.Scott et K.G.Beckm., J. Bryol. 17 (2): 297, 1992 (Beckmann and Scott 1992).
**
Acrobolboideae R.M.Schust. ex Briscoe
*** Acrobolbus Nees, Syn. Hepat. 1: 5, 1844 (Gottsche et al. 1844).
** Acrobolbus africanus (Pearson) Briscoe, Phytotaxa 202 (1): 59, 2015 (Briscoe et al.
2015). Bas.: Tylimanthus africanus Pearson, Forh. Vidensk.-Selsk. Kristiania 1887
(9): 14, 1887 (Pearson 1887b).
64 Schistochilopsis nakanishii is possibly conspeciic with Schistochilopsis incisa.
65 Schistochilopsis opacifolia is sometimes treated as a subspecies of Schistochilopsis incisa (e.g. Bisang 1991).
94
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Acrobolbus anisodontus (Hook.f. et Taylor) Briscoe, Phytotaxa 202 (1): 59, 2015
(Briscoe et al. 2015). Bas.: Jungermannia anisodonta Hook.f. et Taylor, London J.
Bot. 4: 79, 1845 (Hooker and Taylor 1845).
** Acrobolbus antillanus R.M.Schust., J. Hattori Bot. Lab. 90: 143, 2001 (Schuster 2001a).
** Acrobolbus azoricus (Grolle et Perss.) Briscoe, Phytotaxa 202 (1): 59, 2015 (Briscoe
et al. 2015). Bas.: Tylimanthus azoricus Grolle et Perss., Svensk Bot. Tidskr. 60 (1):
169, 1966 (Grolle and Persson 1966).
*** Acrobolbus caducifolius R.M.Schust., J. Hattori Bot. Lab. 90: 154, 2001 (Schuster
2001a).
*** Acrobolbus ciliatus (Mitt.) Schifn., Hepat. (Engl.-Prantl): 86, 1893 (Schifner
1893b). Bas.: Gymnanthe ciliata Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 100, 1860
[1861] (Mitten 1860c).
*** Acrobolbus cinerascens (Lehm. et Lindenb.) Bastow, Pap. & Proc. Roy. Soc. Tasmania 1887: 242, 1888 (Bastow 1888). Bas.: Jungermannia cinerascens Lehm. et
Lindenb., Nov. Stirp. Pug. 4: 46, 1832 (Lehmann 1832).
*** Acrobolbus concinnus (Mitt.) Steph., Trans. & Proc. New Zealand Inst. 24: 399,
1892 (Colenso 1892). Bas.: Gymnanthe concinna Mitt., Bot. antarct. voy. III (Fl.
Tasman. 2): 230, 1860 (Mitten 1860b).
** Acrobolbus cuneifolius (Steph.) Briscoe, Phytotaxa 202 (1): 59, 2015 (Briscoe et al.
2015). Bas.: Tylimanthus cuneifolius Steph., Bull. Herb. Boissier (sér. 2) 5 (12):
1138 (10), 1905 (Stephani 1905b).
*** Acrobolbus diversifolius R.M.Schust., J. Hattori Bot. Lab. 90: 150, 2001 (Schuster 2001a).
*** Acrobolbus epiphytus (Colenso) Briscoe, Phytotaxa 202 (1): 59, 2015 (Briscoe et al.
2015). Bas.: Marsupidium epiphytum Colenso, Trans. & Proc. New Zealand Inst.
21: 64, 1889 (Colenso 1889).
*** Acrobolbus lavicans (J.J.Engel et Grolle) Briscoe et J.J.Engel, Phytotaxa 202 (1):
59, 2015 (Briscoe et al. 2015). Bas.: Marsupidium lavicans J.J.Engel et Grolle, J.
Hattori Bot. Lab. 34: 438, 1971 (Engel and Grolle 1971).
*** Acrobolbus gradsteinii (Grolle) Briscoe, Phytotaxa 202 (1): 59, 2015 (Briscoe et al.
2015). Bas.: Marsupidium gradsteinii Grolle, J. Hattori Bot. Lab. 66: 337, 1989
(Grolle 1989a).
** Acrobolbus integrifolius (A.Evans) Briscoe, Phytotaxa 202 (1): 59, 2015 (Briscoe et
al. 2015). Bas.: Tylimanthus integrifolius A.Evans, Trans. Connecticut Acad. Arts
8 (15): 259, 1891 (Evans 1891).
*** Acrobolbus knightii (Mitt.) Briscoe, Phytotaxa 202 (1): 59, 2015 (Briscoe et al.
2015). Bas.: Marsupidium knightii Mitt., Handb. N. Zeal. l. 2: 753, 1867 (Hooker 1867).
** Acrobolbus kunkelii (Hässel et Solari) Briscoe et J.J.Engel, Phytotaxa 202 (1): 60,
2015 (Briscoe et al. 2015). Bas.: Tylimanthus kunkelii Hässel et Solari, Darwiniana
17: 574, 1972 (Hässel and Solari 1972).
*** Acrobolbus laxus (Lehm. et Lindenb.) Briscoe, Phytotaxa 202 (1): 60, 2015 (Briscoe et al. 2015). Bas.: Plagiochila laxa Lehm. et Lindenb., Sp. Hepat. (Lindenberg)
2-4: 68, 1840 (Lindenberg 1840).
World checklist of hornworts and liverworts
95
*** Acrobolbus limbatus (Steph.) Briscoe et J.J.Engel, Phytotaxa 202 (1): 60, 2015
(Briscoe et al. 2015). Bas.: Tylimanthus limbatus Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9): 25, 1911 (Stephani 1911b).
*** Acrobolbus lophocoleoides (Mitt.) Mitt., Handb. N. Zeal. l. 2: 753, 1867 (Hooker
1867). Bas.: Gymnanthe lophocoleoides Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2):
144, 1854 (Mitten 1854).
** Acrobolbus madeirensis (Grolle et Perss.) Briscoe, Phytotaxa 202 (1): 60, 2015
(Briscoe et al. 2015). Bas.: Tylimanthus madeirensis Grolle et Perss., Svensk Bot.
Tidskr. 60 (1): 166, 1966 (Grolle and Persson 1966).
** Acrobolbus mittenii Steph., Bull. Herb. Boissier (sér. 2) 2 (5): 460 (179), 1902
(Stephani 1902f).
*** Acrobolbus ochrophyllus (Hook.f. et Taylor) R.M.Schust., Rev. Bryol. Lichénol.
30 (1/2): 64, 1961 (Schuster 1961a). Bas.: Jungermannia ochrophylla Hook.f. et
Taylor, London J. Bot. 3: 368, 1844 (Hooker and Taylor 1844a).
*** Acrobolbus papillosus (J.J.Engel et Glenny) Briscoe, Phytotaxa 202 (1): 60, 2015
(Briscoe et al. 2015). Bas.: Marsupidium papillosum J.J.Engel et Glenny, Nova
Hedwigia 87 (3/4): 289, 2008 (Engel and Glenny 2008c).
*** Acrobolbus perpusillus (Colenso) Briscoe, Phytotaxa 202 (1): 60, 2015 (Briscoe
et al. 2015). Bas.: Tylimanthus perpusillus Colenso, Trans. & Proc. New Zealand
Inst. 19: 286, 1887 (Colenso 1887).
*** Acrobolbus perpusillus var. denticulatus (J.J.Engel et Glenny) Briscoe, Phytotaxa
202 (1): 60, 2015 (Briscoe et al. 2015). Bas.: Marsupidium perpusillum var. denticulatum J.J.Engel et Glenny, Nova Hedwigia 87 (3/4): 284, 2008 (Engel and
Glenny 2008c).
*** Acrobolbus plagiochiloides (J.J.Engel et Glenny) Briscoe, Phytotaxa 202 (1): 60,
2015 (Briscoe et al. 2015). Bas.: Marsupidium plagiochiloides J.J.Engel et Glenny,
Nova Hedwigia 87 (3/4): 284, 2008 (Engel and Glenny 2008c).
*** Acrobolbus pseudosaccatus (Grolle) Briscoe, Phytotaxa 202 (1): 60, 2015 (Briscoe et
al. 2015). Bas.: Tylimanthus pseudosaccatus Grolle, Nova Hedwigia 6 (3/4): 391,
1963 (Grolle 1963c).
*** Acrobolbus renifolius (Hässel et Solari) Briscoe et J.J.Engel, Phytotaxa 202 (1): 60,
2015 (Briscoe et al. 2015). Bas.: Tylimanthus renifolius Hässel et Solari, Darwiniana
17: 583, 1972 (Hässel and Solari 1972).
** Acrobolbus ruwenzorensis (S.W.Arnell) Briscoe, Phytotaxa 202 (1): 60, 2015 (Briscoe et al. 2015). Bas.: Tylimanthus ruwenzorensis S.W.Arnell, Ark. Bot. (n.ser.) 3
(16): 560, 1956 (Arnell 1956e).
*** Acrobolbus saccatus (Hook.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 423, 1877 (Trevisan 1877). Bas.: Jungermannia saccata Hook.,
Musci Exot. 1: tab. 16, 1818 (Hooker 1818).
*** Acrobolbus setulosus (Mitt.) Briscoe, Phytotaxa 202 (1): 60, 2015 (Briscoe et al.
2015). Bas.: Gymnanthe setulosa Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 144,
1854 (Mitten 1854).
*** Acrobolbus spinifolius R.M.Schust., J. Hattori Bot. Lab. 90: 137, 2001 (Schuster 2001a).
96
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Acrobolbus sumatranus (Schifn.) Briscoe, Phytotaxa 202 (1): 61, 2015 (Briscoe
et al. 2015). Bas.: Lophozia sumatrana Schifn., Denkschr. Kaiserl. Akad. Wiss.,
Math.-Naturwiss. Kl. 67: 203, 1898 (Schifner 1898a).
*** Acrobolbus surculosus (Nees) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 423, 1877 (Trevisan 1877). Bas.: Scapania surculosa Nees, Syn.
Hepat. 1: 62, 1844 (Gottsche et al. 1844).
*** Acrobolbus tenellus (Taylor) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 423, 1877 (Trevisan 1877). Bas.: Gymnanthe tenella Taylor, Nov.
Stirp. Pug. 8: 1, 1844 (Lehmann 1844).
*** Acrobolbus tenellus var. diversifolius (E.A.Hodgs.) Briscoe, Phytotaxa 202 (1): 61,
2015 (Briscoe et al. 2015). Bas.: Tylimanthus diversifolius E.A.Hodgs., Trans. Roy.
Soc. New Zealand 85 (4): 575, 1958 (Hodgson 1958).
*** Acrobolbus urvilleanus (Mont.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 423, 1877 (Trevisan 1877). Bas.: Plagiochila urvilleana Mont.,
Ann. Sci. Nat. Bot. (sér. 2) 19: 247, 1843 (Montagne 1843).
*** Acrobolbus viridis (Mitt.) Briscoe et J.J.Engel, Phytotaxa 202 (1): 61, 2015 (Briscoe et al. 2015). Bas.: Tylimanthus viridis Mitt., J. Linn. Soc., Bot. 15 (84): 197,
1876 (Mitten 1876b).
*** Acrobolbus wilsonii Nees, Syn. Hepat. 1: 5, 1844 (Gottsche et al. 1844).
** Acrobolbus wilsonii var. andinus Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:
522, 1885 (Spruce 1885).
*
**
Excluded from the genus
Acrobolbus bispinosus (J.B.Jack et Steph.) Steph., Bull. Herb. Boissier (sér. 2) 2 (5):
459 (178), 1902 (Stephani 1902f). Bas.: Tylimanthus bispinosus J.B.Jack et Steph.,
Hedwigia 31 (1): 26, 1892 (Jack and Stephani 1892). 66
Austrolophozioideae R.M.Schust. ex Crand.-Stotl., Váňa et Stotler
** Austrolophozia R.M.Schust., J. Hattori Bot. Lab. 26: 282, 1963 (Schuster 1963b).
** Austrolophozia andina R.M.Schust., Nova Hedwigia 15: 495, 1968 (Schuster 1968b).
*** Austrolophozia camensis (Steph.) Grolle ex Hässel et Solari, Revista Mus. Argent.
Ci. Nat., Bernardino Rivadavia Inst. Nac. Invest. Ci. Nat. Bot. 3 (6): 240, 1970
(Hässel and Solari 1970). Bas.: Tylimanthus camensis Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9): 24, 1911 (Stephani 1911b).
*** Austrolophozia paradoxa R.M.Schust., J. Hattori Bot. Lab. 26: 282, 1963 (Schuster
1963b).
66 Acrobolbus bispinosus is a Plagiochila species (Burghardt and Gradstein 2008).
World checklist of hornworts and liverworts
97
*** Goebelobryum Grolle, J. Hattori Bot. Lab. 25: 135, 1962 (Grolle 1962b).
*** Goebelobryum grossitextum (Steph.) Grolle, J. Hattori Bot. Lab. 25: 137, 1962
(Grolle 1962b). Bas.: Marsupidium grossitextum Steph., Sp. Hepat. (Stephani) 6:
446, 1924 (Stephani 1924).
*** Goebelobryum unguiculatum (Hook.f. et Taylor) Grolle, J. Hattori Bot. Lab. 25:
137, 1962 (Grolle 1962b). Bas.: Jungermannia unguiculata Hook.f. et Taylor,
London J. Bot. 5: 279, 1846 (Taylor 1846a).
*** Goebelobryum vermiculare J.J.Engel et Glenny, Nova Hedwigia 95 (3/4): 320,
2012 (Engel and Glenny 2012).
**
Lethocoleoideae Grolle
*** Lethocolea Mitt., Handb. N. Zeal. l. 2: 751, 1867 (Hooker 1867) nom. conserv.
** Lethocolea congesta (Lehm.) S.W.Arnell, Bot. Not. 108: 311, 1955 (Arnell 1955b).
Bas.: Jungermannia congesta Lehm., Linnaea 4: 365, 1829 (Lehmann 1829).
*** Lethocolea glossophylla (Spruce) Grolle, Bot. Mag. (Tokyo) 78 (921): 83, 1965
(Grolle 1965c). Bas.: Symphyomitra glossophylla Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 503, 1885 (Spruce 1885).
** Lethocolea indica G.Asthana et Maurya, Natl. Acad. Sci. Lett. 37 (6): 535, 2014
(Asthana and Maurya 2014).
*** Lethocolea javanica (Schifn.) Grolle, Bot. Mag. (Tokyo) 78 (921): 83, 1965 (Grolle 1965c). Bas.: Symphyomitra javanica Schifn., Denkschr. Kaiserl. Akad. Wiss.,
Math.-Naturwiss. Kl. 67: 193, 1898 (Schifner 1898a).
** Lethocolea naruto-toganensis Furuki, Bryologist 104 (2): 306, 2001 (Furuki 2001).
*** Lethocolea pansa (Taylor) G.A.M.Scott et K.G.Beckm., Symp. Biol. Hung. 35:
212, 1987 (Scott and Beckmann 1987). Bas.: Jungermannia pansa Taylor, London
J. Bot. 5: 275, 1846 (Taylor 1846a).
*** Lethocolea radicosa (Lehm. et Lindenb.) Grolle, Bot. Mag. (Tokyo) 78 (921): 83,
1965 (Grolle 1965c). Bas.: Jungermannia radicosa Lehm. et Lindenb., Nov. Stirp.
Pug. 6: 35, 1834 (Lehmann 1834).
* Lethocolea repens S.Winkl., Mitt. Inst. Colombo-Alemán Invest. Ci. 3: 67, 1969
(Winkler 1969).
**
Saccogynidioideae Crand.-Stotl., Váňa et Stotler
*** Saccogynidium Grolle, J. Hattori Bot. Lab. 23: 43, 1960 [1961] (Grolle 1960d).
** sect. Decurvum Grolle, J. Hattori Bot. Lab. 23: 59, 1960 (Grolle 1960d).
*** Saccogynidium decurvum (Mitt.) Grolle, J. Hattori Bot. Lab. 23: 59, 1960 [1961]
(Grolle 1960d). Bas.: Lophocolea decurva Mitt., Bot. antarct. voy. III (Fl. Tasman.
2): 227, 1860 (Mitten 1860b).
98
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** sect. Jugata Grolle, J. Hattori Bot. Lab. 23: 55, 1960 (Grolle 1960d).
** Saccogynidium rigidulum (Nees) Grolle, J. Hattori Bot. Lab. 23: 52, 1960 [1961]
(Grolle 1960d). Bas.: Jungermannia rigidula Nees, Enum. Pl. Crypt. Javae: 25,
1830 (Nees 1830).
** sect. Saccogynidium
*** Saccogynidium australe (Mitt.) Grolle, J. Hattori Bot. Lab. 23: 49, 1960 [1961]
(Grolle 1960d). Bas.: Saccogyna australis Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel.
2): 145, 1854 (Mitten 1854).
** Saccogynidium caldense (Ångstr.) Grolle, J. Hattori Bot. Lab. 23: 44, 1960 [1961]
(Grolle 1960d). Bas.: Chiloscyphus caldensis Ångstr., Öfvers. Kongl. Vetensk.-Akad.
Förh. 33 (7): 80, 1876 [1877] (Ångström 1876).
** Saccogynidium goebelii (Herzog) Grolle, J. Hattori Bot. Lab. 23: 52, 1960 [1961]
(Grolle 1960d). Bas.: Leioscyphus goebelii Herzog, Ann. Bryol. 5: 89, 1932 (Herzog
1932a).
** Saccogynidium muricellum (De Not.) Grolle, J. Hattori Bot. Lab. 36: 80, 1972
[1973] (Grolle and Schultze-Motel 1972). Bas.: Chiloscyphus muricellus De Not.,
Epat. Borneo: 24, 1874 (De Notaris 1874).
*** Saccogynidium vasculosum (Hook.f. et Taylor) Grolle, J. Hattori Bot. Lab. 23: 46,
1960 [1961] (Grolle 1960d). Bas.: Jungermannia vasculosa Hook.f. et Taylor, London J. Bot. 3: 461, 1844 (Hooker and Taylor 1844b).
Incertae sedis
** Saccogynidium chiloscyphoides R.M.Schust., J. Hattori Bot. Lab. 26: 272, 1963
(Schuster 1963b).
** Saccogynidium irregularospinum C.Gao, T.Cao et M.J.Lai, Bryologist 104 (1):
129, 2001 (Gao et al. 2001).
***
Antheliaceae R.M.Schust.
*** Anthelia (Dumort.) Dumort., Recueil Observ. Jungerm.: 18, 1835 (Dumortier
1835). Bas.: Jungermannia sect. Anthelia Dumort., Syll. Jungerm. Europ.: 63,
1831 (Dumortier 1831).
*** Anthelia julacea (L.) Dumort., Recueil Observ. Jungerm.: 18, 1835 (Dumortier
1835). Bas.: Jungermannia julacea L., Sp. Pl., ed. 2: 1601, 1763 (Linnaeus 1763).
** Anthelia juratzkana (Limpr.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 416, 1877 (Trevisan 1877). Bas.: Jungermannia juratzkana Limpr., Hedwigia 15 (2): 18, 1876 (Limpricht 1876).
World checklist of hornworts and liverworts
***
99
Arnelliaceae Nakai
Crandall-Stotler et al. (2009) placed Stephaniella and Stephaniellidium in Arnelliaceae
following de Roo et al. (2007). However, Váňa et al. (2012e) argued based on further
molecular evidence that Arnelliaceae should be retained as a monotypic family with a
single species.
*** Arnellia Lindb., Kongl. Svenska Vetensk.-Akad. Handl. (n.ser.) 23 (5): 35, 1889
(Lindberg and Arnell 1889).
*** Arnellia fennica (Gottsche) Lindb., Kongl. Svenska Vetensk.-Akad. Handl. (n.ser.)
23 (5): 35, 1889 (Lindberg and Arnell 1889). Bas.: Jungermannia fennica Gottsche,
Hepat. Eur., Leberm. 42-44: no. 418, 1868 (Gottsche and Rabenhorst 1868).
***
Balantiopsidaceae H.Buch
by J.J. Engel with contribution by J. Váňa (Neesioscyphus)
he placement of Pseudoisotachis in Balantiopsidaceae is only preliminary (cf. Váňa 2013).
*** Acroscyphella N.Kitag. et Grolle, Acta Phytotax. Geobot. 36 (1/3): 58, 1985
(Kitagawa and Grolle 1985). Nom. nov. pro Acroscyphus N.Kitag., Acta Phytotax.
Geobot. 35 (1/3): 1, 1984 (Kitagawa 1984).
*** Acroscyphella iwatsukii (N.Kitag.) N.Kitag. et Grolle, Acta Phytotax. Geobot. 36
(1/3): 58, 1985 (Kitagawa and Grolle 1985). Bas.: Acroscyphus iwatsukii N.Kitag.,
Acta Phytotax. Geobot. 35 (1/3): 3, 1984 (Kitagawa 1984).
*** Acroscyphella phoenicorhiza (Grolle) N.Kitag. et Grolle, Acta Phytotax. Geobot.
36 (1/3): 58, 1985 (Kitagawa and Grolle 1985). Bas.: Neesioscyphus phoenicorhizus
Grolle, Österr. Bot. Z. 111 (1): 27, 1964 (Grolle 1964e).
*** Acroscyphella tjiwideiensis (Sande Lac.) N.Kitag. et Grolle, Acta Phytotax. Geobot.
36 (1/3): 58, 1985 (Kitagawa and Grolle 1985). Bas.: Chiloscyphus tjiwideiensis
Sande Lac., Ned. Kruidk. Arch. 3: 418, 1854 [1855] (Sande Lacoste 1854).
** Pseudoisotachis Váňa, Polish Bot. J. 58 (1): 55, 2013 (Váňa 2013).
** Pseudoisotachis pocsii Váňa, Polish Bot. J. 58 (1): 55, 2013 (Váňa 2013).
***
Balantiopsidoideae J.J.Engel et Váňa
*** Balantiopsis Mitt., Handb. N. Zeal. l. 2: 751, 1867 (Hooker 1867).
*** Balantiopsis asymmetrica (Herzog) J.J.Engel, Nova Hedwigia 16: 93, 1968 (Engel
1968). Bas.: Balantiopsis latifolia var. asymmetrica Herzog, Rev. Bryol. Lichénol.
23 (1/2): 53, 1954 (Herzog 1954).
100
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Balantiopsis bisbiida (Steph.) Steph., Sp. Hepat. (Stephani) 4: 101, 1910 (Stephani 1910b). Bas.: Isotachis bisbiida Steph., Bih. Kongl. Svenska Vetensk.-Akad.
Handl. 26 (III, 17): 24, 1901 (Stephani 1901b).
*** Balantiopsis brasiliensis Steph., Sp. Hepat. (Stephani) 4: 104, 1910 (Stephani 1910b).
*** Balantiopsis cancellata (Nees) Steph., Sp. Hepat. (Stephani) 4: 103, 1910 (Stephani
1910b). Bas.: Ptilidium cancellatum Nees, Syn. Hepat. 2: 251, 1845 (Gottsche et
al. 1845a).
*** Balantiopsis ciliaris S.Hatt., J. Jap. Bot. 41 (5): 129, 1966 (Hattori 1966b).
** Balantiopsis ciliaris subsp. novoguineensis S.Hatt., J. Jap. Bot. 41 (5): 131, 1966
(Hattori 1966b).
*** Balantiopsis convexiuscula Berggr., New Zealand Hepat.: 44, 1898 (Berggren 1898).
*** Balantiopsis crocea Herzog, Beih. Bot. Centralbl. 60B (1/2): 12, 1939 (Herzog 1939c).
*** Balantiopsis diplophylla (Hook.f. et Taylor) Mitt., Handb. N. Zeal. l. 2: 753, 1867
(Hooker 1867). Bas.: Jungermannia diplophylla Hook.f. et Taylor, London J. Bot.
3: 377, 1844 (Hooker and Taylor 1844a).
** Balantiopsis diplophylla var. hockenii (Berggr.) J.J.Engel et G.L.Merr., Fieldiana,
Bot. (n.ser.) 37: 11, 1997 (Engel and Smith Merrill 1997). Bas.: Balantiopsis hockenii Berggr., New Zealand Hepat.: 46, 1898 (Berggren 1898).
*** Balantiopsis erinacea (Hook.f. et Taylor) Mitt., Handb. N. Zeal. l. 2: 753, 1867
(Hooker 1867). Bas.: Jungermannia erinacea Hook.f. et Taylor, London J. Bot. 3:
462, 1844 (Hooker and Taylor 1844b).
*** Balantiopsis lingulata R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 11 (1): 26,
1968 (Schuster 1968a).
*** Balantiopsis montana (Colenso) J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 37:
12, 1997 (Engel and Smith Merrill 1997). Bas.: Chiloscyphus montanus Colenso,
Trans. & Proc. New Zealand Inst. 21: 62, 1889 (Colenso 1889).
** Balantiopsis neocaledonica Pearson, J. Linn. Soc., Bot. 46 (305): 28, 1922 (Pearson
1922b).
** Balantiopsis paucidens Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26 (III,
17): 29, 1901 (Stephani 1901b).
*** Balantiopsis purpurata Mitt., Rep. Challenger, Bot. 1 (3, 1): 86, 1884 (Mitten 1884b).
*** Balantiopsis rosea Berggr., New Zealand Hepat.: 43, 1898 (Berggren 1898).
*** Balantiopsis splendens (Steph.) J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 37:
55, 1997 (Engel and Smith Merrill 1997). Bas.: Isotachis splendens Steph., Hedwigia 34 (2): 49, 1895 (Stephani 1895c).
*** Balantiopsis tumida Berggr., New Zealand Hepat.: 45, 1898 (Berggren 1898).
*** Balantiopsis verrucosa J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 37: 16, 1997
(Engel and Smith Merrill 1997).
World checklist of hornworts and liverworts
***
101
Isotachidoideae Grolle
*** Isotachis Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 148, 1854 (Mitten 1854).
*** Isotachis armata (Nees) Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 121, 1864
(Gottsche 1864). Bas.: Jungermannia armata Nees, Syn. Hepat. 1: 129, 1844
(Gottsche et al. 1844).
*** Isotachis aubertii (Schwägr.) Mitt., J. Linn. Soc., Bot. 22 (146): 322, 1886 (Mitten
1886b). Bas.: Jungermannia aubertii Schwägr., Hist. Musc. Hepat. Prodr.: 19,
1814 (Schwägrichen 1814).
* Isotachis boliviensis Gottsche, Sp. Hepat. (Stephani) 3: 670, 1909 (Stephani 1909a).
** Isotachis chinensis C.Gao, T.Cao et J.Sun, Bryologist 105 (4): 694, 2002 [2003]
(Gao et al. 2002).
** Isotachis erythrorhiza (Lehm. et Lindenb.) Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 338, 1885 (Spruce 1885). Bas.: Jungermannia erythrorhiza Lehm. et
Lindenb., Nov. Stirp. Pug. 4: 44, 1832 (Lehmann 1832).
*** Isotachis fragilis Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9): 67,
1911 (Stephani 1911b).
** Isotachis grandis Carrington et Pearson, Proc. Linn. Soc. New South Wales (ser. 2)
2 (4): 1041, 1888 (Carrington and Pearson 1888a).
*** Isotachis grossidens Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9):
69, 1911 (Stephani 1911b).
** Isotachis hastatistipula (Steph.) J.J.Engel, Phytotaxa 183 (4): 299, 2014 (Engel et
al. 2014). Bas.: Balantiopsis hastatistipula Steph., J. & Proc. Roy. Soc. New South
Wales 48 (1/2): 98, 1914 (Stephani and Watts 1914).
* Isotachis hians Steph., Sp. Hepat. (Stephani) 3: 665, 1909 (Stephani 1909a).
*** Isotachis humectata (Hook.f. et Taylor) Steph., Sp. Hepat. (Stephani) 3: 654, 1909
(Stephani 1909a). Bas.: Jungermannia humectata Hook.f. et Taylor, London J.
Bot. 3: 462, 1844 (Hooker and Taylor 1844b).
** Isotachis inlata Steph., Arch. Mus. Nac. Rio de Janeiro 13: 113, 1905 (Stephani 1905c).
*** Isotachis intortifolia (Hook.f. et Taylor) Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1:
121, 1864 (Gottsche 1864). Bas.: Jungermannia intortifolia Hook.f. et Taylor,
London J. Bot. 3: 374, 1844 (Hooker and Taylor 1844a).
** Isotachis japonica Steph., Sp. Hepat. (Stephani) 3: 652, 1909 (Stephani 1909a).
* Isotachis lacustris Herzog, Hedwigia 74 (2): 94, 1934 (Herzog 1934a).
*** Isotachis lopezii (R.M.Schust.) Gradst., Mem. New York Bot. Gard. 84: 66, 1999
(Gradstein 1999). Bas.: Ruizanthus lopezii R.M.Schust., Phytologia 39 (4): 241,
1978 (Schuster 1978a).
*** Isotachis lyallii Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 149, 1854 (Mitten 1854).
*** Isotachis minima Pearson, Univ. Calif. Publ. Bot. 10 (4): 322, 1923 (Pearson 1923).
*** Isotachis montana Colenso, Trans. & Proc. New Zealand Inst. 21: 68, 1889
(Colenso 1889).
102
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*** Isotachis multiceps (Lindenb. et Gottsche) Gottsche, Mexik. Leverm.: 105, 1863
(Gottsche 1863). Bas.: Jungermannia multiceps Lindenb. et Gottsche, Syn. Hepat.
5: 687, 1847 (Gottsche et al. 1847). 67
** Isotachis multiceps var. fendleri Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 125, 1864
(Gottsche 1864).
** Isotachis nigella Herzog, Memoranda Soc. Fauna Fl. Fennica 27: 107, 1952 (Herzog
1952c).
*** Isotachis obtusa Steph., Sp. Hepat. (Stephani) 6: 354, 1922 (Stephani 1922).
** Isotachis olivacea R.M.Schust., J. Hattori Bot. Lab. 83: 207, 1997 (Schuster and
Engel 1997).
*** Isotachis plicata J.J.Engel, J. Hattori Bot. Lab. 83: 210, 1997 (Schuster and Engel 1997).
** Isotachis pusilla Steph., Sp. Hepat. (Stephani) 3: 655, 1909 (Stephani 1909a).
** Isotachis riparia Rodway, Tasm. Bryoph.: 63, 1917 (Rodway 1917b).
*** Isotachis serrulata (Sw.) Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 121, 1864
(Gottsche 1864). Bas.: Jungermannia serrulata Sw., Prodr. (Swartz): 143, 1788
(Swartz 1788).
*** Isotachis spegazziniana C.Massal., Nuovo Giorn. Bot. Ital. 17 (3): 220, 1885 (Massalongo 1885).
* Isotachis sprucei Beauverd, Sp. Hepat. (Stephani) 6: 572, 1924 (Stephani 1924).
Nom. nov. pro Isotachis triida Steph., Sp. Hepat. (Stephani) 6: 356, 1922 (Stephani
1922), nom. illeg.
*** Isotachis stephanii E.S.Salmon, Rev. Bryol. 28 (4): 75, 1901 (Salmon 1901).
* Isotachis vexans Steph., Sp. Hepat. (Stephani) 3: 662, 1909 (Stephani 1909a).
*** Isotachis westlandica (E.A.Hodgs.) R.M.Schust., Nova Hedwigia 15: 455, 1968
(Schuster 1968b). Bas.: Rhizocaulia westlandica E.A.Hodgs., Trans. Roy. Soc.
New Zealand, Bot. 3 (4): 78, 1965 (Hodgson 1965).
*** Neesioscyphus Grolle, Österr. Bot. Z. 111 (1): 19, 1964 (Grolle 1964e).
*** Neesioscyphus allionii (Steph.) Grolle, Rev. Bryol. Lichénol. 34 (1/2): 185, 1966
(Grolle 1966c). Bas.: Isotachis allionii Steph., Sp. Hepat. (Stephani) 6: 350, 1922
(Stephani 1922).
*** Neesioscyphus argillaceus (Nees) Grolle, Österr. Bot. Z. 111 (1): 24, 1964 (Grolle 1964e).
Bas.: Jungermannia argillacea Nees, Fl. Bras. (Martius) 1 (1): 338, 1833 (Nees 1833a).
*** Neesioscyphus bicuspidatus (Steph.) Grolle, Rev. Bryol. Lichénol. 34 (1/2): 182,
1966 (Grolle 1966c). Bas.: Isotachis bicuspidata Steph., Symb. Antill. 2: 471, 1901
(Stephani 1901f).
*** Neesioscyphus carneus (Nees) Grolle, Österr. Bot. Z. 111 (1): 20, 1964 (Grolle 1964e).
Bas.: Jungermannia carnea Nees, Fl. Bras. (Martius) 1 (1): 337, 1833 (Nees 1833a).
*** Neesioscyphus homophyllus (Nees) Grolle, Österr. Bot. Z. 111 (2/3): 188, 1964
(Grolle 1964f). Bas.: Jungermannia homophylla Nees, Fl. Bras. (Martius) 1 (1):
336, 1833 (Nees 1833a).
67 Isotachis multiceps is sometimes treated as a Hypoisotachis, but recent molecular studies (Forrest et al.
2006, Cooper et al. 2012b) point towards inclusion in Isotachis.
World checklist of hornworts and liverworts
**
103
Ruizanthoideae R.M.Schust. ex J.J.Engel et G.L.Merr.
*** Ruizanthus R.M.Schust., Phytologia 39 (4): 240, 1978 (Schuster 1978a).
*** Ruizanthus venezuelanus R.M.Schust., Phytologia 39 (4): 240, 1978 (Schuster 1978a).
***
Blepharidophyllaceae R.M.Schust. ex J.J.Engel
by J.J. Engel
*** Blepharidophyllum Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 30 (5): 151,
1873 (Ångström 1873).
*** Blepharidophyllum densifolium (Hook.) Ångstr. ex C.Massal., Nuovo Giorn. Bot.
Ital. 17 (3): 208, 1885 (Massalongo 1885). Bas.: Jungermannia densifolia Hook.,
Musci Exot. 1: tab. 36, 1818 (Hooker 1818).
*** Blepharidophyllum vertebrale (Gottsche) Ångstr. ex C.Massal., Nuovo Giorn. Bot.
Ital. 17 (3): 208, 1885 (Massalongo 1885). Bas.: Scapania vertebralis Gottsche,
Syn. Hepat. 1: 72, 1844 (Gottsche et al. 1844).
*** Clandarium (Grolle) R.M.Schust., New Manual Bryol. 1: 541, 1983 [1984]
(Schuster 1983a). Bas.: Blepharidophyllum subg. Clandarium Grolle, J. Hattori
Bot. Lab. 28: 65, 1965 (Grolle 1965a).
*** Clandarium clandestinum (Mont.) R.M.Schust., New Manual Bryol. 1: 541, 1983
[1984] (Schuster 1983a). Bas.: Plagiochila clandestina Mont., Ann. Sci. Nat. Bot.
(sér. 2) 19: 247, 1843 (Montagne 1843).
*** Clandarium gottscheanum (Grolle) R.M.Schust., New Manual Bryol. 1: 541, 1983
[1984] (Schuster 1983a). Bas.: Blepharidophyllum gottscheanum Grolle, J. Hattori
Bot. Lab. 28: 69, 1965 (Grolle 1965a).
*** Clandarium xiphophyllum (Grolle) R.M.Schust., Phytologia 56 (2): 68, 1984
(Schuster 1984). Bas.: Blepharidophyllum xiphophyllum Grolle, J. Hattori Bot. Lab.
28: 65, 1965 (Grolle 1965a).
***
Calypogeiaceae Arnell
by M.A.M. Renner
Calypogeiaceae is shown to be monophyletic (Masuzaki et al. 2010b) and the classiication follows Schuster (2000a) and Masuzaki et al. (2010b).
*** Calypogeia Raddi, Jungermanniogr. Etrusca: 31, 1818 (Raddi 1818a) nom. conserv.
104
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** subg. Asperifoliae (Warnst.) R.M.Schust., Hepat. Anthocerotae N. Amer. 2:
115, 1969 (Schuster 1969b). Bas.: Calypogeia [unranked] Asperifoliae Warnst.,
Bryol. Z. 1 (7): 111, 1917 (Warnstorf 1917).
*** Calypogeia arguta Nees et Mont., Naturgesch. Eur. Leberm. 3: 24, 1838 (Nees 1838b).
** Calypogeia sullivantii Austin, Hepat. bor.-amer.: 19, 1873 (Austin 1873).
**
***
**
**
***
***
**
***
**
**
**
***
***
***
***
**
**
**
subg. Calypogeia
Calypogeia andicola Bischl., Candollea 18: 79, 1962 (Bischler 1962a).
Calypogeia annabonensis Steph., Sp. Hepat. (Stephani) 6: 447, 1924 (Stephani 1924).
Calypogeia azorica Bischl., Rev. Bryol. Lichénol. 37 (1): 116, 1970 (Bischler 1970).
Calypogeia azurea Stotler et Crotz, Taxon 32 (1): 74, 1983 (Stotler and Crotz 1983).
Calypogeia bidentula (F.Weber) Nees, Syn. Hepat. 2: 199, 1845 (Gottsche et al.
1845a). Bas.: Jungermannia bidentula F.Weber, Hist. Musc. Hepat. Prodr.: 38,
1815 (Weber 1815).
Calypogeia falcata Bischl., Candollea 18: 112, 1962 (Bischler 1962c).
Calypogeia issa (L.) Raddi, Jungermanniogr. Etrusca: 33, 1818 (Raddi 1818a).
Bas.: Mnium issum L., Sp. Pl. 1: 1114, 1753 (Linnaeus 1753), nom. conserv. 68
Calypogeia issa subsp. neogaea R.M.Schust., Hepat. Anthocerotae N. Amer. 2:
169, 1969 (Schuster 1969b).
Calypogeia goebelii (Schifn.) Steph., Bull. Herb. Boissier (sér. 2) 8 (9): 677 (409),
1908 (Stephani 1908d). Bas.: Kantius goebelii Schifn., Nova Acta Acad. Caes.
Leop.-Carol. German. Nat. Cur. 60 (2): 260, 1893 (Schifner 1893a).
Calypogeia goebelii var. siamensis N.Kitag., Beih. Nova Hedwigia 90: 165, 1988
(Kitagawa 1988).
Calypogeia grandistipula (Steph.) Steph., Bull. Herb. Boissier (sér. 2) 8 (9): 669
(401), 1908 (Stephani 1908d). Bas.: Kantius grandistipulus Steph., Hedwigia 34
(2): 52, 1895 (Stephani 1895c).
Calypogeia integristipula Steph., Bull. Herb. Boissier (sér. 2) 8 (9): 662 (394), 1908
(Stephani 1908d).
Calypogeia laxa Gottsche et Lindenb., Syn. Hepat. 5: 713, 1847 (Gottsche et al. 1847).
Calypogeia lechleri (Steph.) Steph., Bull. Herb. Boissier (sér. 2) 8 (9): 680 (412),
1908 (Stephani 1908d). Bas.: Kantius lechleri Steph., Hedwigia 34 (2): 53, 1895
(Stephani 1895c). 69
Calypogeia lechleri var. densifolia (Steph.) Bischl., Candollea 18: 101, 1962
(Bischler 1962c). Bas.: Kantius densifolius Steph., Hedwigia 34 (2): 52, 1895
(Stephani 1895c).
Calypogeia longifolia Steph., Sp. Hepat. (Stephani) 6: 449, 1924 (Stephani 1924).
Calypogeia lophocoleoides Steph., Bull. Herb. Boissier (sér. 2) 8 (9): 677 (409),
1908 (Stephani 1908d).
68 Calypogeia issa contains two well separated taxa (Buczkowska et al. 2011), but a formal taxonomic
treatment is not yet available.
69 Calypogeia lechleri was treated as conspeciic with Calypogeia leptoloma by Bischler (1962c) with hesitation
(type specimen not found).
World checklist of hornworts and liverworts
105
** Calypogeia mascarenensis Bischl., Rev. Bryol. Lichénol. 37 (1): 89, 1970 (Bischler
1970).
** Calypogeia microstipula (Steph.) Steph., Bull. Herb. Boissier (sér. 2) 8 (9): 670
(402), 1908 (Stephani 1908d). Bas.: Kantius microstipulus Steph., Hedwigia 34
(2): 53, 1895 (Stephani 1895c).
*** Calypogeia miquelii Mont. ex Gottsche, Lindenb. et Nees, Syn. Hepat. 2: 200,
1845 (Gottsche et al. 1845a).
*** Calypogeia muelleriana (Schifn.) Müll.Frib., Beih. Bot. Centralbl. 10 (4/5): 217,
1901 (Müller 1901b). Bas.: Kantius muellerianus Schifn., Sitzungsber. deutsch.
naturwiss.-med. Vereins Böhmen “Lotos” Prag 48: 342, 1900 (Schifner 1900d). 70
** Calypogeia muelleriana subsp. blomquistii R.M.Schust., Hepat. Anthocerotae N.
Amer. 2: 187, 1969 (Schuster 1969b).
*** Calypogeia neesiana (C.Massal. et Carestia) Müll.Frib., Verh. Bot. Vereins Prov.
Brandenburg 47: 320, 1905 (Loeske 1905). Bas.: Kantius trichomanis var. neesianus C.Massal. et Carestia, Nuovo Giorn. Bot. Ital. 12 (4): 351, 1880 (Massalongo and Carestia 1880).
** Calypogeia neesiana subsp. subalpina (Inoue) Inoue, Mem. Natl. Sci. Mus. (Tokyo) 4: 58, 1971 (Inoue 1971a). Bas.: Calypogeia subalpina Inoue, J. Jap. Bot. 37
(4): 103, 1962 (Inoue 1962b).
*** Calypogeia oblata Herzog, Svensk Bot. Tidskr. 51 (1): 189, 1957 (Herzog 1957a).
*** Calypogeia peruviana Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 9: 47, 1838
(Montagne 1838).
*** Calypogeia rhombifolia (Spruce) Steph., Bull. Herb. Boissier (sér. 2) 8 (9): 667
(399), 1908 (Stephani 1908d). Bas.: Kantius rhombifolius Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 413, 1885 (Spruce 1885).
** Calypogeia rhombifolia var. colombiana Bischl., Candollea 18: 104, 1962 (Bischler
1962c).
*** Calypogeia sphagnicola (Arnell et J.Perss.) Warnst. et Loeske, Verh. Bot. Vereins
Prov. Brandenburg 47: 320, 1905 (Loeske 1905). Bas.: Kantius sphagnicola Arnell
et J.Perss., Rev. Bryol. 29 (2): 26, 1902 (Arnell 1902). 71
*** Calypogeia subintegra (Gottsche, Lindenb. et Nees) Bischl., Candollea 18: 75,
1962 (Bischler 1962a). Bas.: Calypogeia peruviana β subintegra Gottsche, Lindenb.
et Nees, Syn. Hepat. 5: 712, 1847 (Gottsche et al. 1847).
** Calypogeia subintegra var. dussiana (Steph.) Bischl., Candollea 18: 77, 1962
(Bischler 1962a). Bas.: Calypogeia dussiana Steph., Bull. Herb. Boissier (sér. 2) 8
(9): 672 (404), 1908 (Stephani 1908d).
70 Calypogeia muelleriana contains two well separated taxa (Buczkowska 2010, Buczkowska and
Bączkiewicz 2011). One corresponds well to the type of the species while the name of the other is
unknown.
71 Calypogeia sphagnicola is a species complex including at least three genetically and morphologically
well separated taxa (Buczkowska et al. 2012b, 2012a). hey all need to be compared with types of
many existing names in the genus before the correct name can be assigned.
106
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Calypogeia suecica (Arnell et J.Perss.) Müll.Frib., Beih. Bot. Centralbl. 17 (2): 224,
1904 (Müller 1904). Bas.: Kantius suecicus Arnell et J.Perss., Rev. Bryol. 29 (2):
29, 1902 (Arnell 1902).
*** Calypogeia tenax (Spruce) Steph., Bull. Herb. Boissier (sér. 2) 8 (9): 664 (396),
1908 (Stephani 1908d). Bas.: Kantius tenax Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 416, 1885 (Spruce 1885).
** Calypogeia uncinulatula Herzog, Hedwigia 67 (6): 250, 1927 (Herzog 1927).
***
*
**
**
**
**
**
**
*
**
**
**
**
**
**
Incertae sedis
Calypogeia aeruginosa Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 107, 1860 [1861]
(Mitten 1860c).
Calypogeia amazonica (Spruce) Steph., Bull. Herb. Boissier (sér. 2) 8 (9): 680
(412), 1908 (Stephani 1908d). Bas.: Kantius amazonicus Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 415, 1885 (Spruce 1885). 72
Calypogeia angusta Steph., Bull. Herb. Boissier (sér. 2) 8 (9): 663 (395), 1908
(Stephani 1908d).
Calypogeia apiculata (Steph.) Steph., Bull. Herb. Boissier (sér. 2) 8 (9): 668 (400),
1908 (Stephani 1908d). Bas.: Kantius apiculatus Steph., Hedwigia 34 (2): 51,
1895 (Stephani 1895c).
Calypogeia asakawana S.Hatt. ex Inoue, J. Jap. Bot. 39 (4): 107, 1964 (Inoue
1964a).
Calypogeia ceylanica S.Hatt. et Mizut., Candollea 23: 288, 1968 (Hattori 1968).
Calypogeia contracta Inoue, Bull. Natl. Sci. Mus. Tokyo, B 1 (4): 139, 1975 (Inoue
1975b).
Calypogeia cuspidata (Steph.) Steph., Bull. Herb. Boissier (sér. 2) 8 (9): 669 (401),
1908 (Stephani 1908d). Bas.: Kantius cuspidatus Steph., Bull. Herb. Boissier 5
(10): 846, 1897 (Stephani 1897c).
Calypogeia decurrens (Steph.) Steph., Bull. Herb. Boissier (sér. 2) 8 (9): 675 (407),
1908 (Stephani 1908d). Bas.: Kantius decurrens Steph., Hedwigia 34 (2): 52, 1895
(Stephani 1895c).
Calypogeia formosana Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 186,
1934 (Horikawa 1934).
Calypogeia fujisana Inoue, Bull. Natl. Sci. Mus. Tokyo, B 1 (4): 135, 1975 (Inoue
1975b).
Calypogeia granulata Inoue, J. Jap. Bot. 43 (10/11): 468, 1968 (Inoue 1968b).
Calypogeia japonica Steph., Sp. Hepat. (Stephani) 6: 448, 1924 (Stephani 1924).
Calypogeia khasiana Ajit P.Singh et V.Nath, Taiwania 52 (4): 320, 2007 (Singh
and Nath 2007a).
Calypogeia latissima Steph., Sp. Hepat. (Stephani) 6: 449, 1924 (Stephani 1924).
72 Calypogeia amazonica was treated as conspeciic with Calypogeia miquelii in Gradstein et al. (1994),
but it has been accepted by later authors (e.g. Churchill et al. 2008, Söderström et al. 2013e).
World checklist of hornworts and liverworts
107
*** Calypogeia lunata Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 107, 1860 [1861] (Mitten
1860c).
** Calypogeia marginella Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 106, 1860 [1861]
(Mitten 1860c).
** Calypogeia obovata R.M.Schust., Phytologia 39 (4): 242, 1978 (Schuster 1978a).
* Calypogeia steyermarkii Fulford, Mem. New York Bot. Gard. 11 (3): 305, 1968
(Fulford 1968).
** Calypogeia tosana (Steph.) Steph., Bull. Herb. Boissier (sér. 2) 8 (9): 678 (410),
1908 (Stephani 1908d). Bas.: Kantius tosanus Steph., Hedwigia 34 (2): 54, 1895
(Stephani 1895c).
** Calypogeia udarii Sudipa Das et D.K.Singh, Nelumbo 53: 194, 2011 (Das and
Singh 2011).
** Eocalypogeia (R.M.Schust.) R.M.Schust., Fragm. Florist. Geobot. 40 (2): 861,
1995 (Schuster 1995a). Bas.: Metacalypogeia subg. Eocalypogeia R.M.Schust.,
Hepat. Anthocerotae N. Amer. 2: 107, 1969 (Schuster 1969b).
** Eocalypogeia quelpaertensis (S.Hatt. et Inoue) R.M.Schust., Fragm. Florist. Geobot.
40 (2): 861, 1995 (Schuster 1995a). Bas.: Metacalypogeia quelpaertensis S.Hatt. et
Inoue, J. Hattori Bot. Lab. 25: 129, 1962 (Hattori et al. 1962).
** Eocalypogeia schusterana (S.Hatt. et Mizut.) R.M.Schust., Fragm. Florist. Geobot.
40 (2): 861, 1995 (Schuster 1995a). Bas.: Metacalypogeia schusterana S.Hatt. et
Mizut., Misc. Bryol. Lichenol. 4 (8): 121, 1967 (Hattori and Mizutani 1967).
** Metacalypogeia (S.Hatt.) Inoue, J. Hattori Bot. Lab. 21: 231, 1959 (Inoue
1959b). Bas.: Calypogeia subg. Metacalypogeia S.Hatt., J. Hattori Bot. Lab. 18: 83,
1957 (Hattori 1957b).
*** Metacalypogeia alternifolia (Nees) Grolle, Österr. Bot. Z. 111 (2/3): 185, 1964
(Grolle 1964f). Bas.: Mastigobryum alternifolium Nees, Syn. Hepat. 2: 216, 1845
(Gottsche et al. 1845a).
** Metacalypogeia cordifolia (Steph.) Inoue, J. Hattori Bot. Lab. 21: 233, 1959 (Inoue
1959b). Bas.: Calypogeia cordifolia Steph., Bull. Herb. Boissier (sér. 2) 8 (9): 661
(393), 1908 (Stephani 1908d).
** Mizutania Furuki et Z.Iwats., J. Hattori Bot. Lab. 67: 291, 1989 (Furuki and
Iwatsuki 1989).
*** Mizutania riccardioides Furuki et Z.Iwats., J. Hattori Bot. Lab. 67: 291, 1989
(Furuki and Iwatsuki 1989).
*** Mnioloma Herzog, Ann. Bryol. 3: 115, 1930 (Herzog 1930a).
** subg. Caracoma (Bischl.) R.M.Schust., Fragm. Florist. Geobot. 40 (2): 833,
1995 (Schuster 1995a). Bas.: Calypogeia subg. Caracoma Bischl., Candollea 18:
26, 1962 (Bischler 1962b).
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** Mnioloma bolivianum (Fulford) R.M.Schust., Beih. Nova Hedwigia 118: 509,
2000 (Schuster 2000a). Bas.: Calypogeia boliviana Fulford, Mem. New York Bot.
Gard. 11 (3): 291, 1968 (Fulford 1968).
*** Mnioloma caespitosum (Spruce) R.M.Schust., Fragm. Florist. Geobot. 40 (2): 839,
1995 (Schuster 1995a). Bas.: Kantius caespitosus Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 412, 1885 (Spruce 1885).
*** Mnioloma cellulosum (Spreng.) R.M.Schust., Fragm. Florist. Geobot. 40 (2): 836,
1995 (Schuster 1995a). Bas.: Jungermannia cellulosa Spreng. Syst. Veg. (ed. 16)
[Sprengel] 4 (1): 232, 1827 (Sprengel 1827a).
*** Mnioloma crenulatum (Bischl.) R.M.Schust., Fragm. Florist. Geobot. 40 (2): 839,
1995 (Schuster 1995a). Bas.: Calypogeia crenulata Bischl., Candollea 18: 35, 1962
(Bischler 1962b).
*** Mnioloma cyclostipum (Spruce) R.M.Schust., Fragm. Florist. Geobot. 40 (2): 843,
1995 (Schuster 1995a). Bas.: Kantius cyclostipus Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 411, 1885 (Spruce 1885).
** Mnioloma elliottii (Steph.) R.M.Schust., Fragm. Florist. Geobot. 40 (2): 841,
1995 (Schuster 1995a). Bas.: Calypogeia elliottii Steph., Bull. Herb. Boissier (sér.
2) 8 (9): 663 (395), 1908 (Stephani 1908d).
** Mnioloma issistipulum (Bischl.) R.M.Schust., Fragm. Florist. Geobot. 40 (2): 847,
1995 (Schuster 1995a). Bas.: Calypogeia issistipula Bischl., Candollea 18: 47, 1962
(Bischler 1962b).
*** Mnioloma fuscum (Lehm.) R.M.Schust., Fragm. Florist. Geobot. 40 (2): 848,
1995 (Schuster 1995a). Bas.: Jungermannia fusca Lehm., Linnaea 4: 360, 1829
(Lehmann 1829).
*** Mnioloma nephrostipum (Spruce) R.M.Schust., Fragm. Florist. Geobot. 40 (2):
847, 1995 (Schuster 1995a). Bas.: Kantius nephrostipus Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 412, 1885 (Spruce 1885).
** Mnioloma novaezelandiae J.J.Engel, Cryptog. Bryol. 27 (1): 111, 2006 (Engel
2006a).
*** Mnioloma parallelogramum (Spruce) R.M.Schust., Fragm. Florist. Geobot. 40 (2):
847, 1995 (Schuster 1995a). Bas.: Kantius parallelogramus Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 413, 1885 (Spruce 1885).
** Mnioloma retusum (Bischl.) R.M.Schust., Fragm. Florist. Geobot. 40 (2): 839,
1995 (Schuster 1995a). Bas.: Calypogeia retusa Bischl., Candollea 18: 33, 1962
(Bischler 1962b).
** Mnioloma stamatotonum M.A.M.Renner et E.A.Br., Fieldiana, Bot. (n.ser.) 47:
173, 2008 (Renner and Brown 2008).
*** Mnioloma venezuelanum (Fulford) R.M.Schust., Beih. Nova Hedwigia 118: 509,
2000 (Schuster 2000a). Bas.: Calypogeia venezuelana Fulford, Mem. New York
Bot. Gard. 11 (3): 287, 1968 (Fulford 1968).
** subg. Mnioloma, Fragm. Florist. Geobot. 40 (2): 833, 1995 (Schuster 1995a).
*** Mnioloma rhynchophyllum Herzog, Ann. Bryol. 3: 120, 1930 (Herzog 1930a).
World checklist of hornworts and liverworts
***
109
Endogemmataceae Konstant., Vilnet et A.V.Troitsky
by N.A. Konstantinova
Vilnet et al. (2011) described the monotypic family Endogemmataceae based on molecular evidence after a re-evaluation of Solenostomataceae.
*** Endogemma Konstant., Vilnet et A.V.Troitsky, Folia Cryptog. Estonica 48:
132, 2011 (Vilnet et al. 2011).
*** Endogemma caespiticia (Lindenb.) Konstant., Vilnet et A.V.Troitsky, Folia Cryptog. Estonica 48: 132, 2011 (Vilnet et al. 2011). Bas.: Jungermannia caespiticia
Lindenb., Syn. hepat. eur: 67, 1829 (Lindenberg 1829).
***
Geocalycaceae H.Klinggr.
Placement of Geocalycaceae in Jungermanniinae follows Shaw et al. (2015).
*** Geocalyx Nees, Naturgesch. Eur. Leberm. 1: 97, 1833 (Nees 1833c).
*** Geocalyx caledonicus Steph., Bull. Herb. Boissier (sér. 2) 8 (3): 205 (265), 1908
(Stephani 1908h).
*** Geocalyx graveolens (Schrad.) Nees, Naturgesch. Eur. Leberm. 2: 397, 1836 (Nees
1836). Bas.: Jungermannia graveolens Schrad., Syst. Samml. Crypt. Gew. 2: 6,
1797 (Schrader 1797).
** Geocalyx lancistipulus (Steph.) S.Hatt., J. Jap. Bot. 28 (8): 234, 1953 (Hattori
1953a). Bas.: Lophocolea lancistipula Steph., Sp. Hepat. (Stephani) 6: 281, 1922
(Stephani 1922).
** Geocalyx orientalis Besch. et Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36:
clxxxix, 1889 [1890] (Bescherelle and Spruce 1889).
***
Gymnomitriaceae H.Klinggr.
by J. Váňa
he treatment of the family follows Váňa et al. (2010b) with some modiications.
Nardioideae is included following Vilnet et al. (2010) and Váňa et al. (2014c). Some
re-arrangements in Apomarsupella, Gymnomitrion and Marsupella were done by Vilnet
et al (2010) and Shaw et al. (2015). Further notes on nomenclature and taxonomy
can be found in Váňa et al. (2013d). Herzogobryum and Nothogymnomitrion were
removed from the family by Váňa et al. (2013e). he placement of Acrolophozia,
Nanomarsupella and Paramomitrion is provisional. Inclusion of Cryptocoleopsis follows
Shaw et al (2015).
110
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*** Acrolophozia R.M.Schust., Rev. Bryol. Lichénol. 34 (1/2): 259, 1966 (Schuster
1966b).
*** Acrolophozia fuegiana R.M.Schust., Nova Hedwigia 15: 499, 1968 (Schuster 1968b).
*** Acrolophozia pectinata R.M.Schust., Rev. Bryol. Lichénol. 34 (1/2): 261, 1966
(Schuster 1966b).
*** Acrolophozia sulcata Hässel, J. Bryol. 11 (1): 108, 1980 (Hässel 1980).
** Nanomarsupella R.M.Schust. ex A.Hagborg, L.Söderstr. et von Konrat, Phytotaxa 112 (1): 16, 2013 (Hagborg et al. 2013). Based on: Marsupella subg. Nanomarsupella R.M.Schust., Phytologia 39 (4): 248, 1978 (Schuster 1978a).
*** Nanomarsupella xenophylla (R.M.Schust.) R.M.Schust. ex A.Hagborg, L.Söderstr.
et von Konrat, Phytotaxa 112 (1): 16, 2013 (Hagborg et al. 2013). Bas.: Marsupella
xenophylla R.M.Schust., Phytologia 39 (4): 248, 1978 (Schuster 1978a).
** Paramomitrion R.M.Schust., J. Hattori Bot. Lab. 80: 134, 1996 (Schuster 1996a).
*** Paramomitrion paradoxum R.M.Schust., J. Hattori Bot. Lab. 80: 135, 1996
(Schuster 1996a).
***
Gymnomitrioideae T.Jensen
** Cryptocoleopsis Amakawa, J. Hattori Bot. Lab. 21: 274, 1959 (Amakawa 1959a).
*** Cryptocoleopsis imbricata Amakawa, J. Hattori Bot. Lab. 21: 274, 1959 (Amakawa
1959a).
*** Gymnomitrion Corda, Gen. hepat.: 651, 1829 (Corda 1829) nom. conserv.
*** Gymnomitrion adustum Nees, Naturgesch. Eur. Leberm. 1: 120, 1833 (Nees 1833c).
*** Gymnomitrion africanum (Steph.) Horik., Acta Phytotax. Geobot. 13: 212, 1943
(Horikawa 1943). Bas.: Acolea africana Steph., Sp. Hepat. (Stephani) 6: 77, 1917
(Stephani 1917a).
*** Gymnomitrion alpinum (Gottsche ex Husn.) Schifn., Österr. Bot. Z. 53 (7): 280,
1903 (Schifner 1903a). Bas.: Sarcocyphos alpinus Gottsche ex Husn., Hepaticol.
gall. 1: 13, 1875 (Husnot 1875).
*** Gymnomitrion asperulatum R.M.Schust., Acta Acad. Ped. Agr., Sect. Biol. 24: 114,
2003 (Váňa 2003).
*** Gymnomitrion atroilum Váňa, J. Hattori Bot. Lab. 41: 411, 1976 (Váňa 1976b).
*** Gymnomitrion bolivianum (Steph.) Váňa, Novon 20 (2): 225, 2010 (Váňa et al.
2010c). Bas.: Anastrophyllum bolivianum Steph., Biblioth. Bot. 87 (2): 186, 1916
(Stephani 1916a).
*** Gymnomitrion brevissimum (Dumort.) Warnst., Hedwigia 53 (3): 196, 1913
(Warnstorf 1913). Bas.: Acolea brevissima Dumort., Syll. Jungerm. Europ.: 76,
1831 (Dumortier 1831).
World checklist of hornworts and liverworts
111
*** Gymnomitrion commutatum (Limpr.) Schifn., Magyar Bot. Lapok 13: 304, 1914
[1915] (Schifner 1914b). Bas.: Sarcocyphos commutatus Limpr., Jahresber. Schles.
Ges. Vaterl. Cult. 57: 314, 1879 [1880] (Limpricht 1879).
*** Gymnomitrion concinnatum (Lightf.) Corda, Gen. hepat.: 651, 1829 (Corda
1829). Bas.: Jungermannia concinnata Lightf., Fl. Scot. 2: 786, 1777 (Lightfoot
1777), nom. conserv.
*** Gymnomitrion corallioides Nees, Naturgesch. Eur. Leberm. 1: 118, 1833 (Nees 1833c).
*** Gymnomitrion crenatilobum Grolle, Khumbu Himal 1 (4): 278, 1966 (Grolle 1966k).
*** Gymnomitrion crenulatum Gottsche ex Carrington, Trans. Bot. Soc. Edinburgh 7
(3): 444, 1863 (Carrington 1863).
*** Gymnomitrion crystallocaulon (Grolle) Váňa, Crand.-Stotl. et Stotler, Syst. Bot. 40
(1): 39, 2015 (Shaw et al. 2015). Bas.: Marsupella crystallocaulon Grolle, Khumbu
Himal 1 (4): 281, 1966 (Grolle 1966k).
** Gymnomitrion incompletum (Gottsche) R.M.Schust. ex Váňa, J. Hattori Bot. Lab.
40: 186, 1976 (Váňa 1976a). Bas.: Jungermannia incompleta Gottsche, Linnaea 28
(5): 551, 1856 [1857] (Gottsche 1856).
*** Gymnomitrion laceratum (Steph.) Horik., Acta Phytotax. Geobot. 13: 212, 1943
(Horikawa 1943). Bas.: Acolea lacerata Steph., Sp. Hepat. (Stephani) 6: 78, 1917
(Stephani 1917a).
*** Gymnomitrion miniatum Lindenb. et Gottsche, Syn. Hepat. 4: 617, 1846 (Gottsche
et al. 1846).
*** Gymnomitrion minutulum (Hässel) Váňa, Novon 20 (2): 225, 2010 (Váňa et al.
2010c). Bas.: Marsupella minutula Hässel, J. Bryol. 11 (1): 123, 1980 (Hässel 1980).
** Gymnomitrion moralesae Váňa, J. Hattori Bot. Lab. 48: 230, 1980 (Váňa 1980).
*** Gymnomitrion mucronulatum (N.Kitag.) N.Kitag., Acta Phytotax. Geobot. 19
(2/3): 53, 1962 (Kitagawa 1962a). Bas.: Gymnomitrion concinnatum var. mucronulatum N.Kitag., Acta Phytotax. Geobot. 18 (2/3): 38, 1959 (Kitagawa 1959).
** Gymnomitrion mucrophorum R.M.Schust., Bryologist 98 (2): 243, 1995 (Schuster
1995d).
*** Gymnomitrion nigrum (Grolle et Váňa) Váňa, Novon 20 (2): 225, 2010 (Váňa et
al. 2010c). Bas.: Marsupella nigra Grolle et Váňa, J. Hattori Bot. Lab. 40: 186,
1976 (Váňa 1976a).
*** Gymnomitrion noguchianum S.Hatt., J. Jap. Bot. 27 (2): 55, 1952 (Hattori 1952b).
*** Gymnomitrion obtusilobum N.Kitag., Bull. Univ. Mus. Univ. Tokyo 8: 229, 1975
(Hattori 1975e).
*** Gymnomitrion obtusum Lindb., Morgonbladet (Helsinki) 1877 (30, 6 Feb): 2,
1877 (Lindberg 1877a).
*** Gymnomitrion paciicum Grolle, Trans. Brit. Bryol. Soc. 5 (1): 92, 1966 (Grolle
1966f).
*** Gymnomitrion revolutum (Nees) H.Philib., Rev. Bryol. 17 (3): 34, 1890 (Philibert
1890). Bas.: Sarcocyphos revolutus Nees, Naturgesch. Eur. Leberm. 2: 419, 1836 (Nees
1836).
112
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Gymnomitrion revolutum subsp. novoguineanensis (R.M.Schust.) Váňa, Crand.-Stotl. et
Stotler, Syst. Bot. 40 (1): 39, 2015 (Shaw et al. 2015). Bas.: Apomarsupella revoluta subsp.
novoguineanensis R.M.Schust., J. Hattori Bot. Lab. 80: 90, 1996 (Schuster 1996a).
*** Gymnomitrion rubidum (Mitt.) Váňa, Crand.-Stotl. et Stotler, Syst. Bot. 40 (1):
39, 2015 (Shaw et al. 2015). Bas.: Jungermannia rubida Mitt., J. Proc. Linn. Soc.,
Bot. 5 (18): 90, 1860 [1861] (Mitten 1860c).
*** Gymnomitrion setaceum Grolle et Váňa, J. Hattori Bot. Lab. 41: 411, 1976 (Váňa
1976b).
*** Gymnomitrion sinense Müll.Frib., Rev. Bryol. Lichénol. 20 (1/2): 176, 1951 (Müller
1951b).
** Gymnomitrion strictum (Berggr.) R.M.Schust., J. Hattori Bot. Lab. 26: 280, 1963
(Schuster 1963b). Bas.: Cesius strictus Berggr., New Zealand Hepat.: 2, 1898
(Berggren 1898).
** Gymnomitrion strictum var. inaequale R.M.Schust., J. Hattori Bot. Lab. 80: 118,
1996 (Schuster 1996a).
*** Gymnomitrion subintegrum (S.W.Arnell) Váňa, Novon 20 (2): 225, 2010 (Váňa et
al. 2010c). Bas.: Marsupella subintegra S.W.Arnell, Ark. Bot. (n.ser.) 3 (16): 545,
1956 (Arnell 1956e).
*** Gymnomitrion truncatoapiculatum Herzog, Hedwigia 74 (2): 81, 1934 (Herzog
1934a).
*** Gymnomitrion verrucosum W.E.Nicholson, Symb. Sin. 5: 10, 1930 (Nicholson et
al. 1930).
*** Marsupella Dumort., Commentat. Bot. (Dumortier): 114, 1822 (Dumortier 1822).
*** Marsupella alata S.Hatt. et N.Kitag., Mem. Coll. Sci. Kyoto Imp. Univ., Ser. B,
Biol. 27: 79, 1960 (Kitagawa 1960b).
*** Marsupella andreaeoides (Lindb.) Müll.Frib., Feddes Repert. Spec. Nov. Regni
Veg. 54 (2/3): 214, 1951 (Müller 1951a). Bas.: Cesius andreaeoides Lindb., Meddel. Soc. Fauna Fl. Fenn. 14: 68, 1887 (Lindberg 1887a).
*** Marsupella apiculata Schifn., Österr. Bot. Z. 53 (6): 249, 1903 (Schifner 1903b).
*** Marsupella aquatica (Lindenb.) Schifn., Sitzungsber. deutsch. naturwiss.-med. Vereins
Böhmen “Lotos” Prag 44 (8): 267, 1896 [1897] (Schifner 1896a). Bas.: Jungermannia
emarginata var. aquatica Lindenb., Syn. hepat. eur: 75, 1829 (Lindenberg 1829).
*** Marsupella arctica (Berggr.) Bryhn et Kaal., Rep. Second Norweg. Arctic Exped.
11: 26, 1906 (Bryhn 1906). Bas.: Sarcocyphos emarginatus var. arcticus Berggr.,
Kongl. Svenska Vetensk.-Akad. Handl. (n.ser.) 13 (7): 96, 1875 (Berggren 1875).
*** Marsupella boeckii (Austin) Lindb. ex Kaal., Nyt Mag. Naturvidensk. 33 (4/5):
409, 1893 (Kaalaas 1893b). Bas.: Sarcocyphos boeckii Austin, Bull. Torrey Bot.
Club 3 (3): 9, 1872 (Austin 1872).
*** Marsupella bolanderi (Austin) Underw., Zoe 1 (12): 365, 1891 (Underwood
1891). Bas.: Sarcocyphos bolanderi Austin, Bull. Torrey Bot. Club 3 (3): 9, 1872
(Austin 1872).
World checklist of hornworts and liverworts
113
*** Marsupella condensata (Ångstr. ex C.Hartm.) Lindb. ex Kaal., Nyt Mag. Naturvidensk. 33 (4/5): 420, 1893 (Kaalaas 1893b). Bas.: Gymnomitrion condensatum
Ångstr. ex C.Hartm., Handb. Skand. l. (ed. 10): 128, 1871 (Hartman 1871).
*** Marsupella disticha Steph., Bull. Herb. Boissier (sér. 2) 1 (2): 164 (25), 1901 (Stephani 1901h).
*** Marsupella emarginata (Ehrh.) Dumort., Recueil Observ. Jungerm.: 24, 1835
(Dumortier 1835). Bas.: Jungermannia emarginata Ehrh., Hannover. Mag. 22 (8):
141, 1784 (Ehrhart 1784).
** Marsupella emarginata subsp. tubulosa (Steph.) N.Kitag., Mem. Coll. Sci. Kyoto
Imp. Univ., Ser. B, Biol. 27: 76, 1960 (Kitagawa 1960b). Bas.: Marsupella tubulosa
Steph., Bull. Herb. Boissier 5 (2): 99, 1897 (Stephani 1897b).
** Marsupella emarginata subsp. tubulosa var. apertifolia (Steph.) N.Kitag., J. Hattori
Bot. Lab. 26: 89, 1963 (Kitagawa 1963b). Bas.: Marsupella apertifolia Steph., Bull.
Herb. Boissier (sér. 2) 1 (2): 162 (23), 1901 (Stephani 1901h).
** Marsupella emarginata subsp. tubulosa var. patens N.Kitag., Mem. Coll. Sci. Kyoto
Imp. Univ., Ser. B, Biol. 27: 77, 1960 (Kitagawa 1960b).
** Marsupella emarginata subsp. tubulosa var. tubulosa (Steph.) N.Kitag. ex Váňa et
L.Söderstr., Phytotaxa 183 (4): 288, 2014 (Váňa et al. 2014c). Bas.: Marsupella
tubulosa Steph., Bull. Herb. Boissier 5 (2): 99, 1897 (Stephani 1897b).
*** Marsupella funckii (F.Weber et D.Mohr) Dumort., Recueil Observ. Jungerm.: 24,
1835 (Dumortier 1835). Bas.: Jungermannia funckii F.Weber et D.Mohr, Bot.
Taschenb. (Weber): 422, 1807 (Weber and Mohr 1807).
*** Marsupella microphylla R.M.Schust., Phytologia 39 (4): 249, 1978 (Schuster 1978a).
** Marsupella minutissima N.Kitag., Mem. Coll. Sci. Kyoto Imp. Univ., Ser. B, Biol.
27: 81, 1960 (Kitagawa 1960b).
*** Marsupella neesii Sande Lac. ex Schifn., Consp. Hepat. Arch. Ind.: 70, 1898
(Schifner 1898b). Based on: Sarcocyphos neesii Nees ex Sande Lac., Ann. Mus.
Bot. Lugduno-Batavi 1: 288, 1864 (Sande Lacoste 1864), nom. inval.
*** Marsupella paroica R.M.Schust., Bryologist 60 (2): 145, 1957 (Schuster 1957b).
*** Marsupella profunda Lindb., Rev. Bryol. 14 (2): 19, 1887 (Lindberg 1887b).
*** Marsupella pseudofunckii S.Hatt., J. Hattori Bot. Lab. 4: 63, 1950 (Hattori 1950).
*** Marsupella sparsifolia (Lindb.) Dumort., Bull. Soc. Roy. Bot. Belgique 13: 128,
1874 (Dumortier 1874). Bas.: Sarcocyphos sparsifolius Lindb., Not. Sällsk. Fauna
Fl. Fenn. Förh. 9: 280, 1868 (Lindberg 1868b).
** Marsupella sparsifolia subsp. childii R.M.Schust., Phytotaxa 183 (4): 288, 2014
(Váňa et al. 2014c). Based on: Marsupella sparsifolia subsp. childii R.M.Schust., J.
Hattori Bot. Lab. 80: 61, 1996 (Schuster 1996a), nom. inval.
*** Marsupella sphacelata (Giesecke ex Lindenb.) Dumort., Recueil Observ. Jungerm.:
24, 1835 (Dumortier 1835). Bas.: Jungermannia sphacelata Giesecke ex Lindenb.,
Syn. hepat. eur: 76, 1829 (Lindenberg 1829).
*** Marsupella spiniloba R.M.Schust. et Damsh., Phytologia 63 (5): 326, 1987 (Schuster and Damsholt 1987).
114
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Marsupella sprucei (Limpr.) Bernet, Cat. hép. Suisse: 33, 1888 (Bernet 1888). Bas.:
Sarcocyphos sprucei Limpr., Flora 64 (5): 72, 1881 (Limpricht 1881).
** Marsupella stableri Spruce, Rev. Bryol. 8 (6): 96, 1881 (Spruce 1881a).
*** Marsupella stoloniformis N.Kitag., J. Hattori Bot. Lab. 30: 201, 1967 (Kitagawa 1967b).
** Marsupella stoloniformis subsp. vermiformis R.M.Schust., J. Hattori Bot. Lab. 80:
72, 1996 (Schuster 1996a).
*** Marsupella yakushimensis (Horik.) S.Hatt., Bull. Tokyo Sci. Mus. 11: 80, 1944
(Hattori 1944d). Bas.: Sphenolobus yakushimensis Horik., J. Sci. Hiroshima Univ.,
Ser. B, Div. 2, Bot. 2: 156, 1934 (Horikawa 1934).
*** Poeltia Grolle, Khumbu Himal 1 (4): 280, 1966 (Grolle 1966k).
*** Poeltia campylata Grolle, Khumbu Himal 1 (4): 280, 1966 (Grolle 1966k).
*** Prasanthus Lindb., Kongl. Svenska Vetensk.-Akad. Handl. (n.ser.) 23 (5): 62,
1889 (Lindberg and Arnell 1889).
** Prasanthus jamalicus Potemkin, Ann. Bot. Fenn. 29 (4): 319, 1992 (Potemkin 1992).
*** Prasanthus suecicus (Gottsche) Lindb., Kongl. Svenska Vetensk.-Akad. Handl. (n.
ser.) 23 (5): 62, 1889 (Lindberg and Arnell 1889). Bas.: Gymnomitrion suecicum
Gottsche, Fl. Danica 16 (48): 20, 1871 (Lange 1871).
***
Nardioideae Váňa
*** Nardia Gray, Nat. Arr. Brit. Pl. 1: 694, 1821 (Gray 1821) nom. conserv.
*** Nardia arnelliana Grolle, Bot. Mag. (Tokyo) 77 (914): 297, 1964 (Grolle 1964b).
*** Nardia assamica (Mitt.) Amakawa, J. Hattori Bot. Lab. 26: 23, 1963 (Amakawa
1963). Bas.: Jungermannia assamica Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 91,
1860 [1861] (Mitten 1860c).
*** Nardia breidleri (Limpr.) Lindb., Helsingf. Dagbl. 1880 (311, 15 Nov.): 2, 1880
(Lindberg 1880b). Bas.: Alicularia breidleri Limpr., Jahresber. Schles. Ges. Vaterl.
Cult. 57: 311, 1879 [1880] (Limpricht 1879).
*** Nardia compressa (Hook.) Gray, Nat. Arr. Brit. Pl. 1: 694, 1821 (Gray 1821). Bas.:
Jungermannia compressa Hook., Brit. Jungermann.: tab. 58, 1813 (Hooker 1813).
*** Nardia lagelliformis Inoue, J. Jap. Bot. 46 (1): 1, 1971 (Inoue 1971b).
*** Nardia geoscyphus (De Not.) Lindb., Not. Sällsk. Fauna Fl. Fenn. Förh. 13: 371,
1874 (Lindberg 1874a). Bas.: Alicularia geoscyphus De Not., Mem. Reale Accad.
Sci. Torino (ser. 2) 18: 486, 1859 (De Notaris 1859).
* Nardia geoscyphus var. dioica Bakalin, Arctoa 18: 87, 2009 [2010] (Bakalin et al. 2009b).
* Nardia geoscyphus var. suberecta (Lindb. ex Kaal.) Váňa, Phytotaxa 76 (3): 37, 2013
(Váňa et al. 2013d). Bas.: Nardia haematosticta var. suberecta Kaal., Nyt Mag. Naturvidensk. 33 (4/5): 395, 1893 (Kaalaas 1893b).
World checklist of hornworts and liverworts
115
*** Nardia grollei Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 491, 2009 (Váňa and
Long 2009).
*** Nardia insecta Lindb., Helsingf. Dagbl. 1878 (315, 18 Nov.): 2, 1878 (Lindberg 1878).
*** Nardia japonica Steph., Bull. Herb. Boissier 5 (2): 101, 1897 (Stephani 1897b).
* Nardia kamtschatica Arnell et C.E.O.Jensen, Hedwigia 67 (1/2): 111, 1927 (Arnell 1927). 73
* Nardia leptocaulis C.Gao, Fl. Hepat. Chin. Boreali-Orient.: 205, 1981 (Gao and
Chang 1981).
*** Nardia lescurii (Austin) Underw., Bull. Illinois State Lab. Nat. Hist. 2 (1): 115,
1884 (Underwood 1884). Bas.: Alicularia lescurii Austin, Hepat. bor.-amer.: 4,
1873 (Austin 1873).
** Nardia minutifolia Furuki, Bryol. Res. 9 (3): 73, 2006 (Furuki 2006b).
** Nardia nuda (Lindenb. et Gottsche) Váňa, Folia Geobot. Phytotax. 8 (2): 193,
1973 (Váňa 1973b). Bas.: Jungermannia nuda Lindenb. et Gottsche, Syn. Hepat.
5: 668, 1847 (Gottsche et al. 1847).
*** Nardia poeltii Váňa, J. Hattori Bot. Lab. 36: 73, 1972 [1973] (Váňa 1972b).
*** Nardia scalaris Gray, Nat. Arr. Brit. Pl. 1: 694, 1821 (Gray 1821).
* Nardia scalaris var. botryoidea (R.M.Schust.) Váňa, Phytotaxa 76 (3): 38, 2013
(Váňa et al. 2013d). Bas.: Nardia scalaris subsp. botryoidea R.M.Schust., Hepat.
Anthocerotae N. Amer. 2: 862, 1969 (Schuster 1969b).
* Nardia scalaris var. harae (Amakawa) Váňa, Phytotaxa 76 (3): 38, 2013 (Váňa et al.
2013d). Bas.: Nardia harae Amakawa, J. Jap. Bot. 32 (2): 38, 1957 (Amakawa 1957c).
*** Nardia subclavata (Steph.) Amakawa, J. Jap. Bot. 32 (2): 40, 1957 (Amakawa
1957c). Bas.: Jungermannia subclavata Steph., Sp. Hepat. (Stephani) 6: 93, 1917
(Stephani 1917a).
*** Nardia succulenta (A.Rich.) Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 519,
1885 (Spruce 1885). Bas.: Jungermannia succulenta A.Rich., Nov. Stirp. Pug. 4:
43, 1832 (Lehmann 1832).
*** Nardia unispiralis Amakawa, J. Jap. Bot. 32 (6): 167, 1957 (Amakawa 1957a).
**
Gyrothyraceae R.M.Schust.
Placement of Gyrothyraceae in Jungermanniinae follows Shaw et al. (2015).
** Gyrothyra M.Howe, Bull. Torrey Bot. Club 24 (4): 201, 1897 (Howe 1897a).
*** Gyrothyra underwoodiana M.Howe, Bull. Torrey Bot. Club 24 (4): 202, 1897
(Howe 1897a).
73 Nardia kamtschatica may be conspeciic with Nardia assamica (Váňa 1976c), but the type specimen
could not be studied.
116
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Harpanthaceae Arnell
Davis (2004) and Hentschel et al. (2006) resolved Harpanthus as an independent lineage
which is recognized here as a monogeneric family as originally construed by Arnell (1928).
*** Harpanthus Nees, Naturgesch. Eur. Leberm. 2: 351, 1836 (Nees 1836).
*** Harpanthus drummondii (Taylor) Grolle, Österr. Bot. Z. 112 (3): 274, 1965
(Grolle 1965g). Bas.: Chiloscyphus drummondii Taylor, London J. Bot. 5: 283,
1846 (Taylor 1846a).
*** Harpanthus lotovianus (Nees) Nees, Naturgesch. Eur. Leberm. 2: 353, 1836
(Nees 1836). Bas.: Jungermannia lotoviana Nees, Flora 16 (26): 408, 1833 (Nees
1833b).
*** Harpanthus scutatus (F.Weber et D.Mohr) Spruce, Trans. Bot. Soc. Edinburgh 3
(1/4): 209, 1850 (Spruce 1850). Bas.: Jungermannia scutata F.Weber et D.Mohr,
Bot. Taschenb. (Weber): 408, 1807 (Weber and Mohr 1807).
**
Hygrobiellaceae Konstant. et Vilnet
by N. Konstantinova
he family Hygrobiellaceae was validated by Konstantinova et al. (2014a).
*** Hygrobiella Spruce, Cephalozia: 73, 1882 (Spruce 1882).
** Hygrobiella intermedia Bakalin et Vilnet, Pl. Syst. Evol. 300 (10): 2286, 2014 (Bakalin and Vilnet 2014).
*** Hygrobiella laxifolia (Hook.) Spruce, Cephalozia: 74, 1882 (Spruce 1882). Bas.:
Jungermannia laxifolia Hook., Brit. Jungermann.: tab. 59, 1813 (Hooker 1813).
** Hygrobiella squamosa Bakalin et Vilnet, Pl. Syst. Evol. 300 (10): 2286, 2014 (Bakalin and Vilnet 2014).
***
Jackiellaceae R.M.Schust.
by J. Váňa
he monogeneric status of Jackiella is supported by a molecular study based on three
loci by Hendry et al. (2007).
*** Jackiella Schifn., Hep. Fl. Buitenzorg: 211, 1900 (Schifner 1900a).
** Jackiella angustifolia Herzog, Trans. Brit. Bryol. Soc. 1 (4): 296, 1950 (Herzog 1950a).
** Jackiella ceylanica Schifn. ex Steph., Bull. Herb. Boissier (sér. 2) 8 (3): 212 (272),
1908 (Stephani 1908h).
World checklist of hornworts and liverworts
117
*** Jackiella curvata E.A.Hodgs. et Allison, Trans. Roy. Soc. New Zealand 85 (4):
571, 1958 (Hodgson 1958).
*** Jackiella javanica Schifn., Hep. Fl. Buitenzorg: 212, 1900 (Schifner 1900a).
** Jackiella javanica var. cavifolia Schifn., Hep. Fl. Buitenzorg: 213, 1900 (Schifner
1900a).
** Jackiella javanica var. cordifolia Schifn., Hep. Fl. Buitenzorg: 213, 1900 (Schifner
1900a).
** Jackiella renifolia Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl.
70: 218, 1900 [1901] (Schifner 1900c).
** Jackiella sinensis (W.E.Nicholson) Grolle, Österr. Bot. Z. 111 (2/3): 186, 1964
(Grolle 1964f). Bas.: Aplozia sinensis W.E.Nicholson, Symb. Sin. 5: 12, 1930
(Nicholson et al. 1930).
** Jackiella singapurensis Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss.
Kl. 70: 218, 1900 [1901] (Schifner 1900c).
** Jackiella singapurensis var. philippinensis N.Kitag., Misc. Bryol. Lichenol. 9 (1): 9,
1981 (Kitagawa 1981b).
Excluded from the genus
* Jackiella unica Steph., Sp. Hepat. (Stephani) 6: 318, 1922 (Stephani 1922). 74
***
Jungermanniaceae Rchb.
by J. Váňa
he treatment of Jungermanniaceae follows Shaw et al. (2015). Many old names in
Jungermannia are still neither synonymized nor transferred and we do not know their
value. Some of them may prove to be older names of currently accepted taxa. We list
those doubtful names in a separate section below.
**
Delavayelloideae Grolle
** Delavayella Steph., Hedwigia 33 (1): 4, 1894 (Stephani 1894a).
** Delavayella serrata Steph., Mém. Soc. Nat. Sci. Nat. Math. Cherbourg 29: 211,
1894 (Stephani 1894b).
** Delavayella serrata var. purpurea P.C.Chen, Feddes Repert. Spec. Nov. Regni Veg.
58: 38, 1955 (Chen 1955).
** Liochlaena Nees, Syn. Hepat. 2: 150, 1845 (Gottsche et al. 1845a).
*** Liochlaena lanceolata Nees, Syn. Hepat. 2: 150, 1845 (Gottsche et al. 1845a).
74 Jackiella unica was identiied as a Colura species before the B herbarium burned (Grolle 1966d).
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*** Liochlaena subulata (A.Evans) Schljakov, Pečen. Mchi Sev. SSSR 4: 71, 1981
(Shliakov 1981). Bas.: Jungermannia subulata A.Evans, Trans. Connecticut Acad.
Arts 8 (15): 258, 1891 (Evans 1891).
**
Jungermannioideae Dumort.
** Eremonotus Lindb. et Kaal. ex Pearson, Hepat. Br. Isl. 1 (6-15): 200, 1900
(Pearson 1900).
*** Eremonotus myriocarpus (Carrington) Lindb. et Kaal. ex Pearson, Hepat. Br. Isl.
1 (6-15): 201, 1900 (Pearson 1900). Bas.: Jungermannia myriocarpa Carrington,
Hepat. Brit. Exsicc. Fasc. II: no. 96, 1879 (Carrington and Pearson 1879).
*** Jungermannia L., Sp. Pl. 1: 1131, 1753 (Linnaeus 1753). 75
*** Jungermannia atrovirens Dumort., Syll. Jungerm. Europ.: 51, 1831 (Dumortier
1831).
*** Jungermannia borealis Damsh. et Váňa, Lindbergia 4 (1/2): 5, 1977 (Damsholt
and Váňa 1977).
* Jungermannia erectii Ajit P.Singh et V.Nath, Hepat. Khasi Jaintia Hills: E. Himal.:
117, 2007 (Singh and Nath 2007b). 76
*** Jungermannia exsertifolia Steph., Sp. Hepat. (Stephani) 6: 86, 1917 (Stephani
1917a).
** Jungermannia exsertifolia subsp. cordifolia (Dumort.) Váňa, Folia Geobot. Phytotax. 8 (3): 268, 1973 (Váňa 1973c). Bas.: Aplozia cordifolia Dumort., Bull. Soc.
Roy. Bot. Belgique 13: 59, 1874 (Dumortier 1874).
*** Jungermannia gollanii Steph., Sp. Hepat. (Stephani) 6: 86, 1917 (Stephani 1917a).
** Jungermannia konstantinovae Bakalin et Vilnet, Arctoa 18: 161, 2009 [2010] (Bakalin and Vilnet 2009).
*** Jungermannia ovatotrigona (Steph.) Grolle, Feddes Repert. 82 (1): 90, 1971 (Grolle 1971b). Bas.: Jamesoniella ovatotrigona Steph., Biblioth. Bot. 87 (2): 184, 1916
(Stephani 1916a).
*** Jungermannia polaris Lindb., Öfvers. Kongl. Vetensk.-Akad. Förh. 23 (10): 560,
1866 [1867] (Lindberg 1866).
*** Jungermannia pumila With., Arr. Brit. Pl., ed. 3, 3: 883, 1796 (Withering 1796).
75 Jungermannia was used for almost all leafy liverworts from Linnaeus (1753) and well into the 19th
century. Some taxa have neither been transferred nor synonymized. As the names often are very old,
they may gain priority over existing names if proven to be a synonym of something. hey are listed
in the “Names in genera not currently accepted” section below.
76 Jungermannia erectii was described on the basis of sterile material and it is not certain if it belongs in
Jungermannia or Solenostoma (Váňa and Long 2009).
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Taxa doubtfully belonging to the genus 77
Jungermannia amentacea Bertol., Mem. Reale Accad. Sci. Ist. Bologna (ser. 2) 1:
19, 1862 (Bertoloni 1862).
Jungermannia brasiliensis Raddi, Critt. Brasil.: 15, 1822 (Raddi 1822). 78
Jungermannia chinensis Osbeck, Dagb. Ostind. Resa: 221, 1757 (Osbeck 1757). 79
Jungermannia cordata Vill., Hist. Pl. Dauphiné (Villars) 3: 923, 1789 (Villars 1789).
Jungermannia crenulata Schmidel, Jungerm. Char.: 20, 1760 (Schmidel 1760).
Jungermannia creutzeri Kremer, Monogr. hépat. Moselle: 26, 1837 (Krémer 1837).
Jungermannia digitata C.F.W.Meissn. ex Spreng. Syst. Veg. (ed. 16) [Sprengel] 4
(2): 326, 1827 (Sprengel 1827b).
Jungermannia dubioides H.A.Mill., Phytologia 47 (4): 322, 1981 (Miller 1981).
Nom. nov. pro Jungermannia dubia Nees, Prodr. Fl. Norfolk.: 5, 1833 (Endlicher
1833), nom. illeg.
Jungermannia fernandeziana Mitt., Rep. Challenger, Bot. 1 (3, 1): 85, 1884 (Mitten 1884b).
Jungermannia hexagona Schwägr., Hist. Musc. Hepat. Prodr.: 18, 1814 (Schwägrichen 1814).
Jungermannia holandriana Kremer, Monogr. hépat. Moselle: 25, 1837 (Krémer 1837).
Jungermannia incerta Gottsche, Abh. Naturwiss. Vereins Bremen 7: 344, 1882
(Gottsche 1882). 80
Jungermannia laterilora Hampe ex Gottsche, Mexik. Leverm.: 82, 1863 (Gottsche 1863).
Jungermannia lescuriana Austin, Rep. (Annual) Regents Univ. State New York
State Cab. Nat. Hist. 19: 67, 1866 (Peck 1866). 81
Jungermannia longiretis Besch. et Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36:
clxxxv, 1889 [1890] (Bescherelle and Spruce 1889). 82
Jungermannia mastigophora Spreng. Neue Entdeck. Planzenk. 2: 99, 1821 (Sprengel 1821). 83
Jungermannia michelii Mérat, Nouv. l. env. Paris (ed. 2) 1: 219, 1821 (Mérat 1821). 84
Jungermannia minima Scop., Fl. Carniol. (ed. 2) 2: 350, 1772 (Scopoli 1772).
Jungermannia odorata With., Bot. arr. veg. Gr. Brit. 2: 693, 1776 (Withering 1776).
Jungermannia peltata Schmidel, Jungerm. Char.: 14, 1760 (Schmidel 1760).
77 Jungermannia was used for almost all leafy hepatics from Linnaeus (1753) and well into the 19th century.
he following names have not been studied recently and are neither accepted nor synonymized. Many of
them may have priority when their identity is determined.
78 Jungermannia brasiliensis is a taxon of uncertain status. Costa (2009) could not ind any authentic
material and did not know what it is.
79 Jungermannia chinensis may be conspeciic with Cephalozia connivens or Cephalozia bicuspidata.
80 Jungermannia incerta is a doubtful taxon (Grolle 1995). he type specimen was burned in B.
81 Jungermannia lescuriana is a Cephaloziella species.
82 Jungermannia longiretis is an Isotachis species.
83 Jungermannia mastigophora is probably a Mastigophora species.
84 Jungermannia michelii is a Fossombronia species.
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Jungermannia quadridigitata Grif., Not. pl. asiat. 2: 314, 1849 (Griith 1849). 85
Jungermannia sauteri De Not. ex Rabenh., Hedwigia 1 (20): 121, 1857 (Rabenhorst 1857).
Jungermannia secunda Hampe ex Gottsche, Mexik. Leverm.: 82, 1863 (Gottsche 1863).
Jungermannia stereocaulis Bory, Voy. Uranie, Bot. 4: 130, 1827 (Gaudichaud 1827). 86
Jungermannia submersa Kremer, Monogr. hépat. Moselle: 36, 1837 (Krémer 1837).
Jungermannia sullivantiana Austin, Rep. (Annual) Regents Univ. State New York
State Cab. Nat. Hist. 19: 66, 1866 (Peck 1866). 87
Jungermannia supina Hofm., Deutschl. Fl., heil 2 (Hofm.): 86, 1795 [1796]
(Hofmann 1795).
Jungermannia tenuis Ehrh., Beitr. Naturk. (Ehrhart) 4: 45, 1789 (Ehrhart 1789). 88
Jungermannia uncifolia Steph., Hedwigia 34 (2): 51, 1895 (Stephani 1895c).
Jungermannia vernicosa Cass. ex Mérat, Nouv. l. env. Paris (ed. 2) 1: 221, 1821
(Mérat 1821).
Mesoptychioideae R.M.Schust.
*** Mesoptychia (Lindb.) A.Evans, Ottawa Naturalist 17: 15, 1903 (Evans 1903a).
Bas.: Jungermannia sect. Mesoptychia Lindb., Kongl. Svenska Vetensk.-Akad.
Handl. (n.ser.) 23 (5): 39, 1889 (Lindberg and Arnell 1889).
*** Mesoptychia badensis (Gottsche ex Rabenh.) L.Söderstr. et Váňa, Phytotaxa 65:
52, 2012 (Váňa et al. 2012b). Bas.: Jungermannia badensis Gottsche ex Rabenh.,
Hepat. Eur., Leberm. 9-10: no. 95, 1859 (Rabenhorst 1859).
*** Mesoptychia bantriensis (Hook.) L.Söderstr. et Váňa, Phytotaxa 65: 52, 2012 (Váňa
et al. 2012b). Bas.: Jungermannia bantriensis Hook., Brit. Jungermann.: tab. 41,
1813 (Hooker 1813).
** Mesoptychia bantriensis subsp. wallischii (Ştefănuţ) L.Söderstr. et Váňa, Phytotaxa 65: 52, 2012 (Váňa et al. 2012b). Bas.: Leiocolea bantriensis subsp. wallischii
Ştefănuţ, Hornwort Liverwort Romania: 21, 2008 (Ştefănuţ 2008).
* Mesoptychia chichibuensis (Inoue) L.Söderstr. et Váňa, Phytotaxa 65: 53, 2012
(Váňa et al. 2012b). Bas.: Lophozia chichibuensis Inoue, J. Jap. Bot. 36 (2): 41,
1961 (Inoue 1961b).
* Mesoptychia collaris (Nees) L.Söderstr. et Váňa, Phytotaxa 65: 53, 2012 (Váňa et al.
2012b). Bas.: Jungermannia collaris Nees, Fl. crypt. erlang.: xv, 1817 (Martius 1817).
85 Jungermannia quadridigitata is conspeciic with Kurzia paucilora in Schuster (1969b), but that species
does not occur in India (Sharma and Srivastava 1993). It is probably some other Kurzia species.
86 Jungermannia stereocaulis is a doubtful taxon. Engel (1990b) did not ind any type specimen and did
not know what it is.
87 Jungermannia sullivantiana is a Cephaloziella species.
88 Jungermannia tenuis is a Lejeunea species.
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121
Mesoptychia itzgeraldiae (Paton et A.R.Perry) L.Söderstr. et Váňa, Phytotaxa 65:
53, 2012 (Váňa et al. 2012b). Bas.: Leiocolea itzgeraldiae Paton et A.R.Perry, J.
Bryol. 18 (3): 470, 1995 (Paton and Perry 1995).
Mesoptychia gillmanii (Austin) L.Söderstr. et Váňa, Phytotaxa 65: 53, 2012 (Váňa
et al. 2012b). Bas.: Jungermannia gillmanii Austin, Bull. Torrey Bot. Club 3 (3):
12, 1872 (Austin 1872).
Mesoptychia heterocolpos (hed. ex Hartm.) L.Söderstr. et Váňa, Phytotaxa 65:
53, 2012 (Váňa et al. 2012b). Bas.: Jungermannia heterocolpos hed. ex Hartm.,
Handb. Skand. l. (ed. 3): 328, 1838 (Hartman 1838).
Mesoptychia heterocolpos var. arctica (S.W.Arnell) L.Söderstr. et Váňa, Phytotaxa
65: 53, 2012 (Váňa et al. 2012b). Bas.: Leiocolea arctica S.W.Arnell, Svensk Bot.
Tidskr. 44 (2): 374, 1950 (Arnell 1950).
Mesoptychia heterocolpos var. harpanthoides (Bryhn et Kaal.) L.Söderstr. et Váňa,
Phytotaxa 65: 53, 2012 (Váňa et al. 2012b). Bas.: Lophozia harpanthoides Bryhn et
Kaal., Rep. Second Norweg. Arctic Exped. 11: 31, 1906 (Bryhn 1906).
Mesoptychia igiana (S.Hatt.) L.Söderstr. et Váňa, Phytotaxa 65: 54, 2012 (Váňa et
al. 2012b). Bas.: Lophozia igiana S.Hatt., J. Jap. Bot. 31 (7): 201, 1956 (Hattori
1956a).
Mesoptychia mamatkulovii (Duda) L.Söderstr. et Váňa, Phytotaxa 65: 54, 2012
(Váňa et al. 2012b). Bas.: Lophozia mamatkulovii Duda, Trans. Brit. Bryol. Soc. 6
(1): 82, 1970 (Duda 1970).
Mesoptychia mayebarae (S.Hatt.) L.Söderstr. et Váňa, Phytotaxa 65: 54, 2012
(Váňa et al. 2012b). Bas.: Cephalozia mayebarae S.Hatt., J. Hattori Bot. Lab. 3:
37, 1948 [1950] (Hattori 1948a).
Mesoptychia morrisoncola (Horik.) L.Söderstr. et Váňa, Phytotaxa 65: 54, 2012
(Váňa et al. 2012b). Bas.: Lophozia morrisoncola Horik., J. Sci. Hiroshima Univ.,
Ser. B, Div. 2, Bot. 2: 150, 1934 (Horikawa 1934).
Mesoptychia polymorpha Stotler, Crand.-Stotl. et Bakalin, Polish Bot. J. 58 (1): 82,
2013 (Crandall-Stotler et al. 2013).
Mesoptychia rutheana (Limpr.) L.Söderstr. et Váňa, Phytotaxa 65: 54, 2012 (Váňa
et al. 2012b). Bas.: Jungermannia rutheana Limpr., Jahresber. Schles. Ges. Vaterl.
Cult. 61: 207, 1884 (Limpricht 1884).
Mesoptychia rutheana var. laxa (Schifn. ex Burrell) L.Söderstr. et Váňa, Phytotaxa
65: 54, 2012 (Váňa et al. 2012b). Bas.: Lophozia schultzii var. laxa Schifn. ex Burrell, J. Bot. 49: 217, 1911 (Burrell 1911).
Mesoptychia sahlbergii (Lindb. et Arnell) A.Evans, Ottawa Naturalist 17: 15, 1903
(Evans 1903a). Bas.: Jungermannia sahlbergii Lindb. et Arnell, Kongl. Svenska
Vetensk.-Akad. Handl. (n.ser.) 23 (5): 40, 1889 (Lindberg and Arnell 1889).
Mesoptychia subcrispa (Herzog) L.Söderstr. et Váňa, Phytotaxa 65: 54, 2012 (Váňa
et al. 2012b). Bas.: Lophozia subcrispa Herzog, Ann. Naturhist. Mus. Wien 53 (1):
362, 1942 [1943] (Herzog 1942b).
122
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*** Mesoptychia turbinata (Raddi) L.Söderstr. et Váňa, Phytotaxa 65: 55, 2012 (Váňa
et al. 2012b). Bas.: Jungermannia turbinata Raddi, Jungermanniogr. Etrusca: 18,
1818 (Raddi 1818a).
* Mesoptychia ussuriensis (Bakalin) L.Söderstr. et Váňa, Phytotaxa 65: 55, 2012
(Váňa et al. 2012b). Bas.: Leiocolea ussuriensis Bakalin, Arctoa 17: 103, 2008
[2009] (Bakalin 2008c).
** Rivulariella D.H.Wagner, Phytoneuron 2013 (10): 2, 2013 (Wagner 2013).
** Rivulariella gemmipara (A.Evans) D.H.Wagner, Phytoneuron 2013 (10): 2, 2013
(Wagner 2013). Bas.: Chiloscyphus gemmiparus A.Evans, Bryologist 41 (3): 50,
1938 (Evans 1938b).
**
Notoscyphaceae Crand.-Stotl., Váňa et Stotler
Shaw et al. (2015) circumscribed Notoscyphaceae as monogeneric based on molecular
and morphological evidence.
*** Notoscyphus Mitt., Fl. vit.: 407, 1871 [1873] (Mitten 1871).
*** Notoscyphus lutescens (Lehm. et Lindenb.) Mitt., Fl. vit.: 407, 1871 [1873] (Mitten 1871). Bas.: Jungermannia lutescens Lehm. et Lindenb., Nov. Stirp. Pug. 4: 16,
1832 (Lehmann 1832).
**
Saccogynaceae Heeg
Shaw et al. (2015) recognized Saccogynaceae as being monogeneric.
*** Saccogyna Dumort., Commentat. Bot. (Dumortier): 113, 1822 (Dumortier
1822) nom. conserv.
* Saccogyna ligulata Steph., Bull. Herb. Boissier (sér. 2) 8 (3): 207 (267), 1908
(Stephani 1908h).
** Saccogyna subacuta Steph., Sp. Hepat. (Stephani) 6: 316, 1922 (Stephani 1922).
*** Saccogyna viticulosa (L.) Dumort., Syll. Jungerm. Europ.: 74, 1831 (Dumortier
1831). Bas.: Jungermannia viticulosa L., Sp. Pl. 1: 1131, 1753 (Linnaeus 1753).
*
Excluded from the genus
Saccogyna tridens Steph., Sp. Hepat. (Stephani) 6: 317, 1922 (Stephani 1922). 89
89 Saccogyna tridens is a Bazzania species, but the ainity is unclear and the type specimen was burned
in B (Grolle and Piippo 1984).
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123
Solenostomataceae Stotler et Crand.-Stotl.
by J. Váňa
Solenostomataceae was irst deined by Crandall-Stotler et al. (2009) and reined to
the current circumscription in Shaw et al (2015). he placement of Aponardia is provisional following Váňa et al. (2012c) and the placement of Arctoscyphus, Cryptocolea and
Diplocolea in the family is also provisional. Taxonomic and nomenclatural notes can
also be found in Váňa and Long (2009) and Váňa et al. (2012d, 2013c).
** Aponardia (R.M.Schust.) Váňa, Phytotaxa 65: 46, 2012 (Váňa et al. 2012c).
Bas.: Nardia subg. Aponardia R.M.Schust., Beih. Nova Hedwigia 119: 360, 2002
(Schuster 2002b).
*** Aponardia huerlimannii (Váňa et Grolle) Váňa, Phytotaxa 65: 46, 2012 (Váňa et
al. 2012c). Bas.: Nardia huerlimannii Váňa et Grolle, Österr. Bot. Z. 118 (3): 233,
1970 (Váňa 1970c).
** Arctoscyphus Hässel, Lindbergia 16 (4): 133, 1990 [1992] (Hässel 1990a).
** Arctoscyphus fuegiensis (C.Massal.) Hässel, Cryptog. Bryol. Lichénol. 17 (3): 164,
1996 (Hässel 1996). Bas.: Leioscyphus repens var. fuegiensis C.Massal., Nuovo
Giorn. Bot. Ital. 17 (3): 212, 1885 (Massalongo 1885).
** Arctoscyphus ronsmithii Hässel, Lindbergia 16 (4): 133, 1990 [1992] (Hässel 1990a).
** Cryptocolea R.M.Schust., Amer. Midl. Naturalist 49 (2): 414, 1953 (Schuster 1953).
*** Cryptocolea imbricata R.M.Schust., Amer. Midl. Naturalist 49 (2): 417, 1953
(Schuster 1953).
** Diplocolea Amakawa, J. Jap. Bot. 37 (9): 274, 1962 (Amakawa 1962).
*** Diplocolea sikkimensis Amakawa, J. Jap. Bot. 37 (9): 274, 1962 (Amakawa 1962).
*** Solenostoma Mitt., J. Proc. Linn. Soc., Bot. 8 (29): 51, 1864 [1865] (Mitten
1864c) nom. conserv. 90
** subg. Eucalyx (Lindb.) Váňa, Crand.-Stotl. et Stotler, Syst. Bot. 40 (1): 38,
2015 (Shaw et al. 2015). Bas.: Nardia sect. Eucalyx Lindb., Acta Soc. Sci. Fenn.
10: 525, 1875 (Lindberg 1875).
*** Solenostoma bilobum (S.Hatt. ex Amakawa) Potemkin et Nyushko, Liverworts and
Hornworts of Russia 1: 286, 2009 (Potemkin and Sofronova 2009). Bas.: Plectocolea biloba S.Hatt. ex Amakawa, J. Jap. Bot. 32 (7): 216, 1957 (Amakawa 1957d).
90 Solenostoma subdivisions are following Shaw et al. (2015), but the subgeneric placement of several
species is preliminary and only based on morphological characteristics. Plectocolea is synonymous
with Solenostoma, but one taxon has neither been transferred nor synonymized. It is listed in the
“Names in genera not currently accepted” section below.
124
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*** Solenostoma emarginatum (Amakawa) Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 136, 2010 (Váňa et al. 2010a). Bas.: Plectocolea emarginata Amakawa, J. Jap. Bot. 33 (11): 340, 1958 (Amakawa 1958b).
*** Solenostoma lagellatum (S.Hatt.) Váňa et D.G.Long, Nova Hedwigia 89 (3/4):
501, 2009 (Váňa and Long 2009). Bas.: Plectocolea lagellata S.Hatt., J. Hattori
Bot. Lab. 3: 13, 1948 [1950] (Hattori 1948b).
*** Solenostoma hokkaidense (Váňa) Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31
(2): 137, 2010 (Váňa et al. 2010a). Bas.: Jungermannia hokkaidensis Váňa, J. Hattori Bot. Lab. 35: 314, 1972 (Váňa 1972a).
*** Solenostoma obovatum (Nees) C.Massal., Epat. erb. critt. ital.: 17, 1903 (Massalongo 1903). Bas.: Jungermannia obovata Nees, Naturgesch. Eur. Leberm. 1: 332,
1833 (Nees 1833c).
*** Solenostoma obscurum (A.Evans) R.M.Schust., Hepat. Anthocerotae N. Amer. 2:
1013, 1969 (Schuster 1969b). Bas.: Nardia obscura A.Evans, Rhodora 21 (249):
159, 1919 (Evans 1919a).
*** Solenostoma schusteranum (J.D.Godfrey et G.Godfrey) Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31 (2): 137, 2010 (Váňa et al. 2010a). Bas.: Jungermannia
schusterana J.D.Godfrey et G.Godfrey, J. Hattori Bot. Lab. 46: 109, 1979 (Godfrey and Godfrey 1979).
*** Solenostoma subtilissimum (Schifn.) R.M.Schust., Hepat. Anthocerotae N. Amer.
2: 1027, 1969 (Schuster 1969b). Bas.: Nardia subtilissima Schifn., Ann. K. K.
Naturhist. Hofmus. 23: 136, 1909 (Schifner 1909b).
** subg. Metasolenostoma Váňa, Crand.-Stotl. et Stotler, Syst. Bot. 40 (1): 38,
2015 (Shaw et al. 2015).
*** Solenostoma fusiforme (Steph.) R.M.Schust., Hepat. Anthocerotae N. Amer. 2:
944, 1969 (Schuster 1969b). Bas.: Nardia fusiformis Steph., Bull. Herb. Boissier 5
(2): 99, 1897 (Stephani 1897b).
*** Solenostoma gracillimum (Sm.) R.M.Schust., Hepat. Anthocerotae N. Amer. 2:
972, 1969 (Schuster 1969b). Bas.: Jungermannia gracillima Sm., Engl. Bot. 32:
tab. 2238, 1811 (Smith and Sowerby 1811).
*** Solenostoma handelii (Schifn.) Müll.Frib., Beitr. Kryptogamenl. Schweiz 10 (2):
38, 1947 (Müller 1947). Bas.: Nardia handelii Schifn., Ann. K. K. Naturhist.
Hofmus. 23: 135, 1909 (Schifner 1909b).
** Solenostoma lignicola (Schifn.) Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31
(2): 137, 2010 (Váňa et al. 2010a). Bas.: Nardia lignicola Schifn., Ann. K. K.
Naturhist. Hofmus. 23: 137, 1909 (Schifner 1909b).
*** Solenostoma limbatifolium (Amakawa) Váňa et D.G.Long, Nova Hedwigia 89
(3/4): 504, 2009 (Váňa and Long 2009). Bas.: Jungermannia limbatifolia Amakawa, J. Hattori Bot. Lab. 31: 112, 1968 (Amakawa 1968b).
** Solenostoma philippinense Váňa, Phytotaxa 152 (1): 43, 2013 (Váňa et al. 2013c).
Based on: Solenostoma gracillimum subsp. camiguinense Bakalin, Polish Bot. J. 58
(1): 134, 2013 (Bakalin 2013).
World checklist of hornworts and liverworts
125
*** Solenostoma rubrum (Gottsche) R.M.Schust., Hepat. Anthocerotae N. Amer. 2:
975, 1969 (Schuster 1969b). Bas.: Jungermannia rubra Gottsche, Bot. Gaz. 13 (5):
113, 1888 (Underwood 1888).
*** Solenostoma suborbiculatum (Amakawa) Váňa et D.G.Long, Nova Hedwigia 89
(3/4): 508, 2009 (Váňa and Long 2009). Bas.: Jungermannia suborbiculata Amakawa, J. Hattori Bot. Lab. 31: 112, 1968 (Amakawa 1968b).
** subg. Plectocolea Mitt., J. Linn. Soc., Bot. 8 (31): 156, 1864 (Mitten 1864a).
*** Solenostoma balfourii (Váňa) Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31
(2): 136, 2010 (Váňa et al. 2010a). Bas.: Jungermannia balfourii Váňa, Folia Geobot. Phytotax. 9 (3): 279, 1974 (Váňa 1974a).
*** Solenostoma borneense (Amakawa) Váňa, Hentschel et Heinrichs, Cryptog. Bryol.
31 (2): 136, 2010 (Váňa et al. 2010a). Bas.: Jungermannia borneensis Amakawa, J.
Hattori Bot. Lab. 33: 160, 1970 (Amakawa 1970).
*** Solenostoma callithrix (Lindenb. et Gottsche) Steph., Bull. Herb. Boissier (sér. 2)
1 (5): 486 (48), 1901 (Stephani 1901a). Bas.: Jungermannia callithrix Lindenb. et
Gottsche, Syn. Hepat. 5: 673, 1847 (Gottsche et al. 1847).
*** Solenostoma champawatense (S.N.Srivast. et Amakawa) Váňa et D.G.Long, Nova
Hedwigia 89 (3/4): 496, 2009 (Váňa and Long 2009). Bas.: Jungermannia champawatensis S.N.Srivast. et Amakawa, Proc. Natl. Acad. Sci. India, B 61 (2): 205,
1991 (Srivastava and Amakawa 1991).
*** Solenostoma comatum (Nees) C.Gao, Fl. Hepat. Chin. Boreali-Orient.: 73, 1981
(Gao and Chang 1981). Bas.: Jungermannia comata Nees, Enum. Pl. Crypt. Javae:
78, 1830 (Nees 1830).
*** Solenostoma comatum var. novae-guineae (Váňa) Váňa, Hentschel et Heinrichs,
Cryptog. Bryol. 31 (2): 136, 2010 (Váňa et al. 2010a). Bas.: Jungermannia comata
var. novae-guineae Váňa, J. Hattori Bot. Lab. 37: 187, 1973 (Váňa 1973a).
*** Solenostoma crenuliforme (Austin) Steph., Bull. Herb. Boissier (sér. 2) 1 (5): 494
(56), 1901 (Stephani 1901a). Bas.: Jungermannia crenuliformis Austin, Bull. Torrey Bot. Club 3 (3): 10, 1872 (Austin 1872).
*** Solenostoma decolor (Schifn.) R.M.Schust. ex Váňa, Hentschel et Heinrichs,
Cryptog. Bryol. 31 (2): 136, 2010 (Váňa et al. 2010a). Bas.: Jungermannia decolor
Schifn., Leberm., Forschungsr. Gazelle 4 (4): 10, 1890 (Schifner 1890).
*** Solenostoma dusenii (Steph.) Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31 (2):
136, 2010 (Váňa et al. 2010a). Bas.: Nardia dusenii Steph., Hedwigia 30 (5): 209,
1891 (Stephani 1891a).
*** Solenostoma erectum (Amakawa) C.Gao, Fl. Hepat. Chin. Boreali-Orient.: 66,
1981 (Gao and Chang 1981). Bas.: Plectocolea erecta Amakawa, J. Jap. Bot. 32
(10): 307, 1957 (Amakawa 1957b).
* Solenostoma lagellalioides C.Gao, Fl. Hepat. Chin. Boreali-Orient.: 205, 1981
(Gao and Chang 1981).
126
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Solenostoma lavialbicans (Amakawa et Grolle) Váňa et D.G.Long, Nova Hedwigia
89 (3/4): 501, 2009 (Váňa and Long 2009). Bas.: Jungermannia lavialbicans Amakawa et Grolle, J. Hattori Bot. Lab. 31: 108, 1968 (Amakawa 1968b).
*** Solenostoma fossombronioides (Austin) R.M.Schust., Hepat. Anthocerotae N.
Amer. 2: 1027, 1969 (Schuster 1969b). Bas.: Jungermannia fossombronioides Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 220, 1869 (Austin 1869).
*** Solenostoma glaucum (Amakawa) Váňa et D.G.Long, Nova Hedwigia 89 (3/4):
502, 2009 (Váňa and Long 2009). Bas.: Jungermannia glauca Amakawa, Fl. E.
Himalaya: 511, 1966 (Hattori 1966c).
* Solenostoma gongshanense (C.Gao et J.Sun) Váňa et D.G.Long, Nova Hedwigia 89
(3/4): 502, 2009 (Váňa and Long 2009). Bas.: Jungermannia gongshanensis C.Gao
et J.Sun, Bull. Bot. Res., Harbin 27 (2): 140, 2007 (Sun and Duan 2007).
*** Solenostoma haskarlianum (Nees) R.M.Schust. ex Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 502, 2009 (Váňa and Long 2009). Bas.: Alicularia haskarliana
Nees, Syn. Hepat. 1: 12, 1844 (Gottsche et al. 1844).
*** Solenostoma hattorianum (Amakawa) Potemkin et Nyushko, Liverworts and Hornworts of Russia 1: 287, 2009 (Potemkin and Sofronova 2009). Bas.: Plectocolea
hattoriana Amakawa, J. Jap. Bot. 33 (11): 341, 1958 (Amakawa 1958b).
*** Solenostoma hirticalyx (Steph.) R.M.Schust. ex Váňa, Hentschel et Heinrichs,
Cryptog. Bryol. 31 (2): 137, 2010 (Váňa et al. 2010a). Bas.: Jungermannia hirticalyx Steph., Sp. Hepat. (Stephani) 6: 87, 1917 (Stephani 1917a).
*** Solenostoma horikawanum (Amakawa) Váňa, Hentschel et Heinrichs, Cryptog.
Bryol. 31 (2): 137, 2010 (Váňa et al. 2010a). Bas.: Plectocolea horikawana Amakawa, J. Jap. Bot. 32 (7): 219, 1957 (Amakawa 1957d).
*** Solenostoma hyalinum (Lyell) Mitt., Nat. hist. Azores: 319, 1870 (Mitten 1870).
Bas.: Jungermannia hyalina Lyell, Brit. Jungermann.: tab. 63, 1814 (Hooker 1814).
*** Solenostoma infuscum (Mitt.) Hentschel, Pl. Syst. Evol. 268 (1/4): 152, 2007
(Hentschel et al. 2007a). Bas.: Plectocolea infusca Mitt., Trans. Linn. Soc. London,
Bot. 3 (3): 196, 1891 (Mitten 1891).
*** Solenostoma infuscum var. ovicalyx (Steph.) Potemkin et Sofronova, Liverworts and
Hornworts of Russia 1: 288, 2009 (Potemkin and Sofronova 2009). Bas.: Solenostoma ovicalyx Steph., Sp. Hepat. (Stephani) 6: 82, 1917 (Stephani 1917a).
** Solenostoma kurilense (Bakalin) Váňa, Phytotaxa 152 (1): 40, 2013 (Váňa et al.
2013c). Bas.: Plectocolea lagellata var. kurilensis Bakalin, Arctoa 18: 90, 2009
[2010] (Bakalin et al. 2009b).
* Solenostoma lixingjiangii (C.Gao et X.L.Bai) Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 504, 2009 (Váňa and Long 2009). Bas.: Jungermannia lixingjiangii
C.Gao et X.L.Bai, Philipp. Scientist 38: 128, 2001 (Gao and Bai 2001).
*** Solenostoma marginatum (S.Hatt.) R.M.Schust., Hepat. Anthocerotae N. Amer. 2:
983, 1969 (Schuster 1969b). Bas.: Plectocolea marginata S.Hatt., J. Hattori Bot.
Lab. 3: 40, 1948 [1950] (Hattori 1948a).
World checklist of hornworts and liverworts
127
*** Solenostoma micranthum (Mitt.) Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31
(2): 137, 2010 (Váňa et al. 2010a). Bas.: Plectocolea micrantha Mitt., Fl. vit.: 405,
1871 [1873] (Mitten 1871).
** Solenostoma montanum (Steph.) Váňa, Phytotaxa 65: 44, 2012 (Váňa et al. 2012d).
Bas.: Nardia montana Steph., Hedwigia 28 (3): 164, 1889 (Stephani 1889d).
* Solenostoma multicarpum (C.Gao et J.Sun) Váňa et D.G.Long, Nova Hedwigia 89
(3/4): 505, 2009 (Váňa and Long 2009). Bas.: Jungermannia multicarpa C.Gao et
J.Sun, Bull. Bot. Res., Harbin 27 (2): 139, 2007 (Sun and Duan 2007).
** Solenostoma nilgiriense (A.Alam, Ad.Kumar et S.C.Srivast.) Váňa et D.G.Long,
Nova Hedwigia 89 (3/4): 505, 2009 (Váňa and Long 2009). Bas.: Jungermannia
nilgiriensis A.Alam, Ad.Kumar et S.C.Srivast., Bull. Bot. Surv. India 49 (1/4): 220,
2007 (Alam et al. 2007).
*** Solenostoma obliquifolium (Schifn.) R.M.Schust. ex Váňa, Hentschel et Heinrichs,
Cryptog. Bryol. 31 (2): 137, 2010 (Váňa et al. 2010a). Bas.: Nardia obliquifolia
Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 67: 191, 1898
(Schifner 1898a).
*** Solenostoma onraedtii (Váňa) Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31
(2): 137, 2010 (Váňa et al. 2010a). Bas.: Jungermannia onraedtii Váňa, Folia Geobot. Phytotax. 9 (3): 282, 1974 (Váňa 1974a).
* Solenostoma orbicularifolium (Piippo ex C.Gao et Bai) Váňa, Phytotaxa 222 (2):
199, 2015 (Söderström et al. 2015d). Bas.: Jungermannia orbicularifolia Piippo ex
C.Gao et Bai, Philipp. Scientist 38: 129, 2001 (Gao and Bai 2001).
*** Solenostoma otianum (S.Hatt.) R.M.Schust., Hepat. Anthocerotae N. Amer. 2:
984, 1969 (Schuster 1969b). Bas.: Plectocolea otiana S.Hatt., J. Jap. Bot. 28 (6):
183, 1953 (Hattori 1953b).
*** Solenostoma ovalifolium (Amakawa) Váňa, Phytotaxa 152 (1): 40, 2013 (Váňa et
al. 2013c). Bas.: Plectocolea infusca var. ovalifolia Amakawa, J. Jap. Bot. 34 (4):
115, 1959 (Amakawa 1959b).
*** Solenostoma paroicum (Schifn.) R.M.Schust., Amer. Midl. Naturalist 49 (2):
402, 1953 (Schuster 1953). Bas.: Nardia paroica Schifn., Lotos 58: 320, 1910
(Schifner 1910d).
*** Solenostoma plagiochilaceum (Grolle) Váňa et D.G.Long, Nova Hedwigia 89 (3/4):
505, 2009 (Váňa and Long 2009). Bas.: Jungermannia plagiochilacea Grolle, J.
Hattori Bot. Lab. 58: 197, 1985 (Grolle 1985b).
*** Solenostoma polyrhizoides (Grolle ex Amakawa) Váňa et D.G.Long, Nova Hedwigia
89 (3/4): 505, 2009 (Váňa and Long 2009). Bas.: Jungermannia polyrhizoides Grolle
ex Amakawa, J. Hattori Bot. Lab. 29: 262, 1966 (Amakawa 1966).
*** Solenostoma radicellosum Mitt., J. Linn. Soc., Bot. 8 (31): 156, 1864 [1865] (Mitten
1864a).
*** Solenostoma renauldii (Steph.) Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31
(2): 137, 2010 (Váňa et al. 2010a). Bas.: Jungermannia renauldii Steph., Bull. Soc.
Roy. Bot. Belgique, Compt. Rend. 30 (2): 201, 1891 [1892] (Stephani 1891b).
128
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Solenostoma rigidulum (S.Hatt.) R.M.Schust., Hepat. Anthocerotae N. Amer. 2:
983, 1969 (Schuster 1969b). Bas.: Plectocolea rigidula S.Hatt., J. Jap. Bot. 27 (2):
53, 1952 (Hattori 1952b).
*** Solenostoma rosulans (Steph.) Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 507,
2009 (Váňa and Long 2009). Bas.: Nardia rosulans Steph., Bull. Herb. Boissier 5
(2): 101, 1897 (Stephani 1897b).
*** Solenostoma rotundatum Amakawa, J. Jap. Bot. 31 (2): 50, 1956 (Amakawa 1956).
*** Solenostoma rubripunctatum (S.Hatt.) R.M.Schust., Hepat. Anthocerotae N.
Amer. 2: 898, 1969 (Schuster 1969b). Bas.: Plectocolea rubripunctata S.Hatt., J.
Hattori Bot. Lab. 3: 41, 1948 [1950] (Hattori 1948a).
* Solenostoma rupicola (Amakawa) Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 507,
2009 (Váňa and Long 2009). Bas.: Jungermannia rupicola Amakawa, J. Hattori
Bot. Lab. 22: 23, 1960 (Amakawa 1960b). 91
*** Solenostoma sikkimense (Schifn. ex Steph.) Váňa et D.G.Long, Nova Hedwigia 89
(3/4): 508, 2009 (Váňa and Long 2009). Bas.: Jungermannia sikkimensis Schifn.
ex Steph., Sp. Hepat. (Stephani) 6: 92, 1917 (Stephani 1917a).
*** Solenostoma tetragonum (Lindenb.) R.M.Schust. ex Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 509, 2009 (Váňa and Long 2009). Bas.: Jungermannia tetragona
Lindenb., Bot. Zeitung (Berlin) 6 (25): 462, 1848 (Meissner 1848).
* Solenostoma tetragonum var. kodaikanalense A.Alam, D.Sharma et So.Yadav, Phytotaxonomy 12: 70, 2012 (Alam et al. 2012).
*** Solenostoma torticalyx (Steph.) C.Gao, Fl. Hepat. Chin. Boreali-Orient.: 69, 1981
(Gao and Chang 1981). Bas.: Jungermannia torticalyx Steph., Sp. Hepat. (Stephani)
6: 94, 1917 (Stephani 1917a).
*** Solenostoma truncatum (Nees) R.M.Schust. ex Váňa et D.G.Long, Nova Hedwigia
89 (3/4): 509, 2009 (Váňa and Long 2009). Bas.: Jungermannia truncata Nees,
Enum. Pl. Crypt. Javae: 29, 1830 (Nees 1830).
** Solenostoma truncatum var. setulosum (Herzog) Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 510, 2009 (Váňa and Long 2009). Bas.: Plectocolea setulosa Herzog,
J. Hattori Bot. Lab. 14: 33, 1955 (Herzog and Noguchi 1955).
*** Solenostoma tuberculiferum (Herzog) Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31 (2): 138, 2010 (Váňa et al. 2010a). Bas.: Aplozia tuberculifera Herzog, Ann.
Bryol. 5: 84, 1932 (Herzog 1932a).
*** Solenostoma unispire (Amakawa) Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31
(2): 138, 2010 (Váňa et al. 2010a). Bas.: Plectocolea unispiris Amakawa, J. Jap. Bot.
29 (6): 178, 1954 (Amakawa 1954).
*** Solenostoma virgatum (Mitt.) Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 510,
2009 (Váňa and Long 2009). Bas.: Plectocolea virgata Mitt., Trans. Linn. Soc.
London, Bot. 3 (3): 197, 1891 (Mitten 1891).
91 Solenostoma rupicola is conspeciic with Solenostoma rosulans in Váňa (1975a), but accepted by Váňa
and Long (2009).
World checklist of hornworts and liverworts
129
*** Solenostoma vulcanicola (Schifn.) Nyushko ex Potemkin et Sofronova, Liverworts
and Hornworts of Russia 1: 289, 2009 (Potemkin and Sofronova 2009). Bas.:
Nardia vulcanicola Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl.
67: 191, 1898 (Schifner 1898a).
* Solenostoma zangmuii (C.Gao et X.L.Bai) Váňa et D.G.Long, Nova Hedwigia 89
(3/4): 510, 2009 (Váňa and Long 2009). Bas.: Jungermannia zangmuii C.Gao et
X.L.Bai, Philipp. Scientist 38: 135, 2001 (Gao and Bai 2001).
** subg. Solenostoma
*** Solenostoma amoenum (Lindenb. et Gottsche) R.M.Schust. ex Váňa, Hentschel et
Heinrichs, Cryptog. Bryol. 31 (2): 136, 2010 (Váňa et al. 2010a). Bas.: Jungermannia amoena Lindenb. et Gottsche, Syn. Hepat. 5: 674, 1847 (Gottsche et al. 1847).
*** Solenostoma amplexifolium (Hampe) Váňa et Schäf.-Verw., Acta Bot. Hung. 51
(3/4): 408, 2009 (Schäfer-Verwimp and Pócs 2009). Bas.: Plagiochila amplexifolia
Hampe, Nov. Stirp. Pug. 7: 6, 1838 (Lehmann 1838).
** Solenostoma appalachianum R.M.Schust. ex Bakalin, Arctoa 23: 127, 2014 (Bakalin 2014). Based on: Solenostoma appalachianum R.M.Schust., Rhodora 60 (717):
246, 1958 (Schuster 1958a), nom. inval. 92
*** Solenostoma appressifolium (Mitt.) Váňa et D.G.Long, Nova Hedwigia 89 (3/4):
494, 2009 (Váňa and Long 2009). Bas.: Jungermannia appressifolia Mitt., J. Proc.
Linn. Soc., Bot. 5 (18): 91, 1860 [1861] (Mitten 1860c). 93
*** Solenostoma appressifolium var. minor (Amakawa) Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 495, 2009 (Váňa and Long 2009). Bas.: Jungermannia clavellata
var. minor Amakawa, J. Hattori Bot. Lab. 26: 25, 1963 (Amakawa 1963).
*** Solenostoma appressifolium var. nigricans (Amakawa) Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 495, 2009 (Váňa and Long 2009). Bas.: Jungermannia decolyana var.
nigricans Amakawa, J. Hattori Bot. Lab. 30: 185, 1967 (Amakawa 1967b).
*** Solenostoma ariadne (Taylor) R.M.Schust. ex Váňa et D.G.Long, Nova Hedwigia
89 (3/4): 495, 2009 (Váňa and Long 2009). Bas.: Jungermannia ariadne Taylor,
Nov. Stirp. Pug. 8: 9, 1844 (Lehmann 1844).
*** Solenostoma atrobrunneum (Amakawa) Váňa et D.G.Long, Nova Hedwigia 89
(3/4): 495, 2009 (Váňa and Long 2009). Bas.: Jungermannia atrobrunnea Amakawa, J. Hattori Bot. Lab. 30: 192, 1967 (Amakawa 1967b).
*** Solenostoma atrorevolutum (Grolle ex Amakawa) Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 496, 2009 (Váňa and Long 2009). Bas.: Jungermannia atrorevoluta
Grolle ex Amakawa, J. Hattori Bot. Lab. 29: 255, 1966 (Amakawa 1966).
*** Solenostoma atrovirens Steph., Bull. Herb. Boissier (sér. 2) 1 (5): 493 (55), 1901
(Stephani 1901a).
92 Solenostoma appalachianum is placed in synonymy with Solenostoma pyrilorum by Schuster (1969b)
but Shaw et al. (2015) shows that the North American specimens difers from the East Asiatic specimens so named.
93 Solenostoma appressifolium seems to be a species complex (Shaw et al. 2015).
130
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*** Solenostoma baueri (Schifn.) Steph., Bull. Herb. Boissier (sér. 2) 1 (5): 495 (57),
1901 (Stephani 1901a). Bas.: Aplozia baueri Schifn., Denkschr. Kaiserl. Akad.
Wiss., Math.-Naturwiss. Kl. 67: 195, 1898 (Schifner 1898a).
*** Solenostoma bengalense (Amakawa) Váňa et D.G.Long, Nova Hedwigia 89 (3/4):
496, 2009 (Váňa and Long 2009). Bas.: Jungermannia bengalensis Amakawa, J.
Hattori Bot. Lab. 31: 112, 1968 (Amakawa 1968b).
** Solenostoma brevilorum Kashyap et R.S.Chopra, Liverworts W. Himal. 2: 85,
1932 (Kashyap and Chopra 1932).
*** Solenostoma caeleste (Inoue et Váňa) Váňa, Hentschel et Heinrichs, Cryptog. Bryol.
31 (2): 136, 2010 (Váňa et al. 2010a). Bas.: Jungermannia caelestis Inoue et Váňa,
Stud. Cryptog. Papua N. Guinea: 16, 1979 (Inoue 1979b).
* Solenostoma caoi (C.Gao et X.L.Bai) Váňa et D.G.Long, Nova Hedwigia 89 (3/4):
496, 2009 (Váňa and Long 2009). Bas.: Jungermannia caoi C.Gao et X.L.Bai,
Philipp. Scientist 38: 152, 2001 (Gao and Bai 2001).
*** Solenostoma caucasicum (Váňa) Konstant., Arctoa 1: 123, 1992 (Konstantinova et
al. 1992). Bas.: Jungermannia caucasica Váňa, Preslia 42: 96, 1970 (Váňa 1970a).
*** Solenostoma chenianum (C.Gao, Y.H.Wu et Grolle) Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 496, 2009 (Váňa and Long 2009). Bas.: Jungermannia cheniana
C.Gao, Y.H.Wu et Grolle, Nova Hedwigia 77 (1/2): 190, 2003 (Gao et al. 2003).
*** Solenostoma clavellatum Mitt. ex Steph., Bull. Herb. Boissier (sér. 2) 1 (5): 491
(53), 1901 (Stephani 1901a).
*** Solenostoma confertissimum (Nees) Schljakov, Novosti Sist. Nizš. Rast. 17: 239,
1980 (Shliakov 1980b). Bas.: Jungermannia confertissima Nees, Naturgesch. Eur.
Leberm. 1: 291, 1833 (Nees 1833c).
*** Solenostoma conilorum (Schifn.) Steph., Bull. Herb. Boissier (sér. 2) 1 (5): 497
(59), 1901 (Stephani 1901a). Bas.: Jungermannia conilora Schifn., Leberm.,
Forschungsr. Gazelle 4 (4): 10, 1890 (Schifner 1890).
*** Solenostoma crassulum (Nees et Mont.) Steph., Bull. Herb. Boissier (sér. 2) 1 (5):
497 (59), 1901 (Stephani 1901a). Bas.: Jungermannia crassula Nees et Mont.,
Ann. Sci. Nat. Bot. (sér. 2) 5: 54, 1836 (Nees and Montagne 1836).
** Solenostoma cryptogynum R.M.Schust. ex J.J.Engel, Novon 17 (3): 311, 2007
(Engel 2007). Based on: Solenostoma cryptogynum R.M.Schust., Beih. Nova Hedwigia 119: 380, 2002 (Schuster 2002b), nom. inval.
*** Solenostoma cyclops (S.Hatt.) R.M.Schust., Hepat. Anthocerotae N. Amer. 2: 945,
1969 (Schuster 1969b). Bas.: Jungermannia cyclops S.Hatt., J. Hattori Bot. Lab. 3:
5, 1948 [1950] (Hattori 1948b).
*** Solenostoma diversiclavellatum (Amakawa et Grolle) R.M.Schust. ex Váňa et
D.G.Long, Nova Hedwigia 89 (3/4): 501, 2009 (Váňa and Long 2009). Bas.:
Jungermannia diversiclavellata Amakawa et Grolle, J. Hattori Bot. Lab. 31: 107,
1968 (Amakawa 1968b).
*** Solenostoma dulongense Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 497, 2009
(Váňa and Long 2009).
World checklist of hornworts and liverworts
131
*** Solenostoma exsertum (A.Evans) Steph., Bull. Herb. Boissier (sér. 2) 1 (5): 490
(52), 1901 (Stephani 1901a). Bas.: Nardia exserta A.Evans, Trans. Connecticut
Acad. Arts 8 (15): 259, 1891 (Evans 1891).
*** Solenostoma faurieanum (Beauverd) R.M.Schust., Hepat. Anthocerotae N. Amer.
2: 945, 1969 (Schuster 1969b). Bas.: Jungermannia faurieana Beauverd, Sp. Hepat.
(Stephani) 6: 571, 1924 (Stephani 1924).
*** Solenostoma lagellare (Amakawa) Váňa et D.G.Long, Nova Hedwigia 89 (3/4):
501, 2009 (Váňa and Long 2009). Bas.: Jungermannia lagellaris Amakawa, J. Hattori Bot. Lab. 29: 258, 1966 (Amakawa 1966).
*** Solenostoma lavorevolutum (Váňa) Váňa et D.G.Long, Nova Hedwigia 89 (3/4):
501, 2009 (Váňa and Long 2009). Bas.: Jungermannia lavorevoluta Váňa, J. Hattori Bot. Lab. 36: 63, 1972 [1973] (Váňa 1972b).
*** Solenostoma grollei (D.G.Long et Váňa) K.Feldberg, Hentschel, Bombosch,
D.G.Long, Váňa et Heinrichs, Pl. Syst. Evol. 280 (3/4): 244, 2009 (Feldberg et
al. 2009). Bas.: Gottschelia grollei D.G.Long et Váňa, J. Bryol. 29 (3): 167, 2007
(Long and Váňa 2007).
*** Solenostoma grosseverrucosum (Amakawa et S.Hatt.) Váňa, Crand.-Stotl. et Stotler,
Syst. Bot. 40 (1): 39, 2015 (Shaw et al. 2015). Bas.: Horikawaella grosseverrucosa
Amakawa et S.Hatt., Bull. Univ. Mus. Univ. Tokyo 8: 216, 1975 (Hattori 1975e).
*** Solenostoma heterolimbatum (Amakawa) Váňa et D.G.Long, Nova Hedwigia 89
(3/4): 503, 2009 (Váňa and Long 2009). Bas.: Jungermannia heterolimbata Amakawa, J. Hattori Bot. Lab. 30: 183, 1967 (Amakawa 1967b).
*** Solenostoma hewsoniae (Amakawa et Grolle) R.M.Schust. ex Váňa, Hentschel et
Heinrichs, Cryptog. Bryol. 31 (2): 136, 2010 (Váňa et al. 2010a). Bas.: Jungermannia hewsoniae Amakawa et Grolle, J. Hattori Bot. Lab. 31: 108, 1968 (Amakawa 1968b).
*** Solenostoma hiugaense Amakawa, J. Jap. Bot. 31 (2): 47, 1956 (Amakawa 1956).
*** Solenostoma indrodayanum (Sushil K.Singh et D.K.Singh) Váňa et D.G.Long,
Nova Hedwigia 89 (3/4): 503, 2009 (Váňa and Long 2009). Bas.: Jungermannia indrodayana Sushil K.Singh et D.K.Singh, Cryptog. Bryol. 28 (2): 103, 2007
(Singh and Singh 2007b).
*** Solenostoma inundatum (Hook.f. et Taylor) Mitt. ex Steph., Bull. Herb. Boissier (sér. 2) 1 (5): 490 (52), 1901 (Stephani 1901a). Bas.: Jungermannia inundata
Hook.f. et Taylor, London J. Bot. 3: 559, 1844 (Hooker and Taylor 1844d).
*** Solenostoma javanicum (Schifn.) Steph., Bull. Herb. Boissier (sér. 2) 1 (5): 494
(56), 1901 (Stephani 1901a). Bas.: Aplozia javanica Schifn., Denkschr. Kaiserl.
Akad. Wiss., Math.-Naturwiss. Kl. 67: 193, 1898 (Schifner 1898a).
** Solenostoma kanaii (Amakawa) Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 503,
2009 (Váňa and Long 2009). Bas.: Jungermannia kanaii Amakawa, J. Hattori Bot.
Lab. 30: 194, 1967 (Amakawa 1967b).
*** Solenostoma kashyapii (S.C.Srivast., S.Srivast. et D.Sharma) Váňa et D.G.Long,
Nova Hedwigia 89 (3/4): 503, 2009 (Váňa and Long 2009). Bas.: Jungermannia
132
***
***
***
*
*
***
***
*
***
***
***
***
***
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
kashyapii S.C.Srivast., S.Srivast. et D.Sharma, Lindbergia 28 (3): 131, 2003 (Srivastava et al. 2003).
Solenostoma lanigerum (Mitt.) Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 503,
2009 (Váňa and Long 2009). Bas.: Jungermannia lanigera Mitt., J. Proc. Linn.
Soc., Bot. 5 (18): 91, 1860 [1861] (Mitten 1860c).
Solenostoma macrocarpum (Schifn. ex Steph.) Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 504, 2009 (Váňa and Long 2009). Bas.: Jungermannia macrocarpa
Schifn. ex Steph., Sp. Hepat. (Stephani) 6: 87, 1917 (Stephani 1917a).
Solenostoma mamatkulovii (Váňa et Zerov) Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31 (2): 137, 2010 (Váňa et al. 2010a). Bas.: Jungermannia mamatkulovii Váňa et Zerov, Preslia 49: 181, 1977 (Váňa 1977).
Solenostoma microphyllum C.Gao, Fl. Hepat. Chin. Boreali-Orient.: 206, 1981
(Gao and Chang 1981).
Solenostoma microrevolutum (C.Gao et X.L.Bai) Váňa et D.G.Long, Nova Hedwigia
89 (3/4): 505, 2009 (Váňa and Long 2009). Bas.: Jungermannia microrevoluta C.Gao
et X.L.Bai, Philipp. Scientist 38: 146, 2001 (Gao and Bai 2001).
Solenostoma mildbraedii (Steph.) R.M.Schust., Beih. Nova Hedwigia 119: 387,
2002 (Schuster 2002b). Bas.: Jungermannia mildbraedii Steph., Wiss. Ergebn.
Deut. Zentr.-Afr. Exped. (1907-08), Bot. 2: 113, 1911 (Stephani 1911a).
Solenostoma niveum (Grolle) R.M.Schust. ex Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31 (2): 137, 2010 (Váňa et al. 2010a). Bas.: Jungermannia nivea Grolle,
Misc. Bryol. Lichenol. 6 (1): 1, 1971 (Grolle 1971c).
Solenostoma novazelandiae R.M.Schust., Beih. Nova Hedwigia 119: 380, 2002
(Schuster 2002b). 94
Solenostoma ohbae (Amakawa) C.Gao, Bryol. Xizang: 495, 1985 (Li 1985). Bas.:
Jungermannia ohbae Amakawa, Bull. Univ. Mus. Univ. Tokyo 8: 218, 1975 (Hattori
1975e).
Solenostoma orbiculatum (Colenso) R.M.Schust., Beih. Nova Hedwigia 119: 380,
2002 (Schuster 2002b). Bas.: Gymnomitrion orbiculatum Colenso, Trans. & Proc.
New Zealand Inst. 18: 236, 1886 (Colenso 1886b).
Solenostoma parvitextum (Amakawa) Váňa et D.G.Long, Nova Hedwigia 89 (3/4):
505, 2009 (Váňa and Long 2009). Bas.: Jungermannia parvitexta Amakawa, J.
Hattori Bot. Lab. 30: 187, 1967 (Amakawa 1967b).
Solenostoma patoniae (Grolle, D.B.Schill et D.G.Long) K.Feldberg, Hentschel,
Bombosch, D.G.Long, Váňa et Heinrichs, Pl. Syst. Evol. 280 (3/4): 244, 2009
(Feldberg et al. 2009). Bas.: Gottschelia patoniae Grolle, D.B.Schill et D.G.Long,
J. Bryol. 25 (1): 3, 2003 (Grolle et al. 2003).
Solenostoma pocsii (Váňa) Bakalin, Arctoa 18: 160, 2009 [2010] (Bakalin and Vilnet 2009). Bas.: Jungermannia pocsii Váňa, Folia Geobot. Phytotax. 10 (4): 365,
1975 (Váňa 1975b).
94 Solenostoma novaezelandiae is possibly conspeciic with Solenostoma orbiculatum (Engel and Glenny
2008a).
World checklist of hornworts and liverworts
133
*** Solenostoma poeltii (Amakawa) Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 505,
2009 (Váňa and Long 2009). Bas.: Jungermannia poeltii Amakawa, J. Hattori Bot.
Lab. 29: 258, 1966 (Amakawa 1966).
*** Solenostoma pseudocyclops (Inoue) Váňa et D.G.Long, Nova Hedwigia 89 (3/4):
506, 2009 (Váňa and Long 2009). Bas.: Jungermannia pseudocyclops Inoue, Bull.
Natl. Sci. Mus. Tokyo (n.ser.) 9 (1): 37, 1966 (Inoue 1966a).
** Solenostoma pseudopyrilorum Bakalin et Vilnet, Arctoa 18: 159, 2009 [2010] (Bakalin and Vilnet 2009).
*** Solenostoma purpuratum (Mitt.) Steph., Bull. Herb. Boissier (sér. 2) 1 (5): 489
(51), 1901 (Stephani 1901a). Bas.: Jungermannia purpurata Mitt., J. Proc. Linn.
Soc., Bot. 5 (18): 91, 1860 [1861] (Mitten 1860c).
*** Solenostoma pyrilorum Steph., Sp. Hepat. (Stephani) 6: 83, 1917 (Stephani 1917a). 95
*** Solenostoma pyrilorum var. gracillimum (Amakawa) Váňa et D.G.Long, Nova
Hedwigia 89 (3/4): 506, 2009 (Váňa and Long 2009). Bas.: Jungermannia pyrilora var. gracillima Amakawa, Fl. E. Himalaya 2: 228, 1971 (Hattori 1971a).
** Solenostoma pyrilorum var. major (S.Hatt.) Bakalin, Arctoa 17: 230, 2008 [2009]
(Bakalin 2008a). Bas.: Jungermannia monticola f. major S.Hatt., J. Hattori Bot.
Lab. 3: 8, 1948 [1950] (Hattori 1948b).
*** Solenostoma pyrilorum var. minutissimum (Amakawa) Bakalin, Arctoa 16: 208,
2007 [2008] (Bakalin 2007). Bas.: Jungermannia pyrilora var. minutissima Amakawa, J. Hattori Bot. Lab. 22: 61, 1960 (Amakawa 1960b). 96
*** Solenostoma raujeanum (Grolle ex Amakawa) Váňa et D.G.Long, Nova Hedwigia
89 (3/4): 507, 2009 (Váňa and Long 2009). Bas.: Jungermannia raujeana Grolle
ex Amakawa, J. Hattori Bot. Lab. 29: 262, 1966 (Amakawa 1966).
*** Solenostoma ricleii Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 507, 2009 (Váňa
and Long 2009). Nom. nov. pro Jungermannia grollei Amakawa, J. Hattori Bot.
Lab. 29: 260, 1966 (Amakawa 1966).
*** Solenostoma sanguinolentum (Grif.) Steph., Bull. Herb. Boissier (sér. 2) 1 (5): 489
(51), 1901 (Stephani 1901a). Bas.: Jungermannia sanguinolenta Grif., Not. pl.
asiat. 2: 302, 1849 (Griith 1849).
*** Solenostoma schaulianum (Steph.) Váňa et D.G.Long, Nova Hedwigia 89 (3/4):
508, 2009 (Váňa and Long 2009). Bas.: Jungermannia schauliana Steph., Sp.
Hepat. (Stephani) 6: 90, 1917 (Stephani 1917a).
*** Solenostoma shimizuanum (S.Hatt. ex Váňa) Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31 (2): 138, 2010 (Váňa et al. 2010a). Bas.: Jungermannia shimizuana
S.Hatt. ex Váňa, J. Hattori Bot. Lab. 35: 315, 1972 (Váňa 1972a).
*** Solenostoma speciosum (Horik.) Hentschel, K.Feldberg, Bombosch, D.G.Long,
Váňa et Heinrichs, Pl. Syst. Evol. 280 (3/4): 244, 2009 (Feldberg et al. 2009).
Bas.: Anastrophyllum speciosum Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2,
Bot. 2: 147, 1934 (Horikawa 1934).
95 Solenostoma pyrilorum is a species complex (Shaw et al. 2015).
96 Solenostoma pyrilorum var. minutissimum was treated as a separate species, Solenostoma rishiriense
Amakawa by Bakalin and Vilnet (2009).
134
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Solenostoma speciosum subsp. villosum (R.M.Schust.) Hentschel, K.Feldberg, Bombosch, D.G.Long, Váňa et Heinrichs, Pl. Syst. Evol. 280 (3/4): 246, 2009 (Feldberg et al. 2009). Bas.: Scaphophyllum speciosum subsp. villosum R.M.Schust., Bryologist 101 (3): 434, 1998 (Schuster 1998b).
*** Solenostoma sphaerocarpum (Hook.) Steph., Bull. Herb. Boissier (sér. 2) 1 (5): 499
(61), 1901 (Stephani 1901a). Bas.: Jungermannia sphaerocarpa Hook., Brit. Jungermann.: tab. 74, 1815 (Hooker 1815).
*** Solenostoma stephanii (Schifn.) Steph., Bull. Herb. Boissier (sér. 2) 1 (5): 496 (58),
1901 (Stephani 1901a). Bas.: Aplozia stephanii Schifn., Denkschr. Kaiserl. Akad.
Wiss., Math.-Naturwiss. Kl. 67: 195, 1898 (Schifner 1898a).
*** Solenostoma stoloniferum (Steph.) S.W.Arnell, Hepat. South Africa: 316, 1963 (Arnell 1963b). Bas.: Nardia stolonifera Steph., Hedwigia 31 (3): 128, 1892 (Stephani
1892d).
*** Solenostoma strictum (Schifn.) Váňa, Hentschel et Heinrichs, Cryptog. Bryol. 31
(2): 138, 2010 (Váňa et al. 2010a). Bas.: Aplozia stricta Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 67: 194, 1898 (Schifner 1898a).
*** Solenostoma subacutum (Herzog) Váňa, Crand.-Stotl. et Stotler, Syst. Bot. 40 (1):
39, 2015 (Shaw et al. 2015). Bas.: Anastrophyllum subacutum Herzog, Ann. Bryol.
12: 75, 1939 (Herzog 1939b).
*** Solenostoma subrubrum (Schifn. ex Steph.) Váňa et D.G.Long, Nova Hedwigia 89
(3/4): 508, 2009 (Váňa and Long 2009). Bas.: Jungermannia subrubra Schifn. ex
Steph., Sp. Hepat. (Stephani) 6: 93, 1917 (Stephani 1917a).
* Solenostoma sunii Bakalin et Vilnet, Bot. Paciica 3 (2): 15, 2014 (Bakalin et al. 2014).
* Solenostoma totopapillosum (E.A.Hodgs.) R.M.Schust., Bryologist 100 (3): 366,
1997 (Schuster 1997a). Bas.: Jungermannia totopapillosa E.A.Hodgs., J. Roy. Soc.
New Zealand 2 (1): 111, 1972 (Hodgson 1972). 97
*** Solenostoma udarii (S.C.Srivast. et P.Singh) Váňa et D.G.Long, Nova Hedwigia
89 (3/4): 510, 2009 (Váňa and Long 2009). Bas.: Jungermannia udarii S.C.Srivast.
et P.Singh, Recent Stud. Indian Bryoph.: 152, 1995 (Srivastava and Singh 1995).
* Solenostoma ventroversum (Grolle) Váňa et D.G.Long, Nova Hedwigia 89 (3/4):
510, 2009 (Váňa and Long 2009). Bas.: Jungermannia ventroversa Grolle, Khumbu Himal 1 (4): 284, 1966 (Grolle 1966k). 98
*** Solenostoma zantenii (Amakawa) R.M.Schust. ex Váňa et D.G.Long, Nova Hedwigia 89 (3/4): 510, 2009 (Váňa and Long 2009). Bas.: Jungermannia zantenii
Amakawa, J. Hattori Bot. Lab. 31: 110, 1968 (Amakawa 1968b).
* Solenostoma zengii (C.Gao et X.L.Bai) Váňa et D.G.Long, Nova Hedwigia 89
(3/4): 510, 2009 (Váňa and Long 2009). Bas.: Jungermannia zengii C.Gao et
X.L.Bai, Philipp. Scientist 38: 151, 2001 (Gao and Bai 2001).
97 Solenostoma totopapillosum is possibly conspeciic with Solenostoma inundatum (Shaw et al. 2015).
98 Solenostoma ventroversum is possibly conspeciic with Solenostoma appressifolium.
World checklist of hornworts and liverworts
***
135
Southbyaceae Váňa, Crand.-Stotl., Stotler et D.G.Long
by J. Váňa
Southbyaceae has traditionally been placed in Arnelliaceae (e.g. Crandall-Stotler et al.
2009) or kept as a separate family (e.g. Vanden Berghen 1957) that has never been
validated. However, it was re-established, rearranged and validated in the study by
Váňa et al. (2012e).
*** Gongylanthus Nees, Naturgesch. Eur. Leberm. 2: 405, 1836 (Nees 1836).
*** Gongylanthus dusenii Steph., Bull. Herb. Boissier (sér. 2) 6 (5): 385 (41), 1906
(Stephani 1906a).
*** Gongylanthus ericetorum (Raddi) Nees, Naturgesch. Eur. Leberm. 2: 407, 1836
(Nees 1836). Bas.: Calypogeia ericetorum Raddi, Jungermanniogr. Etrusca: 31,
1818 (Raddi 1818a).
*** Gongylanthus granatensis (Gottsche) Steph., Bull. Herb. Boissier (sér. 2) 6 (5): 385
(41), 1906 (Stephani 1906a). Bas.: Lindigina granatensis Gottsche, Ann. Sci. Nat.
Bot. (sér. 5) 1: 138, 1864 (Gottsche 1864).
*** Gongylanthus himalayensis Grolle, Khumbu Himal 1 (4): 285, 1966 (Grolle
1966k).
*** Gongylanthus javanicus Grolle, J. Jap. Bot. 40 (7): 206, 1965 (Grolle 1965d).
*** Gongylanthus liebmanianus (Lindenb. et Gottsche) Steph., Bull. Herb. Boissier
(sér. 2) 6 (5): 388 (44), 1906 (Stephani 1906a). Bas.: Gymnanthe liebmaniana
Lindenb. et Gottsche, Syn. Hepat. 5: 712, 1847 (Gottsche et al. 1847).
*** Gongylanthus limbatus (Herzog) Grolle et Váňa, Folia Geobot. Phytotax. 9 (2):
198, 1974 (Váňa 1974c). Bas.: Aplozia limbata Herzog, Beih. Bot. Centralbl. 61B
(3): 561, 1942 (Herzog 1942d).
*** Gongylanthus muelleri (Gottsche) Steph., Bull. Herb. Boissier (sér. 2) 6 (5): 388
(44), 1906 (Stephani 1906a). Bas.: Lindigia muelleri Gottsche, Mexik. Leverm.:
121, 1863 (Gottsche 1863).
** Gongylanthus oniscoides (Spruce) Steph., Bull. Herb. Boissier (sér. 2) 6 (5): 386
(42), 1906 (Stephani 1906a). Bas.: Calypogeia oniscoides Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 448, 1885 (Spruce 1885).
*** Gongylanthus richardsii E.W.Jones, Trans. Brit. Bryol. Soc. 4 (4): 650, 1964 (Jones
1964).
*** Southbya Spruce, Ann. Mag. Nat. Hist. (ser. 2) 3 (18): 501, 1849 (Spruce 1849).
*** Southbya gollanii Steph., Bull. Herb. Boissier (sér. 2) 6 (5): 381 (37), 1906 (Stephani
1906a).
*** Southbya nigrella (De Not.) Henriq., Bol. Soc. Brot. 4: 244, 1886 [1887] (Henriques 1886). Bas.: Jungermannia nigrella De Not., Mem. Reale Accad. Sci. Torino
(ser. 2) 1: 315, 1838 (De Notaris 1838).
136
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Southbya organensis Herzog, Memoranda Soc. Fauna Fl. Fennica 25: 53, 1950
(Herzog 1950c).
*** Southbya tophacea (Spruce) Spruce, Ann. Mag. Nat. Hist. (ser. 2) 3 (18): 501,
1849 (Spruce 1849). Bas.: Jungermannia tophacea Spruce, Hep. Pyr. Exsic.: no.
23, 1847 (Spruce 1847).
**
Stephaniellaceae R.M.Schust.
by J. Váňa
In the absence of molecular data, the phylogenetic ainities of Stephaniella and Stephaniellidium remain equivocal, but there is no evidence to support their placement in
either the Arnelliaceae or Southbyaceae where they have been placed recently. hus the
family as construed by Schuster (2002b) is retained.
*** Stephaniella J.B.Jack, Hedwigia 33 (1): 11, 1894 (Jack 1894).
* Stephaniella boliviensis Steph., Biblioth. Bot. 87 (2): 182, 1916 (Stephani 1916a).
*** Stephaniella hamata Steph., Bull. Herb. Boissier (sér. 2) 1 (10): 1024 (87), 1901
(Stephani 1901c).
*** Stephaniella paraphyllina J.B.Jack, Hedwigia 33 (1): 11, 1894 (Jack 1894).
*** Stephaniella rostrata U.Schmitt, Österr. Bot. Z. 115 (2): 124, 1968 (Schmitt and
Winkler 1968).
*** Stephaniella uncifolia S.Winkl., Österr. Bot. Z. 115 (2): 124, 1968 (Schmitt and
Winkler 1968).
** Stephaniellidium S.Winkl. ex Grolle, Acta Bot. Fenn. 121: 38, 1983 (Grolle
1983b). Based on: Stephaniellidium S.Winkl., Mitt. Inst. Colombo-Alemán Invest. Ci. 3: 60, 1969 (Winkler 1969).
*** Stephaniellidium sleumeri (Müll.Frib.) S.Winkl. ex Grolle, Acta Bot. Fenn. 121:
38, 1983 (Grolle 1983b). Bas.: Stephaniella sleumeri Müll.Frib., Rev. Bryol. Lichénol. 20 (1/2): 177, 1951 (Müller 1951b).
***
Trichotemnomataceae R.M.Schust.
*** Trichotemnoma R.M.Schust., Nova Hedwigia 15: 440, 1968 (Schuster 1968b).
*** Trichotemnoma corrugatum (Steph.) R.M.Schust., Nova Hedwigia 15: 440, 1968
(Schuster 1968b). Bas.: Blepharostoma corrugatum Steph., Hedwigia 32 (5): 315,
1893 (Stephani 1893d).
World checklist of hornworts and liverworts
137
Lophocoleineae Schljakov
***
Blepharostomataceae W.Frey et M.Stech
he recognition of Blepharostomataceae follows Frey and Stech (2008) following reconsideration of molecular and morphological data available at that time.
*** Blepharostoma (Dumort.) Dumort., Recueil Observ. Jungerm.: 18, 1835 (Dumortier 1835). Bas.: Jungermannia sect. Blepharostoma Dumort., Syll. Jungerm.
Europ.: 65, 1831 (Dumortier 1831).
** Blepharostoma arachnoideum M.Howe, Mem. Torrey Bot. Club 7: 140, 1899
(Howe 1899).
** Blepharostoma indicum G.Asthana, M.Saxena et Maurya, J. Bryol. 35 (4): 267,
2013 (Asthana et al. 2013).
** Blepharostoma minor Horik., Hikobia 1 (2): 104, 1951 [1952] (Horikawa 1951b).
*** Blepharostoma trichophyllum (L.) Dumort., Recueil Observ. Jungerm.: 18, 1835 (Dumortier 1835). Bas.: Jungermannia trichophylla L., Sp. Pl. 1: 1135, 1753 (Linnaeus 1753).
** Blepharostoma trichophyllum subsp. brevirete (Bryhn et Kaal.) R.M.Schust., Bull.
Natl. Mus. Canada 164: 16, 1959 (Schuster et al. 1959). Bas.: Blepharostoma
trichophyllum var. brevirete Bryhn et Kaal., Rep. Second Norweg. Arctic Exped.
11: 46, 1906 (Bryhn 1906).
*** Brevianthaceae J.J.Engel et R.M.Schust.
by L. Söderström, J. Váňa, R. Stotler, B.J. Crandall-Stotler and J.J. Engel
he treatment of Brevianthaceae follows Söderström et al. (2013b).
*** Brevianthus J.J.Engel et R.M.Schust., Phytologia 47 (4): 317, 1981 (Engel and
Schuster 1981).
*** Brevianthus lavus (Grolle) J.J.Engel et R.M.Schust., Phytologia 47 (4): 318, 1981
(Engel and Schuster 1981). Bas.: Jackiella lava Grolle, J. Hattori Bot. Lab. 33:
222, 1970 (Grolle 1970b).
** Brevianthus lavus subsp. crenulatus J.J.Engel, Nova Hedwigia 93 (3/4): 406, 2011
(Engel 2011).
* Brevianthus hypocanthidium M.A.M.Renner et J.J.Engel, PhytoKeys 50: 46, 2015
(Renner et al. 2015).
*** Tetracymbaliella Grolle, Nova Hedwigia 3 (1): 48, 1961 (Grolle 1961b).
** Tetracymbaliella comptonii (Pearson) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 164,
1963 [1964] (Grolle 1963d). Bas.: Chiloscyphus comptonii Pearson, J. Linn. Soc.,
Bot. 46 (305): 23, 1922 (Pearson 1922b).
*** Tetracymbaliella cymbalifera (Hook.f. et Taylor) Grolle, Nova Hedwigia 3 (1):
50, 1961 (Grolle 1961b). Bas.: Jungermannia cymbalifera Hook.f. et Taylor, Bot.
Antarct. Voy. I (Fl. Antarct. 1): 151, 1845 (Taylor and Hooker 1845).
138
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Tetracymbaliella decipiens (Gottsche) Grolle, Nova Hedwigia 3 (1): 49, 1961
(Grolle 1961b). Bas.: Chiloscyphus decipiens Gottsche, Syn. Hepat. 2: 176, 1845
(Gottsche et al. 1845a).
*** Tetracymbaliella subsimplex (Austin) J.J.Engel, Phytotaxa 207 (2): 185, 2015 (Engel 2015b). Bas.: Polyotus subsimplex Austin, Bull. Torrey Bot. Club 6 (7): 46,
1875 (Austin 1875c).
**
Chonecoleaceae R.M.Schust. ex Grolle
** Chonecolea Grolle, Rev. Bryol. Lichénol. 25 (3/4): 294, 1956 [1957] (Grolle 1956).
** Chonecolea acutiloba (Schifn.) R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 321,
1980 (Schuster 1980c). Bas.: Clasmatocolea acutiloba Schifn., Österr. Akad. Wiss.,
Math.-Naturwiss. Kl., Denkschr. 111: 63, 1964 (Schifner and Arnell 1964).
** Chonecolea andina Grolle et Váňa, J. Hattori Bot. Lab. 48: 229, 1980 (Váňa 1980).
** Chonecolea doellingeri (Nees) Grolle, Rev. Bryol. Lichénol. 25 (3/4): 295, 1956
[1957] (Grolle 1956). Bas.: Jungermannia doellingeri Nees, Syn. Hepat. 1: 104,
1844 (Gottsche et al. 1844).
** Chonecolea ruwenzorensis E.W.Jones, J. Bryol. 13 (4): 498, 1986 (Jones 1986).
** Chonecolea schusteri Udar et Ad.Kumar, Bryologist 85 (3): 315, 1982 (Udar and
Kumar 1982a).
** Chonecolea verae Potemkin, Proc. int. meeting 90th anniv. Abramova: 165, 2005
(Potemkin 2005).
**
Grolleaceae Solari ex R.M.Schust.
** Grollea R.M.Schust., Nova Hedwigia 8 (3/4): 288, 1964 (Schuster 1964b).
** Grollea antheliopsis R.M.Schust., Nova Hedwigia 8 (3/4): 288, 1964 (Schuster
1964b).
***
Herbertaceae Müll.Frib. ex Fulford et Hatcher
by D. Bell
*** Herbertus Gray, Nat. Arr. Brit. Pl. 1: 705, 1821 (Gray 1821). 99
*** Herbertus aduncus (Dicks.) Gray, Nat. Arr. Brit. Pl. 1: 705, 1821 (Gray 1821). Bas.:
Jungermannia adunca Dicks., Fasc. Pl. Crypt. Brit. 3: 12, 1793 (Dickson 1793).
99 Herbertus includes Schisma, but a few taxa have neither been transferred nor synonymized. hey are
listed in the “Names in genera not currently accepted” section below.
World checklist of hornworts and liverworts
139
** Herbertus arcticus (Inoue et Steere) Schljakov, Novosti Sist. Nizš. Rast. 19: 209,
1982 (Shliakov 1982). Bas.: Herbertus sakuraii subsp. arcticus Inoue et Steere, J.
Hattori Bot. Lab. 44: 266, 1978 (Steere and Inoue 1978).
** Herbertus armitanus (Steph.) H.A.Mill., J. Hattori Bot. Lab. 28: 324, 1965 (Miller
1965). Bas.: Schisma armitanum Steph., Sp. Hepat. (Stephani) 4: 28, 1909 (Stephani
1909d).
** Herbertus asparus Tixier, Bryophyt. Biblioth. 18: 64, 1979 (Tixier 1979a).
*** Herbertus bivittatus Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 343, 1885
(Spruce 1885).
** Herbertus borealis Crundw., Trans. Brit. Bryol. Soc. 6 (1): 41, 1970 (Crundwell 1970).
** Herbertus buchii Juslén, Ann. Bot. Fenn. 43 (6): 416, 2006 (Juslén 2006a).
** Herbertus ceylanicus (Steph.) Abeyw., Ceylon J. Sci., Biol. Sci. 2 (1): 43, 1959
(Abeywickrama 1959). Bas.: Schisma ceylanicum Steph., Sp. Hepat. (Stephani) 4:
22, 1909 (Stephani 1909d).
** Herbertus circinatus (Steph.) H.A.Mill., J. Hattori Bot. Lab. 28: 318, 1965 (Miller 1965). Bas.: Schisma circinatum Steph., Sp. Hepat. (Stephani) 4: 25, 1909
(Stephani 1909d).
*** Herbertus delavayi (Steph.) Steph., Hedwigia 34 (2): 43, 1895 (Stephani 1895c).
Bas.: Schisma delavayi Steph., Mém. Soc. Nat. Sci. Nat. Math. Cherbourg 29: 228,
1894 (Stephani 1894b). 100
*** Herbertus dicranus (Gottsche, Lindenb. et Nees) Trevis., Mem. Reale Ist. Lombardo
Sci. (Ser. 3), C. Sci. Mat. 4 (13): 397, 1877 (Trevisan 1877). Bas.: Sendtnera dicrana
Gottsche, Lindenb. et Nees, Syn. Hepat. 2: 239, 1845 (Gottsche et al. 1845a).
** Herbertus durandii (Steph.) Herzog, Rev. Bryol. Lichénol. 11 (1): 25, 1938 [1939]
(Herzog 1938a). Bas.: Schisma durandii Steph., Trans. Linn. Soc. London, Bot. 6
(1): 99, 1901 (Stephani 1901e).
* Herbertus evittatus (Steph.) H.A.Mill., J. Hattori Bot. Lab. 28: 327, 1965 (Miller 1965). Bas.: Schisma evittatum Steph., Sp. Hepat. (Stephani) 6: 357, 1922
(Stephani 1922).
** Herbertus gaochienii X.Fu, Fl. Bryoph. Sin. 9: 38, 2003 (Gao 2003).
** Herbertus gracilis (Mont.) Steph., Bull. Herb. Boissier 5 (10): 842, 1897 (Stephani
1897c). Bas.: Mastigophora gracilis Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 254,
1843 (Montagne 1843).
** Herbertus guangdongii P.J.Lin ex Piippo, Bryobrothera 1: 206, 1992 (Lin et al. 1992).
** Herbertus hawaiiensis H.A.Mill., J. Hattori Bot. Lab. 28: 317, 1965 (Miller 1965).
** Herbertus helleri (Steph.) W.E.Nicholson, Rev. Bryol. Lichénol. 13: 143, 1942
(Nicholson 1942). Bas.: Schisma helleri Steph., Sp. Hepat. (Stephani) 4: 29, 1909
(Stephani 1909d).
** Herbertus herpocladioides E.B.Scott et H.A.Mill., Bryologist 62 (2): 116, 1959
(Scott and Miller 1959).
100 Herbertus delavayi is a species complex that requires further studies (Bell et al. 2012). It also includes
Herbertus borealis and may include several, partly cryptic species.
140
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Herbertus hutchinsiae (Gottsche et Rabenh.) A.Evans, Bull. Torrey Bot. Club 44
(4): 214, 1917 (Evans 1917c). Bas.: Sendtnera adunca var. hutchinsiae Gottsche et
Rabenh., Hepat. Eur., Leberm. 21-22: no. 210, 1862 (Rabenhorst 1862).
*** Herbertus juniperoideus (Sw.) Grolle, Rev. Bryol. Lichénol. 30 (1/2): 80, 1961
(Grolle 1961a). Bas.: Jungermannia juniperoidea Sw., Prodr. (Swartz): 144, 1788
(Swartz 1788). 101
*** Herbertus juniperoideus subsp. acanthelius (Spruce) K.Feldberg et Heinrichs, Bot.
J. Linn. Soc. 151: 326, 2006 (Feldberg and Heinrichs 2006). Bas.: Herbertus acanthelius Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 341, 1885 (Spruce 1885).
*** Herbertus juniperoideus subsp. pensilis (Taylor) K.Feldberg et Heinrichs, Bot. J.
Linn. Soc. 151: 329, 2006 (Feldberg and Heinrichs 2006). Bas.: Sendtnera pensilis
Taylor, London J. Bot. 5: 372, 1846 (Taylor 1846b).
** Herbertus kurzii (Steph.) R.S.Chopra, J. Indian Bot. Soc. 22: 247, 1943 (Chopra 1943).
Bas.: Schisma kurzii Steph., Sp. Hepat. (Stephani) 4: 24, 1909 (Stephani 1909d).
** Herbertus leratii (Steph.) H.A.Mill., J. Hattori Bot. Lab. 28: 327, 1965 (Miller 1965).
Bas.: Schisma leratii Steph., Sp. Hepat. (Stephani) 6: 360, 1922 (Stephani 1922).
** Herbertus lonchobasis H.A.Mill., J. Hattori Bot. Lab. 28: 306, 1965 (Miller 1965).
** Herbertus longiissus Steph., Hedwigia 34 (2): 44, 1895 (Stephani 1895c).
** Herbertus longispinus J.B.Jack et Steph., Hedwigia 31 (1): 15, 1892 (Jack and
Stephani 1892).
** Herbertus mauritianus N.G.Hodgetts, J. Bryol. 30 (4): 247, 2008 (Hodgetts 2008).
*** Herbertus norenus D.G.Long, D.Bell et H.H.Blom, Molec. Ecol. Res. 12: 44,
2012 (Bell et al. 2012).
** Herbertus oldieldianus (Steph.) Rodway, Tasm. Bryoph.: 72, 1917 (Rodway
1917b). Bas.: Schisma oldieldianum Steph., Sp. Hepat. (Stephani) 4: 20, 1909
(Stephani 1909d).
** Herbertus pilifer Schifn., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur.
60 (2): 252, 1893 (Schifner 1893a).
** Herbertus pocsii N.G.Hodgetts, J. Bryol. 30 (4): 249, 2008 (Hodgetts 2008).
** Herbertus pumilus Steph., Hedwigia 34 (2): 44, 1895 (Stephani 1895c).
** Herbertus ramosus (Steph.) H.A.Mill., J. Hattori Bot. Lab. 28: 314, 1965 (Miller
1965). Bas.: Schisma ramosum Steph., Sp. Hepat. (Stephani) 4: 23, 1909 (Stephani
1909d).
** Herbertus runcinatus (Taylor) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 397, 1877 (Trevisan 1877). Bas.: Sendtnera runcinata Taylor,
London J. Bot. 5: 372, 1846 (Taylor 1846b).
*** Herbertus sendtneri (Nees) Lindb., Hepat. Scand. Exsicc.: no. 4, 1874 (Lindberg
and Lackström 1874). Bas.: Schisma sendtneri Nees, Naturgesch. Eur. Leberm. 3:
575, 1838 (Nees 1838b).
** Herbertus spicatus N.G.Hodgetts, J. Bryol. 30 (4): 244, 2008 (Hodgetts 2008).
101 Herbertus juniperoideus is here treated with two subspecies, but they may deserve species rank instead.
World checklist of hornworts and liverworts
141
*** Herbertus stramineus (Dumort.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 397, 1877 (Trevisan 1877). Bas.: Schisma stramineum Dumort., Syll. Jungerm. Europ.: 77, 1831 (Dumortier 1831).
** Herbertus streimannii M.L.So, Syst. Bot. 28 (1): 13, 2003 (So 2003).
*** Herbertus tenuis A.Evans, Bull. Torrey Bot. Club 44 (4): 219, 1917 (Evans 1917c).
** Herbertus udarii D.Kumar et N.Manocha, Geophytology 29 (1/2): 105, 1999
[2000] (Kumar and Manocha 1999).
*
Excluded from the genus
Herbertus subrotundatus X.Fu et Y.J.Yi, Acta Phytotax. Sin. 39 (1): 89, 2001 (Yi et
al. 2001). 102
*** Triandrophyllum Fulford et Hatcher, Bryologist 64 (4): 349, 1961 [1962]
(Fulford and Hatcher 1961). Based on: Triandrophyllum Fulford et Hatcher, Bryologist 61 (4): 277, 1958 [1959] (Fulford and Hatcher 1958).
*** Triandrophyllum eophyllum (R.M.Schust.) Gradst., Mem. New York Bot. Gard.
86: 104, 2001 (Gradstein et al. 2001a). Bas.: Olgantha eophylla R.M.Schust., Nova
Hedwigia 63 (3/4): 535, 1996 (Schuster 1996b).
** Triandrophyllum fernandeziense (S.W.Arnell) Grolle ex Fulford et Hatcher, Bryologist 64 (4): 351, 1961 [1962] (Fulford and Hatcher 1961). Bas.: Acromastigum
fernandeziense S.W.Arnell, Ark. Bot. (n.ser.) 4 (1): 10, 1957 (Arnell 1957b).
** Triandrophyllum heterophyllum (Steph.) Grolle, J. Jap. Bot. 39 (8): 238, 1964
(Grolle 1964g). Bas.: Mastigophora heterophylla Steph., Sp. Hepat. (Stephani) 6:
367, 1922 (Stephani 1922).
*** Triandrophyllum subtriidum (Hook.f. et Taylor) Fulford et Hatcher, Bryologist
64 (4): 350, 1961 [1962] (Fulford and Hatcher 1961). Bas.: Jungermannia subtriida Hook.f. et Taylor, London J. Bot. 3: 579, 1844 (Hooker and Taylor 1844c).
** Triandrophyllum subtriidum var. triidum (Gottsche) Solari, Bol. Soc. Argent. Bot.
15 (2/3): 201, 1973 (Solari 1973). Bas.: Sendtnera triida Gottsche, Ann. Sci. Nat.
Bot. (sér. 5) 1: 142, 1864 (Gottsche 1864).
** Triandrophyllum symmetricum J.J.Engel, Haussknechtia, Beih. 9: 115, 1999 (Engel
1999b).
***
Lepicoleaceae R.M.Schust.
by M. von Konrat
Lepicoleaceae has conventionally been considered a monogeneric family only containing Lepicolea (e.g. Crandall-Stotler et al. 2009). However, we expand Lepicoleaceae
here to include the monogeneric family Vetaformaceae. A number of molecular analy102 Herbertus subrotundus does not belong in Herbertus (Juslén 2006a), but it is unclear where it belongs.
142
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
ses have shown that Vetaforma dusenii is sister to Lepicolea (e.g. He-Nygrén et al. 2006,
Juslén 2006b) together forming a robust clade.
*** Lepicolea Dumort., Recueil Observ. Jungerm.: 20, 1835 (Dumortier 1835).
*** Lepicolea attenuata (Mitt.) Steph., J. Linn. Soc., Bot. 29 (201): 276, 1892 (Stephani
1892b). Bas.: Sendtnera attenuata Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 153,
1854 (Mitten 1854).
*** Lepicolea magellanica (Gola) Solari, Lindbergia 9 (2): 86, 1983 (Solari 1983b).
Bas.: Lepicolea scolopendra var. magellanica Gola, Nuovo Giorn. Bot. Ital. (n.ser.)
29 (1/4): 170, 1922 [1923] (Gola 1922).
*** Lepicolea norrisii Piippo, Ann. Bot. Fenn. 25 (1): 55, 1988 (Piippo 1988).
*** Lepicolea ochroleuca (Spreng.) Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 345,
1885 (Spruce 1885). Bas.: Jungermannia ochroleuca Spreng. Syst. Veg. (ed. 16)
[Sprengel] 4 (2): 325, 1827 (Sprengel 1827b).
*** Lepicolea pruinosa (Taylor) Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 345,
1885 (Spruce 1885). Bas.: Sendtnera pruinosa Taylor, London J. Bot. 5: 373, 1846
(Taylor 1846b).
*** Lepicolea ramentiissa Herzog, Biblioth. Bot. 88: 30, 1920 [1921] (Herzog 1920).
*** Lepicolea rara (Steph.) Grolle, Nova Hedwigia 16: 152, 1968 (Grolle 1968d). Bas.:
Lepidozia rara Steph., Sp. Hepat. (Stephani) 3: 618, 1909 (Stephani 1909a). 103
*** Lepicolea rigida (De Not.) E.B.Scott, Nova Hedwigia 2: 148, 1960 (Scott 1960).
Bas.: Sendtnera rigida De Not., Mem. Reale Accad. Sci. Torino (ser. 2) 16: 229,
1857 (De Notaris 1857).
*** Lepicolea scolopendra (Hook.) Dumort. ex Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 398, 1877 (Trevisan 1877). Bas.: Jungermannia scolopendra Hook., Musci Exot. 1: tab. 40, 1818 (Hooker 1818).
*** Lepicolea yakusimensis (S.Hatt.) S.Hatt., J. Hattori Bot. Lab. 10: 42, 1953 (Hattori
1953c). Bas.: Lepicolea scolopendra var. yakusimensis S.Hatt., J. Hattori Bot. Lab. 2:
9, 1947 [1948] (Hattori 1947b).
*** Vetaforma Fulford et J.Taylor, Nova Hedwigia 4 (1/2): 81, 1962 (Fulford 1962b).
*** Vetaforma dusenii (Steph.) Fulford et J.Taylor, Nova Hedwigia 4 (1/2): 82,
1962 (Fulford 1962b). Bas.: Lepidozia dusenii Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26 (III, 6): 52, 1900 (Stephani 1900b).
***
Lepidoziaceae Limpr.
by E.D. Cooper
he classiication within Lepidoziaceae is in a state of lux. he treatment provided here
follows the interim classiication proposed by Cooper (2013) based on recent molecular
phylogenetic studies (Heslewood and Brown 2007, Cooper et al. 2011, 2012a, 2012b)
103 Lepicolea rara is possibly a species complex and warrants further investigation (Schuster 2000a).
World checklist of hornworts and liverworts
143
incorporating nomenclatural changes from Cooper et al. (2013). Several higher taxa are
unlikely to represent monophyletic units, but the phylogenetic data currently available
are insuicient to re-circumscribe the doubtful subfamilies and genera (Cooper 2013).
Infrageneric taxa have been retained where phylogenetic data are absent or inconclusive.
** Meinungeria Frank Müll., Bryologist 110 (3): 494, 2007 (Müller 2007).
** Meinungeria mouensis Frank Müll., Bryologist 110 (3): 494, 2007 (Müller 2007).
***
Bazzanioideae Rodway
*** Acromastigum A.Evans, Bull. Torrey Bot. Club 27 (3): 103, 1900 (Evans 1900b). 104
** subg. Acromastigum
*** Acromastigum caledonicum (Steph.) Grolle, Österr. Bot. Z. 111 (2/3): 243, 1964
(Grolle 1964h). Bas.: Acolea caledonica Steph., Nova Caledonia, Bot. 1: 19, 1914
(Stephani 1914c).
*** Acromastigum cavifolium R.M.Schust., J. Hattori Bot. Lab. 26: 257, 1963 (Schuster
1963b).
** Acromastigum herzogii Grolle, Österr. Bot. Z. 111 (2/3): 250, 1964 (Grolle 1964h).
*** Acromastigum homodictyon (Herzog) Grolle, Österr. Bot. Z. 111 (2/3): 245, 1964
(Grolle 1964h). Bas.: Acromastigum integrifolium var. homodictyon Herzog, Ark.
Bot. (n.ser.) 3 (3): 44, 1953 (Herzog 1953a).
** Acromastigum integrifolium (Austin) A.Evans, Bull. Torrey Bot. Club 27 (3): 103,
1900 (Evans 1900b). Bas.: Mastigobryum integrifolium Austin, Bot. Bull. (Hanover) 1 (7): 32, 1876 (Austin 1876b).
*** Acromastigum stellare N.Kitag., Acta Phytotax. Geobot. 36 (4/6): 109, 1985 (Kitagawa 1985).
** Acromastigum stenophyllum R.M.Schust., Nova Hedwigia 15: 465, 1968 (Schuster
1968b).
*** Acromastigum verticale (Steph.) E.A.Hodgs., Trans. Roy. Soc. New Zealand 82
(1): 18, 1954 (Hodgson 1954). Bas.: Bazzania verticalis Steph., Hedwigia 32 (4):
214, 1893 (Stephani 1893c).
** subg. Inaequilatera (Schifn.) Grolle, J. Hattori Bot. Lab. 44: 2, 1978 (Grolle
1978b). Bas.: Bazzania sect. Inaequilaterae Schifn., Hepat. (Engl.-Prantl): 101,
1893 (Schifner 1893b).
*** Acromastigum adaptatum Hürl., Bauhinia 7 (4): 263, 1983 (Hürlimann 1983).
*** Acromastigum anisostomum (Lehm. et Lindenb.) A.Evans, Ann. Bryol., Suppl. 3:
48, 1934 (Evans 1934). Bas.: Jungermannia anisostoma Lehm. et Lindenb., Nov.
Stirp. Pug. 6: 57, 1834 (Lehmann 1834).
104 Acromastigum is here divided into two subgenera which are supported by recent molecular studies
(Heslewood and Brown 2007, Cooper et al. 2011).
144
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Acromastigum anisostomum var. minutum E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot. 3 (4): 71, 1965 (Hodgson 1965).
** Acromastigum aurescens A.Evans, Ann. Bryol., Suppl. 3: 45, 1934 (Evans 1934).
** Acromastigum bancanum (Sande Lac.) A.Evans, Ann. Bryol., Suppl. 3: 20, 1934
(Evans 1934). Bas.: Mastigobryum bancanum Sande Lac., Ann. Mus. Bot. Lugduno-Batavi 1: 301, 1864 (Sande Lacoste 1864).
** Acromastigum brotheri (Steph.) A.Evans, Ann. Bryol., Suppl. 3: 70, 1934 (Evans
1934). Bas.: Mastigobryum brotheri Steph., Sp. Hepat. (Stephani) 3: 536, 1909
(Stephani 1909a).
** Acromastigum capillare (Steph.) A.Evans, Ann. Bryol., Suppl. 3: 37, 1934 (Evans
1934). Bas.: Mastigobryum capillare Steph., Sp. Hepat. (Stephani) 6: 457, 1924
(Stephani 1924).
*** Acromastigum colensoanum (Mitt.) A.Evans ex Reimers, Hedwigia 73 (3/4): 142,
1933 (Reimers 1933). Bas.: Mastigobryum colensoanum Mitt., Bot. antarct. voy. II
(Fl. Nov.-Zel. 2): 147, 1854 (Mitten 1854).
*** Acromastigum cunninghamii (Steph.) A.Evans, Ann. Bryol., Suppl. 3: 106, 1934
(Evans 1934). Bas.: Bazzania cunninghamii Steph., Hedwigia 32 (4): 205, 1893
(Stephani 1893c).
** Acromastigum curtilobum A.Evans, Ann. Bryol., Suppl. 3: 97, 1934 (Evans 1934).
*** Acromastigum divaricatum (Nees) A.Evans ex Reimers, Hedwigia 73 (3/4): 142,
1933 (Reimers 1933). Bas.: Mastigobryum divaricatum Nees, Syn. Hepat. 2: 219,
1845 (Gottsche et al. 1845a).
** Acromastigum echinatiforme (De Not.) A.Evans, Ann. Bryol., Suppl. 3: 64, 1934
(Evans 1934). Bas.: Mastigobryum echinatiforme De Not., Epat. Borneo: 38, 1874
(De Notaris 1874).
* Acromastigum echinatum (Gottsche) A.Evans, Ann. Bryol., Suppl. 3: 147, 1934
(Evans 1934). Bas.: Mastigobryum echinatum Gottsche, Syn. Hepat. 2: 218, 1845
(Gottsche et al. 1845a). 105
*** Acromastigum exiguum (Steph.) A.Evans, Ann. Bryol., Suppl. 3: 75, 1934 (Evans
1934). Bas.: Mastigobryum exiguum Steph., Hedwigia 25 (1): 6, 1886 (Stephani
1886e).
** Acromastigum exile (Lindenb.) A.Evans, Ann. Bryol., Suppl. 3: 24, 1934 (Evans
1934). Bas.: Mastigobryum exile Lindenb., Syn. Hepat. 2: 217, 1845 (Gottsche et
al. 1845a).
*** Acromastigum ilum (Steph.) A.Evans, Ann. Bryol., Suppl. 3: 31, 1934 (Evans
1934). Bas.: Bazzania ilum Steph., Hedwigia 32 (4): 206, 1893 (Stephani 1893c).
*** Acromastigum ilum var. papillosum N.Kitag., Acta Phytotax. Geobot. 36 (4/6):
112, 1985 (Kitagawa 1985).
** Acromastigum imbriatum (Steph.) A.Evans, Ann. Bryol., Suppl. 3: 153, 1934 (Evans 1934). Bas.: Mastigobryum imbriatum Steph., Sp. Hepat. (Stephani) 3: 538,
1909 (Stephani 1909a).
105 Acromastigum echinatum may be conspeciic with Acromastigum inaequilaterum since all specimens
identiied by Piippo et al. (2002) belong there, but she did not see the type specimen.
World checklist of hornworts and liverworts
145
** Acromastigum furcatifolium (Steph.) E.A.Br., Phytotaxa 65: 58, 2012 (Brown et al.
2012). Bas.: Lepidozia furcatifolia Steph., J. & Proc. Roy. Soc. New South Wales
48 (1/2): 112, 1914 (Stephani and Watts 1914).
*** Acromastigum inaequilaterum (Lehm. et Lindenb.) A.Evans, Ann. Bryol., Suppl.
3: 129, 1934 (Evans 1934). Bas.: Jungermannia inaequilatera Lehm. et Lindenb.,
Nov. Stirp. Pug. 6: 56, 1834 (Lehmann 1834).
** Acromastigum interstisiale E.A.Br. et M.A.M.Renner, Telopea 17: 268, 2014
(Brown and Renner 2014).
** Acromastigum laetevirens (Sande Lac. ex Steph.) A.Evans, Ann. Bryol., Suppl. 3:
94, 1934 (Evans 1934). Bas.: Mastigobryum laetevirens Sande Lac. ex Steph., Hedwigia 25 (4): 133, 1886 (Stephani 1886b).
** Acromastigum laevigatum A.Evans, Ann. Bryol., Suppl. 3: 101, 1934 (Evans 1934).
** Acromastigum linganum (De Not.) A.Evans, Ann. Bryol., Suppl. 3: 118, 1934
(Evans 1934). Bas.: Mastigobryum linganum De Not., Epat. Borneo: 37, 1874 (De
Notaris 1874).
** Acromastigum lobuliferum A.Evans, Ann. Bryol., Suppl. 3: 157, 1934 (Evans 1934).
** Acromastigum longirete Grolle, J. Hattori Bot. Lab. 44: 11, 1978 (Grolle 1978b).
*** Acromastigum mooreanum (Steph.) E.A.Hodgs., Trans. Roy. Soc. New Zealand 82
(1): 19, 1954 (Hodgson 1954). Bas.: Bazzania mooreana Steph., Hedwigia 33 (1):
1, 1894 (Stephani 1894a).
*** Acromastigum moratii N.Kitag., Acta Phytotax. Geobot. 36 (4/6): 119, 1985 (Kitagawa 1985).
** Acromastigum obliquatum (Mitt.) A.Evans, Ann. Bryol., Suppl. 3: 110, 1934 (Evans
1934). Bas.: Bazzania obliquata Mitt., Hedwigia 32 (4): 211, 1893 (Stephani 1893c).
** Acromastigum tenax (Steph.) A.Evans, Ann. Bryol., Suppl. 3: 41, 1934 (Evans
1934). Bas.: Mastigobryum tenax Steph., Sp. Hepat. (Stephani) 6: 483, 1924
(Stephani 1924).
**
**
***
**
**
**
**
**
Incertae sedis
Acromastigum fumosum E.A.Br. et M.A.M.Renner, Telopea 17: 274, 2014 (Brown
and Renner 2014).
Acromastigum leptophyllum Herzog, Trans. Brit. Bryol. Soc. 1 (4): 309, 1950 (Herzog
1950a).
Acromastigum marginatum E.A.Hodgs., Trans. Roy. Soc. New Zealand 82 (1): 22,
1954 (Hodgson 1954).
Acromastigum microstictum A.Evans, Ann. Bryol., Suppl. 3: 115, 1934 (Evans 1934).
Acromastigum prismaticale E.A.Br. et M.A.M.Renner, Telopea 17: 281, 2014
(Brown and Renner 2014).
Acromastigum pusillum N.Kitag., Acta Phytotax. Geobot. 36 (4/6): 116, 1985
(Kitagawa 1985).
Acromastigum rigidum R.M.Schust., Nova Hedwigia 64 (3/4): 617, 1997 (Schuster
1997b).
Acromastigum subechinatiforme Hürl., Bauhinia 7 (4): 266, 1983 (Hürlimann 1983).
146
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Bazzania Gray, Nat. Arr. Brit. Pl. 1: 704, 1821 (Gray 1821) nom. conserv. 106
*** Bazzania acanthostipa Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 381, 1885
(Spruce 1885).
*** Bazzania accreta (Lehm. et Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser.
3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum accretum
Lehm. et Lindenb., Syn. Hepat. 2: 222, 1845 (Gottsche et al. 1845a).
** Bazzania acinaciformis Steph., Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr. 81: 290, 1907 (Stephani 1907a).
** Bazzania acuminata (Lindenb. et Gottsche) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum acuminatum Lindenb. et Gottsche, Syn. Hepat. 5: 719, 1847 (Gottsche et al. 1847).
** Bazzania acutifolia (Steph.) Schifn., Consp. Hepat. Arch. Ind.: 147, 1898
(Schifner 1898b). Bas.: Mastigobryum acutifolium Steph., Hedwigia 24 (5): 214,
1885 (Stephani 1885a).
*** Bazzania adnexa (Lehm. et Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Jungermannia adnexa Lehm. et Lindenb., Nov. Stirp. Pug. 4: 58, 1832 (Lehmann 1832).
** Bazzania adnexa var. aucklandica (Lindenb. et Gottsche) J.J.Engel et G.L.Merr.,
Bryologist 97 (3): 319, 1994 (Engel and Smith Merrill 1994). Bas.: Mastigobryum
novae-hollandiae f. aucklandicum Lindenb. et Gottsche, Sp. Hepat. (Lindenberg)
8-11: 33, 1851 (Lindenberg and Gottsche 1851b).
*** Bazzania ainis (Lindenb. et Gottsche) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 415, 1877 (Trevisan 1877). Bas.: Mastigobryum aine
Lindenb. et Gottsche, Syn. Hepat. 5: 720, 1847 (Gottsche et al. 1847).
** Bazzania albifolia Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 195,
1934 (Horikawa 1934).
** Bazzania ambigua (Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum ambiguum
Lindenb., Syn. Hepat. 2: 217, 1845 (Gottsche et al. 1845a).
*** Bazzania amblyphylla Meagher, Nova Hedwigia 92 (3/4): 487, 2011 (Meagher
2011).
** Bazzania aneityensis (Steph.) Tixier, Bull. Mus. Natl. Hist. Nat. (Sér. 3), Bot. 10
(190): 80, 1973 (Tixier 1973d). Bas.: Mastigobryum aneityense Steph., J. & Proc.
Roy. Soc. New South Wales 48 (1/2): 121, 1914 (Stephani and Watts 1914).
** Bazzania angusta (Steph.) Herzog, Trans. Brit. Bryol. Soc. 1 (4): 307, 1950 (Herzog 1950a). Bas.: Mastigobryum angustum Steph., Sp. Hepat. (Stephani) 6: 453,
1924 (Stephani 1924).
** Bazzania angustifalcata Herzog, Feddes Repert. Spec. Nov. Regni Veg. 57 (1/2):
166, 1955 (Herzog 1955).
106 Bazzania can not be subdivided based on recent molecular studies, so infrageneric taxa are not used
here. he genus includes Mastigobryum, but several taxa have neither been transferred nor synonymized.
hey are listed in the “Names in genera not currently accepted” section below.
World checklist of hornworts and liverworts
147
** Bazzania angustifolia Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 198,
1934 (Horikawa 1934).
** Bazzania angustisedens (Steph.) N.Kitag., J. Hattori Bot. Lab. 36: 445, 1972
[1973] (Kitagawa 1972). Bas.: Mastigobryum angustisedens Steph., Bull. Herb.
Boissier (sér. 2) 8 (10): 745 (429), 1908 (Stephani 1908c).
** Bazzania angustistipula N.Kitag., J. Hattori Bot. Lab. 30: 268, 1967 (Kitagawa 1967a).
*** Bazzania appendiculata (Mitt.) S.Hatt., Fl. E. Himalaya: 505, 1966 (Hattori
1966c). Bas.: Mastigobryum appendiculatum Mitt., J. Proc. Linn. Soc., Bot. 5 (18):
105, 1860 [1861] (Mitten 1860c).
** Bazzania approximata Onr., Bull. Jard. Bot. Natl. Belg. 47 (1/2): 139, 1977 (Onraedt
1977).
** Bazzania arcuata (Lindenb. et Gottsche) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum arcuatum Lindenb. et Gottsche, Syn. Hepat. 5: 718, 1847 (Gottsche et al. 1847).
** Bazzania arcuata var. mamillosa Gradst. et A.R.Benitez, Nova Hedwigia 99 (1/2):
113, 2014 (Gradstein and Benitez 2014).
** Bazzania armatistipula (Steph.) Fulford, Ann. Cryptog. Phytopathol. 3: 114, 1946
(Fulford 1946). Bas.: Mastigobryum armatistipulum Steph., Bull. Herb. Boissier
(sér. 2) 8 (11): 866 (490), 1908 (Stephani 1908a).
** Bazzania asperrima Steph., Rev. Bryol. 34 (3): 48, 1907 (Paris 1907).
* Bazzania asymmetrica (Steph.) N.Kitag., Bull. Nara Univ. Educ., B 28 (2): 77,
1979 (Kitagawa 1979a). Bas.: Mastigobryum asymmetricum Steph., Sp. Hepat.
(Stephani) 6: 454, 1924 (Stephani 1924). 107
** Bazzania aterrima (Steph.) N.Kitag., Bull. Nara Univ. Educ., B 26 (2): 77, 1977
(Kitagawa 1977). Bas.: Mastigobryum aterrimum Steph., Sp. Hepat. (Stephani) 6:
454, 1924 (Stephani 1924).
*** Bazzania aurescens Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 374, 1885
(Spruce 1885).
** Bazzania avia Meagher, Nova Hedwigia 97 (3/4): 529, 2013 (Meagher 2013).
** Bazzania azorica H.Buch et Perss., Bryophyt. Azoren Madeira: 3, 1941 (Buch and
Persson 1941).
** Bazzania baldwinii Austin, Trans. Connecticut Acad. Arts 8 (15): 255, 1891 (Evans
1891).
** Bazzania bernieri (Steph.) Inoue et H.A.Mill., Bull. Natl. Sci. Mus. Tokyo (n.ser.)
8 (2): 142, 1965 (Inoue and Miller 1965). Bas.: Mastigobryum bernieri Steph.,
Bull. Herb. Boissier (sér. 2) 8 (11): 852 (476), 1908 (Stephani 1908a).
*** Bazzania bescherellei Steph., Hedwigia 32 (4): 204, 1893 (Stephani 1893c).
** Bazzania bhutanica N.Kitag. et Grolle, J. Hattori Bot. Lab. 61: 269, 1986 [1987]
(Kitagawa and Grolle 1986).
** Bazzania bicrenata N.Kitag., J. Hattori Bot. Lab. 47: 127, 1980 (Kitagawa 1980).
107 Bazzania asymmetrica is conspeciic with Bazzania macgregorii in Grolle (1968a), but Kitagawa
(1979a) kept them separate.
148
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Bazzania bidens (Gottsche et Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 415, 1877 (Trevisan 1877). Bas.: Mastigobryum bidens Gottsche et Lindenb., Syn. Hepat. 2: 228, 1845 (Gottsche et al. 1845a).
** Bazzania bidens var. heterodonta Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:
372, 1885 (Spruce 1885).
** Bazzania bidentula (Steph.) Yasuda, Shokubutsugaku Kakuron: 711, 1911 (Yasuda 1911). Bas.: Pleuroschisma bidentulum Steph., Mém. Soc. Nat. Sci. Nat. Math.
Cherbourg 29: 222, 1894 (Stephani 1894b).
*** Bazzania bilobata N.Kitag., J. Hattori Bot. Lab. 30: 257, 1967 (Kitagawa 1967a).
** Bazzania borneensis N.Kitag., J. Hattori Bot. Lab. 37: 263, 1973 (Kitagawa 1973).
Nom. nov. pro Mastigobryum borneense Steph., Bull. Herb. Boissier (sér. 2) 8 (11):
839 (463), 1908 (Stephani 1908a), nom. illeg.
** Bazzania brasiliensis (Gottsche et Lindenb.) Trevis., Mem. Reale Ist. Lombardo
Sci. (Ser. 3), C. Sci. Mat. 4 (13): 415, 1877 (Trevisan 1877). Bas.: Mastigobryum
brasiliense Gottsche et Lindenb., Syn. Hepat. 2: 227, 1845 (Gottsche et al. 1845a).
** Bazzania brighamii (Austin) A.Evans, Trans. Connecticut Acad. Arts 8 (15): 255,
1891 (Evans 1891). Bas.: Mastigobryum brighamii Austin, Bull. Torrey Bot. Club
5 (3): 16, 1874 (Austin 1874).
** Bazzania cadens N.Kitag., J. Hattori Bot. Lab. 47: 129, 1980 (Kitagawa 1980).
*** Bazzania calcarata (Sande Lac.) Schifn., Consp. Hepat. Arch. Ind.: 149, 1898
(Schifner 1898b). Bas.: Mastigobryum calcaratum Sande Lac., Ann. Mus. Bot.
Lugduno-Batavi 1: 304, 1864 (Sande Lacoste 1864).
** Bazzania callida (Sande Lac. ex Steph.) Abeyw., Ceylon J. Sci., Biol. Sci. 2 (1): 45,
1959 (Abeywickrama 1959). Bas.: Mastigobryum callidum Sande Lac. ex Steph.,
Hedwigia 24 (6): 246, 1885 (Stephani 1885b).
*** Bazzania canelensis (Steph.) Fulford, Ann. Cryptog. Phytopathol. 3: 152, 1946
(Fulford 1946). Bas.: Mastigobryum canelense Steph., Sp. Hepat. (Stephani) 3:
518, 1909 (Stephani 1909a).
** Bazzania caudata (Steph.) Herzog, Trans. Brit. Bryol. Soc. 1 (4): 301, 1950 (Herzog 1950a). Bas.: Mastigobryum caudatum Steph., Bull. Herb. Boissier (sér. 2) 8
(12): 947 (497), 1908 (Stephani 1908b).
*** Bazzania caudistipula (Steph.) Inoue et H.A.Mill., Bull. Natl. Sci. Mus. Tokyo
(n.ser.) 8 (2): 141, 1965 (Inoue and Miller 1965). Bas.: Mastigobryum caudistipulum Steph., Bull. Herb. Boissier (sér. 2) 8 (12): 945 (495), 1908 (Stephani 1908b).
** Bazzania ceylanica (Mitt.) Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 306, 1896 (Stephani 1896a). Bas.: Mastigobryum ceylanicum Mitt., J. Proc. Linn. Soc., Bot. 5 (18):
105, 1860 [1861] (Mitten 1860c).
** Bazzania chilensis (Steph.) Fulford, Ann. Cryptog. Phytopathol. 3: 51, 1946
(Fulford 1946). Bas.: Mastigobryum chilense Steph., Hedwigia 24 (6): 247, 1885
(Stephani 1885b).
*** Bazzania chimantensis Fulford, Bryologist 63 (2): 89, 1960 (Fulford 1960).
World checklist of hornworts and liverworts
149
** Bazzania cincinnata (De Not.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum cincinnatum De
Not., Epat. Borneo: 34, 1874 (De Notaris 1874).
*** Bazzania citharodes Meagher, Nova Hedwigia 86 (3/4): 481, 2008 (Meagher 2008).
** Bazzania combinata (J.B.Jack et Steph.) Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 306,
1896 (Stephani 1896a). Bas.: Mastigobryum combinatum J.B.Jack et Steph., Bot.
Centralbl. 60 (4): 102, 1894 (Jack and Stephani 1894).
** Bazzania commutata (Lindenb. et Gottsche) Schifn., Consp. Hepat. Arch. Ind.: 149,
1898 (Schifner 1898b). Bas.: Mastigobryum commutatum Lindenb. et Gottsche, Sp.
Hepat. (Lindenberg) 8-11: 97, 1851 (Lindenberg and Gottsche 1851b).
* Bazzania comorensis Steph., Bull. Soc. Roy. Bot. Belgique, Compt. Rend. 30 (2):
197, 1891 [1892] (Stephani 1891b).
** Bazzania confertifolia (Steph.) Herzog, Trans. Brit. Bryol. Soc. 1 (4): 298, 1950
(Herzog 1950a). Bas.: Mastigobryum confertifolium Steph., Bull. Herb. Boissier
(sér. 2) 8 (11): 858 (482), 1908 (Stephani 1908a).
** Bazzania conistipula (Steph.) H.A.Mill., Phytologia 47 (4): 320, 1981 (Miller
1981). Bas.: Mastigobryum conistipulum Steph., Sp. Hepat. (Stephani) 6: 458,
1924 (Stephani 1924).
** Bazzania conophylla (Sande Lac.) Schifn., Consp. Hepat. Arch. Ind.: 150, 1898
(Schifner 1898b). Bas.: Mastigobryum conophyllum Sande Lac., Ann. Mus. Bot.
Lugduno-Batavi 1: 304, 1864 (Sande Lacoste 1864).
** Bazzania consanguinea (Hampe et Lindenb.) Trevis., Mem. Reale Ist. Lombardo
Sci. (Ser. 3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum
consanguineum Hampe et Lindenb., Linnaea 20 (3): 327, 1847 (Hampe 1847).
** Bazzania consociata (Steph.) H.A.Mill., Phytologia 47 (4): 320, 1981 (Miller
1981). Bas.: Mastigobryum consociatum Steph., Sp. Hepat. (Stephani) 6: 458, 1924
(Stephani 1924).
* Bazzania corbieri (Steph.) Meagher, Nova Hedwigia 86 (3/4): 483, 2008 (Meagher 2008). Bas.: Mastigobryum corbieri Steph., J. & Proc. Roy. Soc. New South
Wales 48 (1/2): 122, 1914 (Stephani and Watts 1914). 108
** Bazzania crassidentata Fulford, Bull. Torrey Bot. Club 86 (5): 338, 1959 (Fulford
1959b).
** Bazzania crassitexta Steph., Hedwigia 32 (4): 205, 1893 (Stephani 1893c).
* Bazzania crenata (Trevis.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 415, 1877 (Trevisan 1877). Bas.: Pleuroschisma crenatum Trevis.,
Herb. Crypt. Trev. 2: 30, 1853 (Trevisan 1853; non vidi). 109
** Bazzania cubensis (Gottsche ex Steph.) Pagán, Bryologist 42 (2): 38, 1939 (Pagán
1939b). Bas.: Mastigobryum cubense Gottsche ex Steph., Hedwigia 24 (6): 248,
1885 (Stephani 1885b).
108 Bazzania corbieri may be conspeciic with Bazzania crenata (Meagher 2008).
109 Bazzania crenata from Europe has neither been recognized in any recent treatment nor synonymized.
he name may have priority once the identity is established.
150
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Bazzania cucullata Onr., Bull. Jard. Bot. Natl. Belg. 47 (1/2): 142, 1977 (Onraedt
1977).
*** Bazzania cuneistipula (Gottsche et Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum cuneistipulum Gottsche et Lindenb., Syn. Hepat. 2: 225, 1845 (Gottsche et al. 1845a).
* Bazzania curvidens Steph., Bull. Soc. Roy. Bot. Belgique, Compt. Rend. 30 (2):
197, 1891 [1892] (Stephani 1891b).
** Bazzania debilis N.Kitag., J. Hattori Bot. Lab. 30: 256, 1967 (Kitagawa 1967a).
** Bazzania deciduifolia Onr., Bull. Jard. Bot. Natl. Belg. 47 (1/2): 144, 1977 (Onraedt
1977).
*** Bazzania decrescens (Lehm. et Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Jungermannia decrescens Lehm. et Lindenb., Nov. Stirp. Pug. 4: 57, 1832 (Lehmann 1832). 110
** Bazzania decrescens var. dentistipula Kiaer et Pearson, Forh. Vidensk.-Selsk. Kristiania 1892 (14): 6, 1893 (Pearson 1893).
** Bazzania decrescens subsp. molleri (Steph.) E.W.Jones, J. Bryol. 8 (3): 303, 1975
(Jones 1975). Bas.: Mastigobryum molleri Steph., Bot. Jahrb. Syst. 8 (2): 84, 1886
(Stephani 1886d).
** Bazzania decrescens subsp. pumila (Mitt.) Pócs, Trop. Bryol. 9: 129, 1994 (Pócs 1994c).
Bas.: Bazzania pumila Mitt., J. Linn. Soc., Bot. 22 (146): 322, 1886 (Mitten 1886b).
** Bazzania densa (Sande Lac.) Schifn., Consp. Hepat. Arch. Ind.: 151, 1898
(Schifner 1898b). Bas.: Mastigobryum densum Sande Lac., Ned. Kruidk. Arch. 3:
418, 1854 [1855] (Sande Lacoste 1854).
** Bazzania densa var. connata (Sande Lac.) Schifn., Consp. Hepat. Arch. Ind.: 151,
1898 (Schifner 1898b). Bas.: Mastigobryum densum β connatum Sande Lac., Ann.
Mus. Bot. Lugduno-Batavi 1: 302, 1864 (Sande Lacoste 1864).
*** Bazzania denticulata (Lindenb. et Gottsche) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum denticulatum Lindenb. et Gottsche, Syn. Hepat. 5: 718, 1847 (Gottsche et al. 1847).
** Bazzania denticulifera Mägd., Nova Hedwigia 38: 53, 1983 (Mägdefrau 1983).
** Bazzania denudata (Lindenb. et Gottsche) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum denudatum Lindenb. et Gottsche, Syn. Hepat. 2: 216, 1845 (Gottsche et al. 1845a).
*** Bazzania deplanchei (Gottsche) Jovet-Ast, Rev. Bryol. Lichénol. 18 (1/2): 83, 1949
(Jovet-Ast 1949b). Bas.: Mastigobryum deplanchei Gottsche, Bull. Herb. Boissier
(sér. 2) 8 (12): 955 (505), 1908 (Stephani 1908b).
** Bazzania deplanchei var. ilamentosa Tixier, Cryptog. Bryol. Lichénol. 6 (2): 179,
1985 (Tixier 1985b).
** Bazzania desciscens (Steph.) Meijer, Blumea 10 (2): 382, 1960 (Meijer 1960). Bas.:
Mastigobryum desciscens Steph., Bull. Herb. Boissier (sér. 2) 8 (11): 862 (486),
1908 (Stephani 1908a).
110 Bazzania decrescens is a species complex also including also Bazzania comorensis, Bazzania curvidens
and Bazzania mascarena (Grolle 1995).
World checklist of hornworts and liverworts
151
** Bazzania didericiana (Gottsche ex Steph.) Steph., Bull. Herb. Boissier 5 (10): 841,
1897 (Stephani 1897c). Bas.: Mastigobryum didericianum Gottsche ex Steph.,
Hedwigia 24 (6): 249, 1885 (Stephani 1885b).
** Bazzania diminuta Herzog, Trans. Brit. Bryol. Soc. 1 (4): 305, 1950 (Herzog 1950a).
* Bazzania distans (Nees) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum distans Nees, Syn.
Hepat. 2: 216, 1845 (Gottsche et al. 1845a). 111
*** Bazzania diversicuspis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 373, 1885
(Spruce 1885).
** Bazzania drepanophylla Herzog, Trans. Brit. Bryol. Soc. 1 (4): 301, 1950 (Herzog
1950a).
** Bazzania dulitensis Herzog, Trans. Brit. Bryol. Soc. 1 (4): 299, 1950 (Herzog
1950a).
** Bazzania dulongensis L.P.Zhou et Li Zhang, J. Bryol. 34 (1): 22, 2012 (Zhou et al.
2012).
** Bazzania eggersiana (Steph.) Pagán, Bryologist 42 (2): 39, 1939 (Pagán 1939b).
Bas.: Mastigobryum eggersianum Steph., Bull. Herb. Boissier (sér. 2) 8 (11): 844
(468), 1908 (Stephani 1908a).
** Bazzania elmeri (Steph.) N.Kitag., J. Hattori Bot. Lab. 36: 446, 1972 [1973] (Kitagawa 1972). Bas.: Mastigobryum elmeri Steph., Leal. Philipp. Bot. 6: 2289, 1914
(Stephani 1914a).
** Bazzania elongata Fulford, Bull. Torrey Bot. Club 86 (5): 337, 1959 (Fulford 1959b).
** Bazzania emarginata (Steph.) C.M.Cooke, Trans. Connecticut Acad. Arts 12
(1): 17, 1904 (Cooke 1904). Bas.: Mastigobryum didericianum var. emarginatum
Steph., Hedwigia 24 (6): 249, 1885 (Stephani 1885b).
** Bazzania engelii Glenny, Fieldiana, Bot. (n.ser.) 47: 176, 2007 (Glenny and Bartlett
2007).
*** Bazzania erosa (Reinw., Blume et Nees) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 415, 1877 (Trevisan 1877). Bas.: Jungermannia erosa
Reinw., Blume et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat.
Cur. 12 (1): 230, 1824 [1825] (Reinwardt et al. 1824a).
** Bazzania erosa var. pulopenangensis (Lindenb. et Gottsche) Schifn., Consp. Hepat.
Arch. Ind.: 157, 1898 (Schifner 1898b). Bas.: Mastigobryum erosum δ pulopenangense Lindenb. et Gottsche, Sp. Hepat. (Lindenberg) 8-11: 99, 1851 (Lindenberg and Gottsche 1851b).
*** Bazzania exempta J.J.Engel, J. Hattori Bot. Lab. 99: 197, 2006 (Engel 2006c).
*** Bazzania falcata (Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 415, 1877 (Trevisan 1877). Bas.: Mastigobryum falcatum Lindenb., Syn. Hepat. 2: 231, 1845 (Gottsche et al. 1845a).
** Bazzania falcifolia (Steph.) H.A.Mill., Phytologia 47 (4): 320, 1981 (Miller 1981).
Bas.: Mastigobryum falcifolium Steph., Akad. Wiss. Wien, Math.-Naturwiss. Kl.,
Denkschr. 88: 33, 1911 (Stephani 1911d).
111 Bazzania distans is probably conspeciic with Bazzania loricata (Meijer 1960).
152
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Bazzania fallax (Sande Lac.) Schifn., Consp. Hepat. Arch. Ind.: 158, 1898
(Schifner 1898b). Bas.: Mastigobryum fallax Sande Lac., Ann. Mus. Bot. Lugduno-Batavi 1: 304, 1864 (Sande Lacoste 1864).
Bazzania fasciculata (Steph.) Meagher, Australas. Bryol. Newslett. 46: 6, 2002
(Meagher 2002). Bas.: Mastigobryum fasciculatum Steph., Bull. Herb. Boissier (sér.
2) 8 (10): 748 (432), 1908 (Stephani 1908c).
Bazzania fauriana (Steph.) S.Hatt., Bot. Mag. (Tokyo) 59 (693/694): 27, 1946
(Hattori 1946). Bas.: Mastigobryum faurianum Steph., Bull. Herb. Boissier (sér. 2)
8 (11): 843 (467), 1908 (Stephani 1908a).
Bazzania iliformis Steph., Hedwigia 28 (2): 131, 1889 (Stephani 1889a).
Bazzania laccida (Dumort.) Grolle, Lindbergia 1 (3/4): 200, 1972 [1973] (Grolle
1972b). Bas.: Pleuroschisma laccidum Dumort., Syll. Jungerm. Europ.: 71, 1831
(Dumortier 1831).
Bazzania lavescens (Sande Lac. ex Steph.) Schifn., Consp. Hepat. Arch. Ind.:
158, 1898 (Schifner 1898b). Bas.: Mastigobryum lavescens Sande Lac. ex Steph.,
Hedwigia 25 (1): 6, 1886 (Stephani 1886e).
Bazzania leischeri (Steph.) Abeyw., Ceylon J. Sci., Biol. Sci. 2 (1): 45, 1959 (Abeywickrama 1959). Bas.: Mastigobryum leischeri Steph., Bull. Herb. Boissier (sér. 2)
8 (10): 773 (457), 1908 (Stephani 1908c).
Bazzania francana (Steph.) N.Kitag., J. Hattori Bot. Lab. 36: 446, 1972 [1973]
(Kitagawa 1972). Bas.: Mastigobryum francanum Steph., Sp. Hepat. (Stephani) 6:
463, 1924 (Stephani 1924).
Bazzania friabilis N.Kitag. et T.Kodama, J. Hattori Bot. Lab. 39: 67, 1975 (Kitagawa
and Kodama 1975b).
Bazzania fuhreri Meagher, Nova Hedwigia 92 (3/4): 488, 2011 (Meagher 2011).
Bazzania fuscescens A.Evans, Pap. Michigan Acad. Sci. 17: 85, 1932 [1933] (Evans
1932b).
Bazzania gamscottii Meagher, Nova Hedwigia 92 (3/4): 492, 2011 (Meagher 2011).
Bazzania gedeana (Steph.) Meijer, Blumea 10 (2): 378, 1960 (Meijer 1960). Bas.:
Mastigobryum gedeanum Steph., Sp. Hepat. (Stephani) 3: 540, 1909 (Stephani
1909a).
Bazzania gracilis (Hampe et Gottsche) Steph., Hedwigia 27 (11/12): 279, 1888
(Stephani 1888c). Bas.: Mastigobryum gracile Hampe et Gottsche, Linnaea 25 (3):
346, 1852 [1853] (Hampe and Gottsche 1852).
Bazzania grandiretis (Steph.) Herzog, Trans. Brit. Bryol. Soc. 1 (4): 298, 1950
(Herzog 1950a). Bas.: Mastigobryum grandirete Steph., Bull. Herb. Boissier (sér. 2)
8 (10): 747 (431), 1908 (Stephani 1908c).
Bazzania griithiana (Steph.) Mizut., J. Hattori Bot. Lab. 30: 82, 1967 (Mizutani
1967). Bas.: Mastigobryum griithianum Steph., Bull. Herb. Boissier (sér. 2) 8
(12): 959 (509), 1908 (Stephani 1908b).
Bazzania gunniana (Steph.) H.A.Mill., Phytologia 47 (4): 320, 1981 (Miller
1981). Bas.: Mastigobryum gunnianum Steph., J. & Proc. Roy. Soc. New South
Wales 48 (1/2): 123, 1914 (Stephani and Watts 1914).
World checklist of hornworts and liverworts
153
** Bazzania hainanensis L.P.Zhou et Li Zhang, J. Bryol. 34 (1): 25, 2012 (Zhou et
al. 2012).
** Bazzania halconiensis (Steph.) N.Kitag., J. Hattori Bot. Lab. 36: 447, 1972 [1973]
(Kitagawa 1972). Bas.: Mastigobryum halconiense Steph., Bull. Herb. Boissier (sér.
2) 8 (10): 759 (443), 1908 (Stephani 1908c).
** Bazzania hamatifolia (Steph.) H.A.Mill., Phytologia 47 (4): 320, 1981 (Miller
1981). Bas.: Mastigobryum hamatifolium Steph., Bull. Herb. Boissier (sér. 2) 8
(11): 862 (486), 1908 (Stephani 1908a).
** Bazzania harpago (De Not.) Schifn., Consp. Hepat. Arch. Ind.: 159, 1898
(Schifner 1898b). Bas.: Mastigobryum harpago De Not., Epat. Borneo: 29, 1874
(De Notaris 1874).
** Bazzania hebridensis (Steph.) H.A.Mill., Phytologia 47 (4): 320, 1981 (Miller
1981). Bas.: Mastigobryum hebridense Steph., J. & Proc. Roy. Soc. New South
Wales 48 (1/2): 124, 1914 (Stephani and Watts 1914).
** Bazzania herminieri (Gottsche ex Steph.) Pagán, Bryologist 45 (4): 90, 1942
(Pagán 1942b). Bas.: Mastigobryum herminieri Gottsche ex Steph., Hedwigia 25
(1): 8, 1886 (Stephani 1886e).
** Bazzania herzogiana Meijer, Blumea 10 (2): 371, 1960 (Meijer 1960). Nom. nov.
pro Bazzania remotifolia Herzog, Trans. Brit. Bryol. Soc. 1 (4): 304, 1950 (Herzog
1950a), nom. illeg.
** Bazzania heterostipa (Steph.) Fulford, Bull. Torrey Bot. Club 86 (6): 410, 1959
(Fulford 1959a). Bas.: Mastigobryum heterostipum Steph., Sp. Hepat. (Stephani) 3:
532, 1909 (Stephani 1909a).
*** Bazzania himalayana (Mitt.) Schifn., Österr. Bot. Z. 49 (4): 132, 1899 (Schifner
1899b). Bas.: Mastigobryum himalayanum Mitt., J. Proc. Linn. Soc., Bot. 5 (18):
105, 1860 [1861] (Mitten 1860c).
*** Bazzania hochstetteri (Reichardt) E.A.Hodgs., Trans. Roy. Soc. New Zealand 82
(1): 11, 1954 (Hodgson 1954). Bas.: Mastigobryum hochstetteri Reichardt, Verh.
K.K. Zool.-Bot. Ges. Wien 16: 959, 1866 (Reichardt 1866).
*** Bazzania hookeri (Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum hookeri Lindenb., Syn. Hepat. 2: 226, 1845 (Gottsche et al. 1845a).
** Bazzania horridula Schifn., Nova Acta Acad. Caes. Leop.-Carol. German. Nat.
Cur. 60 (2): 258, 1893 (Schifner 1893a).
** Bazzania inaequabilis Steph., Trans. Connecticut Acad. Arts 12 (1): 21, 1904
(Cooke 1904).
** Bazzania inaequitexta Steph., Hedwigia 32 (4): 208, 1893 (Stephani 1893c).
** Bazzania incrassata (Steph.) N.Kitag., J. Hattori Bot. Lab. 36: 448, 1972 [1973]
(Kitagawa 1972). Bas.: Mastigobryum incrassatum Steph., Sp. Hepat. (Stephani) 6:
469, 1924 (Stephani 1924).
** Bazzania indica (Gottsche et Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 415, 1877 (Trevisan 1877). Bas.: Mastigobryum indicum Gottsche et Lindenb., Syn. Hepat. 2: 230, 1845 (Gottsche et al. 1845a).
154
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Bazzania indigenarum (Steph.) N.Kitag., Bull. Nara Univ. Educ., B 26 (2): 82,
1977 (Kitagawa 1977). Bas.: Mastigobryum indigenarum Steph., J. & Proc. Roy.
Soc. New South Wales 48 (1/2): 124, 1914 (Stephani and Watts 1914).
** Bazzania insignis (De Not.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum insigne De Not.,
Epat. Borneo: 26, 1874 (De Notaris 1874).
* Bazzania intermedia (Gottsche et Lindenb.) Trevis., Mem. Reale Ist. Lombardo
Sci. (Ser. 3), C. Sci. Mat. 4 (13): 415, 1877 (Trevisan 1877). Bas.: Mastigobryum
intermedium Gottsche et Lindenb., Sp. Hepat. (Lindenberg) 8-11: 82, 1851 (Lindenberg and Gottsche 1851b).
** Bazzania intermedia var. sarawakiana (De Not.) Schifn., Consp. Hepat. Arch.
Ind.: 162, 1898 (Schifner 1898b). Bas.: Mastigobryum intermedium var. sarawakianum De Not., Epat. Borneo: 32, 1874 (De Notaris 1874).
** Bazzania involuta (Mont.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Herpetium involutum Mont., Ann.
Sci. Nat. Bot. (sér. 2) 19: 253, 1843 (Montagne 1843).
** Bazzania involuta var. submutica (Lindenb. et Gottsche) J.J.Engel et G.L.Merr.,
Bryologist 97 (3): 314, 1994 (Engel and Smith Merrill 1994). Bas.: Mastigobryum
novae-hollandiae γ3 submuticum Lindenb. et Gottsche, Sp. Hepat. (Lindenberg)
8-11: 33, 1851 (Lindenberg and Gottsche 1851b).
** Bazzania involutiformis (De Not.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum involutiforme
De Not., Epat. Borneo: 28, 1874 (De Notaris 1874).
** Bazzania irregularis (Steph.) Schifn., Consp. Hepat. Arch. Ind.: 163, 1898
(Schifner 1898b). Bas.: Mastigobryum irregulare Steph., Hedwigia 25 (4): 133,
1886 (Stephani 1886b).
*** Bazzania jamaicensis (Lehm. et Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Herpetium jamaicense Lehm. et Lindenb., Nov. Stirp. Pug. 7: 7, 1838 (Lehmann 1838).
** Bazzania japonica (Sande Lac.) Lindb., Acta Soc. Sci. Fenn. 10: 224, 1872 [1873]
(Lindberg 1872b). Bas.: Mastigobryum japonicum Sande Lac., Ann. Mus. Bot.
Lugduno-Batavi 1: 303, 1864 (Sande Lacoste 1864).
** Bazzania japonica var. sumatrana Herzog, Ann. Naturhist. Mus. Wien 53 (1):
366, 1942 [1943] (Herzog 1942b).
** Bazzania javanica (Sande Lac.) Schifn., Consp. Hepat. Arch. Ind.: 163, 1898
(Schifner 1898b). Bas.: Mastigobryum javanicum Sande Lac., Ned. Kruidk. Arch.
3: 418, 1854 [1855] (Sande Lacoste 1854).
** Bazzania kernii Steph., Hedwigia 32 (4): 208, 1893 (Stephani 1893c).
** Bazzania kokawana N.Kitag. et T.Kodama, J. Jap. Bot. 50 (1): 11, 1975 (Kitagawa and Kodama 1975a).
** Bazzania latifolia Steph., Hedwigia 32 (4): 209, 1893 (Stephani 1893c).
World checklist of hornworts and liverworts
*
**
**
**
**
**
**
***
**
***
**
***
***
**
155
Bazzania lehmanniana (Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum lehmannianum Lindenb., Syn. Hepat. 2: 223, 1845 (Gottsche et al. 1845a). 112
Bazzania leratii (Beauverd) H.A.Mill., Phytologia 47 (4): 320, 1981 (Miller 1981).
Bas.: Mastigobryum leratii Beauverd, Sp. Hepat. (Stephani) 6: 477, 1924 (Stephani
1924).
Bazzania lessonii (Steph.) H.A.Mill., Phytologia 47 (4): 321, 1981 (Miller 1981).
Bas.: Mastigobryum lessonii Steph., Sp. Hepat. (Stephani) 3: 531, 1909 (Stephani
1909a).
Bazzania levieri (Steph.) N.Kitag., J. Hattori Bot. Lab. 36: 448, 1972 [1973] (Kitagawa 1972). Bas.: Mastigobryum levieri Steph., Bull. Herb. Boissier (sér. 2) 8
(12): 944 (494), 1908 (Stephani 1908b).
Bazzania liebmaniana (Lindenb. et Gottsche) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum liebmanianum Lindenb. et Gottsche, Syn. Hepat. 5: 719, 1847 (Gottsche et al. 1847).
Bazzania linearis Herzog, Trans. Brit. Bryol. Soc. 1 (4): 303, 1950 (Herzog 1950a).
Bazzania linguiformis (Sande Lac.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser.
3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum linguiforme Sande Lac., Plagiochila Sandei: 8, 1856 (Sande Lacoste 1856c).
Bazzania longa (Nees) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 415, 1877 (Trevisan 1877). Bas.: Jungermannia longa Nees, Linnaea
6 (4): 623, 1831 (Nees 1831).
Bazzania longa var. papillata (Steph.) Fulford, Ann. Cryptog. Phytopathol. 3: 94,
1946 (Fulford 1946). Bas.: Mastigobryum papillatum Steph., Sp. Hepat. (Stephani) 3: 526, 1909 (Stephani 1909a).
Bazzania longicaulis (Sande Lac.) Schifn., Consp. Hepat. Arch. Ind.: 165, 1898
(Schifner 1898b). Bas.: Mastigobryum longicaule Sande Lac., Ann. Mus. Bot. Lugduno-Batavi 1: 303, 1864 (Sande Lacoste 1864).
Bazzania longicaulis var. latiareata Herzog, Memoranda Soc. Fauna Fl. Fennica
26: 46, 1950 [1951] (Herzog 1950b).
Bazzania longistipula (Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 415, 1877 (Trevisan 1877). Bas.: Mastigobryum longistipulum
Lindenb., Syn. Hepat. 2: 228, 1845 (Gottsche et al. 1845a).
Bazzania loricata (Reinw., Blume et Nees) Trevis., Mem. Reale Ist. Lombardo
Sci. (Ser. 3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Jungermannia
loricata Reinw., Blume et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol.
Nat. Cur. 12 (1): 233, 1824 [1825] (Reinwardt et al. 1824a).
Bazzania lowii (Sande Lac. ex Steph.) Schifn., Consp. Hepat. Arch. Ind.: 166,
1898 (Schifner 1898b). Bas.: Mastigobryum lowii Sande Lac. ex Steph., Hedwigia
25 (5): 204, 1886 (Stephani 1886f).
112 Bazzania lehmanniana is possibly conspeciic with Bazzania arcuata or Bazzania longa (Meagher 2012).
156
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Bazzania luzonensis (Steph.) Del Ros., Philipp. J. Sci. 100 (3/4): 231, 1971 (Del
Rosario 1971). Bas.: Mastigobryum luzonense Steph., Sp. Hepat. (Stephani) 6: 472,
1924 (Stephani 1924).
** Bazzania macgregorii Steph., Hedwigia 32 (4): 210, 1893 (Stephani 1893c).
** Bazzania macrostipula Fulford, Bull. Torrey Bot. Club 86 (6): 407, 1959 (Fulford
1959a).
** Bazzania magna Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 197, 1934
(Horikawa 1934).
** Bazzania magnistipula N.Kitag., J. Hattori Bot. Lab. 47: 132, 1980 (Kitagawa
1980).
** Bazzania malaccensis (Steph.) Tixier, Gard. Bull. Singapore 25 (3): 342, 1971
(Tixier 1971). Bas.: Mastigobryum malaccense Steph., Bull. Herb. Boissier (sér. 2)
8 (12): 944 (494), 1908 (Stephani 1908b).
** Bazzania manillana (Gottsche ex Steph.) S.Hatt., Bot. Mag. (Tokyo) 64 (755/756):
113, 1951 (Hattori 1951c). Bas.: Mastigobryum manillanum Gottsche ex Steph.,
Hedwigia 25 (5): 204, 1886 (Stephani 1886f).
* Bazzania marginata (Steph.) N.Kitag., J. Hattori Bot. Lab. 36: 449, 1972 [1973]
(Kitagawa 1972). Bas.: Mastigobryum marginatum Steph., Rev. Bryol. 35 (2): 31,
1908 (Stephani 1908l).
** Bazzania marginella (Herzog) N.Kitag. et T.Kodama, Bull. Osaka Mus. Nat. Hist.
27: 17, 1973 (Kitagawa and Kodama 1973). Bas.: Mastigobryum marginellum Herzog,
Ann. Bryol. 5: 91, 1932 (Herzog 1932a).
* Bazzania mascarena (Steph.) Herzog, Bot. Not. 100 (4): 334, 1947 (Herzog
1947). Bas.: Mastigobryum mascarenum Steph., Hedwigia 25 (5): 205, 1886 (Stephani 1886f).
** Bazzania mayebarae S.Hatt., J. Hattori Bot. Lab. 19: 91, 1958 (Hattori and
Mizutani 1958).
* Bazzania menzelii E.D.Cooper, Phytotaxa 97 (2): 52, 2013 (Cooper et al. 2013).
Nom. nov. pro Acromastigum emarginatum Herzog, Trans. Brit. Bryol. Soc. 1 (4):
309, 1950 (Herzog 1950a).
** Bazzania merrillana (Steph.) Inoue ex Bonner, Index Hepat. 3: 359, 1963 (Bonner 1963). Bas.: Mastigobryum merrillanum Steph., Bull. Herb. Boissier (sér. 2) 8
(12): 944 (494), 1908 (Stephani 1908b).
** Bazzania minuta (Austin) A.Evans, Trans. Connecticut Acad. Arts 8 (15): 255,
1891 (Evans 1891). Bas.: Mastigobryum minutum Austin, Bull. Torrey Bot. Club
5 (3): 17, 1874 (Austin 1874).
** Bazzania minutidens (Steph.) Inoue et H.A.Mill., Bull. Natl. Sci. Mus. Tokyo
(n.ser.) 8 (2): 142, 1965 (Inoue and Miller 1965). Bas.: Mastigobryum minutidens
Steph., Sp. Hepat. (Stephani) 6: 474, 1924 (Stephani 1924).
** Bazzania minutiserra (Steph.) N.Kitag., Bull. Nara Univ. Educ., B 26 (2): 77,
1977 (Kitagawa 1977). Bas.: Mastigobryum minutiserrum Steph., Sp. Hepat.
(Stephani) 6: 474, 1924 (Stephani 1924).
World checklist of hornworts and liverworts
157
** Bazzania missionum (Herzog) Jovet-Ast, Rev. Bryol. Lichénol. 20 (1/2): 96, 1951
(Jovet-Ast 1951). Bas.: Mastigobryum missionum Herzog, Ann. Bryol. 5: 93, 1932
(Herzog 1932a).
** Bazzania mittenii (Steph.) Steph., Hedwigia 28 (2): 132, 1889 (Stephani 1889a).
Bas.: Mastigobryum mittenii Steph., Hedwigia 25 (6): 245, 1886 (Stephani 1886c).
*** Bazzania monilinervis (Lehm. et Lindenb.) Trevis., Mem. Reale Ist. Lombardo
Sci. (Ser. 3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Jungermannia
monilinervis Lehm. et Lindenb., Nov. Stirp. Pug. 4: 56, 1832 (Lehmann 1832).
** Bazzania morokensis (Steph.) Grolle, J. Hattori Bot. Lab. 31: 1, 1968 (Grolle
1968a). Bas.: Mastigobryum morokense Steph., Bull. Herb. Boissier (sér. 2) 8 (10):
765 (449), 1908 (Stephani 1908c).
*** Bazzania nitida (F.Weber) Grolle, Rev. Bryol. Lichénol. 29 (3/4): 210, 1960
[1961] (Grolle 1960a). Bas.: Jungermannia nitida F.Weber, Hist. Musc. Hepat.
Prodr.: 43, 1815 (Weber 1815).
*** Bazzania novae-zelandiae (Mitt.) Besch. et C.Massal., Miss. sci. Cape Horn, Bot.
5: 233, 1889 (Bescherelle and Massalongo 1889). Bas.: Mastigobryum novae-zelandiae Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 148, 1854 (Mitten 1854).
** Bazzania nudicaulis A.Evans, Bryologist 26 (6): 62, 1923 [1924] (Evans 1923b).
** Bazzania nuuanuensis C.M.Cooke, Trans. Connecticut Acad. Arts 12 (1): 15,
1904 (Cooke 1904).
** Bazzania obcuneata (Steph.) H.A.Mill., Phytologia 47 (4): 321, 1981 (Miller
1981). Bas.: Mastigobryum obcuneatum Steph., Bull. Herb. Boissier (sér. 2) 8 (11):
863 (487), 1908 (Stephani 1908a).
** Bazzania obtusata (Mitt.) Abeyw., Ceylon J. Sci., Biol. Sci. 2 (1): 45, 1959 (Abeywickrama 1959). Bas.: Mastigobryum obtusatum Mitt., J. Proc. Linn. Soc., Bot. 5
(18): 106, 1860 [1861] (Mitten 1860c).
** Bazzania okaritana Meagher et Glenny, J. Bryol. 29 (1): 60, 2007 (Meagher and
Glenny 2007).
** Bazzania orbanii Pócs, Acta Biol. Pl. Agr. 1: 16, 2010 [2011] (Pócs 2010c).
** Bazzania ovistipula (Steph.) Abeyw., Ceylon J. Sci., Biol. Sci. 2 (1): 45, 1959
(Abeywickrama 1959). Bas.: Mastigobryum ovistipulum Steph., Bull. Herb. Boissier (sér. 2) 8 (10): 760 (444), 1908 (Stephani 1908c).
** Bazzania pallidevirens (Steph.) Fulford, Ann. Cryptog. Phytopathol. 3: 42, 1946
(Fulford 1946). Bas.: Mastigobryum pallidevirens Steph., Bull. Herb. Boissier (sér.
2) 8 (11): 849 (473), 1908 (Stephani 1908a).
** Bazzania papillosa S.W.Arnell, Svensk Bot. Tidskr. 59 (1): 67, 1965 (Arnell 1965).
** Bazzania paradoxa (Sande Lac.) Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 307, 1896
(Stephani 1896a). Bas.: Mastigobryum paradoxum Sande Lac., Ned. Kruidk. Arch.
3: 419, 1854 [1855] (Sande Lacoste 1854).
*** Bazzania parisii (Steph.) N.Kitag., J. Hattori Bot. Lab. 47: 135, 1980 (Kitagawa
1980). Bas.: Mastigobryum parisii Steph., Bull. Herb. Boissier (sér. 2) 8 (10): 769
(453), 1908 (Stephani 1908c).
158
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Bazzania parvitexta Steph., Hedwigia 32 (4): 211, 1893 (Stephani 1893c).
** Bazzania patens (Mont.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Herpetium patens Mont., Ann. Sci.
Nat. Bot. (sér. 2) 20: 295, 1844 (Montagne 1844b).
** Bazzania patentistipa (Sande Lac.) Schifn., Consp. Hepat. Arch. Ind.: 168, 1898
(Schifner 1898b). Bas.: Mastigobryum patentistipum Sande Lac., Ann. Mus. Bot.
Lugduno-Batavi 1: 302, 1864 (Sande Lacoste 1864).
** Bazzania paucidens (Steph.) H.A.Mill., Phytologia 47 (4): 321, 1981 (Miller
1981). Bas.: Mastigobryum paucidens Steph., Bull. Herb. Boissier (sér. 2) 8 (11):
860 (484), 1908 (Stephani 1908a).
** Bazzania pearsonii Steph., Hedwigia 32 (4): 212, 1893 (Stephani 1893c).
*** Bazzania pectinata (Lindenb. et Gottsche) Schifn., Nova Acta Acad. Caes. Leop.Carol. German. Nat. Cur. 60 (2): 259, 1893 (Schifner 1893a). Bas.: Mastigobryum pectinatum Lindenb. et Gottsche, Sp. Hepat. (Lindenberg) 8-11: 84, 1851
(Lindenberg and Gottsche 1851b).
** Bazzania perfalcata N.Kitag., J. Hattori Bot. Lab. 47: 135, 1980 (Kitagawa 1980).
** Bazzania perrotana E.W.Jones, J. Bryol. 8 (3): 310, 1975 (Jones 1975). Nom.
nov. pro Mastigobryum perrotanum Steph., Sp. Hepat. (Stephani) 6: 476, 1924
(Stephani 1924), nom. illeg.
** Bazzania peruviana (Nees) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum peruvianum Nees,
Syn. Hepat. 2: 220, 1845 (Gottsche et al. 1845a).
*** Bazzania phyllobola Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 372, 1885
(Spruce 1885).
*** Bazzania placophylla (Taylor) Grolle, Rev. Bryol. Lichénol. 27 (1/2): 54, 1958
(Grolle 1958). Bas.: Jungermannia placophylla Taylor, London J. Bot. 5: 276, 1846
(Taylor 1846a).
** Bazzania platycnema (Schwägr. ex Steph.) H.A.Mill., Bryologist 63 (2): 121, 1960
(Miller 1960). Bas.: Mastigobryum platycnemum Schwägr. ex Steph., Bull. Herb.
Boissier (sér. 2) 8 (10): 776 (460), 1908 (Stephani 1908c).
** Bazzania pompeana (Sande Lac.) Mitt., Trans. Linn. Soc. London, Bot. 3 (3): 200,
1891 (Mitten 1891). Bas.: Mastigobryum pompeanum Sande Lac., Ann. Mus. Bot.
Lugduno-Batavi 1: 304, 1864 (Sande Lacoste 1864).
*** Bazzania praerupta (Reinw., Blume et Nees) Trevis., Mem. Reale Ist. Lombardo
Sci. (Ser. 3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Jungermannia
praerupta Reinw., Blume et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 229, 1824 [1825] (Reinwardt et al. 1824a).
* Bazzania praerupta var. obliquata (Nees) Schifn., Consp. Hepat. Arch. Ind.: 170,
1898 (Schifner 1898b). Bas.: Jungermannia obliquata Nees, Enum. Pl. Crypt.
Javae: 62, 1830 (Nees 1830).
** Bazzania praerupta var. thermalis Schifn., Arch. Hydrobiol., suppl. 21 (3/4): 396,
1955 (Schifner 1955).
World checklist of hornworts and liverworts
159
** Bazzania pseudovittata N.Kitag. et T.Kodama, J. Hattori Bot. Lab. 39: 69, 1975
(Kitagawa and Kodama 1975b).
** Bazzania pusilla (Mitt.) Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 307, 1896 (Stephani
1896a). Bas.: Mastigobryum pusillum Mitt., Fl. vit.: 406, 1871 [1873] (Mitten 1871).
** Bazzania pycnophylla (Taylor) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum pycnophyllum
Taylor, London J. Bot. 5: 371, 1846 (Taylor 1846b).
** Bazzania quadratistipula H.A.Mill., Phytologia 47 (4): 321, 1981 (Miller 1981).
Nom. nov. pro Mastigobryum quadratum Steph., Sp. Hepat. (Stephani) 6: 477,
1924 (Stephani 1924), nom. illeg.
** Bazzania rabenhorstii (Steph.) Abeyw., Ceylon J. Sci., Biol. Sci. 2 (1): 45, 1959
(Abeywickrama 1959). Bas.: Mastigobryum rabenhorstii Steph., Bull. Herb. Boissier (sér. 2) 8 (10): 774 (458), 1908 (Stephani 1908c).
** Bazzania recurva (Mont.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Herpetium recurvum Mont., Ann.
Sci. Nat. Bot. (sér. 2) 19: 253, 1843 (Montagne 1843).
* Bazzania recurva var. major (Sande Lac.) Schifn., Consp. Hepat. Arch. Ind.: 172,
1898 (Schifner 1898b). Bas.: Mastigobryum recurvum var. majus Sande Lac., Ann.
Mus. Bot. Lugduno-Batavi 1: 304, 1864 (Sande Lacoste 1864).
** Bazzania relexa (Gottsche) Steph., Rev. Bryol. 18 (4): 56, 1891 (Renauld and
Cardot 1891). Bas.: Mastigobryum relexum Gottsche, Abh. Naturwiss. Vereins
Bremen 7: 347, 1882 (Gottsche 1882).
** Bazzania reinwardtii (Sande Lac.) Schifn., Consp. Hepat. Arch. Ind.: 172, 1898
(Schifner 1898b). Bas.: Mastigobryum reinwardtii Sande Lac., Pl. Ind. Batav. Orient.: 22, 1856 (Sande Lacoste 1856a).
** Bazzania renistipula Steph., Hedwigia 32 (4): 212, 1893 (Stephani 1893c).
** Bazzania revoluta (Steph.) N.Kitag., J. Hattori Bot. Lab. 36: 450, 1972 [1973]
(Kitagawa 1972). Bas.: Mastigobryum revolutum Steph., Bull. Herb. Boissier (sér.
2) 8 (12): 961 (511), 1908 (Stephani 1908b).
*** Bazzania rimosa Meagher, Nova Hedwigia 86 (3/4): 489, 2008 (Meagher 2008).
** Bazzania roccatii Gola, Ann. Bot. (Rome) 6 (2): 273, 1907 (Gola 1907).
*** Bazzania roraimensis (Steph.) Fulford, Ann. Cryptog. Phytopathol. 3: 27, 1946
(Fulford 1946). Bas.: Mastigobryum roraimense Steph., Trans. Linn. Soc. London,
Bot. 6 (1): 97, 1901 (Stephani 1901e).
** Bazzania sandvicensis (Gottsche ex Steph.) Steph., Bull. Herb. Boissier 5 (10):
841, 1897 (Stephani 1897c). Bas.: Mastigobryum sandvicense Gottsche ex Steph.,
Hedwigia 25 (5): 207, 1886 (Stephani 1886f).
** Bazzania sauropoda Meagher, Austrobaileya 7 (1): 129, 2005 (Meagher 2005a).
*** Bazzania scalaris Meagher, Telopea 11 (3): 247, 2006 (Meagher 2006).
*** Bazzania schlimiana (Gottsche) Fulford, Bull. Torrey Bot. Club 86 (6): 401, 1959
(Fulford 1959a). Bas.: Mastigobryum schlimianum Gottsche, Ann. Sci. Nat. Bot.
(sér. 5) 1: 140, 1864 (Gottsche 1864).
** Bazzania schultze-motelii N.Kitag., J. Hattori Bot. Lab. 47: 138, 1980 (Kitagawa 1980).
160
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Bazzania schusterana N.Kitag., J. Hattori Bot. Lab. 47: 139, 1980 (Kitagawa 1980).
*** Bazzania schwaneckiana (Hampe et Gottsche) Trevis., Mem. Reale Ist. Lombardo
Sci. (Ser. 3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum
schwaneckianum Hampe et Gottsche, Linnaea 25 (3): 345, 1852 [1853] (Hampe
and Gottsche 1852).
* Bazzania scutigera (Nees et Mont.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser.
3), C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Herpetium scutigerum
Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 9: 44, 1838 (Montagne 1838).
** Bazzania semicordata (Lindenb. et Gottsche) Kuntze, Revis. Gen. Pl. 2: 832, 1891
(Kuntze 1891). Bas.: Mastigobryum semicordatum Lindenb. et Gottsche, Syn. Hepat. 5: 720, 1847 (Gottsche et al. 1847).
* Bazzania serpentina (Nees) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 415, 1877 (Trevisan 1877). Bas.: Jungermannia serpentina Nees,
Enum. Pl. Crypt. Javae: 62, 1830 (Nees 1830). 113
** Bazzania serrapiculata Inoue et H.A.Mill., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 8
(2): 141, 1965 (Inoue and Miller 1965).
** Bazzania serrata Fulford, Bull. Torrey Bot. Club 86 (5): 321, 1959 (Fulford 1959b).
** Bazzania serrulatoides Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 200,
1934 (Horikawa 1934).
* Bazzania sikkimensis (Steph.) Herzog, Ann. Bryol. 12: 78, 1939 (Herzog 1939b).
Bas.: Mastigobryum sikkimense Steph., Hedwigia 44 (2): 73, 1905 (Stephani
1905h). 114
** Bazzania spinosa S.Okamura, J. Coll. Sci. Imp. Univ. Tokyo 38 (4): 2, 1916 (Okamura
1916).
** Bazzania spiralis (Reinw., Blume et Nees) Meijer, Blumea 10 (2): 381, 1960 (Meijer 1960). Bas.: Jungermannia spiralis Reinw., Blume et Nees, Nova Acta Phys.-Med.
Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 231, 1824 [1825] (Reinwardt et al. 1824a).
** Bazzania spruceana Steph., Hedwigia 32 (4): 213, 1893 (Stephani 1893c).
** Bazzania squarrosa (Steph.) H.A.Mill., Phytologia 47 (4): 321, 1981 (Miller
1981). Bas.: Mastigobryum squarrosum Steph., Bull. Herb. Boissier (sér. 2) 8 (10):
776 (460), 1908 (Stephani 1908c).
*** Bazzania stolonifera (Sw.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 415, 1877 (Trevisan 1877). Bas.: Jungermannia stolonifera Sw.,
Prodr. (Swartz): 144, 1788 (Swartz 1788).
* Bazzania stolonifera var. granatensis (Gottsche) Fulford, Ann. Cryptog. Phytopathol. 3: 51, 1946 (Fulford 1946). Bas.: Mastigobryum stoloniferum var. granatense Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 141, 1864 (Gottsche 1864).
** Bazzania stresemannii (Herzog) N.Kitag., Bull. Nara Univ. Educ., B 28 (2): 77,
1979 (Kitagawa 1979a). Bas.: Mastigobryum stresemannii Herzog, Beih. Bot. Centralbl. 38 (2): 324, 1921 (Herzog 1921).
113 Bazzania serpentina is possibly conspeciic with Bazzania bidens (Frahm et al. 1990).
114 Bazzania sikkimensis is possibly conspeciic with Bazzania sumbavensis (Long and Grolle 1990).
World checklist of hornworts and liverworts
161
** Bazzania subacuta (Mitt.) Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 307, 1896 (Stephani
1896a). Bas.: Mastigobryum subacutum Mitt., Fl. vit.: 406, 1871 [1873] (Mitten 1871).
** Bazzania subaequitexta (Steph.) N.Kitag., Bull. Nara Univ. Educ., B 28 (2): 81,
1979 (Kitagawa 1979a). Bas.: Mastigobryum subaequitextum Steph., Bull. Herb.
Boissier (sér. 2) 8 (11): 859 (483), 1908 (Stephani 1908a).
** Bazzania subintegra (Steph.) L.Söderstr. et A.Hagborg, Phytotaxa 202 (1): 69,
2015 (Söderström et al. 2015c). Bas.: Mastigobryum subintegrum Steph., Bull.
Herb. Boissier (sér. 2) 8 (10): 775 (459), 1908 (Stephani 1908c).
** Bazzania sublonga Fulford, Bull. Torrey Bot. Club 86 (5): 334, 1959 (Fulford 1959b).
** Bazzania subserrifolia (Beauverd) H.A.Mill., Phytologia 47 (4): 321, 1981 (Miller
1981). Bas.: Mastigobryum subserrifolium Beauverd, Sp. Hepat. (Stephani) 6: 480,
1924 (Stephani 1924).
** Bazzania subserrulata A.Evans, Pap. Michigan Acad. Sci. 17: 97, 1932 [1933]
(Evans 1932b).
*** Bazzania subtilis (Sande Lac.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum subtile Sande
Lac., Ann. Mus. Bot. Lugduno-Batavi 1: 302, 1864 (Sande Lacoste 1864).
** Bazzania succulenta N.Kitag., J. Hattori Bot. Lab. 47: 141, 1980 (Kitagawa 1980).
** Bazzania sumatrana (Sande Lac. ex Steph.) Steph., Hedwigia 32 (4): 209, 1893
(Stephani 1893c). Bas.: Mastigobryum sumatranum Sande Lac. ex Steph., Hedwigia
25 (6): 234, 1886 (Stephani 1886c).
* Bazzania sumbavensis (Gottsche ex Steph.) Steph., Hedwigia 32 (4): 204, 1893
(Stephani 1893c). Bas.: Mastigobryum sumbavense Gottsche ex Steph., Hedwigia
25 (6): 236, 1886 (Stephani 1886c). 115
*** Bazzania taleana (Gottsche) Fulford, Ann. Cryptog. Phytopathol. 3: 54, 1946
(Fulford 1946). Bas.: Mastigobryum taleanum Gottsche, Mexik. Leverm.: 131,
1863 (Gottsche 1863).
*** Bazzania tayloriana (Mitt.) Kuntze, Revis. Gen. Pl. 2: 832, 1891 (Kuntze 1891).
Bas.: Mastigobryum taylorianum Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 147,
1854 (Mitten 1854).
** Bazzania temariana (Steph.) H.A.Mill., Phytologia 47 (4): 321, 1981 (Miller
1981). Bas.: Mastigobryum temarianum Steph., Sp. Hepat. (Stephani) 3: 532,
1909 (Stephani 1909a).
*** Bazzania tessellata Meagher, Nova Hedwigia 92 (3/4): 492, 2011 (Meagher 2011).
** Bazzania tiaoloensis Mizut. et K.C.Chang, J. Hattori Bot. Lab. 60: 432, 1986
(Mizutani and Chang 1986).
*** Bazzania tricrenata (Wahlenb.) Lindb., Musci Fenn. Exsic., fasc. 2: [2 (adnot.)],
1872 (Brotherus 1872). Bas.: Jungermannia tricrenata Wahlenb., Fl. Carpat.
Princ.: 364, 1814 (Wahlenberg 1814).
** Bazzania tricrenata var. fulfordiae W.S.Hong, Bryologist 91 (4): 331, 1988 (Hong
1988).
115 Bazzania sumbavensis is possibly conspeciic with Bazzania japonica (Söderström et al. 2010a).
162
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Bazzania tridens (Reinw., Blume et Nees) Trevis., Mem. Reale Ist. Lombardo
Sci. (Ser. 3), C. Sci. Mat. 4 (13): 415, 1877 (Trevisan 1877). Bas.: Jungermannia
tridens Reinw., Blume et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol.
Nat. Cur. 12 (1): 228, 1824 [1825] (Reinwardt et al. 1824a).
*** Bazzania tridens var. assamica (Steph.) Pócs, J. Hattori Bot. Lab. 32: 86, 1969
(Pócs 1969). Bas.: Mastigobryum assamicum Steph., Hedwigia 24 (5): 216, 1885
(Stephani 1885a).
** Bazzania tridens var. cornutistipula (Steph.) Pócs, J. Hattori Bot. Lab. 32: 83, 1969
(Pócs 1969). Bas.: Mastigobryum cornutistipulum Steph., Rev. Bryol. 35 (2): 35,
1908 (Stephani 1908l).
*** Bazzania trilobata (L.) Gray, Nat. Arr. Brit. Pl. 1: 704, 1821 (Gray 1821). Bas.:
Jungermannia trilobata L., Sp. Pl. 1: 1133, 1753 (Linnaeus 1753).
** Bazzania trilobata var. depauperata (Müll.Frib.) Grolle, Lindbergia 1 (3/4): 197,
1972 [1973] (Grolle 1972b). Bas.: Pleuroschisma trilobatum var. depauperatum
Müll.Frib., Lebermoose 2 (18): 266, 1913 (Müller 1913b).
*** Bazzania uncigera (Reinw., Blume et Nees) Trevis., Mem. Reale Ist. Lombardo
Sci. (Ser. 3), C. Sci. Mat. 4 (13): 415, 1877 (Trevisan 1877). Bas.: Jungermannia
uncigera Reinw., Blume et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol.
Nat. Cur. 12 (1): 230, 1824 [1825] (Reinwardt et al. 1824a).
** Bazzania uncigera var. brevifolia Herzog, Trans. Brit. Bryol. Soc. 1 (4): 301, 1950
(Herzog 1950a).
** Bazzania uncigera var. gibba (Sande Lac.) Meijer, Blumea 10 (2): 378, 1960 (Meijer 1960). Bas.: Mastigobryum gibbum Sande Lac., Plagiochila Sandei: 8, 1856
(Sande Lacoste 1856c).
** Bazzania undulata Herzog, Trans. Brit. Bryol. Soc. 1 (4): 306, 1950 (Herzog 1950a).
** Bazzania vietnamica Pócs, J. Hattori Bot. Lab. 32: 90, 1969 (Pócs 1969).
** Bazzania vitiana Mitt., Hedwigia 32 (4): 214, 1893 (Stephani 1893c).
*** Bazzania vittata (Gottsche) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum vittatum Gottsche,
Syn. Hepat. 2: 216, 1845 (Gottsche et al. 1845a).
** Bazzania vittata var. luxurians (De Not.) Schifn., Consp. Hepat. Arch. Ind.: 178,
1898 (Schifner 1898b). Bas.: Mastigobryum vittatum var. luxurians De Not., Epat.
Borneo: 29, 1874 (De Notaris 1874).
** Bazzania wallichiana (Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 415, 1877 (Trevisan 1877). Bas.: Mastigobryum wallichianum
Lindenb., Syn. Hepat. 2: 229, 1845 (Gottsche et al. 1845a). 116
** Bazzania watanabei Inoue, J. Jap. Bot. 34 (9): 269, 1959 (Inoue 1959a).
* Bazzania wattsiana (Steph.) Meagher, Australas. Bryol. Newslett. 50: 8, 2005
(Meagher 2005b). Bas.: Mastigobryum wattsianum Steph., Bull. Herb. Boissier
(sér. 2) 8 (11): 850 (474), 1908 (Stephani 1908a). 117
116 Bazzania wallichiana seems to be a species complex also including Bazzania francana, Bazzania
intermedia and Bazzania marginata (Meagher 2010).
117 Bazzania wattsiana is possibly conspeciic with Bazzania tridens (Meagher 2010).
World checklist of hornworts and liverworts
163
** Bazzania wiltensii (Sande Lac. ex Steph.) Schifn., Consp. Hepat. Arch. Ind.: 180,
1898 (Schifner 1898b). Bas.: Mastigobryum wiltensii Sande Lac. ex Steph., Hedwigia 25 (6): 237, 1886 (Stephani 1886c).
** Bazzania wooroonooran Meagher, Nova Hedwigia 100 (3/4): 549, 2015 (Meagher
2015).
** Bazzania wrightii (Gottsche ex Steph.) Steph., Hedwigia 27 (11/12): 279, 1888
(Stephani 1888c). Bas.: Mastigobryum wrightii Gottsche ex Steph., Hedwigia 25
(6): 237, 1886 (Stephani 1886c).
** Bazzania yoshinagana (Steph.) Yasuda, Shokubutsugaku Kakuron: 711, 1911
(Yasuda 1911). Bas.: Mastigobryum yoshinaganum Steph., Bull. Herb. Boissier (sér.
2) 8 (11): 866 (490), 1908 (Stephani 1908a).
** Bazzania zollingeri (Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 414, 1877 (Trevisan 1877). Bas.: Mastigobryum zollingeri Lindenb., Bot. Zeitung (Berlin) 6 (25): 462, 1848 (Meissner 1848).
*** Bazzania zonulata Meagher, Nova Hedwigia 86 (3/4): 491, 2008 (Meagher 2008).
* Mastigopelma Mitt., Fl. vit.: 406, 1871 [1873] (Mitten 1871). 118
** Mastigopelma fragile (Steph.) N.Kitag., J. Hattori Bot. Lab. 36: 454, 1972 [1973]
(Kitagawa 1972). Bas.: Mastigobryum fragile Steph., Sp. Hepat. (Stephani) 6: 463,
1924 (Stephani 1924).
** Mastigopelma pulvinulatum (De Not.) Grolle, J. Hattori Bot. Lab. 33: 39, 1970
(Grolle 1970c). Bas.: Mastigobryum pulvinulatum De Not., Epat. Borneo: 40,
1874 (De Notaris 1874).
** Mastigopelma simplex Mitt., Fl. vit.: 406, 1871 [1873] (Mitten 1871).
** Mastigopelma subissum Grolle, J. Hattori Bot. Lab. 33: 36, 1970 (Grolle 1970c).
***
Drucelloideae R.M.Schust.
*** Drucella E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot. 2 (3): 45, 1962
(Hodgson 1962b).
*** Drucella integristipula (Steph.) E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot. 2
(3): 45, 1962 (Hodgson 1962b). Bas.: Lepidozia integristipula Steph., Sp. Hepat.
(Stephani) 6: 331, 1922 (Stephani 1922).
***
Lembidioideae R.M.Schust.
** Dendrolembidium Herzog, Ark. Bot. (n.ser.) 1 (13): 497, 1951 (Herzog 1951b).
*** Dendrolembidium dendroides (Carrington et Pearson) Herzog, Ark. Bot. (n.ser.) 1
(13): 500, 1951 (Herzog 1951b). Bas.: Lembidium dendroides Carrington et Pearson,
118 Mastigopelma may be better placed in Bazzania, but it is retained here until further studied.
164
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Proc. Linn. Soc. New South Wales (ser. 2) 2 (4): 1047, 1888 (Carrington and Pearson
1888a).
*** Dendrolembidium tenax (Grev.) Herzog, Ark. Bot. (n.ser.) 1 (13): 499, 1951
(Herzog 1951b). Bas.: Jungermannia tenax Grev., Ann. Lyceum Nat. Hist. New
York 1 (2): 277, 1825 (Greville 1825).
** Hygrolembidium R.M.Schust., J. Hattori Bot. Lab. 26: 277, 1963 (Schuster 1963b).
*** Hygrolembidium acrocladum (Berggr.) R.M.Schust., J. Hattori Bot. Lab. 26: 277,
1963 (Schuster 1963b). Bas.: Aplozia acroclada Berggr., New Zealand Hepat.: 9,
1898 (Berggren 1898).
** Hygrolembidium andinum (Herzog) R.M.Schust., Nova Hedwigia 10 (1/2): 23,
1965 (Schuster 1965b). Bas.: Lembidium andinum Herzog, Rev. Bryol. Lichénol.
23 (1/2): 48, 1954 (Herzog 1954).
*** Hygrolembidium australe (Steph.) Grolle, J. Jap. Bot. 41 (8): 229, 1966 (Grolle
1966d). Bas.: Hygrobiella australis Steph., Bull. Herb. Boissier (sér. 2) 8 (8): 574
(358), 1908 (Stephani 1908e).
*** Hygrolembidium boschianum (Sande Lac.) R.M.Schust., J. Hattori Bot. Lab. 26:
277, 1963 (Schuster 1963b). Bas.: Jungermannia boschiana Sande Lac., Ned.
Kruidk. Arch. 3: 521, 1855 (Sande Lacoste 1855).
** Hygrolembidium isophyllum R.M.Schust., Nova Hedwigia 15: 467, 1968 (Schuster
1968b).
*** Hygrolembidium rigidum R.M.Schust. et J.J.Engel, Phytologia 62 (1): 9, 1987
(Schuster and Engel 1987a).
*** Hygrolembidium triquetrum J.J.Engel et R.M.Schust., Phytologia 62 (1): 11, 1987
(Schuster and Engel 1987a).
** Hygrolembidium ventrosum (Mitt.) Grolle, Marion Prince Edw. Is: 233, 1971
(Grolle 1971d). Bas.: Lembidium ventrosum Mitt., J. Linn. Soc., Bot. 15 (82): 69,
1876 (Mitten 1876a).
** Isolembidium R.M.Schust., Nova Hedwigia 15: 466, 1968 (Schuster 1968b).
*** Isolembidium anomalum (Rodway) Grolle, J. Bryol. 10 (3): 264, 1979 (Grolle
1979b). Bas.: Lembidium anomalum Rodway, Tasm. Bryoph.: 70, 1917 (Rodway
1917b).
** Isolembidium anomalum var. cucullatum (E.A.Hodgs.) J.J.Engel et R.M.Schust.,
J. Hattori Bot. Lab. 63: 268, 1987 (Schuster and Engel 1987b). Bas.: Lembidium
cucullatum E.A.Hodgs., Rec. Domin. Mus. 4 (11): 110, 1962 (Hodgson 1962a).
*** Kurzia G.Martens, Flora 53 (27): 417, 1870 (0).
** subg. Kurzia
*** Kurzia capillaris (Sw.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 173, 1963 [1964]
(Grolle 1963b). Bas.: Jungermannia capillaris Sw., Prodr. (Swartz): 144, 1788
(Swartz 1788).
World checklist of hornworts and liverworts
165
*** Kurzia capillaris subsp. capillaris var. verrucosa (Steph.) Pócs, Proc. hird Meeting
Bryol. C. & E. Europe: 111, 1984 (Pócs 1984b). Bas.: Lepidozia verrucosa Steph.,
Hedwigia 24 (4): 167, 1885 (Stephani 1885f).
*** Kurzia capillaris subsp. paramicola Pócs, Acta Biol. Pl. Agr. 2: 102, 2012 (Pócs
2012a).
*** Kurzia capillaris subsp. stephanii (Renauld ex Steph.) Pócs, Proc. hird Meeting
Bryol. C. & E. Europe: 111, 1984 (Pócs 1984b). Bas.: Lepidozia stephanii Renauld
ex Steph., Bot. Gaz. 15 (11): 287, 1890 (Stephani 1890c).
*** Kurzia gonyotricha (Sande Lac.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 167, 1963
[1964] (Grolle 1963b). Bas.: Lepidozia gonyotricha Sande Lac., Ned. Kruidk. Arch.
3: 521, 1855 (Sande Lacoste 1855).
** Kurzia nemoides (Hook.f. et Taylor) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 173,
1963 [1964] (Grolle 1963b). Bas.: Jungermannia nemoides Hook.f. et Taylor, London J. Bot. 4: 84, 1845 (Hooker and Taylor 1845).
** subg. Micrisophylla (Fulford) J.J.Engel ex R.M.Schust., J. Hattori Bot. Lab. 48:
347, 1980 (Schuster 1980a). Bas.: Micrisophylla Fulford, Brittonia 14 (1): 124,
1962 (Fulford 1962a).
** Kurzia saddlensis (Besch. et C.Massal.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 174,
1963 [1964] (Grolle 1963b). Bas.: Lepidozia saddlensis Besch. et C.Massal., Bull.
Mens. Soc. Linn. Paris 1 (79): 637, 1886 (Bescherelle and Massalongo 1886).
** subg. Microlepidozia (Spruce) R.M.Schust., J. Hattori Bot. Lab. 48: 355, 1980
(Schuster 1980a). Bas.: Lepidozia subg. Microlepidozia Spruce, J. Bot. 14: 165,
1876 (Spruce 1876b).
** Kurzia calcarata (Steph.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 178, 1963 [1964]
(Grolle 1963b). Bas.: Lepidozia calcarata Steph., Sp. Hepat. (Stephani) 3: 592,
1909 (Stephani 1909a).
** Kurzia compacta (Steph.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 178, 1963 [1964]
(Grolle 1963b). Bas.: Lepidozia compacta Steph., Sp. Hepat. (Stephani) 3: 592,
1909 (Stephani 1909a).
** Kurzia helophila R.M.Schust., J. Hattori Bot. Lab. 48: 368, 1980 (Schuster 1980a).
** Kurzia helophila var. laccida R.M.Schust. ex J.J.Engel, Novon 17 (3): 310, 2007
(Engel 2007).
*** Kurzia hippurioides (Hook.f. et Taylor) Grolle, Rev. Bryol. Lichénol. 32 (3/4):
178, 1963 [1964] (Grolle 1963b). Bas.: Jungermannia hippurioides Hook.f. et
Taylor, London J. Bot. 3: 287 [387], 1844 (Hooker and Taylor 1844a).
** Kurzia hippurioides var. ornata J.J.Engel et G.L.Merr., J. Hattori Bot. Lab. 80:
228, 1996 (Engel and Smith Merrill 1996).
** Kurzia irregularis (Steph.) Grolle, J. Hattori Bot. Lab. 36: 548, 1972 [1973]
(Grolle 1972a). Bas.: Lepidozia irregularis Steph., Wiss. Ergebn. Deut. Zentr.-Afr.
Exped. (1907-08), Bot. 2: 120, 1911 (Stephani 1911a).
** Kurzia moniliformis J.J.Engel, Cryptog. Bryol. 26 (1): 73, 2005 (Engel 2005).
166
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*** Kurzia paucilora (Dicks.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 171, 1963
[1964] (Grolle 1963b). Bas.: Jungermannia paucilora Dicks., Fasc. Pl. Crypt. Brit.
2: 15, 1790 (Dickson 1790).
Incertae sedis
** Kurzia abbreviata Mizut., J. Hattori Bot. Lab. 38: 379, 1974 (Mizutani 1974).
** Kurzia abietinella (Herzog) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 170, 1963
[1964] (Grolle 1963b). Bas.: Lepidozia abietinella Herzog, Trans. Brit. Bryol. Soc.
1 (4): 311, 1950 (Herzog 1950a).
*** Kurzia bisetula (Steph.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 170, 1963 [1964]
(Grolle 1963b). Bas.: Lepidozia bisetula Steph., Sp. Hepat. (Stephani) 6: 323,
1922 (Stephani 1922).
** Kurzia borneensis Mizut., J. Hattori Bot. Lab. 38: 377, 1974 (Mizutani 1974).
** Kurzia brasiliensis (Steph.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 174, 1963
[1964] (Grolle 1963b). Bas.: Psiloclada brasiliensis Steph., Sp. Hepat. (Stephani)
3: 550, 1909 (Stephani 1909a).
** Kurzia brevicalycina (Steph.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 175, 1963
[1964] (Grolle 1963b). Bas.: Lepidozia brevicalycina Steph., Sp. Hepat. (Stephani)
3: 580, 1909 (Stephani 1909a).
* Kurzia caduciloba R.M.Schust., Beih. Nova Hedwigia 118: 273, 2000 (Schuster
2000a).
* Kurzia cucullifolia (Steph.) R.M.Schust., J. Hattori Bot. Lab. 48: 350, 1980 (Schuster 1980a). Bas.: Lepidozia cucullifolia Steph., Bih. Kongl. Svenska Vetensk.-Akad.
Handl. 26 (III, 6): 51, 1900 (Stephani 1900b). 119
*** Kurzia lagellifera (Steph.) Grolle, J. Jap. Bot. 39 (3): 80, 1964 (Grolle 1964c).
Bas.: Lepidozia lagellifera Steph., Sp. Hepat. (Stephani) 3: 571, 1909 (Stephani
1909a).
*** Kurzia fragilifolia R.M.Schust., J. Hattori Bot. Lab. 48: 364, 1980 (Schuster
1980a).
** Kurzia fragillima (Herzog) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 174, 1963
[1964] (Grolle 1963b). Bas.: Lepidozia fragillima Herzog, Nat. Hist. Juan Fernandez (Botany) 2 (5): 726, 1942 (Herzog 1942a), nom. illeg.
** Kurzia geniculata Mizut., J. Hattori Bot. Lab. 38: 383, 1974 (Mizutani 1974).
** Kurzia hawaica (C.M.Cooke) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 170, 1963
[1964] (Grolle 1963b). Bas.: Lepidozia hawaica C.M.Cooke, Trans. Connecticut
Acad. Arts 12 (1): 8, 1904 (Cooke 1904).
** Kurzia hispida (Steph.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 170, 1963 [1964]
(Grolle 1963b). Bas.: Lepidozia hispida Steph., Sp. Hepat. (Stephani) 3: 607, 1909
(Stephani 1909a).
119 Kurzia cucullifolia is of doubtful status (Engel 1978).
World checklist of hornworts and liverworts
167
** Kurzia lateconica (Steph.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 175, 1963 [1964]
(Grolle 1963b). Bas.: Lepidozia lateconica Steph., J. & Proc. Roy. Soc. New South
Wales 48 (1/2): 113, 1914 (Stephani and Watts 1914).
** Kurzia lineariloba Mizut., J. Hattori Bot. Lab. 38: 382, 1974 (Mizutani 1974).
*** Kurzia longicaulis Piippo, Acta Bot. Fenn. 131: 174, 1985 (Piippo 1985b).
** Kurzia makinoana (Steph.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 171, 1963
[1964] (Grolle 1963b). Bas.: Lepidozia makinoana Steph., Bull. Herb. Boissier 5
(2): 94, 1897 (Stephani 1897b).
** Kurzia mauiensis (H.A.Mill.) H.A.Mill., J. Hattori Bot. Lab. 30: 274, 1967
(Miller 1967). Bas.: Microlepidozia mauiensis H.A.Mill., Ark. Bot. (n.ser.) 5 (2):
496, 1963 (Miller 1963).
** Kurzia mollis (Steph.) J.J.Engel et R.M.Schust., Bryologist 79 (4): 514, 1976
[1977] (Engel 1976b). Bas.: Lepidozia mollis Steph., Sp. Hepat. (Stephani) 3: 601,
1909 (Stephani 1909a).
*** Kurzia nivicola (R.M.Schust.) E.D.Cooper, Phytotaxa 97 (2): 52, 2013 (Cooper
et al. 2013). Bas.: Telaranea nivicola R.M.Schust., Nova Hedwigia 15: 460, 1968
(Schuster 1968b).
*** Kurzia pallescens Grolle, Rev. Bryol. Lichénol. 32 (3/4): 177, 1963 [1964] (Grolle
1963b).
** Kurzia pallida Piippo, Acta Bot. Fenn. 131: 178, 1985 (Piippo 1985b).
*** Kurzia quinquespina J.J.Engel et G.L.Merr., J. Hattori Bot. Lab. 80: 217, 1996
(Engel and Smith Merrill 1996).
** Kurzia reversa (Carrington et Pearson) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 175,
1963 [1964] (Grolle 1963b). Bas.: Lepidozia reversa Carrington et Pearson, J. Bot.
27: 225, 1889 (Carrington and Pearson 1889).
** Kurzia setiformis (De Not.) J.J.Engel et R.M.Schust., Bryologist 79 (4): 514, 1976
[1977] (Engel 1976b). Bas.: Lepidozia setiformis De Not., Mem. Reale Accad. Sci.
Torino (ser. 2) 16: 225, 1857 (De Notaris 1857).
** Kurzia sexida (Steph.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 178, 1963 [1964]
(Grolle 1963b). Bas.: Lepidozia sexida Steph., Sp. Hepat. (Stephani) 3: 582, 1909
(Stephani 1909a).
** Kurzia sinensis K.C.Chang, Bull. Bot. Res., Harbin 4 (3): 83, 1984 (Chang and
Gao 1984).
** Kurzia sylvatica (A.Evans) Grolle, Herzogia 3: 77, 1973 (Grolle 1973a). Bas.: Lepidozia sylvatica A.Evans, Rhodora 6 (69): 186, 1904 (Evans 1904b).
*** Kurzia tasmanica (Steph.) E.D.Cooper, Phytotaxa 97 (2): 52, 2013 (Cooper et
al. 2013). Bas.: Lepidozia tasmanica Steph., Sp. Hepat. (Stephani) 3: 580, 1909
(Stephani 1909a).
** Kurzia tayloriana (H.A.Mill.) H.A.Mill., J. Hattori Bot. Lab. 30: 274, 1967 (Miller 1967). Bas.: Microlepidozia tayloriana H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 497,
1963 (Miller 1963).
168
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** Kurzia tenerrima (Mitt.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 171, 1963 [1964]
(Grolle 1963b). Bas.: Lepidozia tenerrima Mitt., Sp. Hepat. (Stephani) 3: 607,
1909 (Stephani 1909a).
** Kurzia touwii N.Kitag., Acta Phytotax. Geobot. 29 (1/5): 56, 1978 (Kitagawa 1978).
** Kurzia trichoclados (Müll.Frib.) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 171, 1963
[1964] (Grolle 1963b). Bas.: Lepidozia trichoclados Müll.Frib., Hedwigia 38 (4):
197, 1899 (Müller 1899).
*** Kurzia trilobata (R.M.Schust.) R.M.Schust., Beih. Nova Hedwigia 118: 270,
2000 (Schuster 2000a). Bas.: Kurzia quadriseta var. trilobata R.M.Schust., J. Hattori Bot. Lab. 48: 363, 1980 (Schuster 1980a).
** Kurzia verticellata (Carrington) Grolle, Rev. Bryol. Lichénol. 32 (3/4): 178, 1963
[1964] (Grolle 1963b). Bas.: Lepidozia verticellata Carrington, Pap. & Proc. Roy.
Soc. Tasmania 1887: 3, 1888 (Carrington and Pearson 1888b).
** Lembidium Mitt., Handb. N. Zeal. l. 2: 754, 1867 (Hooker 1867) nom. conserv.
*** Lembidium berggrenii Herzog, Ark. Bot. (n.ser.) 1 (13): 485, 1951 (Herzog 1951b).
*** Lembidium longifolium R.M.Schust., Phytologia 45 (5): 420, 1980 (Schuster
1980b).
*** Lembidium nutans (Hook.f. et Taylor) Mitt., Handb. N. Zeal. Fl. 2: 754, 1867
(Mitten 1867). Bas.: Jungermannia nutans Hook.f. et Taylor, London J. Bot. 3:
289 [389], 1844 (Hooker and Taylor 1844a).
** Lembidium nutans var. lagelliferum E.A.Hodgs., Rec. Domin. Mus. 4 (11): 110,
1962 (Hodgson 1962a).
** Megalembidium R.M.Schust., J. Hattori Bot. Lab. 26: 258, 1963 (Schuster 1963b).
*** Megalembidium insulanum (W.Martin et E.A.Hodgs.) R.M.Schust., J. Hattori
Bot. Lab. 26: 258, 1963 (Schuster 1963b). Bas.: Lembidium insulanum W.Martin et E.A.Hodgs., Trans. & Proc. Roy. Soc. New Zealand 78 (4): 497, 1950
(Martin 1950).
** Pseudocephalozia R.M.Schust., Nova Hedwigia 10 (1/2): 21, 1965 (Schuster
1965b).
** sect. Lobulatae R.M.Schust., J. Hattori Bot. Lab. 36: 371, 1972 (Schuster 1972).
** Pseudocephalozia cucullata J.J.Engel et R.M.Schust., J. Hattori Bot. Lab. 38: 694,
1974 (Schuster and Engel 1974).
*** Pseudocephalozia lobulata (Herzog) R.M.Schust., J. Hattori Bot. Lab. 36: 371,
1972 [1973] (Schuster 1972). Bas.: Lembidium lobulatum Herzog, Arch. Esc.
Fárm. Fac. Ci. Méd. Córdoba 7: 24, 1938 (Herzog and Hosseus 1938).
*** Pseudocephalozia quadriloba (Steph.) R.M.Schust., J. Hattori Bot. Lab. 36: 371,
1972 [1973] (Schuster 1972). Bas.: Isotachis quadriloba Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26 (III, 6): 54, 1900 (Stephani 1900b).
World checklist of hornworts and liverworts
169
** sect. Pseudocephalozia
** Pseudocephalozia lepidozioides R.M.Schust., Nova Hedwigia 10 (1/2): 22, 1965
(Schuster 1965b).
** Pseudocephalozia leptodictyon R.M.Schust., J. Hattori Bot. Lab. 36: 369, 1972
[1973] (Schuster 1972).
*** Pseudocephalozia paludicola R.M.Schust., Nova Hedwigia 10 (1/2): 21, 1965
(Schuster 1965b).
***
Lepidozioideae Müll.Frib.
*** Ceramanus E.D.Cooper, Phytotaxa 97 (2): 53, 2013 (Cooper et al. 2013).
*** Ceramanus centipes (Lindenb. et Gottsche) E.D.Cooper, Phytotaxa 97 (2): 54,
2013 (Cooper et al. 2013). Bas.: Lepidozia centipes Lindenb. et Gottsche, Syn.
Hepat. 2: 204, 1845 (Gottsche et al. 1845a).
*** Ceramanus clatritexta (Steph.) E.D.Cooper, Phytotaxa 97 (2): 54, 2013 (Cooper
et al. 2013). Bas.: Lepidozia clatritexta Steph., Sp. Hepat. (Stephani) 3: 583, 1909
(Stephani 1909a).
*** Ceramanus elegans (Colenso) E.D.Cooper, Phytotaxa 97 (2): 54, 2013 (Cooper et
al. 2013). Bas.: Lepidozia elegans Colenso, Trans. & Proc. New Zealand Inst. 21:
65, 1889 (Colenso 1889).
*** Ceramanus grossiseta (Steph.) E.D.Cooper, Phytotaxa 97 (2): 54, 2013 (Cooper
et al. 2013). Bas.: Lepidozia grossiseta Steph., Sp. Hepat. (Stephani) 3: 584, 1909
(Stephani 1909a).
*** Ceramanus perfragilis (J.J.Engel et G.L.Merr.) E.D.Cooper, Phytotaxa 97 (2): 54,
2013 (Cooper et al. 2013). Bas.: Telaranea perfragilis J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 72, 2004 (Engel and Smith Merrill 2004).
*** Ceramanus pruinosa (Herzog) E.D.Cooper, Phytotaxa 97 (2): 54, 2013 (Cooper
et al. 2013). Bas.: Lepidozia pruinosa Herzog, Memoranda Soc. Fauna Fl. Fennica
27: 93, 1952 (Herzog 1952c).
*** Ceramanus tuberifera (J.J.Engel et R.M.Schust.) E.D.Cooper, Phytotaxa 97 (2):
54, 2013 (Cooper et al. 2013). Bas.: Telaranea tuberifera J.J.Engel et R.M.Schust.,
Fieldiana, Bot. (n.ser.) 14: 2, 1983 (Engel and Schuster 1983).
*** Lepidozia (Dumort.) Dumort., Recueil Observ. Jungerm.: 19, 1835 (Dumortier
1835) nom. conserv. Bas.: Pleuroschisma sect. Lepidozia Dumort., Syll. Jungerm.
Europ.: 69, 1831 (Dumortier 1831). 120
** Lepidozia acantha J.J.Engel, Fieldiana, Bot. (n.ser.) 42: 71, 2001 (Engel and
Schuster 2001).
120 Lepidozia seems to be a monophyletic genus, but its infrataxonomy still has to be tested using molecular data. Grolle and Piippo (1984) could not study Lepidozia cordistipula and Lepidozia palmicola
since the types were destroyed in B.
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** Lepidozia aequiloba Steph., Sp. Hepat. (Stephani) 6: 319, 1922 (Stephani 1922).
** Lepidozia africana Steph., Sp. Hepat. (Stephani) 6: 320, 1922 (Stephani 1922).
*** Lepidozia alstonii Fulford, Mem. New York Bot. Gard. 11 (2): 211, 1966 (Fulford
1966).
** Lepidozia ambigua De Not., Epat. Borneo: 25, 1874 (De Notaris 1874).
** Lepidozia andicola Beauverd, Sp. Hepat. (Stephani) 6: 572, 1924 (Stephani 1924).
Nom. nov. pro Lepidozia appendiculata Steph., Biblioth. Bot. 87 (2): 225, 1916
(Stephani 1916a), nom. illeg.
** Lepidozia appressifolia Steph., Sp. Hepat. (Stephani) 3: 583, 1909 (Stephani 1909a).
*** Lepidozia armata Steph., Sp. Hepat. (Stephani) 3: 567, 1909 (Stephani 1909a).
** Lepidozia asymmetrica Steph., Sp. Hepat. (Stephani) 3: 586, 1909 (Stephani 1909a).
** Lepidozia auriculata Mitt., Sp. Hepat. (Stephani) 3: 579, 1909 (Stephani 1909a).
** Lepidozia australis (Lehm. et Lindenb.) Mitt., Fl. vit.: 406, 1871 [1873] (Mitten
1871). Bas.: Jungermannia australis Lehm. et Lindenb., Nov. Stirp. Pug. 6: 28,
1834 (Lehmann 1834).
** Lepidozia bidens J.J.Engel, Fieldiana, Bot. (n.ser.) 42: 79, 2001 (Engel and Schuster
2001).
** Lepidozia biloba Herzog, Ann. Bryol. 4: 83, 1931 (Herzog 1931b).
*** Lepidozia bisbiida Steph., Sp. Hepat. (Stephani) 3: 593, 1909 (Stephani 1909a).
** Lepidozia borneensis Steph., Sp. Hepat. (Stephani) 3: 625, 1909 (Stephani 1909a).
** Lepidozia bragginsiana E.D.Cooper et M.A.M.Renner, Phytotaxa 173 (2): 118,
2014 (Cooper and Renner 2014).
* Lepidozia brasiliensis Steph., Sp. Hepat. (Stephani) 3: 571, 1909 (Stephani 1909a). 121
** Lepidozia brevidentata Mitt., Fl. vit.: 406, 1871 [1873] (Mitten 1871).
** Lepidozia brevifolia Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 104, 1860 [1861] (Mitten 1860c).
** Lepidozia brevifolia var. planifolia Schifn., Ann. Bryol. 8: 155, 1935 (Verdoorn
1935).
** Lepidozia brotheri Steph., Sp. Hepat. (Stephani) 3: 623, 1909 (Stephani 1909a).
** Lepidozia bufalona Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2): 111,
1914 (Stephani and Watts 1914).
** Lepidozia bursifera S.Hatt. et Grolle, J. Hattori Bot. Lab. 30: 115, 1967 (Grolle 1967b).
** Lepidozia caespitosa Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 362, 1885
(Spruce 1885).
** Lepidozia caledonica Steph., Rev. Bryol. 35 (2): 31, 1908 (Stephani 1908l).
** Lepidozia caledonica var. tenuisecta Hürl., Bauhinia 8 (2): 109, 1985 (Hürlimann
1985).
* Lepidozia ceramensis Herzog, Hedwigia 66 (6): 340, 1926 (Herzog 1926).
** Lepidozia cherydrion Hürl., Bauhinia 8 (2): 109, 1985 (Hürlimann 1985).
** Lepidozia chiloensis Steph., Sp. Hepat. (Stephani) 6: 322, 1922 (Stephani 1922).
121 Lepidozia brasiliensis is conspeciic with Kurzia brasiliensis in Pócs (1984b), possibly conspeciic with
Lepidozia cupressina in Gradstein and Costa (2003), but it has later been accepted by many authors
(e.g. Schäfer-Verwimp and Pócs 2009)
World checklist of hornworts and liverworts
171
** Lepidozia chordulifera Taylor, London J. Bot. 5: 371, 1846 (Taylor 1846b).
*** Lepidozia cladorhiza (Reinw., Blume et Nees) Nees, Syn. Hepat. 2: 210, 1845
(Gottsche et al. 1845a). Bas.: Jungermannia cladorhiza Reinw., Blume et Nees,
Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 203, 1824
[1825] (Reinwardt et al. 1824a).
*** Lepidozia coilophylla Taylor, London J. Bot. 5: 370, 1846 (Taylor 1846b).
** Lepidozia coilophylla var. apiculiloba (Steph.) Fulford, Mem. New York Bot. Gard.
11 (2): 194, 1966 (Fulford 1966). Bas.: Lepidozia apiculiloba Steph., Sp. Hepat.
(Stephani) 6: 321, 1922 (Stephani 1922).
** Lepidozia communis Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2): 111,
1914 (Stephani and Watts 1914).
*** Lepidozia concinna Colenso, Trans. & Proc. New Zealand Inst. 18: 244, 1886
(Colenso 1886b).
** Lepidozia cordata Lindenb., Syn. Hepat. 2: 207, 1845 (Gottsche et al. 1845a).
* Lepidozia cordistipula Steph., Sp. Hepat. (Stephani) 6: 345, 1922 (Stephani 1922).
** Lepidozia crassitexta Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2): 111,
1914 (Stephani and Watts 1914).
*** Lepidozia cupressina (Sw.) Lindenb., Syn. Hepat. 2: 207, 1845 (Gottsche et al. 1845a).
Bas.: Jungermannia cupressina Sw., Prodr. (Swartz): 144, 1788 (Swartz 1788).
** Lepidozia cupressina subsp. natalensis (Steph.) Pócs, Proc. hird Meeting Bryol.
C. & E. Europe: 109, 1984 (Pócs 1984b). Bas.: Lepidozia natalensis Steph., Sp.
Hepat. (Stephani) 3: 562, 1909 (Stephani 1909a).
* Lepidozia cupressina subsp. pinnata (Hook.) Pócs, Proc. hird Meeting Bryol. C.
& E. Europe: 109, 1984 (Pócs 1984b). Bas.: Jungermannia reptans var. pinnata
Hook., Brit. Jungermann.: tab. 75, 1815 (Hooker 1815).
* Lepidozia cupressina subsp. quinqueida (Steph.) Pócs, Proc. hird Meeting Bryol.
C. & E. Europe: 109, 1984 (Pócs 1984b). Bas.: Lepidozia quinqueida Steph.,
Wiss. Ergebn. Deut. Zentr.-Afr. Exped. (1907-08), Bot. 2: 123, 1911 (Stephani
1911a).
** Lepidozia decaisnei Steph., Sp. Hepat. (Stephani) 3: 588, 1909 (Stephani 1909a).
** Lepidozia dendritica Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 362, 1885
(Spruce 1885).
** Lepidozia densa Herzog, Repert. Spec. Nov. Regni Veg. 21 (1/7): 26, 1925 (Herzog
1925a).
*** Lepidozia digitata Herzog, Trans. & Proc. Roy. Soc. New Zealand 68 (1): 45,
1938 (Herzog 1938c).
*** Lepidozia eenii S.W.Arnell, Svensk Bot. Tidskr. 57 (2): 190, 1963 (Arnell 1963a).
** Lepidozia elobata R.M.Schust., Fieldiana, Bot. (n.ser.) 42: 74, 2001 (Engel and
Schuster 2001).
** Lepidozia erosa Steph., Sp. Hepat. (Stephani) 3: 621, 1909 (Stephani 1909a).
** Lepidozia erronea Herzog, Nat. Hist. Juan Fernandez (Botany) 2 (5): 725, 1942
(Herzog 1942a). Nom. nov. pro Lepidozia fernandeziensis Steph., Sp. Hepat.
(Stephani) 6: 326, 1922 (Stephani 1922), nom. illeg.
172
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Lepidozia everettii Steph., Sp. Hepat. (Stephani) 3: 622, 1909 (Stephani 1909a).
Lepidozia everettii var. javensis Herzog, Ann. Bryol. 5: 78, 1932 (Herzog 1932b).
Lepidozia fauriana Steph., Sp. Hepat. (Stephani) 3: 631, 1909 (Stephani 1909a).
Lepidozia ferdinandi-muelleri Steph., Sp. Hepat. (Stephani) 3: 614, 1909 (Stephani
1909a).
Lepidozia ilamentosa (Lehm. et Lindenb.) Lehm. et Lindenb., Syn. Hepat. 2: 206,
1845 (Gottsche et al. 1845a). Bas.: Jungermannia ilamentosa Lehm. et Lindenb.,
Nov. Stirp. Pug. 6: 29, 1834 (Lehmann 1834).
Lepidozia istulosa Mitt., Fl. vit.: 406, 1871 [1873] (Mitten 1871).
Lepidozia lexuosa Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 103, 1860 [1861] (Mitten
1860c).
Lepidozia fuegiensis Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9):
63, 1911 (Stephani 1911b).
Lepidozia fugax J.J.Engel, Fieldiana, Bot. (n.ser.) 42: 63, 2001 (Engel and Schuster
2001).
Lepidozia gedena Steph., Sp. Hepat. (Stephani) 6: 327, 1922 (Stephani 1922).
Lepidozia glaucescens J.J.Engel, Fieldiana, Bot. (n.ser.) 42: 101, 2001 (Engel and
Schuster 2001).
Lepidozia glaucophylla (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 2: 207, 1845 (Gottsche et al. 1845a). Bas.: Jungermannia glaucophylla Hook.f.
et Taylor, London J. Bot. 3: 580, 1844 (Hooker and Taylor 1844c).
Lepidozia grandifolia Steph., Sp. Hepat. (Stephani) 3: 625, 1909 (Stephani 1909a).
Lepidozia griseola Herzog, Hedwigia 66 (6): 340, 1926 (Herzog 1926).
Lepidozia groenlandica Lehm., Nov. Stirp. Pug. 10: 7, 1857 (Lehmann 1857).
Lepidozia gwamii Piippo, Ann. Bot. Fenn. 21 (4): 311, 1984 (Piippo 1984b).
Lepidozia hampeana Lindenb., Syn. Hepat. 2: 208, 1845 (Gottsche et al. 1845a).
Lepidozia haskarliana (Gottsche, Lindenb. et Nees) Steph., Sp. Hepat. (Stephani)
3: 614, 1909 (Stephani 1909a). Bas.: Lepidozia supradecomposita β haskarliana
Gottsche, Lindenb. et Nees, Syn. Hepat. 2: 202, 1845 (Gottsche et al. 1845a).
Lepidozia hastatistipula Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2):
113, 1914 (Stephani and Watts 1914).
Lepidozia hirta Steph., Sp. Hepat. (Stephani) 3: 599, 1909 (Stephani 1909a).
Lepidozia holorhiza (Reinw., Blume et Nees) Nees, Syn. Hepat. 2: 210, 1845
(Gottsche et al. 1845a). Bas.: Jungermannia holorhiza Reinw., Blume et Nees,
Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 204, 1824
[1825] (Reinwardt et al. 1824a).
Lepidozia holorhiza var. laxa (Nees) Schifn., Consp. Hepat. Arch. Ind.: 186, 1898
(Schifner 1898b). Bas.: Jungermannia holorhiza β laxa Nees, Enum. Pl. Crypt.
Javae: 14, 1830 (Nees 1830).
Lepidozia inaequalis (Lehm. et Lindenb.) Lehm. et Lindenb., Syn. Hepat. 2: 209,
1845 (Gottsche et al. 1845a). Bas.: Jungermannia inaequalis Lehm. et Lindenb.,
Nov. Stirp. Pug. 5: 1, 1833 (Lehmann 1833).
Lepidozia incurvata Lindenb., Syn. Hepat. 2: 203, 1845 (Gottsche et al. 1845a).
World checklist of hornworts and liverworts
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173
Lepidozia infuscata Mitt., Fl. vit.: 406, 1871 [1873] (Mitten 1871).
Lepidozia integrifolia Doei, J. Hattori Bot. Lab. 63: 421, 1987 (Doei 1987).
Lepidozia jamaicensis Steph., Sp. Hepat. (Stephani) 3: 568, 1909 (Stephani 1909a).
Lepidozia kashyapii D.Singh et D.K.Singh, Nova Hedwigia 94 (1/2): 222, 2012
(Singh and Singh 2012).
Lepidozia kinabaluensis Mizut., J. Hattori Bot. Lab. 38: 372, 1974 (Mizutani 1974).
Lepidozia kirkii Steph., Sp. Hepat. (Stephani) 3: 598, 1909 (Stephani 1909a).
Lepidozia lacerifolia Steph., Sp. Hepat. (Stephani) 6: 332, 1922 (Stephani 1922).
Lepidozia laevifolia (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 2:
208, 1845 (Gottsche et al. 1845a). Bas.: Jungermannia laevifolia Hook.f. et Taylor,
London J. Bot. 3: 285 [385], 1844 (Hooker and Taylor 1844a).
Lepidozia laevifolia var. acutiloba J.J.Engel, Fieldiana, Bot. (n.ser.) 42: 61, 2001
(Engel and Schuster 2001).
Lepidozia laevifolia var. alpina R.M.Schust. et J.J.Engel, Fieldiana, Bot. (n.ser.) 42:
62, 2001 (Engel and Schuster 2001).
Lepidozia lindigiana Steph., Sp. Hepat. (Stephani) 3: 573, 1909 (Stephani 1909a).
Lepidozia loheri Steph., Sp. Hepat. (Stephani) 3: 621, 1909 (Stephani 1909a).
Lepidozia longifolia Steph., Sp. Hepat. (Stephani) 3: 606, 1909 (Stephani 1909a).
Lepidozia loriana Steph., Sp. Hepat. (Stephani) 6: 333, 1922 (Stephani 1922).
Lepidozia macrocolea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 363, 1885
(Spruce 1885).
Lepidozia massartiana Schifn. ex Steph., Sp. Hepat. (Stephani) 3: 611, 1909
(Stephani 1909a). Based on: Lepidozia massartiana Schifn., Hedwigia 39 (4): 196,
1900 (Schifner 1900b), nom. inval. 122
Lepidozia microphylla (Hook.) Lindenb., Syn. Hepat. 2: 202, 1845 (Gottsche et
al. 1845a). Bas.: Jungermannia microphylla Hook., Musci Exot. 1: tab. 80, 1818
(Hooker 1818).
Lepidozia microstipula Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2):
114, 1914 (Stephani and Watts 1914).
Lepidozia minima Steph., Sp. Hepat. (Stephani) 6: 335, 1922 (Stephani 1922).
Lepidozia minor (Gottsche, Lindenb. et Nees) Solari, Boll. Mus. Civico Storia Nat.
Verona 10: 203, 1983 [1985] (Solari and Hässel 1983). Bas.: Lepidozia truncatella
β minor Gottsche, Lindenb. et Nees, Syn. Hepat. 2: 209, 1845 (Gottsche et al.
1845a). 123
Lepidozia miqueliana Sande Lac., Ann. Mus. Bot. Lugduno-Batavi 1: 301, 1864
(Sande Lacoste 1864).
Lepidozia montana Steph., Sp. Hepat. (Stephani) 3: 587, 1909 (Stephani 1909a).
Lepidozia newtonii Steph., Sp. Hepat. (Stephani) 3: 623, 1909 (Stephani 1909a).
122 Lepidozia massartiana has not been studied recently. Stephani (1909a) confused it with Telaranea
cuneifolia (Söderström et al. 2010a).
123 Lepidozia minor may be conspeciic with a known taxon. Lepidozia minor sensu Solari and Hässel
(1983) is probably another taxon perhaps without name.
174
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Lepidozia nova Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2): 115, 1914
(Stephani and Watts 1914).
Lepidozia novae-zelandiae Steph., Sp. Hepat. (Stephani) 3: 595, 1909 (Stephani
1909a).
Lepidozia novae-zelandiae var. heterostipa R.M.Schust., Fieldiana, Bot. (n.ser.) 42:
70, 2001 (Engel and Schuster 2001).
Lepidozia novae-zelandiae var. minima R.M.Schust., Fieldiana, Bot. (n.ser.) 42:
71, 2001 (Engel and Schuster 2001).
Lepidozia obtusiloba Steph., Sp. Hepat. (Stephani) 3: 598, 1909 (Stephani 1909a).
Lepidozia obtusiloba var. parvula J.J.Engel, Fieldiana, Bot. (n.ser.) 42: 48, 2001
(Engel and Schuster 2001).
Lepidozia omeiensis P.C.Chen ex Mizut. et K.C.Chang, J. Hattori Bot. Lab. 60:
421, 1986 (Mizutani and Chang 1986).
Lepidozia ornata J.J.Engel, Fieldiana, Bot. (n.ser.) 42: 49, 2001 (Engel and Schuster
2001).
Lepidozia pallida Steph., Sp. Hepat. (Stephani) 3: 604, 1909 (Stephani 1909a). 124
Lepidozia palmicola Steph., Sp. Hepat. (Stephani) 6: 346, 1922 (Stephani 1922).
Lepidozia paschalis Steph., Sp. Hepat. (Stephani) 6: 336, 1922 (Stephani 1922).
Lepidozia patens Lindenb., Syn. Hepat. 2: 202, 1845 (Gottsche et al. 1845a).
Lepidozia paucifolia Steph., Sp. Hepat. (Stephani) 3: 610, 1909 (Stephani 1909a). 125
Lepidozia paupercula Steph., Sp. Hepat. (Stephani) 6: 337, 1922 (Stephani 1922).
Lepidozia pearsonii Spruce, J. Bot. 19: 34, 1881 (Spruce 1881b).
Lepidozia pendulina (Hook.) Lindenb., Syn. Hepat. 2: 208, 1845 (Gottsche et
al. 1845a). Bas.: Jungermannia pendulina Hook., Musci Exot. 1: tab. 60, 1818
(Hooker 1818).
Lepidozia peruviensis Steph., Sp. Hepat. (Stephani) 3: 575, 1909 (Stephani 1909a).
Lepidozia pinnaticruris Spruce ex Steph., Sp. Hepat. (Stephani) 3: 579, 1909
(Stephani 1909a).
Lepidozia plumula Herzog, Beih. Bot. Centralbl. 38 (2): 331, 1921 (Herzog 1921).
Lepidozia portoricensis Fulford, Mem. New York Bot. Gard. 11 (2): 187, 1966
(Fulford 1966).
Lepidozia procera Mitt., Bot. antarct. voy. III (Fl. Tasman. 2): 231, 1860 (Mitten
1860b).
Lepidozia pseudocupressina Schifn., Krit. Bemerk. Eur. Lebermoose 14: 9, 1919
(Schifner 1919).
Lepidozia pumila J.J.Engel, Fieldiana, Bot. (n.ser.) 42: 76, 2001 (Engel and Schuster
2001).
124 Lepidozia pallida is possibly Lepidozia chordulifera since all specimens cited by Stephani (1911b) and
examined by Engel (1978) belong to this species, except the supposed type specimen which does not
contain any Lepidozia material.
125 Lepidozia paucifolia is conspeciic with Lepidozia fauriana in del Rosario (1977), but Tan and Engel
(1986) kept them apart.
World checklist of hornworts and liverworts
175
*** Lepidozia quadridens (Nees) Nees, Syn. Hepat. 2: 209, 1845 (Gottsche et al.
1845a). Bas.: Jungermannia quadridens Nees, Enum. Pl. Crypt. Javae: 18, 1830
(Nees 1830).
** Lepidozia quadriida Lindenb., Syn. Hepat. 2: 203, 1845 (Gottsche et al. 1845a).
*** Lepidozia reptans (L.) Dumort., Recueil Observ. Jungerm.: 19, 1835 (Dumortier
1835). Bas.: Jungermannia reptans L., Sp. Pl. 1: 1133, 1753 (Linnaeus 1753).
** Lepidozia richardsii Herzog, Trans. Brit. Bryol. Soc. 1 (4): 313, 1950 (Herzog
1950a).
** Lepidozia rigida Steph., Sp. Hepat. (Stephani) 6: 340, 1922 (Stephani 1922).
** Lepidozia robusta Steph., Mém. Soc. Nat. Sci. Nat. Math. Cherbourg 29: 217,
1894 (Stephani 1894b).
* Lepidozia rufescens Steph., Biblioth. Bot. 87 (2): 226, 1916 (Stephani 1916a).
** Lepidozia sandvicensis Lindenb., Syn. Hepat. 2: 201, 1845 (Gottsche et al. 1845a).
** Lepidozia schwabei Herzog, Rev. Bryol. Lichénol. 23 (1/2): 49, 1954 (Herzog 1954).
** Lepidozia sellingiana H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 494, 1963 (Miller 1963).
*** Lepidozia septemida Steph., Sp. Hepat. (Stephani) 3: 588, 1909 (Stephani 1909a).
** Lepidozia serpens Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 364, 1885
(Spruce 1885).
*** Lepidozia serrulata J.J.Engel, J. Hattori Bot. Lab. 96: 273, 2004 (Engel 2004a).
*** Lepidozia setigera Steph., Sp. Hepat. (Stephani) 3: 599, 1909 (Stephani 1909a).
** Lepidozia sikkimensis Steph., Sp. Hepat. (Stephani) 6: 341, 1922 (Stephani 1922).
*** Lepidozia spinosissima (Hook.f. et Taylor) Mitt., Bot. antarct. voy. II (Fl. Nov.Zel. 2): 146, 1854 (Mitten 1854). Bas.: Sendtnera spinosissima Hook.f. et Taylor,
London J. Bot. 5: 373, 1846 (Taylor 1846b).
* Lepidozia squamifolia Steph., Sp. Hepat. (Stephani) 6: 341, 1922 (Stephani 1922).
*** Lepidozia squarrosa Steph., Sp. Hepat. (Stephani) 3: 573, 1909 (Stephani 1909a).
** Lepidozia stahlii Steph., Sp. Hepat. (Stephani) 3: 616, 1909 (Stephani 1909a).
** Lepidozia stuhlmannii Steph., Bot. Jahrb. Syst. 20 (3): 308, 1895 (Stephani 1895a).
** Lepidozia stuhlmannii var. carnosa (Steph.) Pócs, Lidia 4 (1): 22, 1997 (Lye and
Pócs 1997). Bas.: Lepidozia carnosa Steph., Wiss. Ergebn. Deut. Zentr.-Afr. Exped.
(1907-08), Bot. 2: 122, 1911 (Stephani 1911a).
** Lepidozia stuhlmannii subsp. pulvinata (Steph.) Pócs, Trop. Bryol. 9: 127, 1994
(Pócs 1994c). Bas.: Lepidozia pulvinata Steph., Wiss. Ergebn. Deut. Zentr.-Afr.
Exped. (1907-08), Bot. 2: 121, 1911 (Stephani 1911a).
** Lepidozia subdichotoma Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 361,
1885 (Spruce 1885).
** Lepidozia subintegra Lindenb., Syn. Hepat. 2: 201, 1845 (Gottsche et al. 1845a).
** Lepidozia subtransversa Steph., Bull. Herb. Boissier 5 (2): 95, 1897 (Stephani 1897b).
** Lepidozia subtrichodes Steph., Sp. Hepat. (Stephani) 3: 615, 1909 (Stephani 1909a).
** Lepidozia succida Mitt., Trans. Linn. Soc. London 23 (1): 57, 1860 (Mitten 1860a).
** Lepidozia supradecomposita Lindenb., Syn. Hepat. 2: 202, 1845 (Gottsche et al. 1845a).
* Lepidozia supradecomposita var. falcifolia Herzog, Mitt. Inst. Allg. Bot. Hamburg 7
(3): 191, 1931 (Herzog 1931a).
176
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Lepidozia suyungii C.Gao et X.L.Bai, J. Hattori Bot. Lab. 92: 192, 2002 (Gao and
Bai 2002).
** Lepidozia terricola Steph., Sp. Hepat. (Stephani) 3: 585, 1909 (Stephani 1909a).
** Lepidozia triangulifolia Steph., Sp. Hepat. (Stephani) 6: 344, 1922 (Stephani 1922).
*** Lepidozia trichodes (Reinw., Blume et Nees) Nees, Syn. Hepat. 2: 203, 1845
(Gottsche et al. 1845a). Bas.: Jungermannia trichodes Reinw., Blume et Nees, Nova
Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 199, 1824 [1825]
(Reinwardt et al. 1824a).
* Lepidozia tricuspidata Steph., Sp. Hepat. (Stephani) 6: 344, 1922 (Stephani 1922).
** Lepidozia tunguraguae Steph., Sp. Hepat. (Stephani) 6: 345, 1922 (Stephani 1922).
** Lepidozia ubangiensis Steph., Sp. Hepat. (Stephani) 3: 561, 1909 (Stephani
1909a).
*** Lepidozia udarii S.C.Srivast., D.Kumar et D.Sharma, Cryptog. Bryol. Lichénol. 9
(3): 237, 1988 (Srivastava et al. 1988).
*** Lepidozia ulothrix (Schwägr.) Lindenb., Syn. Hepat. 2: 210, 1845 (Gottsche et
al. 1845a). Bas.: Jungermannia ulothrix Schwägr., Hist. Musc. Hepat. Prodr.: 21,
1814 (Schwägrichen 1814).
** Lepidozia vitrea Steph., Bull. Herb. Boissier 5 (2): 96, 1897 (Stephani 1897b).
** Lepidozia wattsiana Steph., Sp. Hepat. (Stephani) 3: 586, 1909 (Stephani 1909a).
** Lepidozia weymouthiana Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2):
117, 1914 (Stephani and Watts 1914).
*
*
Excluded from the genus
Lepidozia hexiloba Pearson, Ann. Cryptog. Exot. 4 (2): 67, 1931 (Pearson 1931a). 126
Lepidozia parvistipa Taylor, London J. Bot. 5: 370, 1846 (Taylor 1846b). 127
*** Neolepidozia Fulford et J.Taylor, Brittonia 11 (2): 81, 1959 (Fulford and Taylor
1959a).
*** Neolepidozia aubertii (Jovet-Ast) E.D.Cooper, Phytotaxa 97 (2): 55, 2013 (Cooper et
al. 2013). Bas.: Lepidozia aubertii Jovet-Ast, Candollea 11: 35, 1947 (Jovet-Ast 1947a).
*** Neolepidozia autoica (J.J.Engel et G.L.Merr.) E.D.Cooper, Phytotaxa 97 (2): 55,
2013 (Cooper et al. 2013). Bas.: Telaranea autoica J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 124, 2004 (Engel and Smith Merrill 2004).
*** Neolepidozia capilligera (Schwägr.) Fulford et J.Taylor, Brittonia 11 (2): 84, 1959
(Fulford and Taylor 1959a). Bas.: Jungermannia capilligera Schwägr., Hist. Musc.
Hepat. Prodr.: 21, 1814 (Schwägrichen 1814).
*** Neolepidozia consobrina (J.J.Engel et G.L.Merr.) E.D.Cooper, Phytotaxa 97 (2):
55, 2013 (Cooper et al. 2013). Bas.: Telaranea consobrina J.J.Engel et G.L.Merr.,
Novon 9 (3): 339, 1999 (Engel and Smith Merrill 1999a).
126 Lepidozia hexiloba is possibly an Arachniopsis or Telaranea species (Wigginton and Grolle 1996).
127 Lepidozia parvistipa is possibly conspeciic with Ceramanus clatritexta.
World checklist of hornworts and liverworts
177
*** Neolepidozia cuneifolia (Steph.) Fulford et J.Taylor, Brittonia 11 (2): 85, 1959
(Fulford and Taylor 1959a). Bas.: Lepidozia cuneifolia Steph., Sp. Hepat. (Stephani) 3: 618, 1909 (Stephani 1909a).
*** Neolepidozia disparata (J.J.Engel et G.L.Merr.) E.D.Cooper, Phytotaxa 97 (2): 55,
2013 (Cooper et al. 2013). Bas.: Telaranea disparata J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 147, 2004 (Engel and Smith Merrill 2004).
*** Neolepidozia disticha (Steph.) Fulford et J.Taylor, Brittonia 11 (2): 85, 1959
(Fulford and Taylor 1959a). Bas.: Lepidozia disticha Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9): 62, 1911 (Stephani 1911b).
*** Neolepidozia gibbsiana (Steph.) E.D.Cooper, Phytotaxa 97 (2): 55, 2013 (Cooper
et al. 2013). Bas.: Lepidozia gibbsiana Steph., Sp. Hepat. (Stephani) 6: 328, 1922
(Stephani 1922).
*** Neolepidozia heterotexta (Steph.) E.D.Cooper, Phytotaxa 97 (2): 55, 2013 (Cooper
et al. 2013). Bas.: Lepidozia heterotexta Steph., Sp. Hepat. (Stephani) 6: 329, 1922
(Stephani 1922).
*** Neolepidozia hodgsoniae (J.J.Engel et G.L.Merr.) E.D.Cooper, Phytotaxa 97 (2):
55, 2013 (Cooper et al. 2013). Bas.: Telaranea hodgsoniae J.J.Engel et G.L.Merr.,
Phytologia 79 (3): 251, 1995 [1996] (Engel and Smith Merrill 1995).
*** Neolepidozia longitudinalis (Herzog) E.D.Cooper, Phytotaxa 97 (2): 56, 2013
(Cooper et al. 2013). Bas.: Lepidozia longitudinalis Herzog, Trans. Brit. Bryol.
Soc. 1 (4): 312, 1950 (Herzog 1950a).
*** Neolepidozia mamillosa (Schifn.) E.D.Cooper, Phytotaxa 97 (2): 56, 2013 (Cooper et al. 2013). Bas.: Lepidozia mamillosa Schifn., Nova Acta Acad. Caes. Leop.Carol. German. Nat. Cur. 60 (2): 254, 1893 (Schifner 1893a).
*** Neolepidozia meridiana (E.A.Hodgs.) E.D.Cooper, Phytotaxa 97 (2): 56, 2013
(Cooper et al. 2013). Bas.: Lepidozia meridiana E.A.Hodgs., Trans. Roy. Soc.
New Zealand 83 (4): 611, 1956 (Hodgson 1956).
*** Neolepidozia oligophylla (Lehm. et Lindenb.) Fulford et J.Taylor, Brittonia 11 (2):
84, 1959 (Fulford and Taylor 1959a). Bas.: Jungermannia oligophylla Lehm. et
Lindenb., Nov. Stirp. Pug. 6: 26, 1834 (Lehmann 1834).
*** Neolepidozia ophiria (Gottsche) E.D.Cooper, Phytotaxa 97 (2): 56, 2013 (Cooper
et al. 2013). Bas.: Lepidozia ophiria Gottsche, Sp. Hepat. (Stephani) 3: 611, 1909
(Stephani 1909a).
*** Neolepidozia palmata (J.J.Engel et G.L.Merr.) E.D.Cooper, Phytotaxa 97 (2): 56,
2013 (Cooper et al. 2013). Bas.: Telaranea palmata J.J.Engel et G.L.Merr., Novon
9 (3): 344, 1999 (Engel and Smith Merrill 1999a).
*** Neolepidozia paludicola (E.A.Hodgs.) E.D.Cooper, Phytotaxa 97 (2): 56, 2013
(Cooper et al. 2013). Bas.: Lepidozia meridiana var. paludicola E.A.Hodgs., Trans.
Roy. Soc. New Zealand 83 (4): 611, 1956 (Hodgson 1956).
*** Neolepidozia papulosa (Steph.) Fulford et J.Taylor, Brittonia 11 (2): 85, 1959 (Fulford and Taylor 1959a). Bas.: Lepidozia papulosa Steph., Sp. Hepat. (Stephani) 3:
609, 1909 (Stephani 1909a).
178
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Neolepidozia parvifolia (Steph.) Fulford et J.Taylor, Brittonia 11 (2): 85, 1959
(Fulford and Taylor 1959a). Bas.: Lepidozia parvifolia Steph., Sp. Hepat. (Stephani) 6: 337, 1922 (Stephani 1922).
*** Neolepidozia patentissima (Hook.f. et Taylor) E.D.Cooper, Phytotaxa 97 (2): 56,
2013 (Cooper et al. 2013). Bas.: Jungermannia patentissima Hook.f. et Taylor,
London J. Bot. 3: 286 [386], 1844 (Hooker and Taylor 1844a).
*** Neolepidozia patentissima var. ampliata (J.J.Engel et G.L.Merr.) E.D.Cooper, Phytotaxa 97 (2): 56, 2013 (Cooper et al. 2013). Bas.: Telaranea patentissima var.
ampliata J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 48, 2004 (Engel and
Smith Merrill 2004).
*** Neolepidozia patentissima var. zebrina (J.J.Engel et G.L.Merr.) E.D.Cooper, Phytotaxa 97 (2): 57, 2013 (Cooper et al. 2013). Bas.: Telaranea patentissima var.
zebrina J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 50, 2004 (Engel and
Smith Merrill 2004).
*** Neolepidozia pennata (J.J.Engel et G.L.Merr.) E.D.Cooper, Phytotaxa 97 (2): 57,
2013 (Cooper et al. 2013). Bas.: Telaranea pennata J.J.Engel et G.L.Merr., Phytologia 79 (3): 252, 1995 [1996] (Engel and Smith Merrill 1995).
*** Neolepidozia planifolia (Steph.) E.D.Cooper, Phytotaxa 97 (2): 57, 2013 (Cooper
et al. 2013). Bas.: Lepidozia planifolia Steph., Sp. Hepat. (Stephani) 3: 629, 1909
(Stephani 1909a).
*** Neolepidozia praenitens (Lehm. et Lindenb.) E.D.Cooper, Phytotaxa 97 (2): 57,
2013 (Cooper et al. 2013). Bas.: Jungermannia praenitens Lehm. et Lindenb., Nov.
Stirp. Pug. 6: 27, 1834 (Lehmann 1834).
*** Neolepidozia praenitens var. dentifolia (J.J.Engel et G.L.Merr.) E.D.Cooper, Phytotaxa 97 (2): 57, 2013 (Cooper et al. 2013). Bas.: Telaranea praenitens var. dentifolia J.J.Engel et G.L.Merr., Phytologia 79 (3): 253, 1995 [1996] (Engel and
Smith Merrill 1995).
*** Neolepidozia quadristipula (Steph.) E.D.Cooper, Phytotaxa 97 (2): 57, 2013
(Cooper et al. 2013). Bas.: Lepidozia quadristipula Steph., J. & Proc. Roy. Soc.
New South Wales 48 (1/2): 115, 1914 (Stephani and Watts 1914).
*** Neolepidozia rectangularis (R.M.Schust.) E.D.Cooper, Phytotaxa 97 (2): 57, 2013
(Cooper et al. 2013). Bas.: Telaranea rectangularis R.M.Schust., Phytologia 39 (4):
241, 1978 (Schuster 1978a).
*** Neolepidozia seriatitexta (Steph.) Fulford, Mem. New York Bot. Gard. 11 (2): 215,
1966 (Fulford 1966). Bas.: Lepidozia seriatitexta Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26 (III, 6): 53, 1900 (Stephani 1900b).
*** Neolepidozia tetrapila (Hook.f. et Taylor) E.D.Cooper, Phytotaxa 97 (2): 57, 2013
(Cooper et al. 2013). Bas.: Lepidozia tetrapila Hook.f. et Taylor, London J. Bot. 5:
370, 1846 (Taylor 1846b).
*** Neolepidozia tetrapila var. cancellata (Colenso) E.D.Cooper, Phytotaxa 97 (2): 57,
2013 (Cooper et al. 2013). Bas.: Lepidozia cancellata Colenso, Trans. & Proc.
New Zealand Inst. 18: 244, 1886 (Colenso 1886b).
World checklist of hornworts and liverworts
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*** Neolepidozia tetrapila var. roseana (Steph.) E.D.Cooper, Phytotaxa 97 (2): 58,
2013 (Cooper et al. 2013). Bas.: Lepidozia roseana Steph., Sp. Hepat. (Stephani)
3: 590, 1909 (Stephani 1909a).
*** Neolepidozia tridactylis (Lehm. et Lindenb.) E.D.Cooper, Phytotaxa 97 (2): 58,
2013 (Cooper et al. 2013). Bas.: Jungermannia tridactylis Lehm. et Lindenb., Nov.
Stirp. Pug. 4: 41, 1832 (Lehmann 1832).
*** Neolepidozia triida (Steph.) E.D.Cooper, Phytotaxa 97 (2): 58, 2013 (Cooper et
al. 2013). Bas.: Lepidozia triida Steph., Wiss. Ergebn. Deut. Zentr.-Afr. Exped.
(1907-08), Bot. 2: 120, 1911 (Stephani 1911a).
*** Neolepidozia verruculosa (J.J.Engel et G.L.Merr.) E.D.Cooper, Phytotaxa 97 (2):
58, 2013 (Cooper et al. 2013). Bas.: Telaranea verruculosa J.J.Engel et G.L.Merr.,
Fieldiana, Bot. (n.ser.) 44: 197, 2004 (Engel and Smith Merrill 2004).
*** Neolepidozia wallichiana (Gottsche) Fulford et J.Taylor, Brittonia 11 (2): 84, 1959
(Fulford and Taylor 1959a). Bas.: Lepidozia wallichiana Gottsche, Syn. Hepat. 2:
204, 1845 (Gottsche et al. 1845a).
*** Tricholepidozia (R.M.Schust.) E.D.Cooper, Phytotaxa 97 (2): 58, 2013 (Cooper et al. 2013). Bas.: Telaranea subg. Tricholepidozia R.M.Schust., J. Hattori Bot.
Lab. 26: 256, 1963 (Schuster 1963b).
*** Tricholepidozia chaetocarpa (Pearson) E.D.Cooper, Phytotaxa 97 (2): 58, 2013
(Cooper et al. 2013). Bas.: Lepidozia chaetocarpa Pearson, J. Linn. Soc., Bot. 46
(305): 27, 1922 (Pearson 1922b).
*** Tricholepidozia fernandeziensis (Steph.) E.D.Cooper, Phytotaxa 97 (2): 58, 2013
(Cooper et al. 2013). Bas.: Lepidozia fernandeziensis Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9): 63, 1911 (Stephani 1911b).
*** Tricholepidozia ferruginea (J.J.Engel et G.L.Merr.) E.D.Cooper, Phytotaxa 97 (2):
58, 2013 (Cooper et al. 2013). Bas.: Telaranea ferruginea J.J.Engel et G.L.Merr.,
Fieldiana, Bot. (n.ser.) 44: 159, 2004 (Engel and Smith Merrill 2004).
*** Tricholepidozia issifolia (Steph.) E.D.Cooper, Phytotaxa 97 (2): 58, 2013 (Cooper
et al. 2013). Bas.: Lepidozia issifolia Steph., Sp. Hepat. (Stephani) 3: 610, 1909
(Stephani 1909a).
*** Tricholepidozia jowettiana (H.A.Mill.) E.D.Cooper, Phytotaxa 97 (2): 59, 2013
(Cooper et al. 2013). Bas.: Telaranea jowettiana H.A.Mill., J. Bryol. 14 (2): 235,
1986 (Miller 1986).
*** Tricholepidozia kaindina (Grolle) E.D.Cooper, Phytotaxa 97 (2): 59, 2013 (Cooper et al. 2013). Bas.: Telaranea kaindina Grolle, J. Hattori Bot. Lab. 31: 9, 1968
(Grolle 1968a).
*** Tricholepidozia kogiana (Steph.) E.D.Cooper, Phytotaxa 97 (2): 59, 2013 (Cooper
et al. 2013). Bas.: Lepidozia kogiana Steph., Sp. Hepat. (Stephani) 6: 332, 1922
(Stephani 1922).
*** Tricholepidozia lawesii (Steph.) E.D.Cooper, Phytotaxa 97 (2): 59, 2013 (Cooper
et al. 2013). Bas.: Lepidozia lawesii Steph., Hedwigia 28 (4): 264, 1889 (Stephani
1889c).
180
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*** Tricholepidozia leratii (Steph.) E.D.Cooper, Phytotaxa 167 (2): 218, 2014 (Cooper et al. 2014). Bas.: Lepidozia leratii Steph., Sp. Hepat. (Stephani) 6: 333, 1922
(Stephani 1922).
*** Tricholepidozia lindenbergii (Gottsche) E.D.Cooper, Phytotaxa 97 (2): 59, 2013
(Cooper et al. 2013). Bas.: Lepidozia lindenbergii Gottsche, Syn. Hepat. 2: 213,
1845 (Gottsche et al. 1845a).
*** Tricholepidozia lindenbergii var. complanata (J.J.Engel et G.L.Merr.) E.D.Cooper,
Phytotaxa 97 (2): 59, 2013 (Cooper et al. 2013). Bas.: Telaranea lindenbergii var.
complanata J.J.Engel et G.L.Merr., Phytologia 79 (3): 252, 1995 [1996] (Engel
and Smith Merrill 1995).
*** Tricholepidozia lindenbergii var. mellea (J.J.Engel et G.L.Merr.) E.D.Cooper, Phytotaxa 97 (2): 59, 2013 (Cooper et al. 2013). Bas.: Telaranea lindenbergii var. mellea J.J.Engel et G.L.Merr., Phytologia 79 (3): 252, 1995 [1996] (Engel and Smith
Merrill 1995).
*** Tricholepidozia lindenbergii var. papillata (J.J.Engel et G.L.Merr.) E.D.Cooper,
Phytotaxa 97 (2): 59, 2013 (Cooper et al. 2013). Bas.: Telaranea lindenbergii var.
papillata J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 83, 2004 (Engel and
Smith Merrill 2004).
*** Tricholepidozia marginata (J.J.Engel et G.L.Merr.) E.D.Cooper, Phytotaxa 97 (2):
59, 2013 (Cooper et al. 2013). Bas.: Telaranea marginata J.J.Engel et G.L.Merr.,
Fieldiana, Bot. (n.ser.) 44: 166, 2004 (Engel and Smith Merrill 2004).
*** Tricholepidozia martinii (E.A.Hodgs.) E.D.Cooper, Phytotaxa 97 (2): 59, 2013
(Cooper et al. 2013). Bas.: Lepidozia martinii E.A.Hodgs., Trans. Roy. Soc. New
Zealand 83 (4): 602, 1956 (Hodgson 1956).
*** Tricholepidozia melanesica (H.A.Mill.) E.D.Cooper, Phytotaxa 97 (2): 60, 2013
(Cooper et al. 2013). Bas.: Telaranea melanesica H.A.Mill., J. Bryol. 14 (2): 237,
1986 (Miller 1986).
*** Tricholepidozia neesii (Lindenb.) E.D.Cooper, Phytotaxa 97 (2): 60, 2013 (Cooper
et al. 2013). Bas.: Lepidozia neesii Lindenb., Syn. Hepat. 2: 212, 1845 (Gottsche et
al. 1845a).
*** Tricholepidozia octoloba (Del Ros.) E.D.Cooper, Phytotaxa 97 (2): 60, 2013
(Cooper et al. 2013). Bas.: Telaranea octoloba Del Ros., Philipp. J. Sci. 100 (3/4):
239, 1971 (Del Rosario 1971).
*** Tricholepidozia plumulosa (Lehm. et Lindenb.) E.D.Cooper, Phytotaxa 97 (2): 60,
2013 (Cooper et al. 2013). Bas.: Jungermannia plumulosa Lehm. et Lindenb., Nov.
Stirp. Pug. 6: 30, 1834 (Lehmann 1834).
*** Tricholepidozia pulcherrima (Steph.) E.D.Cooper, Phytotaxa 97 (2): 60, 2013
(Cooper et al. 2013). Bas.: Lepidozia pulcherrima Steph., Sp. Hepat. (Stephani) 3:
600, 1909 (Stephani 1909a).
*** Tricholepidozia pulcherrima var. mooreana (Steph.) E.D.Cooper, Phytotaxa 97
(2): 60, 2013 (Cooper et al. 2013). Bas.: Lepidozia mooreana Steph., Sp. Hepat.
(Stephani) 3: 585, 1909 (Stephani 1909a).
World checklist of hornworts and liverworts
181
*** Tricholepidozia quadriseta (Steph.) E.D.Cooper, Phytotaxa 97 (2): 60, 2013
(Cooper et al. 2013). Bas.: Lepidozia quadriseta Steph., Sp. Hepat. (Stephani) 3:
582, 1909 (Stephani 1909a).
*** Tricholepidozia remotifolia (E.A.Hodgs.) E.D.Cooper, Phytotaxa 97 (2): 60, 2013
(Cooper et al. 2013). Bas.: Telaranea remotifolia E.A.Hodgs., Rec. Domin. Mus. 4
(11): 107, 1962 (Hodgson 1962a).
*** Tricholepidozia semperiana (Steph.) E.D.Cooper, Phytotaxa 97 (2): 60, 2013
(Cooper et al. 2013). Bas.: Lepidozia semperiana Steph., Sp. Hepat. (Stephani) 3:
612, 1909 (Stephani 1909a).
*** Tricholepidozia tetradactyla (Hook.f. et Taylor) E.D.Cooper, Phytotaxa 97 (2):
61, 2013 (Cooper et al. 2013). Bas.: Jungermannia tetradactyla Hook.f. et Taylor,
London J. Bot. 3: 286 [386], 1844 (Hooker and Taylor 1844a).
*** Tricholepidozia trichocoleoides (Herzog) E.D.Cooper, Phytotaxa 97 (2): 61, 2013
(Cooper et al. 2013). Bas.: Lepidozia trichocoleoides Herzog, Trans. Brit. Bryol.
Soc. 1 (4): 314, 1950 (Herzog 1950a).
**
Micropterygioideae Grolle
** Micropterygium Gottsche, Lindenb. et Nees, Syn. Hepat. 2: 233, 1845 (Gottsche
et al. 1845a).
** Micropterygium angustistipulum Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:
385, 1885 (Spruce 1885).
*** Micropterygium bialatum Fulford, Mem. New York Bot. Gard. 11 (2): 268, 1966
(Fulford 1966).
*** Micropterygium bolivarense Fulford, Mem. New York Bot. Gard. 11 (2): 263, 1966
(Fulford 1966).
*** Micropterygium campanense Spruce, Hedwigia 73 (3/4): 157, 1933 (Reimers 1933).
*** Micropterygium carinatum (Grev.) Reimers, Hedwigia 76 (3): 166, 1936 (Reimers
1936). Bas.: Jungermannia carinata Grev., Ann. Lyceum Nat. Hist. New York 1
(2): 276, 1825 (Greville 1825).
*** Micropterygium conchifolium Reimers, Hedwigia 73 (3/4): 155, 1933 (Reimers 1933).
*** Micropterygium duidae Reimers, Hedwigia 73 (3/4): 147, 1933 (Reimers 1933).
** Micropterygium exalatum Steph., Sp. Hepat. (Stephani) 3: 547, 1909 (Stephani
1909a).
** Micropterygium grandistipulum Steph., Trans. Linn. Soc. London, Bot. 6 (1): 98,
1901 (Stephani 1901e).
** Micropterygium laeve H.Rob., Bol. Soc. Venez. Ci. Nat. 32 (132/133): 252, 1976
(Robinson 1976b).
** Micropterygium lechleri Reimers, Hedwigia 73 (3/4): 184, 1933 (Reimers 1933).
*** Micropterygium leiophyllum Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 386,
1885 (Spruce 1885).
182
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Micropterygium longicellulatum Uribe et E.L.Linares, Phytotaxa 213 (3): 297,
2015 (Uribe and Linares 2015).
** Micropterygium parvistipulum Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:
383, 1885 (Spruce 1885).
*** Micropterygium pterygophyllum (Nees) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 413, 1877 (Trevisan 1877). Bas.: Jungermannia pterygophylla Nees, Fl. Bras. (Martius) 1 (1): 377, 1833 (Nees 1833a).
* Micropterygium pterygophyllum var. lancifolium Reimers, Hedwigia 73 (3/4): 176,
1933 (Reimers 1933).
** Micropterygium reimersianum Herzog, Hedwigia 81 (5/6): 226, 1943 (Herzog 1943b).
*** Micropterygium steyermarkii Fulford, Mem. New York Bot. Gard. 11 (2): 270,
1966 (Fulford 1966).
*** Micropterygium tatei Reimers, Hedwigia 73 (3/4): 150, 1933 (Reimers 1933).
*** Micropterygium tenax (Steph.) Grolle, J. Bryol. 10 (3): 265, 1979 (Grolle 1979b).
Bas.: Harpalejeunea tenax Steph., Trans. Linn. Soc. London, Bot. 6 (1): 100, 1901
(Stephani 1901e).
*** Micropterygium trachyphyllum Reimers, Hedwigia 73 (3/4): 186, 1933 (Reimers
1933). 128
* Micropterygium trachyphyllum var. brasiliense Reimers, Hedwigia 73 (3/4): 195,
1933 (Reimers 1933).
* Micropterygium trachyphyllum var. cubense Reimers, Hedwigia 73 (3/4): 188, 1933
(Reimers 1933).
* Micropterygium trachyphyllum var. guadeloupense Reimers, Hedwigia 73 (3/4):
190, 1933 (Reimers 1933).
* Micropterygium trachyphyllum var. jamaicense Reimers, Hedwigia 73 (3/4): 190,
1933 (Reimers 1933).
*** Micropterygium tumidulum Fulford, Mem. New York Bot. Gard. 11 (2): 272,
1966 (Fulford 1966).
** Mytilopsis Spruce, Cephalozia: 90, 1882 (Spruce 1882).
*** Mytilopsis albifrons Spruce, Cephalozia: 91, 1882 (Spruce 1882).
**
Protocephalozioideae R.M.Schust.
** Protocephalozia (Spruce) K.I.Goebel, Flora 77 (2): 83, 1893 (Goebel 1893b).
Bas.: Cephalozia subg. Protocephalozia Spruce, Cephalozia: 24, 1882 (Spruce 1882).
*** Protocephalozia ephemeroides (Spruce) K.I.Goebel, Flora 77 (2): 83, 1893 (Goebel
1893b). Bas.: Cephalozia ephemeroides Spruce, Cephalozia: 24, 1882 (Spruce 1882).
128 Micropterygium trachyphyllum was split into several varieties by Reimers (1933), but they do not
seem to have been accepted by any later author and Söderström et al. (2011a) were not sure about
their status.
World checklist of hornworts and liverworts
***
183
Zoopsidoideae R.M.Schust.
** Amazoopsis J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 242, 2004 (Engel
and Smith Merrill 2004).
*** Amazoopsis diplopoda (Pócs) J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44:
245, 2004 (Engel and Smith Merrill 2004). Bas.: Arachniopsis diplopoda Pócs,
Proc. hird Meeting Bryol. C. & E. Europe: 114, 1984 (Pócs 1984b).
*** Amazoopsis dissotricha (Spruce) J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44:
247, 2004 (Engel and Smith Merrill 2004). Bas.: Arachniopsis dissotricha Spruce,
Cephalozia: 86, 1882 (Spruce 1882).
*** Amazoopsis gracilis J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 246, 2004
(Engel and Smith Merrill 2004).
Hyalolepidozia S.W.Arnell ex Grolle, Rev. Bryol. Lichénol. 32 (3/4): 179, 1963
[1964] (Grolle 1963b). Based on: Hyalolepidozia S.W.Arnell, Bot. Not. 115: 203,
1962 (Arnell 1962a).
** Hyalolepidozia bicuspidata (C.Massal.) S.W.Arnell ex Grolle, Rev. Bryol. Lichénol.
32 (3/4): 179, 1963 [1964] (Grolle 1963b). Bas.: Lepidozia bicuspidata C.Massal.,
Nuovo Giorn. Bot. Ital. 17 (3): 239, 1885 (Massalongo 1885).
*
Monodactylopsis (R.M.Schust.) R.M.Schust., Nova Hedwigia 56 (1/2): 45,
1993 (Schuster 1993a). Bas.: Arachniopsis subg. Monodactylopsis R.M.Schust.,
Nova Hedwigia 10 (1/2): 24, 1965 (Schuster 1965b).
** Monodactylopsis monodactyla (Spruce) R.M.Schust., Nova Hedwigia 69 (3/4):
523, 1999 (Schuster 1999a). Bas.: Cephalozia monodactyla Spruce, Cephalozia:
28, 1882 (Spruce 1882).
*
** Neogrollea E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot. 3 (4): 70, 1965
(Hodgson 1965).
*** Neogrollea notabilis E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot. 3 (4): 70,
1965 (Hodgson 1965).
*
*
Odontoseries Fulford, Mem. New York Bot. Gard. 11 (3): 364, 1968 (Fulford 1968).
Odontoseries chimantana Fulford, Mem. New York Bot. Gard. 11 (3): 366, 1968
(Fulford 1968).
*** Paracromastigum Fulford et J.Taylor, Brittonia 13 (4): 336, 1961 (Fulford and
Taylor 1961).
*** Paracromastigum denticulatum (Steph.) E.D.Cooper, Phytotaxa 97 (2): 61, 2013
(Cooper et al. 2013). Bas.: Lembidium denticulatum Steph., Sp. Hepat. (Stephani)
6: 444, 1924 (Stephani 1924).
184
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Paracromastigum drucei (R.M.Schust.) R.M.Schust., J. Hattori Bot. Lab. 38: 700,
1974 (Schuster and Engel 1974). Bas.: Pseudocephalozia drucei R.M.Schust., Nova
Hedwigia 10 (1/2): 22, 1965 (Schuster 1965b).
*** Paracromastigum dusenii (Steph.) R.M.Schust., J. Hattori Bot. Lab. 38: 700, 1974
(Schuster and Engel 1974). Bas.: Alobiella dusenii Steph., Bih. Kongl. Svenska
Vetensk.-Akad. Handl. 26 (III, 6): 48, 1900 (Stephani 1900b).
** Paracromastigum iordlandiae R.M.Schust. et J.J.Engel, Brittonia 48 (2): 167,
1996 (Schuster and Engel 1996).
*** Paracromastigum furcifolium (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 276,
1963 (Schuster 1963b). Bas.: Cephalozia furcifolia Steph., Bull. Herb. Boissier
(sér. 2) 8 (7): 485 (315), 1908 (Stephani 1908f).
** Paracromastigum granatense (Gottsche) R.M.Schust., J. Hattori Bot. Lab. 48: 341,
1980 (Schuster 1980a). Bas.: Lepidozia granatensis Gottsche, Ann. Sci. Nat. Bot.
(sér. 5) 1: 139, 1864 (Gottsche 1864).
** Paracromastigum longiscyphum (Taylor) R.M.Schust. et J.J.Engel, Brittonia 48
(2): 167, 1996 (Schuster and Engel 1996). Bas.: Jungermannia longiscypha Taylor,
London J. Bot. 5: 280, 1846 (Taylor 1846a).
** Paracromastigum macrostipum (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 276,
1963 (Schuster 1963b). Bas.: Cephalozia macrostipa Steph., Hedwigia 32 (5): 315,
1893 (Stephani 1893d).
** Paracromastigum micromera (Spruce) R.M.Schust., J. Hattori Bot. Lab. 26: 276,
1963 (Schuster 1963b). Bas.: Cephalozia micromera Spruce, Cephalozia: 32, 1882
(Spruce 1882).
** Paracromastigum microphyllum (R.M.Schust. ex J.J.Engel) E.D.Cooper, Phytotaxa 97 (2): 61, 2013 (Cooper et al. 2013). Bas.: Hyalolepidozia microphylla
R.M.Schust. ex J.J.Engel, Novon 17 (3): 310, 2007 (Engel 2007).
*** Paracromastigum pachyrhizum (Nees) Fulford, Mem. New York Bot. Gard. 11 (3):
390, 1968 (Fulford 1968). Bas.: Jungermannia pachyrhiza Nees, Fl. Bras. (Martius) 1 (1): 339, 1833 (Nees 1833a).
** Paracromastigum ryszardii Váňa, Bedn.-Ochyra et Cykowska, Nova Hedwigia 89
(1/2): 122, 2009 (Váňa et al. 2009).
** Paracromastigum stipulatum (Herzog) Fulford, Mem. New York Bot. Gard. 11 (3):
390, 1968 (Fulford 1968). Bas.: Cephalozia stipulata Herzog, Memoranda Soc.
Fauna Fl. Fennica 27: 105, 1952 (Herzog 1952c).
** Paracromastigum subsimplex (Steph.) Fulford et J.Taylor, Brittonia 13 (4): 336,
1961 (Fulford and Taylor 1961). Bas.: Lepidozia subsimplex Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9): 66, 1911 (Stephani 1911b).
** Paracromastigum succulentum (Sim) J.J.Engel et G.L.Merr., Bryologist 104 (1):
151, 2001 (Engel and Smith Merrill 2001). Bas.: Lepidozia succulenta Sim, Trans.
Roy. Soc. South Africa 15 (1): 90, 1926 (Sim 1926).
** Paracromastigum tristanianum (R.M.Schust.) J.J.Engel et R.M.Schust., J. Hattori
Bot. Lab. 38: 700, 1974 (Schuster and Engel 1974). Bas.: Pseudocephalozia tristaniana R.M.Schust., Nova Hedwigia 10 (1/2): 23, 1965 (Schuster 1965b).
World checklist of hornworts and liverworts
185
** Paracromastigum vastilobum (Steph.) J.J.Engel et G.L.Merr., Fieldiana, Bot.
(n.ser.) 44: 255, 2004 (Engel and Smith Merrill 2004). Bas.: Lepidozia vastiloba
Steph., Sp. Hepat. (Stephani) 3: 581, 1909 (Stephani 1909a).
** Psiloclada Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 143, 1854 (Mitten 1854).
*** Psiloclada clandestina Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 143, 1854
(Mitten 1854).
** Psiloclada clandestina subsp. melanesica R.M.Schust., J. Hattori Bot. Lab. 48: 411,
1980 (Schuster 1980a).
** Psiloclada clandestina subsp. spinosa (S.W.Arnell) R.M.Schust., J. Hattori Bot.
Lab. 48: 410, 1980 (Schuster 1980a). Bas.: Lepidozia spinosa S.W.Arnell, Bot.
Not. 107: 427, 1954 (Arnell 1954c).
** Pteropsiella Spruce, J. Bot. 14: 161, 1876 (Spruce 1876b).
*** Pteropsiella frondiformis Spruce, J. Bot. 14: 161, 1876 (Spruce 1876b).
** Pteropsiella metzgeriiformis R.M.Schust., Nova Hedwigia 10 (1/2): 25, 1965
(Schuster 1965b).
** Telaranea Spruce ex Schifn., Hepat. (Engl.-Prantl): 103, 1893 (Schifner 1893b)
nom. conserv. Based on: Telaranea Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 365, 1885 (Spruce 1885). 129
* sect. Telaranea
*** Telaranea apiahyna (Steph.) Fulford, Brittonia 15 (1): 71, 1963 (Fulford 1963).
Bas.: Lepidozia apiahyna Steph., Sp. Hepat. (Stephani) 3: 572, 1909 (Stephani
1909a).
*** Telaranea bicruris (Steph.) M.Howe, Bull. Torrey Bot. Club 29 (5): 287, 1902
(Howe 1902). Bas.: Lepidozia bicruris Steph., Hedwigia 24 (4): 166, 1885 (Stephani 1885f).
*** Telaranea blepharostoma (Steph.) Fulford, Brittonia 15 (1): 73, 1963 (Fulford
1963). Bas.: Lepidozia blepharostoma Steph., Bih. Kongl. Svenska Vetensk.-Akad.
Handl. 26 (III, 17): 22, 1901 (Stephani 1901b).
*** Telaranea breviseta (Herzog) J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44:
131, 2004 (Engel and Smith Merrill 2004). Bas.: Lepidozia sejuncta var. breviseta
Herzog, Nat. Hist. Juan Fernandez (Botany) 2 (5): 723, 1942 (Herzog 1942a).
*** Telaranea chaetophylla (Spruce) Schifn., Hepat. (Engl.-Prantl): 103, 1893 (Schifner 1893b). Bas.: Lepidozia chaetophylla Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 365, 1885 (Spruce 1885).
*** Telaranea europaea J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 150, 2004
(Engel and Smith Merrill 2004).
129 Telaranea is a heterogeneous genus and several segregates are warranted. Additional studies are needed to clarify the relationships of the species retained here (Cooper 2013).
186
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Telaranea fragilis Mizut., J. Hattori Bot. Lab. 40: 449, 1976 (Mizutani 1976a).
*** Telaranea granulata J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 103, 2004
(Engel and Smith Merrill 2004).
*** Telaranea longifolia (M.Howe) J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44:
163, 2004 (Engel and Smith Merrill 2004). Bas.: Telaranea nematodes var. longifolia M.Howe, Bull. Torrey Bot. Club 29 (5): 286, 1902 (Howe 1902).
*** Telaranea nematodes (Gottsche ex Austin) M.Howe, Bull. Torrey Bot. Club 29
(5): 284, 1902 (Howe 1902). Bas.: Cephalozia nematodes Gottsche ex Austin, Bull.
Torrey Bot. Club 6 (52): 302, 1879 (Austin 1879).
*** Telaranea panchoi Del Ros., Philipp. J. Sci. 100 (3/4): 238, 1971 (Del Rosario 1971).
*** Telaranea pellucida J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 179, 2004
(Engel and Smith Merrill 2004).
*** Telaranea pseudozoopsis (Herzog) Fulford, Brittonia 15 (1): 71, 1963 (Fulford
1963). Bas.: Lepidozia pseudozoopsis Herzog, Nat. Hist. Juan Fernandez (Botany)
2 (5): 723, 1942 (Herzog 1942a).
*** Telaranea redacta (Steph.) J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44:
186, 2004 (Engel and Smith Merrill 2004). Bas.: Lepidozia redacta Steph., Wiss.
Ergebn. Deut. Zentr.-Afr. Exped. (1907-08), Bot. 2: 119, 1911 (Stephani 1911a).
*** Telaranea rosarioana H.A.Mill., J. Bryol. 14 (2): 240, 1986 (Miller 1986).
*** Telaranea setosa J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 192, 2004
(Engel and Smith Merrill 2004).
*** Telaranea trisetosa (Steph.) Grolle, J. Hattori Bot. Lab. 29: 280, 1966 (Grolle 1966g). Bas.: Lepidozia trisetosa Steph., Sp. Hepat. (Stephani) 3: 607, 1909
(Stephani 1909a).
*
***
**
***
***
***
***
sect. Tenuifoliae (R.M.Schust.) J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.)
44: 112, 2004 (Engel and Smith Merrill 2004). Bas.: Arachniopsis sect. Tenuifoliae
R.M.Schust., Beih. Nova Hedwigia 118: 461, 2000 (Schuster 2000a).
Telaranea anomala R.M.Schust. ex J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.)
44: 121, 2004 (Engel and Smith Merrill 2004).
Telaranea bischleriana Pócs, Acta Bot. Hung. 48 (1/2): 120, 2006 (Pócs 2006c).
Telaranea coactilis (Spruce) J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 140,
2004 (Engel and Smith Merrill 2004). Bas.: Arachniopsis coactilis Spruce, Cephalozia: 85, 1882 (Spruce 1882).
Telaranea confervoides J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 143, 2004
(Engel and Smith Merrill 2004). Based on: Arachniopsis pecten var. confervoides
R.M.Schust., Beih. Nova Hedwigia 118: 455, 2000 (Schuster 2000a), nom. inval.
Telaranea diacantha (Mont.) J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44:
145, 2004 (Engel and Smith Merrill 2004). Bas.: Jungermannia diacantha Mont.,
Ann. Sci. Nat. Bot. (sér. 4) 5: 349, 1856 (Montagne 1856c).
Telaranea herzogii (E.A.Hodgs.) E.A.Hodgs., Rec. Domin. Mus. 4 (11): 106,
1962 (Hodgson 1962a). Bas.: Lepidozia herzogii E.A.Hodgs., Trans. & Proc. Roy.
Soc. New Zealand 78 (4): 500, 1950 (Martin 1950).
World checklist of hornworts and liverworts
187
*** Telaranea inaequalis R.M.Schust. ex J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.)
44: 117, 2004 (Engel and Smith Merrill 2004).
** Telaranea maorensis Pócs, Acta Bot. Hung. 48 (1/2): 124, 2006 (Pócs 2006c).
*** Telaranea microstipulata R.M.Schust., Phytologia 39 (4): 241, 1978 (Schuster 1978a).
*** Telaranea monocera (Mitt. ex R.M.Schust. et Grolle) J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 168, 2004 (Engel and Smith Merrill 2004). Bas.: Arachniopsis monocera Mitt. ex R.M.Schust. et Grolle, Nova Hedwigia 10 (1/2): 25, 1965
(Schuster 1965b).
*** Telaranea pecten (Spruce) J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 178,
2004 (Engel and Smith Merrill 2004). Bas.: Arachniopsis pecten Spruce, Cephalozia: 85, 1882 (Spruce 1882).
*** Telaranea sejuncta (Ångstr.) S.W.Arnell, Bot. Not. 110 (1): 18, 1957 (Arnell
1957a). Bas.: Blepharostoma sejunctum Ångstr., Öfvers. Kongl. Vetensk.-Akad.
Förh. 33 (7): 78, 1876 [1877] (Ångström 1876).
*** Telaranea tenuifolia J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 193, 2004
(Engel and Smith Merrill 2004). Based on: Arachniopsis tenuifolia R.M.Schust.,
Beih. Nova Hedwigia 118: 461, 2000 (Schuster 2000a), nom. inval.
Incertae sedis
** Telaranea azorica (H.Buch et Perss.) Pócs in Schumacker et Váňa, Identif. keys
liverw. hornw. Europe: 160, 2005 (Schumacker and Váňa 2005). Bas.: Lepidozia
azorica H.Buch et Perss., Bryophyt. Azoren Madeira: 4, 1941 (Buch and Persson
1941).
** Telaranea indica (S.C.Srivast. et P.K.Verma) A.E.D.Daniels et P.Daniel, Bull.
Bot. Surv. India 49 (1/4): 231, 2007 (Daniels and Daniel 2007). Bas.: Arachniopsis indica S.C.Srivast. et P.K.Verma, Natl. Acad. Sci. Lett. 27 (7/8): 270, 2004
[2006] (Srivastava and Verma 2004).
** Telaranea major (Herzog) J.J.Engel et G.L.Merr., Fieldiana, Bot. (n.ser.) 44: 165,
2004 (Engel and Smith Merrill 2004). Bas.: Arachniopsis major Herzog, Trans.
Brit. Bryol. Soc. 1 (4): 294, 1950 (Herzog 1950a).
** Zoopsidella R.M.Schust., Nova Hedwigia 10 (1/2): 24, 1965 (Schuster 1965b).
** Zoopsidella antillana (Steph.) H.Rob., Bol. Soc. Venez. Ci. Nat. 32 (132/133):
254, 1976 (Robinson 1976b). Bas.: Zoopsis antillana Steph., Bull. Herb. Boissier
(sér. 2) 8 (4): 268 (282), 1908 (Stephani 1908j).
** Zoopsidella antillana subsp. jamaicensis R.M.Schust., Nova Hedwigia 69 (1/2):
132, 1999 (Schuster 1999b).
*** Zoopsidella caledonica (Steph.) R.M.Schust., Taxon 18 (1): 57, 1969 (Schuster 1969c). Bas.: Zoopsis caledonica Steph., Sp. Hepat. (Stephani) 6: 318, 1922
(Stephani 1922).
** Zoopsidella cynosurandra (Steph.) R.M.Schust., Nova Hedwigia 10 (1/2): 24, 1965
(Schuster 1965b). Bas.: Zoopsis cynosurandra Steph., Bull. Herb. Boissier (sér. 2) 8
(4): 269 (283), 1908 (Stephani 1908j).
188
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*** Zoopsidella integrifolia (Spruce) R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 12
(3): 666, 1969 (Schuster 1969a). Bas.: Cephalozia integrifolia Spruce, Cephalozia:
29, 1882 (Spruce 1882).
** Zoopsidella macella (Spruce) R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 12
(3): 666, 1969 (Schuster 1969a). Bas.: Cephalozia macella Spruce, Cephalozia: 29,
1882 (Spruce 1882).
*** Zoopsidella serra (Spruce) R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 12 (3):
666, 1969 (Schuster 1969a). Bas.: Cephalozia serra Spruce, Cephalozia: 32, 1882
(Spruce 1882).
** Zoopsis Hook.f. ex Gottsche, Lindenb. et Nees, Syn. Hepat. 4: 473, 1846
(Gottsche et al. 1846).
** subg. Eozoopsis R.M.Schust., J. Hattori Bot. Lab. 36: 373, 1972 (Schuster 1972).
*** Zoopsis leitgebiana (Carrington et Pearson) Bastow, Pap. & Proc. Roy. Soc. Tasmania 1887: 269, 1888 (Bastow 1888). Bas.: Cephalozia leitgebiana Carrington et
Pearson, Pap. & Proc. Roy. Soc. Tasmania 1887: 3, 1888 (Carrington and Pearson 1888b).
*** Zoopsis macrophylla R.M.Schust., Nova Hedwigia 68 (1/2): 14, 1999 (Schuster
1999d).
** subg. Zoopsis
*** Zoopsis argentea (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 4:
473, 1846 (Gottsche et al. 1846). Bas.: Jungermannia argentea Hook.f. et Taylor,
London J. Bot. 3: 400, 1844 (Hooker and Taylor 1844a).
** Zoopsis argentea var. lagelliformis (Colenso) R.M.Schust., Nova Hedwigia 68
(1/2): 38, 1999 (Schuster 1999d). Bas.: Zoopsis lagelliformis Colenso, Trans. &
Proc. New Zealand Inst. 18: 250, 1886 (Colenso 1886b).
*** Zoopsis bicruris Glenny et E.A.Br., J. Bryol. 28 (4): 332, 2006 (Renner et al. 2006).
** Zoopsis liukiuensis Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 1: 65, 1931
(Horikawa 1931a).
*** Zoopsis matawaia M.A.M.Renner, J. Bryol. 28 (4): 334, 2006 (Renner et al. 2006).
*** Zoopsis nitida Glenny, Braggins et R.M.Schust., J. Bryol. 19 (4): 776, 1997
(Glenny et al. 1997).
*** Zoopsis setulosa Leitg., Mitt. Naturwiss. Vereins Steiermark 13: 24, 1876 (Leitgeb
1876).
Incertae sedis
*** Zoopsis ceratophylla (Spruce) Hamlin, Rec. Domin. Mus. 7: 311, 1972 (Hamlin
1972). Bas.: Cephalozia ceratophylla Spruce, Cephalozia: 32, 1882 (Spruce 1882).
World checklist of hornworts and liverworts
189
*
Zoopsis ciliata Colenso, Trans. & Proc. New Zealand Inst. 20: 253, 1888 (Colenso
1888). 130
* Zoopsis martinicensis Steph., Bull. Herb. Boissier (sér. 2) 8 (4): 268 (282), 1908
(Stephani 1908j).
** Zoopsis setigera K.I.Goebel, Flora 77 (2): 93, 1893 (Goebel 1893b).
** Zoopsis uleana Steph., Hedwigia 44 (4): 225, 1905 (Stephani 1905a).
***
Lophocoleaceae Vanden Berghen
by B.J. Crandall-Stotler, R. Stotler, J. Váňa, J.J.Engel and L. Söderström
Söderström et al. (2013b) outlined the current status of Lophocoleaceae noting that
several taxonomic problems in delimitating genera remain. he placement of several
species is also unclear. Nomenclatural and taxonomic notes can be found in Söderström et al. (2013b, 2013f ).
** Bragginsella R.M.Schust., Bryologist 100 (3): 363, 1997 (Schuster 1997a).
*** Bragginsella anomala R.M.Schust., Bryologist 100 (3): 363, 1997 (Schuster
1997a).
*** Chiloscyphus Corda, Gen. hepat.: 651, 1829 (Corda 1829) nom. conserv.
** Chiloscyphus kashyapii A.Srivast. et S.C.Srivast., Indian Geocalyc.: 34, 2002 (Srivastava and Srivastava 2002).
*** Chiloscyphus pallescens (Ehrh.) Dumort., Syll. Jungerm. Europ.: 67, 1831 (Dumortier 1831). Bas.: Jungermannia pallescens Ehrh., Deutschl. Fl., heil 2 (Hofm.):
87, 1795 [1796] (Hofmann 1795).
* Chiloscyphus pallescens var. fragilis (Roth) Müll.Frib., Ber. Deutsch. Bot. Ges. 59
(10): 429, 1942 (Müller 1942). Bas.: Jungermannia fragilis Roth, Tent. Fl. Germ.
3: 370, 1800 (Roth 1800). 131
*** Chiloscyphus polyanthos (L.) Corda, Gen. hepat.: 651, 1829 (Corda 1829). Bas.:
Jungermannia polyanthos L., Sp. Pl. 1: 1131, 1753 (Linnaeus 1753).
* Chiloscyphus polyanthos var. rivularis (Schrad.) Lindb. et Arnell, Kongl. Svenska
Vetensk.-Akad. Handl. (n.ser.) 23 (5): 24, 1889 (Lindberg and Arnell 1889). Bas.:
Jungermannia pallescens var. rivularis Schrad., Syst. Samml. Crypt. Gew. 2: 7,
1797 (Schrader 1797). 132
130 Zoopsis ciliata was not studied by Engel and Glenny (2008a) and they did not know what it may be.
131 Chiloscyphus pallescens var. fragilis is a problematic taxon sometimes treated as a species, sometimes as
conspeciic with Chiloscyphus pallescens, sometimes as conspeciic with Chiloscyphus polyanthos.
132 Chiloscyphus polyanthos var. rivularis is a problematic taxon sometimes treated as a species, sometimes
as conspeciic with Chiloscyphus polyanthos.
190
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Incertae sedis 133
Chiloscyphus acutus Steph., Sp. Hepat. (Stephani) 6: 302, 1922 (Stephani 1922).
Chiloscyphus alpicola J.J.Engel, Phytotaxa 207 (2): 181, 2015 (Engel 2015b).
Chiloscyphus beesleyanus Pearson, J. Linn. Soc., Bot. 46 (305): 22, 1922 (Pearson
1922b).
Chiloscyphus biidus Schifn., Hep. Fl. Buitenzorg: 200, 1900 (Schifner 1900a). 134
Chiloscyphus breviculus B.Y.Yang et W.C.Lee, Bot. Bull. Acad. Sin. (n.ser.) 5 (2):
190, 1964 (Yang and Lee 1964).
Chiloscyphus brevistipulus Steph., Sp. Hepat. (Stephani) 6: 303, 1922 (Stephani
1922).
Chiloscyphus chinnarensis Manju, K.P.Rajesh et Madhus., Acta Bot. Hung. 53
(1/2): 152, 2011 (Manju et al. 2011).
Chiloscyphus confertifolius Steph., Sp. Hepat. (Stephani) 6: 304, 1922 (Stephani 1922).
Chiloscyphus confertus Steph., Sp. Hepat. (Stephani) 6: 305, 1922 (Stephani 1922).
Chiloscyphus cornutistipulus Steph., Sp. Hepat. (Stephani) 6: 303, 1922 (Stephani 1922).
Chiloscyphus durus (Steph.) Hässel, Revista Mus. Argent. Ci. Nat. (n.ser.) 1 (2):
122, 1999 (Hässel 1999). Bas.: Lophocolea dura Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9): 43, 1911 (Stephani 1911b). 135
Chiloscyphus ernstianus Steph., Sp. Hepat. (Stephani) 6: 306, 1922 (Stephani 1922).
Chiloscyphus etesseanus Steph., Bull. Herb. Boissier (sér. 2) 7 (10): 845 (217), 1907
(Stephani 1907b).
Chiloscyphus francanus Steph., Sp. Hepat. (Stephani) 6: 306, 1922 (Stephani 1922).
Chiloscyphus graefeanus Steph., Sp. Hepat. (Stephani) 6: 307, 1922 (Stephani 1922).
Chiloscyphus greenwelliae (H.A.Mill.) H.A.Mill., J. Hattori Bot. Lab. 30: 275,
1967 (Miller 1967). Bas.: Lophocolea greenwelliae H.A.Mill., Ark. Bot. (n.ser.) 5
(2): 504, 1963 (Miller 1963).
Chiloscyphus hookeri J.J.Engel, J. Hattori Bot. Lab. 36: 150, 1972 [1973] (Engel 1972).
Chiloscyphus hookeri var. constantifolius J.J.Engel, J. Hattori Bot. Lab. 36: 155,
1972 [1973] (Engel 1972).
Chiloscyphus integerrimus Schifn., Hep. Fl. Buitenzorg: 197, 1900 (Schifner 1900a).
Chiloscyphus kehdingianus (Steph.) N.Kitag., Hikobia, Suppl. 1: 68, 1981 (Kitagawa 1981a). Bas.: Lophocolea kehdingiana Steph., Sp. Hepat. (Stephani) 6: 278,
1922 (Stephani 1922).
Chiloscyphus kilauensis Steph., Sp. Hepat. (Stephani) 6: 309, 1922 (Stephani 1922).
Chiloscyphus koeppensis (Gottsche) Steph., Bull. Herb. Boissier (sér. 2) 8 (2): 139
(255), 1908 (Stephani 1908i). Bas.: Jungermannia koeppensis Gottsche, Int. Polarforsch., Deutsch. Exped. 2: 452, 1890 (Gottsche 1890).
133 Chiloscyphus is here treated in a very narrow sense (cf. Söderström et al. 2013b), but a large number
of species remain to be assigned to other genera. It is unclear if the taxa included here belong to
Cryptolophocolea, Lophocolea or some other genus in Lophocoleaceae.
134 Chiloscyphus biidus is possibly conspeciic with Heteroscyphus aselliformis (Schifner 1900a).
135 Chiloscyphus durus is conspeciic with Leptoscyphus expansus in Grolle (1962a), but it probably belongs to one of its segregates. It was accepted by Hässel (1999).
World checklist of hornworts and liverworts
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191
Chiloscyphus laceratus Steph., Sp. Hepat. (Stephani) 6: 310, 1922 (Stephani 1922).
Chiloscyphus lambertonii H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 506, 1963 (Miller 1963).
Chiloscyphus latistipus Steph., Sp. Hepat. (Stephani) 6: 309, 1922 (Stephani 1922).
Chiloscyphus lepervanchei (Steph.) J.J.Engel et R.M.Schust., Nova Hedwigia 39:
418, 1984 [1985] (Engel and Schuster 1984). Bas.: Lophocolea lepervanchei Steph.,
Bull. Herb. Boissier (sér. 2) 7 (4): 310 (174), 1907 (Stephani 1907d).
Chiloscyphus longiissus Steph., Sp. Hepat. (Stephani) 6: 310, 1922 (Stephani 1922).
Chiloscyphus propagulifer Schifn., Hep. Fl. Buitenzorg: 208, 1900 (Schifner 1900a).
Chiloscyphus purpureus Steph., Sp. Hepat. (Stephani) 6: 312, 1922 (Stephani 1922).
Chiloscyphus quadricilius Steph., Sp. Hepat. (Stephani) 6: 312, 1922 (Stephani 1922).
Chiloscyphus quadridentatus (Spruce) J.J.Engel et R.M.Schust., Nova Hedwigia
39: 422, 1984 [1985] (Engel and Schuster 1984). Bas.: Lophocolea quadridentata
Spruce, Mem. Torrey Bot. Club 1 (3): 137, 1890 (Spruce 1890).
Chiloscyphus rotundifolius Mitt., Rep. Challenger, Bot. 1 (3, 1): 85, 1884 (Mitten
1884b).
Chiloscyphus rotundiphyllus H.A.Mill., Phytologia 47 (4): 321, 1981 (Miller 1981).
Nom. nov. pro Chiloscyphus rotundifolius Steph., Sp. Hepat. (Stephani) 6: 313,
1922 (Stephani 1922), nom. illeg.
Chiloscyphus scaberulus Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: cc, 1889
[1890] (Spruce 1889).
Chiloscyphus septatus J.J.Engel, Fieldiana, Bot. (n.ser.) 48: 125, 2010 (Engel 2010).
Chiloscyphus similis Steph., Rev. Bryol. 35 (2): 28, 1908 (Stephani 1908l).
Chiloscyphus skottsbergianus H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 508, 1963 (Miller 1963).
Chiloscyphus subacuminatus Herzog, Ark. Bot. (n.ser.) 3 (3): 49, 1953 (Herzog 1953a).
Chiloscyphus subsimilis Steph., Sp. Hepat. (Stephani) 6: 314, 1922 (Stephani 1922).
Chiloscyphus tridens Steph., Sp. Hepat. (Stephani) 6: 315, 1922 (Stephani 1922).
Chiloscyphus trigonifolius Steph., Sp. Hepat. (Stephani) 6: 316, 1922 (Stephani 1922).
Chiloscyphus venustulus Colenso, Trans. & Proc. New Zealand Inst. 21: 60, 1889
(Colenso 1889).
** Clasmatocolea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 440, 1885
(Spruce 1885). 136
*** Clasmatocolea bisexualis Glenny et J.J.Engel, New Zealand J. Bot. 51 (1): 23, 2013
(Glenny and Engel 2013).
*** Clasmatocolea crassiretis (Herzog) Grolle, Nova Acta Leop. (n.ser.) 25 (161): 69,
1962 [1963] (Grolle 1962a). Bas.: Lophocolea crassiretis Herzog, Trans. & Proc.
Roy. Soc. New Zealand 65 (3): 354, 1936 (Herzog 1936b).
*** Clasmatocolea ctenophylla (Schifn.) Grolle, Rev. Bryol. Lichénol. 29 (1/2): 71,
1960 (Grolle 1960c). Bas.: Lophocolea ctenophylla Schifn., Leberm., Forschungsr.
Gazelle 4 (4): 12, 1890 (Schifner 1890).
136 Clasmatocolea is here treated without subdivisions although Engel (1980a, 2015) did so.
192
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Clasmatocolea cucullistipula (Steph.) Grolle, Rev. Bryol. Lichénol. 29 (1/2): 71,
1960 (Grolle 1960c). Bas.: Lophocolea cucullistipula Steph., Bih. Kongl. Svenska
Vetensk.-Akad. Handl. 26 (III, 6): 37, 1900 (Stephani 1900b).
*** Clasmatocolea fasciculata (Nees) Grolle, Rev. Bryol. Lichénol. 29 (1/2): 72, 1960
(Grolle 1960c). Bas.: Jungermannia fasciculata Nees, Horae Phys. Berol.: 46, 1820
(Nees 1820).
*** Clasmatocolea fulvella (Hook.f. et Taylor) Grolle, Rev. Bryol. Lichénol. 29 (1/2):
72, 1960 (Grolle 1960c). Bas.: Jungermannia fulvella Hook.f. et Taylor, London J.
Bot. 3: 464, 1844 (Hooker and Taylor 1844b).
*** Clasmatocolea gayana (Mont.) Grolle, Rev. Bryol. Lichénol. 29 (1/2): 72, 1960
(Grolle 1960c). Bas.: Jungermannia gayana Mont., Ann. Sci. Nat. Bot. (sér. 3) 4:
349, 1845 (Montagne 1845b).
*** Clasmatocolea humilis (Hook.f. et Taylor) Grolle, Rev. Bryol. Lichénol. 29 (1/2):
72, 1960 (Grolle 1960c). Bas.: Jungermannia humilis Hook.f. et Taylor, London
J. Bot. 3: 468, 1844 (Hooker and Taylor 1844b).
*** Clasmatocolea humilis var. polymorpha J.J.Engel, Phytologia 41 (5): 309, 1979
(Engel 1979a).
*** Clasmatocolea humilis var. suspecta (C.Massal.) J.J.Engel, Phytologia 41 (5): 309,
1979 (Engel 1979a). Bas.: Lophocolea puccioana β suspecta C.Massal., Nuovo Giorn. Bot. Ital. 17 (3): 228, 1885 (Massalongo 1885).
*** Clasmatocolea inlexispina (Hook.f. et Taylor) J.J.Engel, Bryologist 94 (4): 436,
1991 (Engel 1991b). Bas.: Jungermannia inlexispina Hook.f. et Taylor, London J.
Bot. 4: 82, 1845 (Hooker and Taylor 1845).
*** Clasmatocolea marginata (Steph.) Grolle, Nova Acta Leop. (n.ser.) 25 (161): 73,
1962 [1963] (Grolle 1962a). Bas.: Leioscyphus marginatus Steph., Bull. Herb.
Boissier (sér. 2) 6 (3): 223 (23), 1906 (Stephani 1906g).
*** Clasmatocolea minutiretis J.J.Engel et Grolle, Phytologia 41 (5): 309, 1979 (Engel 1979a).
*** Clasmatocolea moniliformis J.J.Engel, Phytologia 41 (5): 310, 1979 (Engel 1979a).
*** Clasmatocolea navistipula (Steph.) Grolle, Feddes Repert. 82 (1): 88, 1971 (Grolle
1971b). Bas.: Lophocolea navistipula Steph., Bull. Herb. Boissier (sér. 2) 6 (7): 543
(57), 1906 (Stephani 1906f).
*** Clasmatocolea navistipula var. parceramosa J.J.Engel, Phytologia 41 (5): 311, 1979
(Engel 1979a).
*** Clasmatocolea notophylla (Hook.f. et Taylor) Grolle, J. Jap. Bot. 41 (8): 228, 1966
(Grolle 1966d). Bas.: Jungermannia notophylla Hook.f. et Taylor, London J. Bot.
3: 376, 1844 (Hooker and Taylor 1844a).
*** Clasmatocolea obvoluta (Hook.f. et Taylor) Grolle, Rev. Bryol. Lichénol. 29 (1/2):
72, 1960 (Grolle 1960c). Bas.: Jungermannia obvoluta Hook.f. et Taylor, London
J. Bot. 4: 80, 1845 (Hooker and Taylor 1845).
*** Clasmatocolea obvoluta var. cookiana (C.Massal.) J.J.Engel, Phytologia 41 (5): 311,
1979 (Engel 1979a). Bas.: Lophocolea cookiana C.Massal., Nuovo Giorn. Bot. Ital.
17 (3): 224, 1885 (Massalongo 1885).
World checklist of hornworts and liverworts
193
*** Clasmatocolea puccioana (De Not.) Grolle, Rev. Bryol. Lichénol. 29 (1/2): 72,
1960 (Grolle 1960c). Bas.: Jungermannia puccioana De Not., Mem. Reale Accad.
Sci. Torino (ser. 2) 16: 221, 1857 (De Notaris 1857).
*** Clasmatocolea rigens (Hook.f. et Taylor) J.J.Engel, J. Hattori Bot. Lab. 36: 156,
1972 [1973] (Engel 1972). Bas.: Jungermannia rigens Hook.f. et Taylor, London
J. Bot. 3: 461, 1844 (Hooker and Taylor 1844b).
*** Clasmatocolea strongylophylla (Hook.f. et Taylor) Grolle, Rev. Bryol. Lichénol. 29
(1/2): 73, 1960 (Grolle 1960c). Bas.: Jungermannia strongylophylla Hook.f. et Taylor, London J. Bot. 3: 370, 1844 (Hooker and Taylor 1844a).
*** Clasmatocolea trachyopa (Hook.f. et Taylor) Grolle, Rev. Bryol. Lichénol. 29 (1/2):
73, 1960 (Grolle 1960c). Bas.: Jungermannia trachyopa Hook.f. et Taylor, London
J. Bot. 3: 471, 1844 (Hooker and Taylor 1844b).
*** Clasmatocolea vermicularis (Lehm.) Grolle, Rev. Bryol. Lichénol. 29 (1/2): 78,
1960 (Grolle 1960c). Bas.: Jungermannia vermicularis Lehm., Linnaea 4: 361,
1829 (Lehmann 1829).
*** Clasmatocolea verrucosa J.J.Engel, Bryologist 83 (2): 220, 1980 (Engel 1980b).
*** Conoscyphus Mitt., Fl. vit.: 404, 1871 [1873] (Mitten 1871).
** Conoscyphus koponenii Piippo, Mamontov et Potemkin, Acta Bryolichenol. Asiat.
5: 20, 2014 (Piippo et al. 2014).
*** Conoscyphus trapezioides (Sande Lac.) Schifn., Consp. Hepat. Arch. Ind.: 125,
1898 (Schifner 1898b). Bas.: Chiloscyphus trapezioides Sande Lac., Ned. Kruidk.
Arch. 3: 417, 1854 [1855] (Sande Lacoste 1854).
*** Cryptolophocolea L.Söderstr., Crand.-Stotl., Stotler et Váňa, Phytotaxa 97 (2):
39, 2013 (Söderström et al. 2013b). Based on: Plagiochila sect. Connatae Lindenb., Monogr. hep. gen. Plagiochilae: xxix, 1844 [1843] (Lindenberg 1844).
*** Cryptolophocolea aculeata (Mitt.) L.Söderstr., Phytotaxa 112 (1): 21, 2013 (Söderström et al. 2013f). Bas.: Chiloscyphus aculeatus Mitt., Bot. antarct. voy. II (Fl.
Nov.-Zel. 2): 140, 1854 (Mitten 1854).
** Cryptolophocolea chiloscyphoidea (Lindenb.) L.Söderstr. et Crand.-Stotl., Phytotaxa
112 (1): 18, 2013 (Söderström et al. 2013f). Bas.: Plagiochila chiloscyphoidea Lindenb., Nov. Stirp. Pug. 8: 4, 1844 (Lehmann 1844).
*** Cryptolophocolea ciliolata (Nees) L.Söderstr., Crand.-Stotl., Stotler et Váňa, Phytotaxa 97 (2): 39, 2013 (Söderström et al. 2013b). Bas.: Jungermannia ciliolata Nees,
Enum. Pl. Crypt. Javae: 68, 1830 (Nees 1830).
** Cryptolophocolea compacta (Mitt.) L.Söderstr., Phytotaxa 112 (1): 19, 2013 (Söderström et al. 2013f). Bas.: Lophocolea compacta Mitt., Trans. Linn. Soc. London,
Bot. 3 (3): 198, 1891 (Mitten 1891).
*** Cryptolophocolea connata (Sw.) L.Söderstr. et Váňa, Phytotaxa 112 (1): 19, 2013
(Söderström et al. 2013f). Bas.: Jungermannia connata Sw., Prodr. (Swartz): 143,
1788 (Swartz 1788).
194
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*** Cryptolophocolea connatifolia (J.J.Engel) L.Söderstr., Phytotaxa 112 (1): 21, 2013
(Söderström et al. 2013f). Bas.: Chiloscyphus connatifolius J.J.Engel, Phytologia 83
(1): 42, 1997 [1998] (Engel 1997).
*** Cryptolophocolea costata (Nees) L.Söderstr., Phytotaxa 112 (1): 19, 2013 (Söderström et al. 2013f). Bas.: Jungermannia costata Nees, Enum. Pl. Crypt. Javae: 69,
1830 (Nees 1830).
*** Cryptolophocolea edentata (J.J.Engel) L.Söderstr., Phytotaxa 112 (1): 21, 2013
(Söderström et al. 2013f). Bas.: Chiloscyphus edentatus J.J.Engel, Phytologia 83
(1): 43, 1997 [1998] (Engel 1997).
** Cryptolophocolea explanata (Mitt.) Váňa et Crand.-Stotl., Phytotaxa 202 (1): 69,
2015 (Söderström et al. 2015c). Bas.: Lophocolea explanata Mitt., Fl. vit.: 404,
1871 [1873] (Mitten 1871).
** Cryptolophocolea leischeri (Steph.) L.Söderstr., Phytotaxa 112 (1): 21, 2013 (Söderström et al. 2013f). Bas.: Lophocolea leischeri Steph., Bull. Herb. Boissier (sér.
2) 6 (11): 952 (132), 1906 (Stephani 1906c).
*** Cryptolophocolea guadalupensis (Steph.) L.Söderstr. et Váňa, Phytotaxa 112 (1): 19,
2013 (Söderström et al. 2013f). Bas.: Lophocolea guadalupensis Steph., Bull. Herb.
Boissier (sér. 2) 7 (1): 65 (153), 1907 (Stephani 1907c).
*** Cryptolophocolea helmsiana (Steph.) L.Söderstr., Phytotaxa 112 (1): 19, 2013
(Söderström et al. 2013f). Bas.: Lophocolea helmsiana Steph., Bull. Herb. Boissier
(sér. 2) 6 (9): 794 (94), 1906 (Stephani 1906e).
*** Cryptolophocolea leucophylla (Hook.f. et Taylor) L.Söderstr., Phytotaxa 112 (1):
19, 2013 (Söderström et al. 2013f). Bas.: Jungermannia leucophylla Hook.f. et
Taylor, London J. Bot. 3: 384, 1844 (Hooker and Taylor 1844a).
* Cryptolophocolea levieri (Schifn.) L.Söderstr., Phytotaxa 112 (1): 21, 2013 (Söderström et al. 2013f). Bas.: Lophocolea levieri Schifn., Hep. Fl. Buitenzorg: 182,
1900 (Schifner 1900a). 137
* Cryptolophocolea lilliena (Steph.) L.Söderstr., Phytotaxa 112 (1): 21, 2013 (Söderström et al. 2013f). Bas.: Lophocolea lilliena Steph., Sp. Hepat. (Stephani) 6: 282,
1922 (Stephani 1922). 138
*** Cryptolophocolea martiana (Nees) L.Söderstr., Crand.-Stotl. et Stotler, Phytotaxa
112 (1): 20, 2013 (Söderström et al. 2013f). Bas.: Lophocolea martiana Nees, Syn.
Hepat. 2: 152, 1845 (Gottsche et al. 1845a).
** Cryptolophocolea martiana subsp. bidentula (Nees) L.Söderstr., Crand.-Stotl. et
Stotler, Phytotaxa 112 (1): 20, 2013 (Söderström et al. 2013f). Bas.: Chiloscyphus
bidentulus Nees, Syn. Hepat. 2: 181, 1845 (Gottsche et al. 1845a).
** Cryptolophocolea martiana var. perissodonta (Spruce) Gradst., Phytoneuron 2015
(22): 1, 2015 (Bernal et al. 2015). Bas.: Lophocolea martiana var. perissodonta
Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 432, 1885 (Spruce 1885).
137 Cryptolophocolea levieri is possibly conspeciic with Cryptolophocolea ciliolata (Söderström et al. 2010a).
138 Cryptolophocolea lilliena is possibly conspeciic with Cryptolophocolea martiana (Wigginton and
Grolle 1996).
World checklist of hornworts and liverworts
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195
Cryptolophocolea massalongoana (Schifn.) L.Söderstr., Phytotaxa 112 (1): 20, 2013
(Söderström et al. 2013f). Bas.: Lophocolea massalongoana Schifn., Hep. Fl. Buitenzorg: 183, 1900 (Schifner 1900a). 139
Cryptolophocolea mitteniana (Colenso) L.Söderstr., Phytotaxa 112 (1): 21, 2013
(Söderström et al. 2013f). Bas.: Isotachis mitteniana Colenso, Trans. & Proc. New
Zealand Inst. 21: 69, 1889 (Colenso 1889).
Cryptolophocolea mitteniana var. obtusa (J.J.Engel) L.Söderstr., Phytotaxa 112 (1):
21, 2013 (Söderström et al. 2013f). Bas.: Chiloscyphus mittenianus var. obtusus
J.J.Engel, Phytologia 83 (1): 44, 1997 [1998] (Engel 1997).
Cryptolophocolea mitteniana var. symmetrica (J.J.Engel) L.Söderstr., Phytotaxa 112
(1): 22, 2013 (Söderström et al. 2013f). Bas.: Chiloscyphus mittenianus var. symmetricus J.J.Engel, Phytologia 83 (1): 44, 1997 [1998] (Engel 1997).
Cryptolophocolea pallida (Mitt.) L.Söderstr., Phytotaxa 112 (1): 22, 2013 (Söderström et al. 2013f). Bas.: Lophocolea pallida Mitt., Bot. antarct. voy. II (Fl. Nov.Zel. 2): 135, 1854 (Mitten 1854).
Cryptolophocolea pallidovirens (Hook.f. et Taylor) L.Söderstr., Phytotaxa 112 (1):
20, 2013 (Söderström et al. 2013f). Bas.: Jungermannia pallidovirens Hook.f. et
Taylor, London J. Bot. 3: 473, 1844 (Hooker and Taylor 1844b).
Cryptolophocolea proteus (Herzog) L.Söderstr., Phytotaxa 112 (1): 22, 2013 (Söderström et al. 2013f). Bas.: Lophocolea proteus Herzog, Feddes Repert. Spec. Nov.
Regni Veg. 57 (1/2): 164, 1955 (Herzog 1955).
Cryptolophocolea pycnophylla (Spruce) L.Söderstr., Phytotaxa 112 (1): 22, 2013
(Söderström et al. 2013f). Bas.: Lophocolea pycnophylla Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 434, 1885 (Spruce 1885).
Cryptolophocolea regularis (Steph.) L.Söderstr., Phytotaxa 112 (1): 23, 2013 (Söderström et al. 2013f). Bas.: Chiloscyphus regularis Steph., Hedwigia 32 (5): 325,
1893 (Stephani 1893d).
Cryptolophocolea spinifera (Hook.f. et Taylor) L.Söderstr., Phytotaxa 112 (1): 20,
2013 (Söderström et al. 2013f). Bas.: Jungermannia spinifera Hook.f. et Taylor,
London J. Bot. 3: 381, 1844 (Hooker and Taylor 1844a).
Cryptolophocolea stephanii (Schifn.) L.Söderstr., Phytotaxa 112 (1): 23, 2013
(Söderström et al. 2013f). Bas.: Lophocolea stephanii Schifn., Hep. Fl. Buitenzorg:
181, 1900 (Schifner 1900a). 140
Cryptolophocolea subopposita (J.J.Engel) L.Söderstr., Phytotaxa 112 (1): 23, 2013
(Söderström et al. 2013f). Bas.: Chiloscyphus suboppositus J.J.Engel, Phytologia 83
(1): 45, 1997 [1998] (Engel 1997).
Cryptolophocolea thermarum (Schifn.) L.Söderstr., Phytotaxa 112 (1): 23, 2013
(Söderström et al. 2013f). Bas.: Lophocolea thermarum Schifn., Hep. Fl. Buitenzorg: 180, 1900 (Schifner 1900a). 141
139 Cryptolophocolea massalongoana is possibly conspeciic with Cryptolophocolea costata (Piippo 1985a).
140 Cryptolophocolea stephanii is doubtfully distinct from Cryptolophocolea ciliolata (Söderström et al. 2010a).
141 Cryptolophocolea thermarum is possibly conspeciic with Cryptolophocolea ciliolata (Söderström et al.
2010a).
196
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Cryptolophocolea trialata (Gottsche) L.Söderstr., Phytotaxa 112 (1): 23, 2013 (Söderström et al. 2013f). Bas.: Lophocolea trialata Gottsche, Linnaea 28 (5): 552,
1856 [1857] (Gottsche 1856).
** Cryptolophocolea tricorata (Hässel) Crand.-Stotl. et Stotler, Phytotaxa 112 (1): 23,
2013 (Söderström et al. 2013f). Bas.: Chiloscyphus tricoratus Hässel, Nova Hedwigia 70 (3/4): 456, 2000 (Hässel 2000).
*** Cryptolophocolea tuberculata (J.J.Engel) L.Söderstr., Phytotaxa 112 (1): 23, 2013
(Söderström et al. 2013f). Bas.: Chiloscyphus tuberculatus J.J.Engel, Phytologia 83
(1): 45, 1997 [1998] (Engel 1997).
* Cryptolophocolea whittieriana (Inoue et H.A.Mill.) L.Söderstr., Phytotaxa 112
(1): 24, 2013 (Söderström et al. 2013f). Bas.: Lophocolea whittieriana Inoue et
H.A.Mill., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 8 (2): 143, 1965 (Inoue and Miller
1965). 142
*** Deceptifrons J.J.Engel et Váňa, Mem. New York Bot. Gard. 105: 54, 2013 (Váňa
and Engel 2013).
*** Deceptifrons plagiochiloides J.J.Engel et Váňa, Mem. New York Bot. Gard. 105: 54,
2013 (Váňa and Engel 2013).
*** Evansianthus R.M.Schust. et J.J.Engel, Bryologist 76 (4): 516, 1973 (Schuster
and Engel 1973).
*** Evansianthus georgiensis (Gottsche) R.M.Schust. et J.J.Engel, Bryologist 76 (4):
518, 1973 (Schuster and Engel 1973). Bas.: Lophocolea georgiensis Gottsche, Int.
Polarforsch., Deutsch. Exped. 2: 453, 1890 (Gottsche 1890).
*** Hepatostolonophora J.J.Engel et R.M.Schust., J. Hattori Bot. Lab. 46: 91, 1979
(Engel 1979b). Nom. nov. pro Stolonophora J.J.Engel et R.M.Schust., Fieldiana,
Bot. 36 (11): 111, 1975 (Engel and Schuster 1975).
*** Hepatostolonophora abnormis (Besch. et C.Massal.) J.J.Engel et R.M.Schust., J.
Hattori Bot. Lab. 46: 94, 1979 (Engel 1979b). Bas.: Leioscyphus abnormis Besch.
et C.Massal., Bull. Mens. Soc. Linn. Paris 1 (79): 629, 1886 (Bescherelle and Massalongo 1886).
** Hepatostolonophora conica (Steph.) Hässel, Bol. Soc. Argent. Bot. 22 (1/4): 123,
1983 (Hässel 1983). Bas.: Plagiochila conica Steph., Kungl. Svenska Vetensk.-Akad.
Handl. (n.ser.) 46 (9): 28, 1911 (Stephani 1911b).
*** Hepatostolonophora paucistipula (Rodway) J.J.Engel, J. Hattori Bot. Lab. 46: 103,
1979 (Engel 1979b). Bas.: Lophocolea paucistipula Rodway, Pap. & Proc. Roy.
Soc. Tasmania 1916: 46, 1917 (Rodway 1917a).
*** Hepatostolonophora rotata (Hook.f. et Taylor) J.J.Engel, J. Hattori Bot. Lab. 46:
98, 1979 (Engel 1979b). Bas.: Jungermannia rotata Hook.f. et Taylor, London J.
Bot. 3: 560, 1844 (Hooker and Taylor 1844d).
142 Cryptolophocolea whittieriana is possibly conspeciic with Cryptolophocolea helmsiana (Hodgson 1967).
World checklist of hornworts and liverworts
197
*** Hepatostolonophora rotata var. perssonii (R.M.Schust.) J.J.Engel, J. Hattori Bot.
Lab. 46: 103, 1979 (Engel 1979b). Bas.: Calyptrocolea perssonii R.M.Schust., Rev.
Bryol. Lichénol. 34 (3/4): 699, 1966 [1967] (Schuster 1966a).
*** Heteroscyphus Schifn., Österr. Bot. Z. 60 (5): 171, 1910 (Schifner 1910a) nom.
conserv.
** Heteroscyphus acutangulus (Schifn.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus acutangulus Schifn., Hep. Fl. Buitenzorg:
200, 1900 (Schifner 1900a).
** Heteroscyphus allodontus (Hook.f. et Taylor) J.J.Engel et R.M.Schust., Nova Hedwigia 39: 399, 1984 [1985] (Engel and Schuster 1984). Bas.: Jungermannia allodonta Hook.f. et Taylor, London J. Bot. 3: 382, 1844 (Hooker and Taylor 1844a).
** Heteroscyphus amboinensis (Schifn.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus endlicherianus var. amboinensis Schifn.,
Leberm., Forschungsr. Gazelle 4 (4): 15, 1890 (Schifner 1890).
** Heteroscyphus ammophilus (Colenso) R.M.Schust., Hepat. Anthocerotae N. Amer.
4: 248, 1980 (Schuster 1980c). Bas.: Chiloscyphus ammophilus Colenso, Trans. &
Proc. New Zealand Inst. 21: 59, 1889 (Colenso 1889).
** Heteroscyphus ammophilus var. obtusifolius J.J.Engel et G.L.Merr., Nova Hedwigia
99 (1/2): 158, 2014 (Engel 2014).
*** Heteroscyphus argutus (Reinw., Blume et Nees) Schifn., Österr. Bot. Z. 60 (5):
172, 1910 (Schifner 1910a). Bas.: Jungermannia arguta Reinw., Blume et Nees,
Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 206, 1824
[1825] (Reinwardt et al. 1824a).
** Heteroscyphus argutus var. brevidens (Schifn.) Herzog et Nog., J. Hattori Bot. Lab.
14: 40, 1955 (Herzog and Noguchi 1955). Bas.: Chiloscyphus argutus var. brevidens
Schifn., Hep. Fl. Buitenzorg: 195, 1900 (Schifner 1900a).
*** Heteroscyphus aselliformis (Reinw., Blume et Nees) Schifn., Österr. Bot. Z. 60
(5): 172, 1910 (Schifner 1910a). Bas.: Jungermannia aselliformis Reinw., Blume
et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 412,
1824 [1825] (Reinwardt et al. 1824b).
** Heteroscyphus assurgentifolius J.J.Engel, Nova Hedwigia 99 (1/2): 158, 2014
(Engel 2014).
** Heteroscyphus baduinus (Nees) Schifn., Österr. Bot. Z. 60 (5): 171, 1910 (Schifner 1910a). Bas.: Jungermannia baduina Nees, Enum. Pl. Crypt. Javae: 26, 1830
(Nees 1830).
** Heteroscyphus balnetii (Herzog) Grolle, Acta Bot. Fenn. 125: 68, 1984 (Grolle and Piippo 1984). Bas.: Chiloscyphus balnetii Herzog, Ann. Bryol. 5: 89, 1932 (Herzog 1932a).
*** Heteroscyphus billardierei (Schwägr.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Jungermannia billardierei Schwägr., Hist. Musc. Hepat.
Prodr.: 19, 1814 (Schwägrichen 1814).
** Heteroscyphus billardierei var. clasmatocoleoides (J.J.Engel et G.L.Merr.) J.J.Engel,
Nova Hedwigia 100 (3/4): 565, 2015 (Engel 2015a). Bas.: Heteroscyphus circum-
198
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
dentatus var. clasmatocoleoides J.J.Engel et G.L.Merr., Polish Bot. J. 58 (1): 95,
2013 (Engel 2013b).
Heteroscyphus brassii (Grolle) Grolle, Acta Bot. Fenn. 125: 68, 1984 (Grolle and
Piippo 1984). Bas.: Chiloscyphus brassii Grolle, J. Hattori Bot. Lab. 31: 2, 1968
(Grolle 1968a).
Heteroscyphus caesius (Schifn.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus caesius Schifn., Hep. Fl. Buitenzorg: 207,
1900 (Schifner 1900a). 143
Heteroscyphus caledonicus (Steph.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus caledonicus Steph., Bull. Herb. Boissier (sér.
2) 7 (10): 844 (216), 1907 (Stephani 1907b). 144
Heteroscyphus chlorophyllus (Hook.f. et Taylor) Schifn., Österr. Bot. Z. 60 (5):
172, 1910 (Schifner 1910a). Bas.: Jungermannia chlorophylla Hook.f. et Taylor,
London J. Bot. 3: 562, 1844 (Hooker and Taylor 1844d).
Heteroscyphus ciliatus (Steph.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910 (Schifner
1910a). Bas.: Chiloscyphus ciliatus Steph., Hedwigia 32 (5): 320, 1893 (Stephani
1893d).
Heteroscyphus coalitus (Hook.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910 (Schifner
1910a). Bas.: Jungermannia coalita Hook., Musci Exot. 2: tab. 123, 1820 (Hooker
1820).
Heteroscyphus coalitus var. simplicifolius J.J.Engel, Nova Hedwigia 100 (3/4): 572,
2015 (Engel 2015a).
Heteroscyphus combinatus (Nees) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Jungermannia combinata Nees, Enum. Pl. Crypt. Javae:
22, 1830 (Nees 1830).
Heteroscyphus conjugatus (Mitt.) J.J.Engel et R.M.Schust., Nova Hedwigia 39:
399, 1984 [1985] (Engel and Schuster 1984). Bas.: Chiloscyphus conjugatus Mitt.,
Bot. antarct. voy. III (Fl. Tasman. 2): 227, 1860 (Mitten 1860b).
Heteroscyphus contortuplicatus (Nees et Mont.) Grolle, J. Hattori Bot. Lab. 55:
504, 1984 (Grolle 1984a). Bas.: Jungermannia contortuplicata Nees et Mont., Ann.
Sci. Nat. Bot. (sér. 2) 5: 54, 1836 (Nees and Montagne 1836).
Heteroscyphus cuneistipulus (Steph.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus cuneistipulus Steph., Hedwigia 32 (5): 322,
1893 (Stephani 1893d).
Heteroscyphus darjeelingensis A.Srivast. et S.C.Srivast., Indian Geocalyc.: 115, 2002
(Srivastava and Srivastava 2002).
Heteroscyphus deceptifrons J.J.Engel, Nova Hedwigia 99 (1/2): 162, 2014 (Engel 2014).
Heteroscyphus dentammophilus J.J.Engel et G.L.Merr., Polish Bot. J. 58 (1): 96,
2013 (Engel 2013b).
143 Heteroscyphus caesius is possibly conspeciic with Heteroscyphus splendens (Söderström et al. 2010a).
144 Heteroscyphus caledonicus is possibly conspeciic with Heteroscyphus amboinensis Miller et al. (1983)
with a question mark.
World checklist of hornworts and liverworts
199
** Heteroscyphus denticulatus (Mitt.) Schifn., Beih. Bot. Centralbl. 29: 106, 1912
(Schifner 1912c). Bas.: Chiloscyphus denticulatus Mitt., Nat. hist. Azores: 320,
1870 (Mitten 1870).
** Heteroscyphus deplanchei (Steph.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus deplanchei Steph., Bull. Herb. Boissier (sér.
2) 7 (8): 693 (203), 1907 (Stephani 1907e).
** Heteroscyphus diestianus (Sande Lac.) Piippo, Acta Bot. Fenn. 131: 165, 1985
(Piippo 1985a). Bas.: Chiloscyphus diestianus Sande Lac., Ann. Mus. Bot. Lugduno-Batavi 1: 296, 1864 (Sande Lacoste 1864).
** Heteroscyphus divergenticiliatus (Steph.) Fulford, Mem. New York Bot. Gard.
11 (4): 501, 1976 (Fulford 1976). Bas.: Lophocolea divergenticiliata Steph., Bih.
Kongl. Svenska Vetensk.-Akad. Handl. 26 (III, 6): 38, 1900 (Stephani 1900b).
** Heteroscyphus dubius (Gottsche) Schifn., Österr. Bot. Z. 60 (5): 172, 1910 (Schifner 1910a). Bas.: Chiloscyphus dubius Gottsche, Abh. Naturwiss. Vereins Bremen
7: 346, 1882 (Gottsche 1882).
*** Heteroscyphus echinellus (Lindenb. et Gottsche) J.J.Engel et Xiao L.He, Bryologist
113 (1): 155, 2010 (Engel and He 2010). Bas.: Lophocolea echinella Lindenb. et
Gottsche, Syn. Hepat. 5: 703, 1847 (Gottsche et al. 1847).
** Heteroscyphus echinellus var. hyalinus J.J.Engel, Bryologist 113 (1): 161, 2010 (Engel and He 2010).
** Heteroscyphus elliottii (Steph.) Pagán, Bryologist 42 (1): 7, 1939 (Pagán 1939a).
Bas.: Chiloscyphus elliottii Steph., Bull. Herb. Boissier (sér. 2) 8 (1): 55 (231), 1908
(Stephani 1908k).
* Heteroscyphus elliottii var. portoricensis Fulford, Mem. New York Bot. Gard. 11 (4):
489, 1976 (Fulford 1976).
* Heteroscyphus falcifolius (Steph.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus falcifolius Steph., Bull. Herb. Boissier (sér. 2)
7 (8): 698 (208), 1907 (Stephani 1907e).
*** Heteroscyphus issistipus (Hook.f. et Taylor) Schifn., Österr. Bot. Z. 60 (5): 172,
1910 (Schifner 1910a). Bas.: Jungermannia issistipa Hook.f. et Taylor, London J.
Bot. 3: 384, 1844 (Hooker and Taylor 1844a).
** Heteroscyphus issistipus var. multispinus (E.A.Hodgs. et Allison) J.J.Engel, Nova
Hedwigia 99 (1/2): 165, 2014 (Engel 2014). Bas.: Chiloscyphus multispinus
E.A.Hodgs. et Allison, Trans. & Proc. Roy. Soc. New Zealand 73 (1): 37, 1943
(Hodgson 1943).
** Heteroscyphus issistipus var. repandus J.J.Engel, Nova Hedwigia 99 (1/2): 165,
2014 (Engel 2014).
** Heteroscyphus laccidus (Mitt.) A.Srivast. et S.C.Srivast., Indian Geocalyc.: 85,
2002 (Srivastava and Srivastava 2002). Bas.: Lophocolea laccida Mitt., J. Proc.
Linn. Soc., Bot. 5 (18): 99, 1860 [1861] (Mitten 1860c).
** Heteroscyphus leischeri (Steph.) D.G.Long et Rubas., Ceylon J. Sci., Biol. Sci. 43
(1): 26, 2014 (Long and Rubasinghe 2014). Bas.: Chiloscyphus leischeri Steph., Sp.
Hepat. (Stephani) 6: 306, 1922 (Stephani 1922).
200
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Heteroscyphus fragilicilius (Schifn.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus fragilicilius Schifn., Denkschr. Kaiserl. Akad.
Wiss., Math.-Naturwiss. Kl. 70: 210, 1900 [1901] (Schifner 1900c).
Heteroscyphus furcistipulus (E.A.Hodgs.) J.J.Engel et R.M.Schust., Nova Hedwigia
39: 400, 1984 [1985] (Engel and Schuster 1984). Bas.: Chiloscyphus furcistipulus
E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot. 3 (4): 79, 1965 (Hodgson 1965).
Heteroscyphus giganteus (Steph.) Hürl., Bauhinia 12 (1/2): 114, 1998 (Hürlimann
1998). Bas.: Chiloscyphus giganteus Steph., Sp. Hepat. (Stephani) 6: 307, 1922
(Stephani 1922).
Heteroscyphus graefei (J.B.Jack et Steph.) Grolle, Wiss. Z. Friedrich-Schiller-Univ. Jena,
Math.-Naturwiss. Reihe 29 (4): 645, 1980 (Grolle 1980d). Bas.: Lophocolea graefei
J.B.Jack et Steph., Bot. Centralbl. 60 (4): 101, 1894 (Jack and Stephani 1894).
Heteroscyphus grandilorus (Steph.) Hürl., Bauhinia 12 (1/2): 114, 1998 (Hürlimann 1998). Bas.: Chiloscyphus grandilorus Steph., Sp. Hepat. (Stephani) 6: 307,
1922 (Stephani 1922).
Heteroscyphus grandistipus (Steph.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus grandistipus Steph., Bot. Gaz. 15 (11): 283,
1890 (Stephani 1890c).
Heteroscyphus gunnianus (Mitt.) J.J.Engel et R.M.Schust., Nova Hedwigia 39:
400, 1984 [1985] (Engel and Schuster 1984). Bas.: Chiloscyphus gunnianus Mitt.,
Bot. antarct. voy. III (Fl. Tasman. 2): 228, 1860 (Mitten 1860b).
Heteroscyphus hastatus (E.A.Hodgs.) J.J.Engel et R.M.Schust., Nova Hedwigia 39:
400, 1984 [1985] (Engel and Schuster 1984). Bas.: Chiloscyphus hastatus E.A.Hodgs.,
Trans. Roy. Soc. New Zealand, Bot. 3 (11): 181, 1967 (Hodgson 1967).
Heteroscyphus hebridensis (Steph.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus hebridensis Steph., Hedwigia 32 (5): 323,
1893 (Stephani 1893d).
Heteroscyphus heterophyllus (Steph.) J.J.Engel et R.M.Schust., Nova Hedwigia 39:
400, 1984 [1985] (Engel and Schuster 1984). Bas.: Chiloscyphus heterophyllus Steph., Sp. Hepat. (Stephani) 6: 308, 1922 (Stephani 1922).
Heteroscyphus hyalinus (Steph.) A.Srivast. et S.C.Srivast., Indian Geocalyc.: 118,
2002 (Srivastava and Srivastava 2002). Bas.: Lophocolea hyalina Steph., Bull. Soc.
Roy. Bot. Belgique 38 (1): 46, 1899 (Stephani 1899h).
Heteroscyphus integrifolius (Lehm. et Lindenb.) Fulford, Mem. New York Bot.
Gard. 11 (4): 495, 1976 (Fulford 1976). Bas.: Jungermannia integrifolia Lehm. et
Lindenb., Nov. Stirp. Pug. 6: 32, 1834 (Lehmann 1834).
Heteroscyphus iwatsukii (S.Hatt.) Piippo, Willdenowia 18 (2): 522, 1989 (Piippo
1989a). Bas.: Saccogynidium iwatsukii S.Hatt., J. Jap. Bot. 39 (7): 206, 1964 (Hattori
1964).
Heteroscyphus jackii (Steph.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910 (Schifner
1910a). Bas.: Chiloscyphus jackii Steph., Bot. Centralbl. 60 (4): 102, 1894 (Jack
and Stephani 1894).
World checklist of hornworts and liverworts
201
*** Heteroscyphus knightii (Steph.) Grolle, J. Hattori Bot. Lab. 61: 251, 1986 [1987]
(Grolle 1986a). Bas.: Chiloscyphus knightii Steph., Bull. Herb. Boissier (sér. 2) 8
(2): 129 (245), 1908 (Stephani 1908i).
** Heteroscyphus levieri (Steph.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910 (Schifner
1910a). Bas.: Chiloscyphus levieri Steph., Bull. Herb. Boissier (sér. 2) 8 (2): 132
(248), 1908 (Stephani 1908i).
** Heteroscyphus limosus (Carrington et Pearson) Schifn., Österr. Bot. Z. 60 (5): 172,
1910 (Schifner 1910a). Bas.: Chiloscyphus limosus Carrington et Pearson, Pap. &
Proc. Roy. Soc. Tasmania 1887: 6, 1888 (Carrington and Pearson 1888b).
** Heteroscyphus lingulatus (Colenso) J.J.Engel et R.M.Schust., Nova Hedwigia 39:
400, 1984 [1985] (Engel and Schuster 1984). Bas.: Chiloscyphus lingulatus Colenso, Trans. & Proc. New Zealand Inst. 21: 61, 1889 (Colenso 1889).
** Heteroscyphus lophocoleoides S.Hatt., Bull. Tokyo Sci. Mus. 11: 45, 1944 (Hattori
1944d).
*** Heteroscyphus lyallii (Mitt.) R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 248,
1980 (Schuster 1980c). Bas.: Chiloscyphus lyallii Mitt., Bot. antarct. voy. II (Fl.
Nov.-Zel. 2): 140, 1854 (Mitten 1854).
*** Heteroscyphus magellanicus (Steph.) J.J.Engel et R.M.Schust., Nova Hedwigia
39: 400, 1984 [1985] (Engel and Schuster 1984). Bas.: Chiloscyphus magellanicus
Steph., Bull. Herb. Boissier (sér. 2) 8 (2): 140 (256), 1908 (Stephani 1908i).
** Heteroscyphus mamillatus Piippo, Nova Hedwigia 56 (3/4): 360, 1993 (Piippo
1993a).
** Heteroscyphus marginatus (Steph.) Fulford, Mem. New York Bot. Gard. 11 (4):
491, 1976 (Fulford 1976). Bas.: Lophocolea marginata Steph., Sp. Hepat. (Stephani) 6: 282, 1922 (Stephani 1922).
*** Heteroscyphus menziesii (Mitt.) J.J.Engel, Polish Bot. J. 58 (1): 99, 2013 (Engel
2013b). Bas.: Chiloscyphus menziesii Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2):
139, 1854 (Mitten 1854).
** Heteroscyphus merapiensis (Steph.) Piippo, Ann. Bot. Fenn. 30 (3): 200, 1993
(Piippo 1993c). Bas.: Chiloscyphus merapiensis Steph., Sp. Hepat. (Stephani) 6:
311, 1922 (Stephani 1922).
** Heteroscyphus miradorensis (Steph.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus miradorensis Steph., Bull. Herb. Boissier (sér.
2) 8 (1): 56 (232), 1908 (Stephani 1908k).
*** Heteroscyphus mononuculus J.J.Engel, J. Hattori Bot. Lab. 90: 241, 2001 (Engel 2001).
** Heteroscyphus mononuculus var. ammophilopsis J.J.Engel, Nova Hedwigia 99 (1/2):
166, 2014 (Engel 2014).
** Heteroscyphus mononuculus var. bilobus J.J.Engel, Nova Hedwigia 99 (1/2): 167,
2014 (Engel 2014).
** Heteroscyphus montagnei (Steph.) Fulford, Mem. New York Bot. Gard. 11 (4):
497, 1976 (Fulford 1976). Bas.: Chiloscyphus montagnei Steph., Bull. Herb. Boissier (sér. 2) 8 (2): 141 (257), 1908 (Stephani 1908i).
202
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Heteroscyphus multiidus (Steph.) J.J.Engel et R.M.Schust., Nova Hedwigia 39:
400, 1984 [1985] (Engel and Schuster 1984). Bas.: Chiloscyphus multiidus Steph.,
Bull. Herb. Boissier (sér. 2) 8 (2): 132 (248), 1908 (Stephani 1908i).
** Heteroscyphus multiidus var. subintegerrimus J.J.Engel, Nova Hedwigia 100 (3/4):
562, 2015 (Engel 2015a).
** Heteroscyphus nadeaudii (Steph.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus nadeaudii Steph., Bull. Herb. Boissier (sér. 2)
7 (10): 848 (220), 1907 (Stephani 1907b).
** Heteroscyphus orbiculatus A.Srivast. et S.C.Srivast., Indian Geocalyc.: 140, 2002
(Srivastava and Srivastava 2002).
** Heteroscyphus palniensis A.Srivast. et S.C.Srivast., Indian Geocalyc.: 130, 2002
(Srivastava and Srivastava 2002).
** Heteroscyphus pandei S.C.Srivast. et A.Srivast., Lindbergia 15 (6): 197, 1989
[1991] (Srivastava and Srivastava 1989b).
*** Heteroscyphus parallelifolius J.J.Engel, Polish Bot. J. 58 (1): 99, 2013 (Engel 2013b).
** Heteroscyphus parvulus (Schifn.) Schifn., Österr. Bot. Z. 60 (5): 171, 1910
(Schifner 1910a). Bas.: Chiloscyphus parvulus Schifn., Hep. Fl. Buitenzorg: 206,
1900 (Schifner 1900a).
** Heteroscyphus parvus A.Srivast. et S.C.Srivast., Indian Geocalyc.: 112, 2002 (Srivastava and Srivastava 2002).
** Heteroscyphus perfoliatus (Mont.) Schifn., Österr. Bot. Z. 60 (5): 171, 1910
(Schifner 1910a). Bas.: Lophocolea perfoliata Mont., Ann. Sci. Nat. Bot. (sér. 2)
18: 12, 1842 (Montagne 1842b).
** Heteroscyphus pertusus (Lehm.) Fulford, Mem. New York Bot. Gard. 11 (4): 505,
1976 (Fulford 1976). Bas.: Chiloscyphus pertusus Lehm., Nov. Stirp. Pug. 10: 7,
1857 (Lehmann 1857).
*** Heteroscyphus planiusculus (Hook.f. et Taylor) J.J.Engel, J. Hattori Bot. Lab. 68:
315, 1990 (Engel 1990a). Bas.: Jungermannia planiuscula Hook.f. et Taylor, London J. Bot. 3: 382, 1844 (Hooker and Taylor 1844a).
** Heteroscyphus planus (Mitt.) Schifn., Österr. Bot. Z. 60 (5): 171, 1910 (Schifner 1910a). Bas.: Chiloscyphus planus Mitt., J. Linn. Soc., Bot. 8 (31): 157, 1864
[1865] (Mitten 1864a).
*** Heteroscyphus polyblepharis (Spruce) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus polyblepharis Spruce, Trans. & Proc. Bot.
Soc. Edinburgh 15: 442, 1885 (Spruce 1885).
*** Heteroscyphus polychaetus (Spruce) Hentschel et Heinrichs, Taxon 56 (4): 1139,
2007 (Hentschel et al. 2007c). Bas.: Lophocolea polychaeta Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 436, 1885 (Spruce 1885).
** Heteroscyphus polycladus (Hook.f. et Lév.) R.M.Schust., Hepat. Anthocerotae N.
Amer. 4: 248, 1980 (Schuster 1980c). Bas.: Jungermannia polyclada Hook.f. et
Lév., Choix Pl. Nouv.-Zel.: 8, 1846 (Raoul 1846).
World checklist of hornworts and liverworts
203
** Heteroscyphus rectangulatus (Herzog) Piippo, Ann. Bot. Fenn. 30 (3): 200, 1993
(Piippo 1993c). Bas.: Chiloscyphus rectangulatus Herzog, Ann. Naturhist. Mus.
Wien 53 (1): 364, 1942 [1943] (Herzog 1942b).
** Heteroscyphus saccogynoides Herzog, J. Hattori Bot. Lab. 14: 40, 1955 (Herzog and
Noguchi 1955).
** Heteroscyphus sarawaketanus Piippo, Acta Bot. Fenn. 131: 143, 1985 (Piippo 1985a).
*** Heteroscyphus sinuosus (Hook.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Jungermannia sinuosa Hook., Musci Exot. 2: tab. 113,
1820 (Hooker 1820).
** Heteroscyphus spectabilis (Steph.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus spectabilis Steph., Hedwigia 30 (5): 205,
1891 (Stephani 1891a).
** Heteroscyphus spinifer C.Gao, T.Cao et Y.H.Wu, J. Bryol. 26 (2): 97, 2004 (Gao
et al. 2004).
*** Heteroscyphus splendens (Lehm. et Lindenb.) Grolle, Acta Bot. Fenn. 125: 68, 1984
(Grolle and Piippo 1984). Bas.: Jungermannia splendens Lehm. et Lindenb., Nov.
Stirp. Pug. 4: 22, 1832 (Lehmann 1832).
** Heteroscyphus splendidus (E.A.Hodgs.) J.J.Engel et R.M.Schust., Nova Hedwigia 39:
400, 1984 [1985] (Engel and Schuster 1984). Bas.: Chiloscyphus splendidus E.A.Hodgs.,
Trans. Roy. Soc. New Zealand, Bot. 3 (11): 180, 1967 (Hodgson 1967).
*** Heteroscyphus stolonifer J.J.Engel, Polish Bot. J. 58 (1): 102, 2013 (Engel 2013b).
*** Heteroscyphus succulentus (Gottsche) Schifn., Österr. Bot. Z. 60 (5): 171, 1910
(Schifner 1910a). Bas.: Chiloscyphus succulentus Gottsche, Natuurk. Tijdschr.
Ned.-Indië 4: 576, 1853 (Gottsche 1853).
** Heteroscyphus supinus (Hook.f. et Taylor) R.M.Schust., Hepat. Anthocerotae N.
Amer. 4: 248, 1980 (Schuster 1980c). Bas.: Chiloscyphus supinus Hook.f. et Taylor, London J. Bot. 5: 284, 1846 (Taylor 1846a).
** Heteroscyphus tener (Steph.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910 (Schifner
1910a). Bas.: Chiloscyphus tener Steph., Bull. Herb. Boissier (sér. 2) 7 (8): 695
(205), 1907 (Stephani 1907e).
*** Heteroscyphus thraustus (Spruce) Fulford, Mem. New York Bot. Gard. 11 (4): 495,
1976 (Fulford 1976). Bas.: Lophocolea thrausta Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 437, 1885 (Spruce 1885).
** Heteroscyphus timppae Piippo, Ann. Bot. Fenn. 29 (3): 246, 1992 (Piippo 1992).
*** Heteroscyphus triacanthus (Hook.f. et Lév.) Schifn., Österr. Bot. Z. 60 (5): 172,
1910 (Schifner 1910a). Bas.: Jungermannia triacantha Hook.f. et Lév., Choix Pl.
Nouv.-Zel.: 8, 1846 (Raoul 1846).
** Heteroscyphus triacanthus var. magnistipulatus J.J.Engel, Nova Hedwigia 99 (1/2):
167, 2014 (Engel 2014).
** Heteroscyphus tridentatus (Sande Lac.) Grolle, Acta Bot. Fenn. 125: 68, 1984
(Grolle and Piippo 1984). Bas.: Lophocolea tridentata Sande Lac., Ann. Mus. Bot.
Lugduno-Batavi 1: 296, 1864 (Sande Lacoste 1864).
204
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Heteroscyphus turgidus (Schifn.) Schifn., Österr. Bot. Z. 60 (5): 171, 1910
(Schifner 1910a). Bas.: Chiloscyphus turgidus Schifn., Denkschr. Kaiserl. Akad.
Wiss., Math.-Naturwiss. Kl. 70: 212, 1900 [1901] (Schifner 1900c).
*** Heteroscyphus valdiviensis (Mont.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus valdiviensis Mont., Ann. Sci. Nat. Bot. (sér.
3) 4: 351, 1845 (Montagne 1845b).
** Heteroscyphus varians (Steph.) J.J.Engel, J. Hattori Bot. Lab. 68: 315, 1990 (Engel
1990a). Bas.: Lophocolea varians Steph., J. & Proc. Roy. Soc. New South Wales 48
(1/2): 119, 1914 (Stephani and Watts 1914).
** Heteroscyphus wettsteinii (Schifn.) Schifn., Österr. Bot. Z. 60 (5): 172, 1910
(Schifner 1910a). Bas.: Chiloscyphus wettsteinii Schifn., Hep. Fl. Buitenzorg: 202,
1900 (Schifner 1900a).
** Heteroscyphus zollingeri (Gottsche) Schifn., Österr. Bot. Z. 60 (5): 171, 1910
(Schifner 1910a). Bas.: Chiloscyphus zollingeri Gottsche, Natuurk. Tijdschr.
Ned.-Indië 4: 576, 1853 (Gottsche 1853).
*** Lamellocolea J.J.Engel, J. Hattori Bot. Lab. 70: 65, 1991 (Engel 1991c).
*** Lamellocolea granditexta (Steph.) J.J.Engel, J. Hattori Bot. Lab. 70: 66, 1991 (Engel 1991c). Bas.: Lophocolea granditexta Steph., Bull. Herb. Boissier (sér. 2) 6 (10):
881 (106), 1906 (Stephani 1906d).
*** Lamellocolea integrostia J.J.Engel et Glenny, Bryologist 114 (1): 23, 2011 (Engel
and Glenny 2011).
*** Leptophyllopsis R.M.Schust., J. Hattori Bot. Lab. 26: 269, 1963 (Schuster
1963b).
*** Leptophyllopsis laxa (Mitt.) R.M.Schust. ex Hamlin, Rec. Domin. Mus. 7: 284,
1972 (Hamlin 1972). Bas.: Chiloscyphus laxus Mitt., Bot. antarct. voy. II (Fl.
Nov.-Zel. 2): 142, 1854 (Mitten 1854).
*** Leptoscyphopsis R.M.Schust., Phytologia 39 (4): 246, 1978 (Schuster 1978a).
*** Leptoscyphopsis paradoxa R.M.Schust., Phytologia 39 (4): 246, 1978 (Schuster
1978a).
*** Leptoscyphus Mitt., Hooker’s J. Bot. Kew Gard. Misc. 3: 358, 1851 (Mitten 1851). 145
** subg. Anomylia (R.M.Schust.) R.M.Schust., Hepat. Anthocerotae N. Amer. 4:
272, 1980 (Schuster 1980c). Bas.: Anomylia R.M.Schust., Amer. Midl. Naturalist
62 (1): 51, 1959 (Schuster 1959a).
*** Leptoscyphus cuneifolius (Hook.) Mitt., Hooker’s J. Bot. Kew Gard. Misc. 3: 358,
1851 (Mitten 1851). Bas.: Jungermannia cuneifolia Hook., Brit. Jungermann.: tab.
64, 1814 (Hooker 1814).
145 he subdivision of Leptoscyphus follows Vanderpoorten et al. (2010).
World checklist of hornworts and liverworts
205
** Leptoscyphus cuneifolius subsp. fragilis (J.B.Jack et Steph.) Grolle, Nova Acta Leop.
(n.ser.) 25 (161): 28, 1962 [1963] (Grolle 1962a). Bas.: Leioscyphus fragilis J.B.Jack
et Steph., Hedwigia 31 (1): 20, 1892 (Jack and Stephani 1892).
** subg. Austroleptoscyphus Vanderp., Schäf.-Verw. et D.G.Long, Taxon 59 (1):
183, 2010 (Vanderpoorten et al. 2010).
*** Leptoscyphus antarcticus (C.Massal.) Solari, Cryptog. Bryol. Lichénol. 7 (3): 219,
1986 (Solari 1986). Bas.: Leioscyphus antarcticus C.Massal., Atti Reale Ist. Veneto
Sci. Lett. Arti 87 (2): 229, 1928 (Massalongo 1928).
*** Leptoscyphus australis (Gottsche, Lindenb. et Nees) R.M.Schust., J. Hattori Bot.
Lab. 26: 270, 1963 (Schuster 1963b). Bas.: Chiloscyphus australis Gottsche, Lindenb. et Nees, Syn. Hepat. 2: 189, 1845 (Gottsche et al. 1845a).
*** Leptoscyphus belmoranus (Steph.) J.J.Engel, J. Hattori Bot. Lab. 74: 33, 1993 (Engel 1993). Bas.: Lophocolea belmorana Steph., J. & Proc. Roy. Soc. New South
Wales 48 (1/2): 117, 1914 (Stephani and Watts 1914).
*** Leptoscyphus excipulatus (Steph.) J.J.Engel, J. Hattori Bot. Lab. 74: 33, 1993 (Engel
1993). Bas.: Lophocolea excipulata Steph., Bull. Herb. Boissier (sér. 2) 6 (9): 790
(90), 1906 (Stephani 1906e).
*** Leptoscyphus innovatus (E.A.Hodgs.) J.J.Engel, J. Hattori Bot. Lab. 74: 33, 1993
(Engel 1993). Bas.: Lophocolea innovata E.A.Hodgs., Trans. Roy. Soc. New Zealand 80 (3/4): 347, 1952 [1953] (Hodgson 1952).
*** Leptoscyphus longistipulus (Steph.) J.J.Engel, Bryologist 94 (4): 436, 1991 (Engel
1991b). Bas.: Lophocolea longistipula Steph., Bull. Herb. Boissier (sér. 2) 6 (10):
884 (109), 1906 (Stephani 1906d).
** subg. Leptoscyphus
** sect. Hexagonistipa Grolle, Nova Acta Leop. (n.ser.) 25 (161): 46, 1962 (Grolle
1962a).
*** Leptoscyphus gibbosus (Taylor) Mitt., Hooker’s J. Bot. Kew Gard. Misc. 3: 358,
1851 (Mitten 1851). Bas.: Chiloscyphus gibbosus Taylor, London J. Bot. 5: 283,
1846 (Taylor 1846a).
*** Leptoscyphus gradsteinii Vanderp., Schäf.-Verw. et D.G.Long, Taxon 59 (1): 182,
2010 (Vanderpoorten et al. 2010).
*** Leptoscyphus hexagonus (Nees) Grolle, Nova Acta Leop. (n.ser.) 25 (161): 47, 1962
[1963] (Grolle 1962a). Bas.: Chiloscyphus hexagonus Nees, Syn. Hepat. 2: 177,
1845 (Gottsche et al. 1845a).
*** Leptoscyphus jackii (Steph.) Grolle, Nova Acta Leop. (n.ser.) 25 (161): 48, 1962
[1963] (Grolle 1962a). Bas.: Leioscyphus jackii Steph., Hedwigia 31 (1): 21, 1892
(Jack and Stephani 1892).
*** Leptoscyphus physocalyx (Hampe et Gottsche) Gottsche, Bot. Zeitung (Berlin) Beil.
16: 33, 1858 (Gottsche 1858). Bas.: Jungermannia physocalyx Hampe et Gottsche,
Linnaea 20 (3): 326, 1847 (Hampe 1847).
206
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Leptoscyphus sotiauxii Vanderp., Schäf.-Verw. et D.G.Long, Taxon 59 (1): 183,
2010 (Vanderpoorten et al. 2010).
** sect. Leptoscyphus
*** Leptoscyphus aequatus (Hook.f. et Taylor) Mitt., Hooker’s J. Bot. Kew Gard. Misc.
3: 358, 1851 (Mitten 1851). Bas.: Jungermannia aequata Hook.f. et Taylor, London J. Bot. 3: 465, 1844 (Hooker and Taylor 1844b).
*** Leptoscyphus intermedius Grolle, Nova Acta Leop. (n.ser.) 25 (161): 32, 1962
[1963] (Grolle 1962a).
*** Leptoscyphus lambinonii Vanderp., Schäf.-Verw. et D.G.Long, Taxon 59 (1): 179,
2010 (Vanderpoorten et al. 2010).
*** Leptoscyphus obcordatus (Spruce) Grolle, Nova Acta Leop. (n.ser.) 25 (161): 33,
1962 [1963] (Grolle 1962a). Bas.: Leioscyphus obcordatus Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 446, 1885 (Spruce 1885).
*** Leptoscyphus ovatus (Spruce) Grolle, Nova Acta Leop. (n.ser.) 25 (161): 45, 1962
[1963] (Grolle 1962a). Bas.: Leioscyphus ovatus Spruce, J. Linn. Soc., Bot. 30
(210): 357, 1895 (Gepp 1895b).
*** Leptoscyphus porphyrius (Nees) Grolle, Österr. Bot. Z. 117 (1): 3, 1969 (Grolle
1969a). Bas.: Chiloscyphus porphyrius Nees, Syn. Hepat. 2: 185, 1845 (Gottsche et
al. 1845a).
*** Leptoscyphus porphyrius subsp. azoricus (H.Buch et Perss.) Vanderp. et Heinrichs,
Taxon 59 (1): 181, 2010 (Vanderpoorten et al. 2010). Bas.: Mylia azorica H.Buch
et Perss., Bryophyt. Azoren Madeira: 7, 1941 (Buch and Persson 1941).
** subg. Physoscyphus Grolle, Nova Acta Leop. (n.ser.) 25 (161): 51, 1962 (Grolle 1962a).
** sect. Homaloscyphus Grolle, Nova Acta Leop. (n.ser.) 25 (161): 58, 1962 (Grolle
1962a).
*** Leptoscyphus chilensis (De Not.) Hässel, J. Hattori Bot. Lab. 91: 207, 2001 (Hässel
2001). Bas.: Lophocolea chilensis De Not., Mem. Reale Accad. Sci. Torino (ser. 2)
16: 222, 1857 (De Notaris 1857).
* Leptoscyphus diicilis (Steph.) Fulford, Mem. New York Bot. Gard. 11 (4): 534,
1976 (Fulford 1976). Bas.: Chiloscyphus diicilis Steph., Biblioth. Bot. 87 (2): 222,
1916 (Stephani 1916a). 146
*** Leptoscyphus diversifolius (Gottsche) Grolle, Nova Acta Leop. (n.ser.) 25 (161): 58,
1962 [1963] (Grolle 1962a). Bas.: Lophocolea diversifolia Gottsche, Syn. Hepat. 2:
166, 1845 (Gottsche et al. 1845a).
*** Leptoscyphus expansus (Lehm.) Grolle, Nova Acta Leop. (n.ser.) 25 (161): 60, 1962
[1963] (Grolle 1962a). Bas.: Jungermannia expansa Lehm., Linnaea 4: 361, 1829
(Lehmann 1829).
146 Leptoscyphus diicilis is conspeciic with Leptoscyphus expansus in Grolle (1962a), but it was accepted
by Vanderpoorten et al. (2010).
World checklist of hornworts and liverworts
207
*** Leptoscyphus hedbergii (S.W.Arnell) R.M.Schust., Amer. Midl. Naturalist 62 (1):
13, 1959 (Schuster 1959a). Bas.: Mylia hedbergii S.W.Arnell, Ark. Bot. (n.ser.) 3
(16): 547, 1956 (Arnell 1956e).
* Leptoscyphus huidobroanus (Mont.) Gottsche, Bot. Zeitung (Berlin) Beil. 16: 33,
1858 (Gottsche 1858). Bas.: Chiloscyphus huidobroanus Mont., Ann. Sci. Nat. Bot.
(sér. 3) 4: 352, 1845 (Montagne 1845b). 147
*** Leptoscyphus huonicus Piippo, Acta Bot. Fenn. 131: 152, 1985 (Piippo 1985a).
*** Leptoscyphus magellanicus (Gola) Hässel, J. Hattori Bot. Lab. 91: 214, 2001 (Hässel 2001). Bas.: Lophozia magellanica Gola, Nuovo Giorn. Bot. Ital. (n.ser.) 29
(1/4): 165, 1922 [1923] (Gola 1922).
** sect. Physoscyphus Grolle, Nova Acta Leop. (n.ser.) 25 (161): 51, 1962 (Grolle 1962a).
*** Leptoscyphus amphibolius (Nees) Grolle, Nova Acta Leop. (n.ser.) 25 (161): 54,
1962 [1963] (Grolle 1962a). Bas.: Jungermannia amphibolia Nees, Fl. Bras. (Martius) 1 (1): 334, 1833 (Nees 1833a).
*** Leptoscyphus infuscatus (Mitt.) E.W.Jones ex Grolle, Nova Acta Leop. (n.ser.) 25
(161): 52, 1962 [1963] (Grolle 1962a). Bas.: Leioscyphus infuscatus Mitt., J. Linn.
Soc., Bot. 22 (146): 321, 1886 (Mitten 1886b).
** subg. Spinoscyphus Vanderp., Schäf.-Verw. et D.G.Long, Taxon 59 (1): 183,
2010 (Vanderpoorten et al. 2010).
*** Leptoscyphus cleeii Fulford, Mem. New York Bot. Gard. 11 (4): 534, 1976 (Fulford 1976).
*** Leptoscyphus spectabilis (Steph.) Grolle, J. Bryol. 11 (2): 328, 1980 [1981] (Grolle
1980c). Bas.: Lophocolea spectabilis Steph., Bull. Herb. Boissier (sér. 2) 7 (4): 302
(166), 1907 (Stephani 1907d).
Incertae sedis
*** Leptoscyphus autoicus (J.J.Engel et Gradst.) Vanderp. et Gradst., J. Bryol. 34 (4):
252, 2012 (Vanderpoorten et al. 2012). Bas.: Physotheca autoica J.J.Engel et
Gradst., Taxon 52 (4): 764, 2003 (Engel and Gradstein 2003).
** Leptoscyphus beckettianus (Steph.) R.M.Schust. ex J.J.Engel, Nova Hedwigia 93
(3/4): 402, 2011 (Engel 2011). Bas.: Chiloscyphus beckettianus Steph., Bull. Herb.
Boissier (sér. 2) 8 (1): 59 (235), 1908 (Stephani 1908k).
** Leptoscyphus compactus (Colenso) J.J.Engel, Nova Hedwigia 100 (3/4): 579, 2015
(Engel 2015a). Bas.: Chiloscyphus compactus Colenso, Trans. & Proc. New Zealand Inst. 21: 63, 1889 (Colenso 1889).
147 Leptoscyphus huidobroanus is conspeciic with Leptoscyphus expansus in Váňa and Engel (2013) rejecting the conclusion by Hässel (2001) that it is an independent species, but it was accepted by Vanderpoorten et al. (2010).
208
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Leptoscyphus erraticus (W.Martin et E.A.Hodgs.) J.J.Engel, Nova Hedwigia 99 (1/2):
168, 2014 (Engel 2014). Bas.: Chiloscyphus erraticus W.Martin et E.A.Hodgs.,
Trans. & Proc. Roy. Soc. New Zealand 78 (4): 497, 1950 (Martin 1950).
*** Leptoscyphus horizontalis (Hook.) Kühnem., Revista Centro Estud. Doct. Ci. Nat.
1: 176, 1937 (Kühnemann 1937). Bas.: Jungermannia horizontalis Hook., Musci
Exot. 1: tab. 96, 1818 (Hooker 1818).
** Leptoscyphus normalis (Steph.) J.J.Engel, Nova Hedwigia 100 (3/4): 579, 2015
(Engel 2015a). Bas.: Lophocolea normalis Steph., Sp. Hepat. (Stephani) 6: 285,
1922 (Stephani 1922).
** Leptoscyphus physanthus (Hook.f. et Taylor) J.J.Engel, Nova Hedwigia 99 (1/2):
168, 2014 (Engel 2014). Bas.: Jungermannia physantha Hook.f. et Taylor, London
J. Bot. 3: 561, 1844 (Hooker and Taylor 1844d).
** Leptoscyphus subemarginatus (Hook.f. et Taylor) J.J.Engel, Bryologist 94 (4): 436,
1991 (Engel 1991b). Bas.: Lophocolea subemarginata Hook.f. et Taylor, London J.
Bot. 5: 367, 1846 (Taylor 1846b).
*** Leptoscyphus trapezoïdes (Mont.) L.Söderstr., Phytotaxa 112 (1): 27, 2013 (Söderström et al. 2013f). Bas.: Lophocolea trapezoïdes Mont., Ann. Sci. Nat. Bot. (sér. 2)
19: 251, 1843 (Montagne 1843).
*** Lophocolea (Dumort.) Dumort., Recueil Observ. Jungerm.: 17, 1835 (Dumortier 1835). Bas.: Jungermannia sect. Lophocolea Dumort., Syll. Jungerm. Europ.:
59, 1831 (Dumortier 1831).
*** Lophocolea aberrans Lindenb. et Gottsche, Syn. Hepat. 5: 696, 1847 (Gottsche et
al. 1847).
*** Lophocolea aequifolia Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 5: 55, 1836 (Nees
and Montagne 1836).
* Lophocolea angustistipula Steph., Sp. Hepat. (Stephani) 6: 260, 1922 (Stephani 1922).
*** Lophocolea anisoloba (J.J.Engel et Glenny) L.Söderstr., Phytotaxa 112 (1): 25,
2013 (Söderström et al. 2013f). Bas.: Chiloscyphus anisolobus J.J.Engel et Glenny,
Bryologist 111 (1): 118, 2008 (Engel and Glenny 2008b).
** Lophocolea anomala Steph., Sp. Hepat. (Stephani) 6: 300, 1922 (Stephani 1922).
** Lophocolea anomoda (Mont.) Steph., Hedwigia 32 (5): 327, 1893 (Stephani
1893d). Bas.: Chiloscyphus anomodus Mont., Ann. Sci. Nat. Bot. (sér. 3) 4: 352,
1845 (Montagne 1845b).
** Lophocolea apalachicola R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 195, 1980
(Schuster 1980c).
*** Lophocolea aperticaulis (J.J.Engel) L.Söderstr., Phytotaxa 112 (1): 25, 2013 (Söderström et al. 2013f). Bas.: Chiloscyphus aperticaulis J.J.Engel, J. Hattori Bot. Lab.
95: 229, 2004 (Engel 2004b).
** Lophocolea aphelophylla (Hässel) Váňa, Phytotaxa 112 (1): 25, 2013 (Söderström
et al. 2013f). Bas.: Chiloscyphus aphelophyllus Hässel, J. Hattori Bot. Lab. 98: 123,
2005 (Hässel 2005).
World checklist of hornworts and liverworts
209
** Lophocolea apophylla (Hässel) Váňa, Phytotaxa 112 (1): 25, 2013 (Söderström et
al. 2013f). Bas.: Chiloscyphus apophyllus Hässel, J. Hattori Bot. Lab. 98: 126, 2005
(Hässel 2005).
*** Lophocolea appalachiana R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 208,
1980 (Schuster 1980c).
** Lophocolea ascensionis Steph., Sp. Hepat. (Stephani) 6: 261, 1922 (Stephani 1922).
** Lophocolea asperrima Steph., Bull. Herb. Boissier (sér. 2) 6 (11): 949 (129), 1906
(Stephani 1906c).
** Lophocolea atra Gola, Nuovo Giorn. Bot. Ital. (n.ser.) 29 (1/4): 167, 1922 [1923]
(Gola 1922).
*** Lophocolea attenuata Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26 (III, 6):
34, 1900 (Stephani 1900b).
*** Lophocolea australis Gottsche, Linnaea 28 (5): 553, 1856 [1857] (Gottsche 1856).
** Lophocolea autoica Steph., Sp. Hepat. (Stephani) 6: 262, 1922 (Stephani 1922).
** Lophocolea baldwinii Steph., Bull. Herb. Boissier (sér. 2) 6 (11): 950 (130), 1906
(Stephani 1906c).
** Lophocolea bartlettii H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 506, 1963 (Miller 1963).
** Lophocolea bewsii (Sim) Grolle, Trans. Brit. Bryol. Soc. 3 (4): 588, 1959 (Grolle
1959b). Bas.: Leptoscyphus bewsii Sim, Trans. Roy. Soc. South Africa 15 (1): 103,
1926 (Sim 1926).
** Lophocolea bicuspidata Steph., Sp. Hepat. (Stephani) 6: 263, 1922 (Stephani 1922).
*** Lophocolea bidentata (L.) Dumort., Recueil Observ. Jungerm.: 17, 1835 (Dumortier
1835). Bas.: Jungermannia bidentata L., Sp. Pl. 1: 1132, 1753 (Linnaeus 1753). 148
** Lophocolea biidistipula Steph., Sp. Hepat. (Stephani) 6: 264, 1922 (Stephani 1922).
*** Lophocolea bispinosa (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat.
2: 162, 1845 (Gottsche et al. 1845a). Bas.: Jungermannia bispinosa Hook.f. et
Taylor, London J. Bot. 3: 378, 1844 (Hooker and Taylor 1844a).
** Lophocolea bootanensis Steph., Sp. Hepat. (Stephani) 6: 265, 1922 (Stephani 1922).
** Lophocolea boulyana Steph., Sp. Hepat. (Stephani) 6: 264, 1922 (Stephani 1922).
** Lophocolea bowiena Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2): 117,
1914 (Stephani and Watts 1914).
*** Lophocolea brookwoodiana Paton et Sheahan, J. Bryol. 28 (3): 163, 2006 (Paton
and Sheahan 2006).
** Lophocolea caespitans Steph., Bull. Herb. Boissier (sér. 2) 6 (11): 949 (129), 1906
(Stephani 1906c).
*** Lophocolea calcarea Steph., Bull. Herb. Boissier (sér. 2) 6 (10): 884 (109), 1906
(Stephani 1906d).
** Lophocolea caledonica Steph., Sp. Hepat. (Stephani) 6: 267, 1922 (Stephani 1922).
148 Lophocolea bidentata is a species complex as discussed by Váňa and Engel (2013) and it has a complicated nomenclatural history causing many misunderstandings. At least Lophocolea coadunata and
Lophocolea humifusa also belong to the complex.
210
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Lophocolea canaliculata (Gottsche, Lindenb. et Nees) Steph., Bull. Herb. Boissier
(sér. 2) 6 (9): 786 (86), 1906 (Stephani 1906e). Bas.: Chiloscyphus canaliculatus
Gottsche, Lindenb. et Nees, Syn. Hepat. 5: 710, 1847 (Gottsche et al. 1847).
*** Lophocolea canaliculata var. concava (J.J.Engel) L.Söderstr., Phytotaxa 112 (1):
26, 2013 (Söderström et al. 2013f). Bas.: Chiloscyphus canaliculatus var. concavus
J.J.Engel, Fieldiana, Bot. (n.ser.) 48: 107, 2010 (Engel 2010).
** Lophocolea cervicornis Steph., Biblioth. Bot. 87 (2): 219, 1916 (Stephani 1916a).
** Lophocolea ciliifera Steph., Bull. Herb. Boissier (sér. 2) 6 (8): 660 (76), 1906
(Stephani 1906h).
* Lophocolea coadunata (Sw.) Mont., Voy. Amér. Mérid., Bot. 7 (1): 76, 1839
(Montagne 1839b). Bas.: Jungermannia coadunata Sw., Fl. Ind. Occid. 3: 1850,
1806 (Swartz 1806). 149
** Lophocolea concreta Mont., Ann. Sci. Nat. Bot. (sér. 3) 4: 350, 1845 (Montagne
1845b).
** Lophocolea convexula Mitt., Fl. vit.: 405, 1871 [1873] (Mitten 1871).
** Lophocolea corticola Steph., Sp. Hepat. (Stephani) 6: 268, 1922 (Stephani 1922).
*** Lophocolea decurrens Herzog, Trans. & Proc. Roy. Soc. New Zealand 65 (3): 352,
1936 (Herzog 1936b).
* Lophocolea deningeri Herzog, Beih. Bot. Centralbl. 38 (2): 321, 1921 (Herzog 1921).
** Lophocolea dentilora Steph., Bull. Herb. Boissier (sér. 2) 6 (7): 550 (64), 1906
(Stephani 1906f).
** Lophocolea diformis Nees, Syn. Hepat. 2: 166, 1845 (Gottsche et al. 1845a).
** Lophocolea discedens (Lehm. et Lindenb.) Lehm. et Lindenb., Syn. Hepat. 2: 167,
1845 (Gottsche et al. 1845a). Bas.: Jungermannia discedens Lehm. et Lindenb.,
Nov. Stirp. Pug. 5: 3, 1833 (Lehmann 1833).
* Lophocolea dusenii Steph., Cat. Afr. Pl. (Hiern) 2 (2): 314, 1901 (Stephani 1901d).
*** Lophocolea erosa (J.J.Engel) L.Söderstr., Phytotaxa 112 (1): 26, 2013 (Söderström
et al. 2013f). Bas.: Chiloscyphus erosus J.J.Engel, Phytologia 83 (1): 43, 1997 [1998]
(Engel 1997).
*** Lophocolea excisifolia Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2): 118,
1914 (Stephani and Watts 1914).
*** Lophocolea fertilis (J.J.Engel) L.Söderstr., Phytotaxa 112 (1): 26, 2013 (Söderström et al. 2013f). Bas.: Chiloscyphus fertilis J.J.Engel, Phytologia 83 (1): 43, 1997
[1998] (Engel 1997).
** Lophocolea lavicans Steph., Sp. Hepat. (Stephani) 6: 300, 1922 (Stephani 1922).
*** Lophocolea loribunda Steph., Bull. Herb. Boissier (sér. 2) 6 (10): 886 (111), 1906
(Stephani 1906d).
** Lophocolea foliicola Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 428, 1885
(Spruce 1885).
** Lophocolea fragillima Steph., Sp. Hepat. (Stephani) 6: 273, 1922 (Stephani 1922).
149 Lophocolea coadunata belongs to the Lophocolea bidentata species complex with complicated nomenclature (Váňa and Engel 2013).
World checklist of hornworts and liverworts
211
*** Lophocolea fragmentissima R.M.Schust., Phytologia 39 (4): 245, 1978 (Schuster
1978a).
*** Lophocolea fragrans (Moris et De Not.) Gottsche, Lindenb. et Nees, Syn. Hepat. 2:
166, 1845 (Gottsche et al. 1845a). Bas.: Jungermannia fragrans Moris et De Not.,
Fl. Caprariae: 177, 1839 (Moris and De Notaris 1839).
** Lophocolea glaziovii Steph., Bull. Herb. Boissier (sér. 2) 6 (11): 961 (141), 1906
(Stephani 1906c).
** Lophocolea gollanii (Steph.) Váňa, Phytotaxa 112 (1): 26, 2013 (Söderström et al.
2013f). Bas.: Chiloscyphus gollanii Steph., Bull. Herb. Boissier (sér. 2) 7 (10): 837
(209), 1907 (Stephani 1907b).
** Lophocolea granatensis Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 126, 1864 (Gottsche
1864).
** Lophocolea griithiana Steph., Sp. Hepat. (Stephani) 6: 274, 1922 (Stephani 1922).
** Lophocolea hahnii Steph., Bull. Herb. Boissier (sér. 2) 6 (8): 660 (76), 1906
(Stephani 1906h).
** Lophocolea haskarliana Gottsche, Syn. Hepat. 2: 153, 1845 (Gottsche et al. 1845a).
*** Lophocolea hattorii (J.J.Engel) L.Söderstr., Phytotaxa 112 (1): 26, 2013 (Söderström et al. 2013f). Bas.: Chiloscyphus hattorii J.J.Engel, J. Hattori Bot. Lab. 74:
29, 1993 (Engel 1993).
** Lophocolea hawaica Steph., Bull. Herb. Boissier (sér. 2) 6 (11): 945 (125), 1906
(Stephani 1906c).
** Lophocolea heterodonta Steph., Sp. Hepat. (Stephani) 6: 275, 1922 (Stephani 1922).
** Lophocolea heteromorpha Steph., Sp. Hepat. (Stephani) 6: 275, 1922 (Stephani 1922).
*** Lophocolea heterophylla (Schrad.) Dumort., Recueil Observ. Jungerm.: 17, 1835
(Dumortier 1835). Bas.: Jungermannia heterophylla Schrad., J. Bot. (Schrader) 5:
66, 1802 [1803] (Schrader 1802).
** Lophocolea heterophylla subsp. cladogyna R.M.Schust., Hepat. Anthocerotae N.
Amer. 4: 223, 1980 (Schuster 1980c).
** Lophocolea horikawana S.Hatt., Bull. Tokyo Sci. Mus. 11: 50, 1944 (Hattori 1944d).
** Lophocolea howeana Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2): 118,
1914 (Stephani and Watts 1914).
* Lophocolea humifusa (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat.
5: 695, 1847 (Gottsche et al. 1847). Bas.: Jungermannia humifusa Hook.f. et Taylor, London J. Bot. 3: 472, 1844 (Hooker and Taylor 1844b).
** Lophocolea humistrata (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 5: 701, 1847 (Gottsche et al. 1847). Bas.: Jungermannia humistrata Hook.f. et
Taylor, London J. Bot. 4: 82, 1845 (Hooker and Taylor 1845).
** Lophocolea itoana Inoue, J. Jap. Bot. 31 (11): 340, 1956 (Inoue 1956).
** Lophocolea javanica Schifn., Hep. Fl. Buitenzorg: 178, 1900 (Schifner 1900a).
** Lophocolea koponenii (Piippo) Váňa, Phytotaxa 112 (1): 26, 2013 (Söderström et
al. 2013f). Bas.: Chiloscyphus koponenii Piippo, Ann. Bot. Fenn. 35 (1): 55, 1998
(Piippo 1998).
212
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Lophocolea kurzii Sande Lac., Ann. Mus. Bot. Lugduno-Batavi 1: 296, 1864
(Sande Lacoste 1864).
* Lophocolea kurzii var. siamensis N.Kitag., Acta Phytotax. Geobot. 30 (1/3): 33,
1979 (Kitagawa 1979b).
** Lophocolea laceristipula Steph., Sp. Hepat. (Stephani) 6: 281, 1922 (Stephani 1922).
** Lophocolea latistipula Steph., Sp. Hepat. (Stephani) 6: 281, 1922 (Stephani 1922).
*** Lophocolea lauterbachii Steph., Bull. Herb. Boissier (sér. 2) 6 (11): 938 (118),
1906 (Stephani 1906c).
* Lophocolea laxissima Herzog, Ann. Bryol. 5: 77, 1932 (Herzog 1932b).
** Lophocolea ledermannii Steph., Sp. Hepat. (Stephani) 6: 300, 1922 (Stephani 1922).
*** Lophocolea lenta (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 2:
162, 1845 (Gottsche et al. 1845a). Bas.: Jungermannia lenta Hook.f. et Taylor,
London J. Bot. 3: 379, 1844 (Hooker and Taylor 1844a).
*** Lophocolea leptantha (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat.
5: 694, 1847 (Gottsche et al. 1847). Bas.: Jungermannia leptantha Hook.f. et Taylor, London J. Bot. 3: 471, 1844 (Hooker and Taylor 1844b).
*** Lophocolea liebmanniana Gottsche, Mexik. Leverm.: 113, 1863 (Gottsche 1863).
** Lophocolea lindmannii Steph., Bull. Herb. Boissier (sér. 2) 6 (11): 960 (140), 1906
(Stephani 1906c).
*** Lophocolea longiciliata Herzog, Memoranda Soc. Fauna Fl. Fennica 27: 96, 1952
(Herzog 1952c).
** Lophocolea lucida (Spreng.) Mont., Voy. Amér. Mérid., Bot. 7 (2): 78, 1839
(Montagne 1839a). Bas.: Jungermannia lucida Spreng. Nov. Stirp. Pug. 5: 2, 1833
(Lehmann 1833).
** Lophocolea madagascariensis Gottsche, Abh. Naturwiss. Vereins Bremen 7: 344,
1882 (Gottsche 1882).
** Lophocolea magna (Udar et V.Nath) Váňa, Phytotaxa 183 (4): 291, 2014 (Váňa et
al. 2014a). Bas.: Cephaloziella magna Udar et V.Nath, Geophytology 6 (1): 105,
1976 (Udar and Nath 1976).
*** Lophocolea mediinfrons (J.J.Engel et Braggins) L.Söderstr., Phytotaxa 112 (1): 26,
2013 (Söderström et al. 2013f). Bas.: Chiloscyphus mediinfrons J.J.Engel et Braggins, Fieldiana, Bot. (n.ser.) 48: 119, 2010 (Engel 2010).
** Lophocolea micronesica Inoue et H.A.Mill., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 11
(1): 4, 1968 (Inoue and Miller 1968).
* Lophocolea microstipula Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26
(III, 6): 43, 1900 (Stephani 1900b). 150
*** Lophocolea minor Nees, Naturgesch. Eur. Leberm. 2: 330, 1836 (Nees 1836).
** Lophocolea minutistipula Steph., Sp. Hepat. (Stephani) 6: 283, 1922 (Stephani 1922).
* Lophocolea mollis (Nees) Nees, Syn. Hepat. 2: 158, 1845 (Gottsche et al. 1845a).
Bas.: Jungermannia mollis Nees, Enum. Pl. Crypt. Javae: 24, 1830 (Nees 1830). 151
150 Lophocolea microstipula is possibly conspeciic with Lophocolea sabuletorum (Engel 1978), but it was
accepted by Hässel and Rubies (2009).
151 Lophocolea mollis is possibly conspeciic with Lophocolea diformis (Grolle 1995).
World checklist of hornworts and liverworts
213
* Lophocolea morobeana Piippo, Acta Bot. Fenn. 131: 160, 1985 (Piippo 1985a). 152
** Lophocolea muhavurensis (S.W.Arnell) S.W.Arnell ex Pócs, Acta Bot. Acad. Sci.
Hung. 25 (3/4): 227, 1979 [1980] (Bizot and Pócs 1979). Bas.: Chiloscyphus muhavurensis S.W.Arnell, Ark. Bot. (n.ser.) 3 (16): 526, 1956 (Arnell 1956e).
*** Lophocolea muricata (Lehm.) Nees, Syn. Hepat. 2: 169, 1845 (Gottsche et al. 1845a).
Bas.: Jungermannia muricata Lehm., Linnaea 4: 363, 1829 (Lehmann 1829).
* Lophocolea muricata var. major Pearson, Forh. Vidensk.-Selsk. Kristiania 1892
(14): 10, 1893 (Pearson 1893).
** Lophocolea nakajimae S.Hatt. et Inoue, J. Hattori Bot. Lab. 21: 221, 1959 (Inoue
1959c).
*** Lophocolea novae-zeelandiae (Lehm. et Lindenb.) Nees, Syn. Hepat. 2: 168, 1845
(Gottsche et al. 1845a). Bas.: Jungermannia novae-zeelandiae Lehm. et Lindenb.,
Nov. Stirp. Pug. 6: 33, 1834 (Lehmann 1834).
*** Lophocolea novae-zeelandiae var. meridionalis (Steph.) L.Söderstr., Phytotaxa 112
(1): 26, 2013 (Söderström et al. 2013f). Bas.: Lophocolea meridionalis Steph., Bull.
Herb. Boissier (sér. 2) 6 (10): 888 (113), 1906 (Stephani 1906d).
** Lophocolea orbigniana Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 5: 55, 1836
(Nees and Montagne 1836).
** Lophocolea papulimarginata H.A.Mill., Phytologia 47 (4): 323, 1981 (Miller
1981). Nom. nov. pro Lophocolea papulosa Steph., Sp. Hepat. (Stephani) 6: 286,
1922 (Stephani 1922), nom. illeg.
** Lophocolea parca (Gottsche) Fulford et Sharp, Mem. New York Bot. Gard. 63:
19, 1990 (Fulford and Sharp 1990). Bas.: Jungermannia parca Gottsche, Mexik.
Leverm.: 94, 1863 (Gottsche 1863).
** Lophocolea parva Steph., Sp. Hepat. (Stephani) 6: 287, 1922 (Stephani 1922).
*** Lophocolea parvispinea (J.J.Engel) L.Söderstr., Phytotaxa 112 (1): 26, 2013 (Söderström et al. 2013f). Bas.: Chiloscyphus parvispineus J.J.Engel, Phytologia 83 (1): 44,
1997 [1998] (Engel 1997).
** Lophocolea parvistipula Steph., Sp. Hepat. (Stephani) 6: 287, 1922 (Stephani
1922).
*** Lophocolea patulistipa Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46
(9): 50, 1911 (Stephani 1911b).
*** Lophocolea perpusilla (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat.
2: 163, 1845 (Gottsche et al. 1845a). Bas.: Jungermannia perpusilla Hook.f. et
Taylor, London J. Bot. 3: 380, 1844 (Hooker and Taylor 1844a).
** Lophocolea piacenzai (Gola) Váňa, Phytotaxa 112 (1): 26, 2013 (Söderström et al.
2013f). Bas.: Lophozia piacenzai Gola, Atti Reale Accad. Sci. Torino, Cl. Sci. Fis.
Mat. Nat. 49: 759, 1914 (Gola 1914b).
** Lophocolea pilistipula Steph., Sp. Hepat. (Stephani) 6: 288, 1922 (Stephani 1922).
* Lophocolea pinnatistipula Steph., Biblioth. Bot. 87 (2): 220, 1916 (Stephani 1916a).
152 Lophocolea morobeana is conspeciic with Chiloscyphus kurzii in Piippo (1987), but it was accepted as
very close to Chiloscyphus kurzii by Ariyanti et al. (2009).
214
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Lophocolea platensis C.Massal., Atti Accad. Sci. Med. Nat. Ferrara 80 (3/4): 12,
1906 (Massalongo 1906a).
** Lophocolea purpurea Steph., Sp. Hepat. (Stephani) 6: 289, 1922 (Stephani 1922).
** Lophocolea pusilla Steph., Sp. Hepat. (Stephani) 6: 290, 1922 (Stephani 1922).
** Lophocolea randii S.W.Arnell, Svensk Bot. Tidskr. 47 (3): 420, 1953 (Arnell 1953c).
** Lophocolea rara Steph., Sp. Hepat. (Stephani) 6: 290, 1922 (Stephani 1922).
** Lophocolea rectangularis Herzog, Rev. Bryol. Lichénol. 23 (1/2): 43, 1954 (Herzog
1954).
** Lophocolea rectangulata Mitt., Fl. vit.: 404, 1871 [1873] (Mitten 1871). 153
*** Lophocolea rupicola Steph., Bull. Herb. Boissier (sér. 2) 6 (10): 874 (99), 1906
(Stephani 1906d).
*** Lophocolea sabuletorum (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn.
Hepat. 5: 697, 1847 (Gottsche et al. 1847). Bas.: Jungermannia sabuletorum
Hook.f. et Taylor, London J. Bot. 3: 469, 1844 (Hooker and Taylor 1844b).
** Lophocolea salacensis Steph., Bull. Herb. Boissier (sér. 2) 6 (11): 943 (123), 1906
(Stephani 1906c).
** Lophocolea savesiana Steph., Bull. Herb. Boissier (sér. 2) 6 (11): 942 (122), 1906
(Stephani 1906c).
*** Lophocolea semiteres (Lehm.) Mitt., J. Linn. Soc., Bot. 16 (91): 188, 1877 (Mitten
1877). Bas.: Jungermannia semiteres Lehm., Linnaea 4: 363, 1829 (Lehmann 1829). 154
*** Lophocolea semiteres var. retusa (J.J.Engel) L.Söderstr., Phytotaxa 112 (1): 27, 2013
(Söderström et al. 2013f). Bas.: Chiloscyphus semiteres var. retusus J.J.Engel, Phytologia 83 (1): 44, 1997 [1998] (Engel 1997).
** Lophocolea serrata Mitt., St. Helena: 368, 1875 (Mitten 1875).
** Lophocolea siamensis Steph., Sp. Hepat. (Stephani) 6: 293, 1922 (Stephani 1922).
** Lophocolea sikkimensis (Steph.) Herzog et Grolle, Rev. Bryol. Lichénol. 27 (3/4):
164, 1958 [1959] (Herzog and Grolle 1958). Bas.: Herpocladium sikkimense
Steph., Sp. Hepat. (Stephani) 6: 349, 1922 (Stephani 1922).
* Lophocolea silvestris Gottsche, Abh. Naturwiss. Vereins Bremen 7: 345, 1882
(Gottsche 1882).
** Lophocolea steetziae De Not., Epat. Borneo: 20, 1874 (De Notaris 1874).
*** Lophocolea striatella (C.Massal.) Schifn., Leberm., Forschungsr. Gazelle 4 (4): 13,
1890 (Schifner 1890). Bas.: Chiloscyphus striatellus C.Massal., Nuovo Giorn. Bot.
Ital. 17 (3): 232, 1885 (Massalongo 1885).
** Lophocolea subbidentata Herzog, Rev. Bryol. Lichénol. 23 (1/2): 43, 1954 (Herzog
1954).
** Lophocolea subcostata Steph., Sp. Hepat. (Stephani) 6: 295, 1922 (Stephani 1922).
*** Lophocolea subporosa Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 137, 1854
(Mitten 1854).
153 Lophocolea rectangulata may be a Cryptolophocolea species.
154 Lophocolea semiteres is a species complex also including Lophocolea platensis and Lophocolea undulata.
World checklist of hornworts and liverworts
215
** Lophocolea subporosa var. inlexifolia (Steph.) L.Söderstr., Phytotaxa 112 (1): 27,
2013 (Söderström et al. 2013f). Bas.: Lophocolea inlexifolia Steph., Sp. Hepat.
(Stephani) 6: 278, 1922 (Stephani 1922).
** Lophocolea subviridis (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat.
5: 699, 1847 (Gottsche et al. 1847). Bas.: Jungermannia subviridis Hook.f. et Taylor, London J. Bot. 3: 473, 1844 (Hooker and Taylor 1844b).
** Lophocolea sumatrana Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss.
Kl. 70: 195, 1900 [1901] (Schifner 1900c).
*** Lophocolea sylvatica Mitt., Rep. Challenger, Bot. 1 (3, 1): 84, 1884 (Mitten 1884b).
** Lophocolea tenera Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 33 (7): 79, 1876
[1877] (Ångström 1876).
** Lophocolea tenerrima Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 439, 1885
(Spruce 1885).
** Lophocolea teptepensis Piippo, Acta Bot. Fenn. 131: 163, 1985 (Piippo 1985a).
*** Lophocolea textilis (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 5:
696, 1847 (Gottsche et al. 1847). Bas.: Jungermannia textilis Hook.f. et Taylor,
London J. Bot. 3: 468, 1844 (Hooker and Taylor 1844b).
** Lophocolea textiloidea J.J.Engel, Phytologia 41 (5): 311, 1979 (Engel 1979a). Nom.
nov. pro Chiloscyphus lucidus Mitt., J. Linn. Soc., Bot. 15 (82): 64, 1876 (Mitten
1876a), nom. illeg.
*** Lophocolea trichocoleoides (Glenny, J.J.Engel et He-Nygrén) L.Söderstr., Phytotaxa 112 (1): 27, 2013 (Söderström et al. 2013f). Bas.: Chiloscyphus trichocoleoides
Glenny, J.J.Engel et He-Nygrén, J. Bryol. 31 (2): 100, 2009 (Glenny et al. 2009).
** Lophocolea tricuspidata Herzog, Rev. Bryol. Lichénol. 11 (1): 17, 1938 [1939]
(Herzog 1938a).
*** Lophocolea tristaniana S.W.Arnell, Results Norweg. Sci. Exped. Tristan da Cunha
42: 19, 1958 (Arnell 1958b).
* Lophocolea undulata Mont., Ann. Sci. Nat. Bot. (sér. 3) 4: 351, 1845 (Montagne
1845b).
*** Lophocolea villosa Mitt., Sp. Hepat. (Stephani) 6: 299, 1922 (Stephani 1922).
** Lophocolea wacei (S.W.Arnell ex J.J.Engel et Váňa) Váňa et L.Söderstr., Phytotaxa 112
(1): 27, 2013 (Söderström et al. 2013f). Bas.: Chiloscyphus wacei S.W.Arnell ex J.J.Engel et Váňa, Mem. New York Bot. Gard. 105: 48, 2013 (Váňa and Engel 2013).
*** Lophocolea wambana Piippo, Acta Bot. Fenn. 131: 163, 1985 (Piippo 1985a).
** Lophocolea werthii (J.J.Engel et R.M.Schust.) Váňa et L.Söderstr., Phytotaxa 112
(1): 27, 2013 (Söderström et al. 2013f). Bas.: Chiloscyphus werthii J.J.Engel et
R.M.Schust., Nova Hedwigia 39: 425, 1984 [1985] (Engel and Schuster 1984).
** Lophocolea widgrenii Steph., Bull. Herb. Boissier (sér. 2) 7 (1): 66 (154), 1907
(Stephani 1907c). Nom. nov. pro Lophocolea pallida Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 33 (7): 80, 1876 [1877] (Ångström 1876), nom. illeg.
** Otoscyphus J.J.Engel, Bardat et houvenot, Cryptog. Bryol. 33 (3): 280, 2012
(Engel et al. 2012).
216
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Otoscyphus crassicaulis (Steph.) J.J.Engel, Bardat et houvenot, Cryptog. Bryol. 33
(3): 280, 2012 (Engel et al. 2012). Bas.: Lophocolea crassicaulis Steph., Sp. Hepat.
(Stephani) 6: 268, 1922 (Stephani 1922).
*** Pachyglossa Herzog et Grolle, Rev. Bryol. Lichénol. 27 (3/4): 150, 1958 [1959]
(Herzog and Grolle 1958).
*** Pachyglossa austrigena (Hook.f. et Taylor) L.Söderstr., Phytotaxa 112 (1): 24, 2013
(Söderström et al. 2013f). Bas.: Jungermannia austrigena Hook.f. et Taylor, London J. Bot. 3: 466, 1844 (Hooker and Taylor 1844b).
*** Pachyglossa austrigena subsp. okaritana (Steph.) L.Söderstr., Phytotaxa 112 (1):
24, 2013 (Söderström et al. 2013f). Bas.: Lophocolea okaritana Steph., Bull. Herb.
Boissier (sér. 2) 6 (9): 785 (85), 1906 (Stephani 1906e).
*** Pachyglossa boveana (C.Massal.) L.Söderstr., Phytotaxa 112 (1): 24, 2013 (Söderström et al. 2013f). Bas.: Lophocolea boveana C.Massal., Nuovo Giorn. Bot. Ital.
17 (3): 225, 1885 (Massalongo 1885).
** Pachyglossa dissitifolia Herzog et Grolle, Rev. Bryol. Lichénol. 27 (3/4): 155, 1958
[1959] (Herzog and Grolle 1958).
* Pachyglossa exilis (Herzog et Grolle) Hässel et Solari, Transecta botánica de la Patagonia austral: 324, 1985 (Hässel and Solari 1985). Bas.: Pachyglossa spegazziniana
var. exilis Herzog et Grolle, Rev. Bryol. Lichénol. 27 (3/4): 159, 1958 [1959]
(Herzog and Grolle 1958). 155
** Pachyglossa issa (Mitt.) Herzog et Grolle, Rev. Bryol. Lichénol. 28 (3/4): 346,
1959 [1960] (Grolle 1959c). Bas.: Herpocladium issum Mitt., J. Linn. Soc., Bot.
15 (82): 69, 1876 (Mitten 1876a).
*** Pachyglossa gottscheoides (Besch. et C.Massal.) L.Söderstr., Phytotaxa 112 (1): 25,
2013 (Söderström et al. 2013f). Bas.: Lophocolea gottscheoides Besch. et C.Massal.,
Bull. Mens. Soc. Linn. Paris 1 (79): 631, 1886 (Bescherelle and Massalongo 1886).
** Pachyglossa grolleana Váňa, Cryptog. Bryol. 26 (1): 86, 2005 (Váňa and Gremmen 2005).
** Pachyglossa otiphylla (Hook.f. et Taylor) Váňa, Phytotaxa 112 (1): 25, 2013
(Söderström et al. 2013f). Bas.: Jungermannia otiphylla Hook.f. et Taylor, London
J. Bot. 3: 466, 1844 (Hooker and Taylor 1844b).
** Pachyglossa spegazziniana (C.Massal.) Herzog et Grolle, Rev. Bryol. Lichénol. 27
(3/4): 159, 1958 [1959] (Herzog and Grolle 1958). Bas.: Lophocolea spegazziniana
C.Massal., Nuovo Giorn. Bot. Ital. 17 (3): 225, 1885 (Massalongo 1885).
** Pachyglossa tenacifolia (Hook.f. et Taylor) Herzog et Grolle, Rev. Bryol. Lichénol.
27 (3/4): 153, 1958 [1959] (Herzog and Grolle 1958). Bas.: Jungermannia tenacifolia Hook.f. et Taylor, Bot. Antarct. Voy. I (Fl. Antarct. 1): 152, 1845 (Taylor
and Hooker 1845).
** Perdusenia Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia Inst.
Nac. Invest. Ci. Nat. Bot. 7 (2): 11, 1989 (Hässel 1989b).
155 Pachyglossa exilis may be a Chiloscyphus species.
World checklist of hornworts and liverworts
217
** Perdusenia rheophila Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 7 (2): 11, 1989 (Hässel 1989b).
** Pigafettoa C.Massal., Nuovo Giorn. Bot. Ital. 17 (3): 237, 1885 (Massalongo 1885).
** Pigafettoa crenulata C.Massal., Nuovo Giorn. Bot. Ital. 17 (3): 237, 1885 (Massalongo 1885).
** Platycaulis R.M.Schust., Phytologia 39 (4): 245, 1978 (Schuster 1978a).
*** Platycaulis renifolius R.M.Schust., Phytologia 39 (4): 245, 1978 (Schuster 1978a).
*** Stolonivector J.J.Engel, J. Hattori Bot. Lab. 69: 80, 1991 (Engel 1991a).
** Stolonivector clasmatocoleoides J.J.Engel, Nova Hedwigia 88 (3/4): 339, 2009 (Engel
2009).
*** Stolonivector iordlandiae (E.A.Hodgs.) J.J.Engel, J. Hattori Bot. Lab. 69: 82, 1991
(Engel 1991a). Bas.: Lophocolea iordlandiae E.A.Hodgs., Trans. Roy. Soc. New
Zealand 80 (3/4): 340, 1952 [1953] (Hodgson 1952).
** Stolonivector iordlandiae var. nodulosus J.J.Engel, Nova Hedwigia 93 (3/4): 403,
2011 (Engel 2011).
*** Stolonivector gremmenii (Váňa) Váňa, Phytotaxa 112 (1): 28, 2013 (Söderström et
al. 2013f). Bas.: Chiloscyphus gremmenii Váňa, Cryptog. Bryol. 26 (1): 81, 2005
(Váňa and Gremmen 2005).
** Stolonivector obtusilobus J.J.Engel, Nova Hedwigia 88 (3/4): 337, 2009 (Engel 2009).
** Stolonivector waipouensis J.J.Engel, J. Hattori Bot. Lab. 93: 70, 2003 (Engel 2003).
** Xenocephalozia R.M.Schust., Nova Hedwigia 10 (1/2): 25, 1965 (Schuster 1965b).
** Xenocephalozia navicularis (Steph.) R.M.Schust., Nova Hedwigia 10 (1/2): 25,
1965 (Schuster 1965b). Bas.: Lophocolea navicularis Steph., Bih. Kongl. Svenska
Vetensk.-Akad. Handl. 26 (III, 6): 43, 1900 (Stephani 1900b).
***
Mastigophoraceae R.M.Schust.
** Dendromastigophora R.M.Schust., Mem. New York Bot. Gard. 45: 738, 1987
(Schuster 1987a).
*** Dendromastigophora lagellifera (Hook.) R.M.Schust., Mem. New York Bot. Gard.
45: 738, 1987 (Schuster 1987a). Bas.: Jungermannia lagellifera Hook., Musci
Exot. 1: tab. 59, 1818 (Hooker 1818).
*** Mastigophora Nees, Naturgesch. Eur. Leberm. 3: 89, 1838 (Nees 1838b) nom.
conserv. 156
156 Mastigophora includes (Mastigophora appendiculata, Mastigophora guineensis, Mastigophora pyramidana and Mastigophora valida) which Grolle and Piippo (1984) could not study since the types were
destroyed in B.
218
*
*
**
***
**
**
*
*
*
*
**
**
*
**
***
***
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Mastigophora appendiculata Steph., Sp. Hepat. (Stephani) 6: 368, 1922 (Stephani
1922).
Mastigophora attenuata (Taylor) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 416, 1877 (Trevisan 1877). Bas.: Lepidozia attenuata Taylor,
London J. Bot. 5: 369, 1846 (Taylor 1846b).
Mastigophora caledonica Steph., Rev. Bryol. 35 (2): 31, 1908 (Stephani 1908l).
Mastigophora diclados (Brid. ex F.Weber) Nees, Naturgesch. Eur. Leberm. 3: 18,
1838 (Nees 1838b). Bas.: Jungermannia diclados Brid. ex F.Weber, Hist. Musc.
Hepat. Prodr.: 56, 1815 (Weber 1815).
Mastigophora diclados var. borneensis (De Not.) Schifn., Nova Acta Acad. Caes.
Leop.-Carol. German. Nat. Cur. 60 (2): 251, 1893 (Schifner 1893a). Bas.: Sendtnera diclados var. borneensis De Not., Epat. Borneo: 42, 1874 (De Notaris 1874).
Mastigophora diclados var. ramentiissa Herzog, Trans. Brit. Bryol. Soc. 1 (4): 315,
1950 (Herzog 1950a).
Mastigophora diclados var. villosa Herzog, Ann. Bryol. 5: 82, 1932 (Herzog 1932b).
Mastigophora guineensis Steph., Sp. Hepat. (Stephani) 6: 369, 1923 (Stephani 1923).
Mastigophora humillima (Taylor) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 416, 1877 (Trevisan 1877). Bas.: Lepidozia humillima Taylor,
London J. Bot. 5: 369, 1846 (Taylor 1846b).
Mastigophora pyramidana Steph., Sp. Hepat. (Stephani) 6: 369, 1923 (Stephani 1923).
Mastigophora sepikiana Piippo, Ann. Bot. Fenn. 23 (1): 2, 1986 (Piippo 1986b).
Mastigophora tuberculata D.H.Mill. et H.A.Mill., J. Hattori Bot. Lab. 75: 181,
1994 (Miller and Miller 1994).
Mastigophora valida Steph., Sp. Hepat. (Stephani) 6: 369, 1923 (Stephani 1923).
Mastigophora viridula (Nees) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 416, 1877 (Trevisan 1877). Bas.: Jungermannia viridula Nees,
Flora 6 (2): 30, 1823 (Link 1823).
Mastigophora woodsii (Hook.) Nees, Naturgesch. Eur. Leberm. 3: 95, 1838 (Nees
1838b). Bas.: Jungermannia woodsii Hook., Brit. Jungermann.: tab. 66, 1814
(Hooker 1814).
Plagiochilaceae Müll.Frib.
by L. Söderström
he placement of Pedinophyllopsis in Plagiochilaceae follows He-Nygrén and Piippo
(2003). he inclusion of Pseudolophocolea in the family follows Söderström et al.
(2013b). he subgeneric division of Plagiochila follows the review by Söderström et
al. (2015b).
** Acrochila R.M.Schust., J. Hattori Bot. Lab. 26: 285, 1963 (Schuster 1963b).
World checklist of hornworts and liverworts
219
*** Acrochila biserialis (Lehm. et Lindenb.) Grolle, J. Jap. Bot. 39 (8): 236, 1964
(Grolle 1964g). Bas.: Plagiochila biserialis Lehm. et Lindenb., Sp. Hepat. (Lindenberg) 5: 126, 1843 (Lindenberg 1843).
** Acrochila caledonica (Steph.) Inoue, J. Jap. Bot. 42 (6): 182, 1967 (Inoue 1967d).
Bas.: Plagiochila caledonica Steph., Rev. Bryol. 35 (2): 32, 1908 (Stephani 1908l).
** Chiastocaulon Carl, Flora 126: 58, 1931 (Carl 1931a).
*** Chiastocaulon dendroides (Nees) Carl, Flora 126: 59, 1931 (Carl 1931a). Bas.:
Jungermannia dendroides Nees, Enum. Pl. Crypt. Javae: 77, 1830 (Nees 1830).
** Dinckleria Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci. Mat. 4 (13):
421, 1877 (Trevisan 1877).
*** Dinckleria fruticella (Hook.f. et Taylor) J.J.Engel et Heinrichs, Cryptog. Bryol. 29
(2): 194, 2008 (Engel and Heinrichs 2008). Bas.: Jungermannia fruticella Hook.f.
et Taylor, London J. Bot. 3: 565, 1844 (Hooker and Taylor 1844d).
*** Dinckleria pleurata (Hook.f. et Taylor) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 421, 1877 (Trevisan 1877). Bas.: Jungermannia pleurata Hook.f. et Taylor, London J. Bot. 3: 372, 1844 (Hooker and Taylor 1844a).
** Pedinophyllopsis R.M.Schust. et Inoue, Phytologia 47 (4): 311, 1981 (Schuster
and Engel 1981).
** Pedinophyllopsis abdita (Sull.) R.M.Schust. et Inoue, Phytologia 47 (4): 311, 1981
(Schuster and Engel 1981). Bas.: Plagiochila abdita Sull., Hooker’s J. Bot. Kew
Gard. Misc. 2: 317, 1850 (Sullivant 1850).
** Pedinophyllum Lindb. ex Nordst., Bot. Not. 1874: 156, 1874 (Nordstedt 1874).
*** Pedinophyllum autoicum (Steph.) Inoue, Bull. Natl. Sci. Mus. Tokyo (n.ser.) 9 (4):
575, 1966 (Inoue 1966b). Bas.: Plagiochila autoica Steph., Sp. Hepat. (Stephani)
6: 126, 1917 (Stephani 1917a).
*** Pedinophyllum interruptum (Nees) Kaal., Nyt Mag. Naturvidensk. 33 (1): 190, 1893
(Kaalaas 1893a). Bas.: Jungermannia interrupta Nees, Naturgesch. Eur. Leberm. 1:
165, 1833 (Nees 1833c).
*** Pedinophyllum monoicum (Steph.) Grolle, Nova Hedwigia 2: 287, 1960 (Grolle
1960b). Bas.: Plagiochila monoica Steph., Bull. Herb. Boissier (sér. 2) 3 (4): 331
(315), 1903 (Stephani 1903c).
** Pedinophyllum truncatum (Steph.) Inoue, J. Hattori Bot. Lab. 23: 35, 1960 [1961]
(Inoue 1960). Bas.: Clasmatocolea truncata Steph., Bull. Herb. Boissier 5 (2): 87,
1897 (Stephani 1897b).
*** Plagiochila (Dumort.) Dumort., Recueil Observ. Jungerm.: 14, 1835 (Dumortier 1835) nom. conserv. Bas.: Radula sect. Plagiochila Dumort., Syll. Jungerm.
Europ.: 42, 1831 (Dumortier 1831).
220
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Plagiochila heteromalla (Lehm. et Lindenb.) Lindenb., Sp. Hepat. (Lindenberg)
2-4: 83, 1840 (Lindenberg 1840). Bas.: Jungermannia heteromalla Lehm. et Lindenb., Nov. Stirp. Pug. 6: 62, 1834 (Lehmann 1834).
*** sect. Adianthoideae Lindenb., Monogr. hep. gen. Plagiochilae: xx, 1844 (Lindenberg 1844).
*** Plagiochila adianthoides (Sw.) Lindenb., Sp. Hepat. (Lindenberg) 2-4: 77, 1840
(Lindenberg 1840). Bas.: Jungermannia adianthoides Sw., Prodr. (Swartz): 142,
1788 (Swartz 1788).
* Plagiochila adianthoides var. aspergillifera Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 474, 1885 (Spruce 1885).
*** Plagiochila cristata (Sw.) Lindenb., Sp. Hepat. (Lindenberg) 1: 33, 1839 (Lindenberg 1839). Bas.: Jungermannia cristata Sw., Prodr. (Swartz): 143, 1788 (Swartz
1788).
*** Plagiochila grandicrista Steph., Bull. Herb. Boissier (sér. 2) 5 (10): 931 (581), 1905
(Stephani 1905f).
*** Plagiochila herminieri Steph., Bull. Herb. Boissier (sér. 2) 5 (8): 748 (547), 1905
(Stephani 1905i).
*** sect. Africanae Heinrichs, Taxon 54 (2): 319, 2005 (Heinrichs et al. 2005).
*** Plagiochila barteri Mitt., J. Linn. Soc., Bot. 22 (146): 320, 1886 (Mitten 1886b).
** Plagiochila barteri var. valida (Steph.) Vanden Berghen, Bull. Jard. Bot. Natl. Belg.
51 (1/2): 73, 1981 (Vanden Berghen 1981). Bas.: Plagiochila valida Steph., Bull.
Herb. Boissier (sér. 2) 4 (6): 587 (438), 1904 (Stephani 1904b).
*** Plagiochila colorans Steph., Wiss. Ergebn. Deut. Zentr.-Afr. Exped. (1907-08),
Bot. 2: 116, 1911 (Stephani 1911a).
*** sect. Arrectae Carl, Ann. Bryol., Suppl. 2: 52, 1931 (Carl 1931b).
* Plagiochila arnelliana Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 861 (230), 1902
(Stephani 1902g). 157
*** Plagiochila badia Mitt., Rep. Challenger, Bot. 1 (3, 1): 84, 1884 (Mitten 1884b).
*** Plagiochila bidens Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 8: 322, 1857 (Gottsche 1857).
*** Plagiochila bifaria (Sw.) Lindenb., Sp. Hepat. (Lindenberg) 5: 127, 1843 (Lindenberg
1843). Bas.: Jungermannia bifaria Sw., Prodr. (Swartz): 145, 1788 (Swartz 1788).
** Plagiochila bifaria var. rosea (R.M.Schust.) Heinrichs, Org. Divers. Evol. 4 (1/2):
112, 2004 (Heinrichs et al. 2004). Bas.: Rhodoplagiochila rosea R.M.Schust., Phytologia 39 (4): 247, 1978 (Schuster 1978a).
*** Plagiochila chacabucensis Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.)
46 (9): 27, 1911 (Stephani 1911b).
*** Plagiochila emeiensis Grolle et M.L.So, Bryologist 101 (2): 282, 1998 (Grolle and
So 1998a).
157 Plagiochila arnelliana may be conspeciic with Plagiochila bifaria (Heinrichs et al. 1998).
World checklist of hornworts and liverworts
*
***
***
*
**
***
***
***
***
***
***
***
***
**
***
***
***
***
221
Plagiochila fragilis Taylor, London J. Bot. 7: 198, 1848 (Taylor 1848a). 158
Plagiochila lunata S.W.Arnell, Bot. Not. 115: 204, 1962 (Arnell 1962a).
Plagiochila pachyloma Taylor, London J. Bot. 5: 267, 1846 (Taylor 1846a).
Plagiochila pachyloma var. elatior Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:
480, 1885 (Spruce 1885).
Plagiochila papillifolia Steph., Biblioth. Bot. 87 (2): 207, 1916 (Stephani 1916a).
Plagiochila parviramifera Inoue, J. Hattori Bot. Lab. 46: 317, 1979 (Mizutani 1979a).
Plagiochila pseudoattenuata S.W.Arnell, Bot. Not. 115: 206, 1962 (Arnell 1962a).
Plagiochila punctata (Taylor) Taylor, London J. Bot. 5: 261, 1846 (Taylor 1846a).
Bas.: Jungermannia punctata Taylor, Trans. Bot. Soc. Edinburgh 1: 179, 1844
(Taylor 1844b).
Plagiochila renauldii Steph., Bull. Herb. Boissier (sér. 2) 4 (2): 156 (408), 1904
(Stephani 1904f).
Plagiochila retrorsa Gottsche, Mexik. Leverm.: 67, 1863 (Gottsche 1863).
Plagiochila rubescens (Lehm. et Lindenb.) Lindenb., Sp. Hepat. (Lindenberg) 2-4:
46, 1840 (Lindenberg 1840). Bas.: Jungermannia rubescens Lehm. et Lindenb.,
Nov. Stirp. Pug. 6: 63, 1834 (Lehmann 1834).
Plagiochila sichuanensis Grolle et M.L.So, Bryologist 101 (2): 284, 1998 (Grolle
and So 1998a).
Plagiochila spinulosa (Dicks.) Dumort., Recueil Observ. Jungerm.: 15, 1835 (Dumortier 1835). Bas.: Jungermannia spinulosa Dicks., Fasc. Pl. Crypt. Brit. 2: 14,
1790 (Dickson 1790).
Plagiochila sticticola Mont. et Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 6: 198, 1856
(Montagne 1856a).
Plagiochila stricta Lindenb., Sp. Hepat. (Lindenberg) 1: 20, 1839 (Lindenberg
1839).
Plagiochila tronadoris Herzog, Darwiniana 11 (2): 214, 1957 (Herzog 1957b).
Plagiochila uncialis (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 5:
628, 1847 (Gottsche et al. 1847). Bas.: Jungermannia uncialis Hook.f. et Taylor,
London J. Bot. 3: 459, 1844 (Hooker and Taylor 1844b).
Plagiochila wilmsiana Steph., Sp. Hepat. (Stephani) 6: 240, 1921 (Stephani 1921).
sect. Caducifoliae J.J.Engel et G.L.Merr., Nova Hedwigia 96 (3/4): 407, 2013
(Engel and Smith Merrill 2013).
** Plagiochila caducifolia Inoue et R.M.Schust., J. Hattori Bot. Lab. 34: 71, 1971
(Inoue and Schuster 1971).
*
sect. Cardotiae Inoue, Bull. Natl. Sci. Mus. Tokyo (n.ser.) 8 (3): 386, 1965 (Inoue 1965a).
*** Plagiochila cumingiana Steph., Bull. Herb. Boissier (sér. 2) 4 (1): 32 (404), 1904
(Stephani 1904g).
*
158 Plagiochila fragilis may be conspeciic with Plagiochila bifaria (Heinrichs et al. 1998).
222
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Plagiochila denticulata Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 95, 1860 [1861]
(Mitten 1860c).
*** Plagiochila fragillima Steph., Bull. Herb. Boissier (sér. 2) 3 (6): 522 (326), 1903
(Stephani 1903d).
*** Plagiochila pseudorenitens Schifn., Österr. Bot. Z. 49 (4): 132, 1899 (Schifner
1899b).
*** Plagiochila stevensiana Steph., Bull. Herb. Boissier (sér. 2) 3 (2): 110 (290), 1903
(Stephani 1903b).
* sect. Cobanae Carl, Ann. Bryol., Suppl. 2: 79, 1931 (Carl 1931b).
** Plagiochila cobana Steph., Sp. Hepat. (Stephani) 6: 138, 1918 (Stephani 1918).
** Plagiochila detecta M.L.So et Grolle, Nova Hedwigia 71 (3/4): 391, 2000 (So and
Grolle 2000b).
*** Plagiochila singularis Schifn., Hep. Fl. Buitenzorg: 158, 1900 (Schifner 1900a).
*** Plagiochila tagawae Inoue, J. Hattori Bot. Lab. 38: 561, 1974 (Inoue 1974a).
** Plagiochila tixieri Inoue, Bull. Natl. Sci. Mus. Tokyo, B 1 (3): 87, 1975 (Inoue
1975a).
*** Plagiochila zhuensis Grolle et M.L.So, Bryologist 102 (2): 200, 1999 (Grolle and
So 1999c).
*** sect. Cucullatae Schifn., Hep. Fl. Buitenzorg: 107, 1900 (Schifner 1900a).
*** Plagiochila bantamensis (Reinw., Blume et Nees) Mont., Voy. Amér. Mérid.,
Bot. 7 (2): 82, 1839 (Montagne 1839a). Bas.: Jungermannia bantamensis Reinw.,
Blume et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1):
235, 1824 [1825] (Reinwardt et al. 1824a).
*** Plagiochila blepharophora (Nees) Lindenb., Sp. Hepat. (Lindenberg) 2-4: 102,
1840 (Lindenberg 1840). Bas.: Jungermannia blepharophora Nees, Enum. Pl.
Crypt. Javae: 71, 1830 (Nees 1830).
** Plagiochila chauviniana Mont., Ann. Sci. Nat. Bot. (sér. 3) 11: 34, 1849 (Montagne
1849).
** Plagiochila clavatosaccata Steph., Bull. Herb. Boissier (sér. 2) 4 (1): 25 (397), 1904
(Stephani 1904g).
** Plagiochila grossispina Steph., Sp. Hepat. (Stephani) 6: 162, 1918 (Stephani 1918).
*** Plagiochila integerrima Steph., Bot. Jahrb. Syst. 8 (2): 83, 1886 (Stephani 1886d).
* Plagiochila integrilobula Schifn., Hep. Fl. Buitenzorg: 170, 1900 (Schifner
1900a). 159
** Plagiochila johannis-winkleri Herzog, Mitt. Inst. Allg. Bot. Hamburg 7 (3): 187,
1931 (Herzog 1931a).
* Plagiochila kuhliana Sande Lac., Ann. Mus. Bot. Lugduno-Batavi 1: 292, 1864
(Sande Lacoste 1864). 160
159 Plagiochila integrilobula is possibly conspeciic with Plagiochila blepharophora (Söderström et al. 2010a).
160 Plagiochila kuhliana is possibly conspeciic with Plagiochila sciophila (Söderström et al. 2010a).
World checklist of hornworts and liverworts
223
** Plagiochila kurzii Steph., Bull. Herb. Boissier (sér. 2) 3 (2): 112 (292), 1903
(Stephani 1903b).
** Plagiochila reischeckiana Steph., Bull. Herb. Boissier (sér. 2) 3 (4): 331 (315), 1903
(Stephani 1903c).
*** Plagiochila sandei Dozy ex Sande Lac., Plagiochila Sandei: 5, 1856 (Sande Lacoste
1856c).
*** Plagiochila sciophila Nees, Sp. Hepat. (Lindenberg) 2-4: 100, 1840 (Lindenberg 1840).
** Plagiochila sciophila subsp. ciliigera (R.M.Schust.) L.Söderstr., Phytotaxa 208
(1): 84, 2015 (Söderström et al. 2015b). Bas.: Plagiochila japonica subsp. ciliigera
R.M.Schust., Amer. Midl. Naturalist 62 (2): 354, 1959 (Schuster 1959b).
* Plagiochila stephanii Schifn., Hep. Fl. Buitenzorg: 166, 1900 (Schifner 1900a). 161
** Plagiochila subplana Lindenb., Sp. Hepat. (Lindenberg) 2-4: 73, 1840 (Lindenberg 1840).
** Plagiochila vitiensis Mitt., Bonplandia 9 (24): 367, 1861 (Mitten 1861).
*** sect. Denticulatae Schifn., Hep. Fl. Buitenzorg: 106, 1900 (Schifner 1900a).
*** Plagiochila alternans Lindenb. et Gottsche, Syn. Hepat. 5: 648, 1847 (Gottsche et
al. 1847).
** Plagiochila ansata (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 5:
649, 1847 (Gottsche et al. 1847). Bas.: Jungermannia ansata Hook.f. et Taylor,
London J. Bot. 3: 457, 1844 (Hooker and Taylor 1844b).
*** Plagiochila banksiana Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 8: 329, 1857 (Gottsche
1857).
** Plagiochila banksiana var. echinophora Inoue et R.M.Schust., J. Hattori Bot. Lab.
34: 62, 1971 (Inoue and Schuster 1971).
** Plagiochila chonotica Taylor, London J. Bot. 5: 260, 1846 (Taylor 1846a).
** Plagiochila equitans Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 8: 331, 1857 (Gottsche
1857).
* Plagiochila fragmentissima Inoue et R.M.Schust., J. Hattori Bot. Lab. 34: 155,
1971 (Inoue and Schuster 1971). 162
*** Plagiochila gigantea Lindenb., Sp. Hepat. (Lindenberg) 2-4: 115, 1840 (Lindenberg 1840). Nom. nov. pro Jungermannia gigantea Hook., Musci Exot. 1: tab. 93,
1818 (Hooker 1818), nom. illeg.
** Plagiochila gigantea var. inermis J.J.Engel et G.L.Merr., Nova Hedwigia 91 (3/4):
513, 2010 (Engel and Smith Merrill 2010).
*** Plagiochila gregaria (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 5:
654, 1847 (Gottsche et al. 1847). Bas.: Jungermannia gregaria Hook.f. et Taylor,
London J. Bot. 3: 564, 1844 (Hooker and Taylor 1844d).
** Plagiochila hookeriana Lindenb., Sp. Hepat. (Lindenberg) 2-4: 81, 1840 (Lindenberg
1840).
161 Plagiochila stephanii is possibly conspeciic with Plagiochila blepharophora (Söderström et al. 2010a).
162 Plagiochila fragmentissima may be conspeciic with Plagiochila gregaria (M.L. So and D. Glenny, pers.
comm.).
224
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Plagiochila latifrons Gottsche et Hampe, Linnaea 27 (5): 553, 1854 (Hampe 1854).
** Plagiochila minutula (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat.
5: 652, 1847 (Gottsche et al. 1847). Bas.: Jungermannia minutula Hook.f. et Taylor, London J. Bot. 3: 459, 1844 (Hooker and Taylor 1844b).
*** Plagiochila nobilis Gottsche, Bot. Zeitung (Berlin) Beil. 16: 37, 1858 (Gottsche 1858).
** Plagiochila obovata Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9):
33, 1911 (Stephani 1911b).
*** Plagiochila ovata Lindenb. et Gottsche, Syn. Hepat. 5: 656, 1847 (Gottsche et al. 1847).
*** Plagiochila retrospectans Lindenb., Sp. Hepat. (Lindenberg) 5: 123, 1843 (Lindenberg 1843). Nom. nov. pro Jungermannia retrospectans Nees, Linnaea 6 (4): 619,
1831 (Nees 1831), nom. illeg.
** Plagiochila riparia Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9):
34, 1911 (Stephani 1911b).
** Plagiochila rutlandii Steph., Bull. Herb. Boissier (sér. 2) 4 (8): 776 (454), 1904
(Stephani 1904c).
** Plagiochila subpectinata Besch. et C.Massal., Bull. Mens. Soc. Linn. Paris 1 (79):
628, 1886 (Bescherelle and Massalongo 1886).
*** sect. Durae Carl, Ann. Bryol., Suppl. 2: 123, 1931 (Carl 1931b).
** Plagiochila acanthocaulis Sull., Hooker’s J. Bot. Kew Gard. Misc. 2: 317, 1850
(Sullivant 1850).
** Plagiochila angulata Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26 (III,
6): 26, 1900 (Stephani 1900b).
** Plagiochila bicornuta Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 305, 1896 (Stephani 1896a).
** Plagiochila crozetensis Kaal., Nyt Mag. Naturvidensk. 49 (2/3): 92, 1911 (Kaalaas
1911).
*** Plagiochila deltoidea Lindenb., Sp. Hepat. (Lindenberg) 5: 132, 1843 (Lindenberg
1843).
** Plagiochila deltoidea var. densa J.J.Engel et G.L.Merr., Nova Hedwigia 91 (3/4):
506, 2010 (Engel and Smith Merrill 2010).
** Plagiochila dura De Not., Mem. Reale Accad. Sci. Torino (ser. 2) 16: 214, 1857
(De Notaris 1857).
** Plagiochila heterodonta (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 5: 638, 1847 (Gottsche et al. 1847). Bas.: Jungermannia heterodonta Hook.f.
et Taylor, London J. Bot. 3: 460, 1844 (Hooker and Taylor 1844b).
*** Plagiochila ramosissima (Hook.) Lindenb., Sp. Hepat. (Lindenberg) 2-4: 87, 1840
(Lindenberg 1840). Bas.: Jungermannia ramosissima Hook., Musci Exot. 1: tab.
92, 1818 (Hooker 1818).
*** sect. Duseniae Carl, Ann. Bryol., Suppl. 2: 126, 1931 (Carl 1931b).
** Plagiochila dusenii Steph., Bull. Herb. Boissier (sér. 2) 4 (10): 979 (475), 1904
(Stephani 1904a).
** Plagiochila elata Taylor, London J. Bot. 5: 259, 1846 (Taylor 1846a).
World checklist of hornworts and liverworts
225
** Plagiochila lechleri Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 8: 325, 1857 (Gottsche 1857).
** Plagiochila validissima Steph., Biblioth. Bot. 87 (2): 214, 1916 (Stephani 1916a).
* sect. Flexicaules Carl, Ann. Bryol., Suppl. 2: 127, 1931 (Carl 1931b).
** Plagiochila lexicaulis Mont., Syn. Hepat. 5: 629, 1847 (Gottsche et al. 1847).
*** sect. Fruticosae Inoue, Gen. Plagiochila SE Asia: 50, 1984 (Inoue 1984b).
*** Plagiochila assamica Steph., Sp. Hepat. (Stephani) 6: 125, 1917 (Stephani 1917a).
** Plagiochila benitoi Inoue ex Piippo, J. Hattori Bot. Lab. 72: 122, 1992 (Piippo and
Tan 1992).
*** Plagiochila frondescens (Nees) Lindenb., Sp. Hepat. (Lindenberg) 2-4: 52, 1840
(Lindenberg 1840). Bas.: Jungermannia frondescens Nees, Linnaea 6 (4): 610, 1831
(Nees 1831).
*** Plagiochila fruticosa Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 94, 1860 [1861] (Mitten
1860c).
** Plagiochila mauiensis Steph., Sp. Hepat. (Stephani) 6: 184, 1921 (Stephani 1921).
*** Plagiochila pulcherrima Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 1: 63,
1931 (Horikawa 1931a).
** Plagiochila tamiensis Steph., Sp. Hepat. (Stephani) 6: 235, 1921 (Stephani 1921).
***
***
***
***
**
**
***
***
***
***
***
sect. Fuscoluteae Carl, Ann. Bryol., Suppl. 2: 46, 1931 (Carl 1931b).
Plagiochila aerea Taylor, London J. Bot. 5: 263, 1846 (Taylor 1846a).
Plagiochila dependula Taylor, London J. Bot. 5: 265, 1846 (Taylor 1846a).
Plagiochila fuscolutea Taylor, London J. Bot. 5: 263, 1846 (Taylor 1846a).
Plagiochila heterophylla Lindenb., Nov. Stirp. Pug. 10: 2, 1857 (Lehmann 1857).
Plagiochila heterophylla var. beauverdii (Steph.) Heinrichs, Bryophyt. Biblioth. 58:
148, 2002 (Heinrichs 2002). Bas.: Plagiochila beauverdii Steph., Biblioth. Bot. 87
(2): 191, 1916 (Stephani 1916a).
Plagiochila longiramea Steph., Biblioth. Bot. 87 (2): 204, 1916 (Stephani 1916a).
Plagiochila paraphyllina Herzog, Hedwigia 74 (2): 89, 1934 (Herzog 1934a).
Plagiochila rudischusteri H.Rob., Beih. Nova Hedwigia 90: 199, 1988 (Robinson
1988).
Plagiochila subbidentata Taylor, Ann. Mag. Nat. Hist. 20 (135): 381, 1847 (Taylor 1847a).
Plagiochila tabinensis Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 862 (231), 1902
(Stephani 1902g).
*** sect. Glaucescentes Carl, Ann. Bryol., Suppl. 2: 70, 1931 (Carl 1931b).
*** Plagiochila buchtiniana Steph., Sp. Hepat. (Stephani) 6: 135, 1918 (Stephani 1918).
*** Plagiochila diversifolia Lindenb. et Gottsche, Syn. Hepat. 5: 640, 1847 (Gottsche
et al. 1847).
*** Plagiochila longispina Lindenb. et Gottsche, Syn. Hepat. 5: 642, 1847 (Gottsche et
al. 1847).
226
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** sect. Hylacoetes Carl, Ann. Bryol., Suppl. 2: 50, 1931 (Carl 1931b).
*** Plagiochila amicta Steph., Bull. Herb. Boissier (sér. 2) 5 (9): 895 (561), 1905
(Stephani 1905g).
*** Plagiochila boryana Gottsche, Bull. Soc. Roy. Bot. Belgique 31: 118, 1892 (Stephani
1892c).
*** Plagiochila breuteliana Lindenb., Sp. Hepat. (Lindenberg) 5: 150, 1843 (Lindenberg
1843).
*** Plagiochila canelensis Steph., Bull. Herb. Boissier (sér. 2) 5 (10): 926 (576), 1905
(Stephani 1905f).
*** Plagiochila cucullifolia J.B.Jack et Steph., Hedwigia 31 (1): 24, 1892 (Jack and
Stephani 1892).
*** Plagiochila cucullifolia var. anomala Heinrichs et Gradst., Pl. Syst. Evol. 242 (1/4):
208, 2003 (Heinrichs et al. 2003).
*** Plagiochila dimorpha Lindenb. et Gottsche, Syn. Hepat. 5: 627, 1847 (Gottsche et
al. 1847).
*** Plagiochila dominicensis Taylor, London J. Bot. 5: 270, 1846 (Taylor 1846a).
*** Plagiochila ecuadorica (Inoue) L.Söderstr., Phytotaxa 208 (1): 84, 2015 (Söderström et al. 2015b). Bas.: Steereochila ecuadorica Inoue, Mem. New York Bot.
Gard. 45: 279, 1987 (Inoue 1987a).
*** Plagiochila ensiformis Taylor, London J. Bot. 5: 265, 1846 (Taylor 1846a).
*** Plagiochila labellilora Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 880 (249), 1902
(Stephani 1902g).
** Plagiochila guevarii H.Rob., Bryologist 70 (1): 48, 1967 (Robinson 1967).
** Plagiochila husnotii Steph., Bull. Herb. Boissier (sér. 2) 5 (2): 178 (506), 1905
(Stephani 1905d).
*** Plagiochila macrostachya Lindenb., Sp. Hepat. (Lindenberg) 2-4: 75, 1840 (Lindenberg 1840).
*** Plagiochila patriciae Heinrichs et H.Anton, Bryophyt. Biblioth. 58: 107, 2002
(Heinrichs 2002).
*** Plagiochila superba (Nees ex Spreng.) Mont. et Nees, Voy. Amér. Mérid., Bot.
7 (2): 81, 1839 (Montagne 1839a). Bas.: Jungermannia superba Nees ex Spreng.
Syst. Veg. (ed. 16) [Sprengel] 4 (2): 326, 1827 (Sprengel 1827b).
*** Plagiochila superba var. macrotricha (Spruce) Heinrichs, Bryophyt. Biblioth. 58:
114, 2002 (Heinrichs 2002). Bas.: Plagiochila macrotricha Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 476, 1885 (Spruce 1885).
*** Plagiochila turgida Herzog, Hedwigia 72 (6): 196, 1932 (Herzog 1932c).
*** Plagiochila vincentina Lindenb., Sp. Hepat. (Lindenberg) 2-4: 39, 1840 (Lindenberg
1840).
* sect. Jacquinotiae Hässel, Nova Hedwigia 89 (1/2): 85, 2009 (Hässel 2009).
** Plagiochila jacquinotii Mont., Voy. Pole Sud, Bot. 1: 273, 1845 (Montagne 1845c).
* sect. Kaalaasiae Carl, Ann. Bryol., Suppl. 2: 103, 1931 (Carl 1931b).
** Plagiochila amboynensis Taylor, London J. Bot. 5: 260, 1846 (Taylor 1846a).
World checklist of hornworts and liverworts
227
* sect. Longilorae Carl, Ann. Bryol., Suppl. 2: 130, 1931 (Carl 1931b).
** Plagiochila longilora Mont., Syn. Hepat. 5: 651, 1847 (Gottsche et al. 1847).
* sect. Oligodontes Carl, Ann. Bryol., Suppl. 2: 130, 1931 (Carl 1931b).
** Plagiochila lophocoleoides Mont., Ann. Sci. Nat. Bot. (sér. 3) 4: 348, 1845 (Montagne 1845b).
*** sect. Peculiares Schifn., Hep. Fl. Buitenzorg: 107, 1900 (Schifner 1900a).
** Plagiochila aspericaulis Grolle et M.L.So, Syst. Bot. 24 (3): 307, 1999 (Grolle and
So 1999b).
** Plagiochila caulimammillosa Grolle et M.L.So, J. Bryol. 20 (1): 42, 1998 (Grolle
and So 1998c).
*** Plagiochila devexa Steph., Bull. Herb. Boissier (sér. 2) 3 (4): 340 (324), 1903
(Stephani 1903c). Nom. nov. pro Plagiochila delexa Mitt., J. Proc. Linn. Soc., Bot.
5 (18): 97, 1860 [1861] (Mitten 1860c), nom. illeg.
*** Plagiochila durelii Schifn., Österr. Bot. Z. 49 (4): 131, 1899 (Schifner 1899b).
** Plagiochila durelii subsp. guizhouensis Grolle et M.L.So, Syst. Bot. 24 (3): 304,
1999 (Grolle and So 1999b).
** Plagiochila grollei Inoue, Bull. Natl. Sci. Mus. Tokyo (n.ser.) 8 (3): 384, 1965
(Inoue 1965a).
** Plagiochila hoei Inoue, J. Hattori Bot. Lab. 40: 421, 1976 (Inoue 1976b).
** Plagiochila hyalodermica Grolle et M.L.So, Bryologist 100 (4): 470, 1997 (Grolle
and So 1997b).
** Plagiochila magna Inoue, J. Hattori Bot. Lab. 28: 216, 1965 (Inoue 1965c).
** Plagiochila paraphyllosa Grolle et M.L.So, Syst. Bot. 24 (3): 298, 1999 (Grolle and
So 1999b).
** Plagiochila peculiaris Schifn., Hep. Fl. Buitenzorg: 157, 1900 (Schifner 1900a).
*** Plagiochila perserrata Herzog, Symb. Sin. 5: 19, 1930 (Nicholson et al. 1930).
** Plagiochila philippinensis Steph., Bull. Herb. Boissier (sér. 2) 3 (6): 526 (330),
1903 (Stephani 1903d).
*** Plagiochila pseudopoeltii Inoue, Bull. Natl. Sci. Mus. Tokyo (n.ser.) 8 (3): 382,
1965 (Inoue 1965a).
*** Plagiochila renitens (Nees) Lindenb., Sp. Hepat. (Lindenberg) 2-4: 90, 1840 (Lindenberg 1840). Bas.: Jungermannia renitens Nees, Enum. Pl. Crypt. Javae: 76,
1830 (Nees 1830).
** Plagiochila semidecurrens (Lehm. et Lindenb.) Lindenb., Sp. Hepat. (Lindenberg)
5: 142, 1843 (Lindenberg 1843). Bas.: Jungermannia semidecurrens Lehm. et Lindenb., Nov. Stirp. Pug. 4: 21, 1832 (Lehmann 1832).
** Plagiochila semidecurrens var. alaskana (A.Evans) Inoue, J. Hattori Bot. Lab. 28:
216, 1965 (Inoue 1965c). Bas.: Plagiochila alaskana A.Evans, Bull. Torrey Bot.
Club 41 (12): 590, 1914 [1915] (Evans 1914a).
*** Plagiochila vexans Schifn. ex Steph., Sp. Hepat. (Stephani) 6: 237, 1921
(Stephani 1921).
228
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Plagiochila zangii Grolle et M.L.So, Bryologist 100 (4): 467, 1997 (Grolle and So
1997b).
*** Plagiochila zonata Steph., Mém. Soc. Nat. Sci. Nat. Math. Cherbourg 29: 225,
1894 (Stephani 1894b).
*** sect. Plagiochila
** Plagiochila arctica Bryhn et Kaal., Rep. Second Norweg. Arctic Exped. 11: 41,
1906 (Bryhn 1906).
* Plagiochila arctica var. intermedia R.M.Schust., Amer. Midl. Naturalist 62 (1):
152, 1959 (Schuster 1959a).
*** Plagiochila asplenioides (L.) Dumort., Recueil Observ. Jungerm.: 14, 1835 (Dumortier 1835). Bas.: Jungermannia asplenioides L., Sp. Pl. 1: 1131, 1753 (Linnaeus 1753).
*** Plagiochila bischleriana Grolle et M.L.So, Cryptog. Bryol. Lichénol. 18 (3): 191,
1997 (Grolle and So 1997a).
** Plagiochila britannica Paton, J. Bryol. 10 (3): 245, 1979 (Paton 1979b).
*** Plagiochila chinensis Steph., Mém. Soc. Nat. Sci. Nat. Math. Cherbourg 29: 223,
1894 (Stephani 1894b).
*** Plagiochila circumdentata Steph., Bull. Herb. Boissier (sér. 2) 4 (8): 778 (456),
1904 (Stephani 1904c).
** Plagiochila circumdentata var. carinata J.J.Engel et G.L.Merr., Nova Hedwigia 91
(3/4): 511, 2010 (Engel and Smith Merrill 2010).
** Plagiochila circumserrata Inoue et Grolle, Bull. Natl. Sci. Mus. Tokyo, B 5 (1): 27,
1979 (Inoue 1979c).
** Plagiochila columbiana A.Evans, Bot. Gaz. 21 (4): 189, 1896 (Evans 1896).
*** Plagiochila delavayi Steph., Mém. Soc. Nat. Sci. Nat. Math. Cherbourg 29: 224,
1894 (Stephani 1894b).
*** Plagiochila elegans Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 97, 1860 [1861] (Mitten
1860c).
*** Plagiochila hakkodensis Steph., Bull. Herb. Boissier 5 (2): 103, 1897 (Stephani 1897b).
** Plagiochila korthalsiana Molk., Ned. Kruidk. Arch. 3: 416, 1854 [1855] (Sande
Lacoste 1854).
** Plagiochila loriana Steph., Bull. Herb. Boissier (sér. 2) 3 (7): 608 (354), 1903
(Stephani 1903e).
** Plagiochila microdentata M.L.So, New Zealand J. Bot. 38 (3): 426, 2000 (So 2000b).
** Plagiochila mundaliensis Steph., Bull. Herb. Boissier (sér. 2) 3 (6): 536 (340), 1903
(Stephani 1903d).
** Plagiochila orbicularis (S.Hatt.) S.Hatt., J. Hattori Bot. Lab. 3: 26, 1948 [1950]
(Hattori 1948b). Bas.: Plagiochila ovalifolia var. orbicularis S.Hatt., Bull. Tokyo
Sci. Mus. 11: 61, 1944 (Hattori 1944d).
*** Plagiochila ovalifolia Mitt., Trans. Linn. Soc. London, Bot. 3 (3): 193, 1891 (Mitten
1891).
*** Plagiochila porelloides (Torr. ex Nees) Lindenb., Sp. Hepat. (Lindenberg) 2-4: 61,
1840 (Lindenberg 1840). Bas.: Jungermannia porelloides Torr. ex Nees, Naturgesch. Eur. Leberm. 1: 170, 1833 (Nees 1833c).
World checklist of hornworts and liverworts
229
** Plagiochila porelloides var. norvegica (H.H.Blom et Holten) Schumacker et Váňa,
Identif. keys liverw. hornw. Europe: 131, 2005 (Schumacker and Váňa 2005).
Bas.: Plagiochila norvegica H.H.Blom et Holten, Lindbergia 14 (1): 8, 1988 (Blom
and Holten 1988).
** Plagiochila porelloides var. subarctica (Jørg.) Lammes, Fl. Fenn. 6: 54, 1977 (Koponen et al. 1977). Bas.: Plagiochila asplenioides var. subarctica Jørg., Bergens Mus.
Skr. (n.ser.) 16: 173, 1934 (Jørgensen 1934).
** Plagiochila schoieldiana Inoue, Bull. Natl. Sci. Mus. Tokyo (n.ser.) 15 (1): 183,
1972 (Inoue 1972c).
*** Plagiochila secretifolia Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 98, 1860 [1861]
(Mitten 1860c).
** Plagiochila spinosa M.L.So, New Zealand J. Bot. 38 (3): 428, 2000 (So 2000b).
** Plagiochila sumatrana Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss.
Kl. 70: 183, 1900 [1901] (Schifner 1900c).
** Plagiochila taiwanensis Inoue, Bull. Natl. Sci. Mus. Tokyo, B 8 (4): 136, 1982
(Inoue 1982).
*** Plagiochila trapezoidea Lindenb., Sp. Hepat. (Lindenberg) 2-4: 112, 1840 (Lindenberg 1840). Nom. nov. pro Jungermannia asplenioides β australis Nees, Enum.
Pl. Crypt. Javae: 73, 1830 (Nees 1830).
** Plagiochila uniformis Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 98, 1860 [1861] (Mitten
1860c).
** sect. Poeltiae Inoue, Bull. Natl. Sci. Mus. Tokyo (n.ser.) 8 (3): 394, 1965 (Inoue
1965a).
*** Plagiochila biondiana C.Massal., Hepat. Shen-si: 15, 1897 (Massalongo 1897).
*** Plagiochila carringtonii (Balf. ex Carrington) Grolle, Trans. Brit. Bryol. Soc. 4 (4):
656, 1964 (Grolle 1964i). Bas.: Adelanthus carringtonii Balf. ex Carrington, Trans.
Bot. Soc. Edinburgh 10: 380, 1870 (Carrington 1870).
** Plagiochila carringtonii subsp. lobuchensis Grolle, Trans. Brit. Bryol. Soc. 4 (4):
660, 1964 (Grolle 1964i).
*** Plagiochila duthiana Steph., Bull. Herb. Boissier (sér. 2) 3 (6): 527 (331), 1903
(Stephani 1903d).
** Plagiochila erlangensis M.L.So, Haussknechtia, Beih. 9: 350, 1999 (So 1999).
*** Plagiochila poeltii Inoue et Grolle, Trans. Brit. Bryol. Soc. 4 (4): 656, 1964 (Grolle
1964i).
*** Plagiochila recurvata (W.E.Nicholson) Grolle, Trans. Brit. Bryol. Soc. 4 (4): 654,
1964 (Grolle 1964i). Bas.: Jamesoniella carringtonii var. recurvata W.E.Nicholson,
Symb. Sin. 5: 13, 1930 (Nicholson et al. 1930).
*** Plagiochila retusa Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 96, 1860 [1861] (Mitten
1860c).
*** Plagiochila wallichiana Steph., Bull. Herb. Boissier (sér. 2) 3 (6): 523 (327), 1903
(Stephani 1903d).
** Plagiochila wangii Inoue, J. Jap. Bot. 37 (6): 187, 1962 (Inoue 1962c).
** Plagiochila yulungensis Piippo, Ann. Bot. Fenn. 34 (4): 283, 1997 (Piippo 1997).
230
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**
**
*
***
***
***
***
**
***
***
***
**
**
**
**
***
***
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
sect. Rutilantes Carl, Ann. Bryol., Suppl. 2: 83, 1931 (Carl 1931b).
Plagiochila bicuspidata Gottsche, Mexik. Leverm.: 43, 1863 (Gottsche 1863).
Plagiochila caduciloba H.L.Blomq., Bryologist 42 (5): 114, 1939 (Blomquist 1939).
Plagiochila chimborazensis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 469,
1885 (Spruce 1885).
Plagiochila cuneata Lindenb. et Gottsche, Syn. Hepat. 5: 632, 1847 (Gottsche et
al. 1847). 163
Plagiochila debilis Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 97, 1860 [1861] (Mitten
1860c).
Plagiochila delexa Mont. et Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 6: 192, 1856
(Montagne 1856a).
Plagiochila defolians Grolle et M.L.So, Syst. Bot. 23 (4): 459, 1998 [1999] (Grolle
and So 1998b).
Plagiochila exigua (Taylor) Taylor, London J. Bot. 5: 264, 1846 (Taylor 1846a). Bas.:
Jungermannia exigua Taylor, Trans. Bot. Soc. Edinburgh 1: 179, 1844 (Taylor 1844b).
Plagiochila fracta Pócs, Phytotaxa 195 (2): 183, 2015 (Pócs 2015a).
Plagiochila ghatiensis Steph., Sp. Hepat. (Stephani) 6: 159, 1918 (Stephani 1918).
Plagiochila grossa Grolle et M.L.So, Syst. Bot. 23 (4): 461, 1998 [1999] (Grolle
and So 1998b).
Plagiochila gymnocalycina (Lehm. et Lindenb.) Mont. et Nees, Voy. Amér. Mérid.,
Bot. 7 (2): 81, 1839 (Montagne 1839a). Bas.: Jungermannia gymnocalycina Lehm.
et Lindenb., Nov. Stirp. Pug. 5: 28, 1833 (Lehmann 1833).
Plagiochila gymnocalycina var. surinamensis (Molk.) Heinrichs et D.S.Rycroft, J.
Hattori Bot. Lab. 100: 139, 2006 (Heinrichs et al. 2006). Bas.: Plagiochila surinamensis Molk., Syn. hepat. jav.: 103, 1856 [1857] (Sande Lacoste 1856b).
Plagiochila loriloba Herzog ex Carl, Ann. Bryol., Suppl. 2: 47, 1931 (Carl 1931b).
Plagiochila maderensis Gottsche ex Steph., Bull. Herb. Boissier (sér. 2) 4 (4): 350
(426), 1904 (Stephani 1904e).
Plagiochila oresitropha Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 467, 1885
(Spruce 1885).
Plagiochila pectinata Lindenb., Sp. Hepat. (Lindenberg) 1: 14, 1839 (Lindenberg
1839).
Plagiochila rutilans Lindenb., Sp. Hepat. (Lindenberg) 2-4: 47, 1840 (Lindenberg
1840).
Plagiochila rutilans var. moritziana (Lindenb. et Gottsche) Heinrichs, Bryologist
105 (2): 197, 2002 (Heinrichs et al. 2002). Bas.: Plagiochila moritziana Lindenb.
et Gottsche, Linnaea 20 (3): 323, 1847 (Hampe 1847).
Plagiochila rutilans var. standleyi (Herzog ex Carl) Heinrichs et D.S.Rycroft, Bryologist 104 (3): 357, 2001 (Heinrichs et al. 2001). Bas.: Plagiochila standleyi Herzog ex
Carl, Ann. Bryol., Suppl. 2: 80, 1931 (Carl 1931b).
163 Plagiochila cuneata is conspeciic with Plagiochila bursata in Inoue (1980), but accepted by Grolle and
Heinrichs (1999).
World checklist of hornworts and liverworts
231
* Plagiochila steyermarkii H.Rob., Bryologist 68 (1): 93, 1965 (Robinson 1965). 164
*** Plagiochila trichostoma Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 113, 1864
(Gottsche 1864).
sect. Strombifoliae Inoue et R.M.Schust., J. Hattori Bot. Lab. 34: 130, 1971
(Inoue and Schuster 1971).
** Plagiochila strombifolia Taylor, Nov. Stirp. Pug. 8: 5, 1844 (Lehmann 1844).
*
*** sect. Tayloriae Carl, Ann. Bryol., Suppl. 2: 140, 1931 (Carl 1931b).
*** Plagiochila annotina Lindenb., Sp. Hepat. (Lindenberg) 1: 34, 1839 (Lindenberg 1839).
*** Plagiochila baylisii Inoue et R.M.Schust., J. Hattori Bot. Lab. 34: 150, 1971 (Inoue
and Schuster 1971).
*** Plagiochila bazzanioides J.J.Engel et G.L.Merr., Novon 9 (1): 29, 1999 (Engel and
Smith Merrill 1999b).
*** Plagiochila chenii Grolle et M.L.So, Syst. Bot. 25 (1): 6, 2000 (Grolle and So
2000).
*** Plagiochila circinalis (Lehm. et Lindenb.) Lehm., Sp. Hepat. (Lindenberg) 5: 124,
1843 (Lindenberg 1843). Bas.: Jungermannia circinalis Lehm. et Lindenb., Nov.
Stirp. Pug. 4: 64, 1832 (Lehmann 1832).
** Plagiochila circinalis var. hemicardia (Hook.f. et Taylor) J.J.Engel et G.L.Merr.,
Nova Hedwigia 91 (3/4): 512, 2010 (Engel and Smith Merrill 2010). Bas.: Jungermannia hemicardia Hook.f. et Taylor, London J. Bot. 3: 371, 1844 (Hooker and
Taylor 1844a).
*** Plagiochila colensoi Hook.f. et Taylor, London J. Bot. 5: 269, 1846 (Taylor 1846a).
** Plagiochila colensoi var. quinquespina (Steph.) J.J.Engel et G.L.Merr., Nova Hedwigia
91 (3/4): 504, 2010 (Engel and Smith Merrill 2010). Bas.: Plagiochila quinquespina
Steph., Bull. Herb. Boissier (sér. 2) 3 (4): 328 (312), 1903 (Stephani 1903c).
** Plagiochila corticola Steph., Mém. Soc. Nat. Sci. Nat. Math. Cherbourg 29: 224,
1894 (Stephani 1894b).
** Plagiochila fasciculata Lindenb., Sp. Hepat. (Lindenberg) 1: 7, 1839 (Lindenberg
1839).
*** Plagiochila fusca Sande Lac., Ned. Kruidk. Arch. 3: 417, 1854 [1855] (Sande Lacoste 1854).
** Plagiochila fuscella (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 5:
648, 1847 (Gottsche et al. 1847). Bas.: Jungermannia fuscella Hook.f. et Taylor,
London J. Bot. 3: 373, 1844 (Hooker and Taylor 1844a).
** Plagiochila fuscella var. novae-zelandiae (E.A.Hodgs.) J.J.Engel et G.L.Merr., Nova
Hedwigia 89 (3/4): 294, 2009 (Engel and Smith Merrill 2009). Bas.: Plagiochila
retrospectans var. novae-zelandiae E.A.Hodgs., Trans. & Proc. Roy. Soc. New Zealand 73 (4): 293, 1944 (Hodgson 1944).
164 Plagiochila steyermarkii is conspeciic with Plagiochila aerea in Inoue in Gradstein and Hekking
(1979), but this was rejected by Grolle and Heinrichs (1999).
232
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Plagiochila gracilis Lindenb. et Gottsche, Syn. Hepat. 5: 632, 1847 (Gottsche et al.
1847).
*** Plagiochila gymnoclada Sande Lac., Plagiochila Sandei: 6, 1856 (Sande Lacoste 1856c).
*** Plagiochila himalayana Schifn., Österr. Bot. Z. 49 (4): 131, 1899 (Schifner 1899b).
** Plagiochila incurvicolla (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn.
Hepat. 5: 651, 1847 (Gottsche et al. 1847). Bas.: Jungermannia incurvicolla
Hook.f. et Taylor, London J. Bot. 3: 564, 1844 (Hooker and Taylor 1844d).
** Plagiochila incurvicolla var. lonchoscypha (Herzog) J.J.Engel et G.L.Merr., Nova
Hedwigia 91 (3/4): 505, 2010 (Engel and Smith Merrill 2010). Bas.: Plagiochila
lonchoscypha Herzog, Trans. & Proc. Roy. Soc. New Zealand 68 (1): 42, 1938
(Herzog 1938c).
** Plagiochila microdictyon Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 131, 1854
(Mitten 1854).
** Plagiochila monospiris Inoue et Grolle, J. Hattori Bot. Lab. 36: 489, 1972 [1973]
(Inoue 1972b).
*** Plagiochila nitens Inoue, Willdenowia 18 (2): 561, 1989 (Inoue 1989b).
** Plagiochila pseudocapillaris Inoue, Bull. Natl. Sci. Mus. Tokyo (n.ser.) 11 (3): 302,
1968 (Inoue 1968a).
*** Plagiochila pseudoirma Herzog, Symb. Sin. 5: 17, 1930 (Nicholson et al. 1930).
** Plagiochila radiculosa Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 132, 1854
(Mitten 1854).
** Plagiochila spathulifolia Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 96, 1860 [1861]
(Mitten 1860c).
*** Plagiochila stephensoniana Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 133, 1854
(Mitten 1854).
*** sect. Trabeculatae S.Hatt. ex Inoue, J. Hattori Bot. Lab. 20: 75, 1958 (Inoue 1958).
** Plagiochila austinii A.Evans, Rhodora 16 (184): 68, 1914 (Evans 1914b).
*** Plagiochila lexuosa Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 94, 1860 [1861] (Mitten
1860c).
** Plagiochila sullivantii Gottsche, Bot. Gaz. 21 (4): 191, 1896 (Evans 1896).
** Plagiochila sullivantii var. spinigera R.M.Schust., Amer. Midl. Naturalist 62 (2):
323, 1959 (Schuster 1959b).
*** Plagiochila trabeculata Steph., Bull. Herb. Boissier (sér. 2) 3 (2): 103 (283), 1903
(Stephani 1903b).
*** sect. Vagae Lindenb., Monogr. hep. gen. Plagiochilae: xv, 1844 (Lindenberg 1844).
*** Plagiochila abietina (Nees) Mont. et Nees, Voy. Amér. Mérid., Bot. 7 (2): 81,
1839 (Montagne 1839a). Bas.: Jungermannia abietina Nees, Enum. Pl. Crypt.
Javae: 76, 1830 (Nees 1830).
** Plagiochila abrupta Lehm. et Lindenb., Sp. Hepat. (Lindenberg) 2-4: 106, 1840
(Lindenberg 1840).
World checklist of hornworts and liverworts
233
** Plagiochila aequatorialis Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 8: 334, 1857
(Gottsche 1857).
*** Plagiochila africana Steph., Bull. Herb. Boissier (sér. 2) 2 (12): 973 (263), 1902
(Stephani 1902h).
*** Plagiochila akiyamae Inoue, Bull. Natl. Sci. Mus. Tokyo, B 12 (3): 73, 1986 (Inoue 1986).
*** Plagiochila angusta Lindenb., Sp. Hepat. (Lindenberg) 5: 148, 1843 (Lindenberg
1843).
*** Plagiochila angustitexta Steph., Bull. Herb. Boissier (sér. 2) 2 (12): 977 (267),
1902 (Stephani 1902h).
*** Plagiochila arbuscula (Brid. ex Lehm. et Lindenb.) Lindenb., Sp. Hepat. (Lindenberg) 1: 23, 1839 (Lindenberg 1839). Bas.: Jungermannia arbuscula Brid. ex
Lehm. et Lindenb., Nov. Stirp. Pug. 4: 63, 1832 (Lehmann 1832).
** Plagiochila arbuscula var. rekohuensis J.J.Engel et G.L.Merr., Nova Hedwigia 91
(3/4): 509, 2010 (Engel and Smith Merrill 2010).
* Plagiochila aspera Steph., Sp. Hepat. (Stephani) 6: 125, 1917 (Stephani 1917a). 165
** Plagiochila aspleniformis R.M.Schust., Amer. Midl. Naturalist 63 (1): 51, 1960
(Schuster 1960c).
** Plagiochila beddomei Steph., Bull. Herb. Boissier (sér. 2) 3 (10): 876 (361), 1903
(Stephani 1903f).
*** Plagiochila boivinii Steph., Bull. Herb. Boissier (sér. 2) 2 (12): 987 (277), 1902
(Stephani 1902h).
*** Plagiochila bryopteroides Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 499,
1885 (Spruce 1885).
** Plagiochila contigua Gottsche, Mexik. Leverm.: 30, 1863 (Gottsche 1863).
*** Plagiochila corrugata (Nees) Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 5: 52, 1836
(Nees and Montagne 1836). Bas.: Jungermannia corrugata Nees, Fl. Bras. (Martius) 1 (1): 378, 1833 (Nees 1833a).
*** Plagiochila cuspidata Steph., Sp. Hepat. (Stephani) 6: 144, 1918 (Stephani 1918).
** Plagiochila cymata Inoue et Grolle, Bull. Natl. Sci. Mus. Tokyo, B 1 (3): 95, 1975
(Inoue 1975a).
** Plagiochila delexirama Taylor, London J. Bot. 5: 262, 1846 (Taylor 1846a).
*** Plagiochila dichotoma (P.Beauv.) Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 5: 53,
1836 (Nees and Montagne 1836). Bas.: Carpolepidum dichotomum P.Beauv., Fl.
Oware 1 (3): 23, 1805 (Palisot de Beauvois 1805b).
* Plagiochila dichotoma var. luitans Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:
490, 1885 (Spruce 1885).
*** Plagiochila dissecta Steph., Bull. Herb. Boissier (sér. 2) 3 (7): 600 (346), 1903
(Stephani 1903e).
165 Plagiochila aspera is conspeciic with Plagiochila trigona Steph. (=Plagiochila metcalii) in Inoue
(1970b), but it was accepted by So (2000a, 2001a).
234
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Plagiochila disticha (Lehm. et Lindenb.) Lindenb., Sp. Hepat. (Lindenberg) 2-4:
107, 1840 (Lindenberg 1840). Bas.: Jungermannia disticha Lehm. et Lindenb.,
Nov. Stirp. Pug. 6: 64, 1834 (Lehmann 1834).
** Plagiochila distinctifolia Lindenb., Sp. Hepat. (Lindenberg) 1: 17, 1839 (Lindenberg 1839).
*** Plagiochila divergens Steph., Hedwigia 30 (6): 268, 1891 (Stephani 1891c).
*** Plagiochila drepanophylla Sande Lac., Syn. hepat. jav.: 103, 1856 [1857] (Sande
Lacoste 1856b).
* Plagiochila drepanophylla var. minor Gottsche, Abh. Naturwiss. Vereins Bremen 7:
341, 1882 (Gottsche 1882).
*** Plagiochila efusa Steph., Bot. Jahrb. Syst. 20 (3): 310, 1895 (Stephani 1895a).
* Plagiochila efusa var. decurrens Steph., Bot. Jahrb. Syst. 20 (3): 310, 1895 (Stephani
1895a).
*** Plagiochila ericicola Steph., Bull. Herb. Boissier (sér. 2) 4 (6): 590 (441), 1904
(Stephani 1904b).
** Plagiochila exinnovata Steph., Bull. Herb. Boissier (sér. 2) 3 (7): 600 (346), 1903
(Stephani 1903e).
** Plagiochila fastigiata Lindenb. et Gottsche, Syn. Hepat. 5: 657, 1847 (Gottsche et
al. 1847).
*** Plagiochila labellata Steph., Bot. Jahrb. Syst. 8 (2): 82, 1886 (Stephani 1886d).
** Plagiochila loridana A.Evans, Bot. Gaz. 21 (4): 190, 1896 (Evans 1896).
*** Plagiochila fordiana Steph., Bull. Herb. Boissier (sér. 2) 3 (2): 104 (284), 1903
(Stephani 1903b).
** Plagiochila francana Steph., Sp. Hepat. (Stephani) 6: 157, 1918 (Stephani 1918).
*** Plagiochila furcifolia Mitt., Trans. Linn. Soc. London, Bot. 3 (3): 194, 1891 (Mitten
1891).
*** Plagiochila fusifera Taylor, London J. Bot. 5: 268, 1846 (Taylor 1846a).
*** Plagiochila hampeana Gottsche, Bot. Zeitung (Berlin) Beil. 16: 38, 1858 (Gottsche
1858).
* Plagiochila heterospina Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2):
128, 1914 (Stephani and Watts 1914).
* Plagiochila heterostipa Steph., Hedwigia 31 (3): 129, 1892 (Stephani 1892d). 166
** Plagiochila incerta Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 8: 324, 1857 (Gottsche
1857).
*** Plagiochila indica Mitt. ex Steph., Bull. Herb. Boissier (sér. 2) 3 (6): 532 (336),
1903 (Stephani 1903d).
** Plagiochila invisa (R.M.Schust.) R.M.Schust., Hepat. Anthocerotae N. Amer. 4:
513, 1980 (Schuster 1980c). Bas.: Plagiochila ludoviciana var. invisa R.M.Schust.,
Amer. Midl. Naturalist 63 (1): 101, 1960 (Schuster 1960c).
*** Plagiochila javanica (Sw.) Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 5: 52, 1836
(Nees and Montagne 1836). Bas.: Jungermannia javanica Sw., Meth. Musc.: 35,
1781 (Swartz 1781).
166 Plagiochila heterostipa is possibly conspeciic with Plagiochila corymbulosa (Jones 1962).
World checklist of hornworts and liverworts
235
*** Plagiochila junghuhniana Sande Lac., Ned. Kruidk. Arch. 3: 416, 1854 [1855]
(Sande Lacoste 1854).
*** Plagiochila khasiana Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 95, 1860 [1861] (Mitten
1860c).
** Plagiochila kiaeri Gottsche, Abh. Naturwiss. Vereins Bremen 7: 341, 1882
(Gottsche 1882).
** Plagiochila kiaeri var. capensis (Steph.) M.Wigginton et Grolle, Bryophyt. Biblioth. 50: 182, 1996 (Wigginton and Grolle 1996). Bas.: Plagiochila capensis Steph.,
Bull. Herb. Boissier (sér. 2) 4 (4): 350 (426), 1904 (Stephani 1904e).
** Plagiochila kiaeri var. myriocarpa (Pearson) Pócs, J. E. Afr. Nat. Hist. 96 (1): 38,
2007 (Pócs and Luke 2007). Bas.: Plagiochila myriocarpa Pearson, Ark. Bot. 19
(5): 5, 1924 (Pearson 1924b).
*** Plagiochila kunmingensis Piippo, Ann. Bot. Fenn. 34 (4): 281, 1997 (Piippo 1997).
** Plagiochila kurokawae Inoue, J. Hattori Bot. Lab. 32: 104, 1969 (Inoue 1969).
*** Plagiochila laetevirens Lindenb., Sp. Hepat. (Lindenberg) 2-4: 101, 1840 (Lindenberg
1840).
** Plagiochila lamellistipula Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 491,
1885 (Spruce 1885).
*** Plagiochila lastii Mitt., J. Linn. Soc., Bot. 22 (146): 320, 1886 (Mitten 1886b).
** Plagiochila latifolia Steph., Bull. Herb. Boissier (sér. 2) 5 (8): 742 (541), 1905
(Stephani 1905i).
* Plagiochila loloënsis Steph., Bull. Herb. Boissier (sér. 2) 4 (2): 166 (418), 1904
(Stephani 1904f). 167
** Plagiochila manillana Mont. et Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 6: 189, 1856
(Montagne 1856a).
** Plagiochila massalongoana Schifn., Hep. Fl. Buitenzorg: 136, 1900 (Schifner 1900a).
** Plagiochila mastigophoroides Inoue, J. Hattori Bot. Lab. 32: 99, 1969 (Inoue 1969).
** Plagiochila metcalii Steph., Bull. Herb. Boissier (sér. 2) 3 (6): 533 (337), 1903
(Stephani 1903d).
** Plagiochila micropteryx Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 107, 1864
(Gottsche 1864).
** Plagiochila miradorensis Gottsche, Mexik. Leverm.: 31, 1863 (Gottsche 1863).
** Plagiochila miradorensis var. convoluta R.M.Schust., Amer. Midl. Naturalist 63
(1): 113, 1960 (Schuster 1960c).
* Plagiochila moenkemeyeri Steph., Bull. Herb. Boissier (sér. 2) 4 (2): 160 (412),
1904 (Stephani 1904f). 168
*** Plagiochila montagnei Nees, Ann. Sci. Nat. Bot. (sér. 2) 5: 53, 1836 (Nees and
Montagne 1836).
167 Plagiochila loloënsis is probably an entire-leaved form of Plagiochila moenkemeyeri (Wigginton and
Grolle 1996).
168 Plagiochila moenkemeyeri is possibly conspeciic with Plagiochila winteri (Jones 1962), but he expressed
some doubts on the synonymy.
236
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Plagiochila morobensis Inoue et Piippo, Ann. Bot. Fenn. 26 (2): 203, 1989 (Piippo
1989b).
** Plagiochila multipinnula Herzog et S.Hatt., J. Hattori Bot. Lab. 14: 36, 1955
(Herzog and Noguchi 1955).
*** Plagiochila neckeroidea Mitt., Trans. Linn. Soc. London 23 (1): 57, 1860 (Mitten
1860a).
*** Plagiochila nepalensis Lindenb., Sp. Hepat. (Lindenberg) 2-4: 93, 1840 (Lindenberg
1840).
** Plagiochila norfolkiensis Steph., Bull. Herb. Boissier (sér. 2) 3 (10): 877 (362),
1903 (Stephani 1903f).
*** Plagiochila obtusa Lindenb., Sp. Hepat. (Lindenberg) 2-4: 42, 1840 (Lindenberg 1840).
* Plagiochila owaihiensis Nees et Lindenb., Sp. Hepat. (Lindenberg) 1: 30, 1839
(Lindenberg 1839). 169
** Plagiochila paciica Mitt., Fl. vit.: 407, 1871 [1873] (Mitten 1871).
** Plagiochila parallela Steph., Bull. Herb. Boissier (sér. 2) 2 (8): 686 (223), 1902
(Stephani 1902a).
*** Plagiochila parvifolia Lindenb., Sp. Hepat. (Lindenberg) 1: 28, 1839 (Lindenberg
1839).
*** Plagiochila patentissima Lindenb., Sp. Hepat. (Lindenberg) 2-4: 64, 1840 (Lindenberg
1840).
*** Plagiochila patula (Sw.) Lindenb., Sp. Hepat. (Lindenberg) 1: 21, 1839 (Lindenberg
1839). Bas.: Jungermannia patula Sw., Fl. Ind. Occid. 3: 1844, 1806 (Swartz 1806).
** Plagiochila patula var. brevifolia Gottsche, Mexik. Leverm.: 10, 1863 (Gottsche 1863).
** Plagiochila paucidens Steph., Bull. Herb. Boissier (sér. 2) 3 (2): 117 (297), 1903
(Stephani 1903b).
** Plagiochila pensilis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 497, 1885
(Spruce 1885).
** Plagiochila peradenyensis Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 70: 172, 1900 [1901] (Schifner 1900c).
** Plagiochila perdentata M.L.So et Grolle, Syst. Bot. 26 (3): 460, 2001 (So and
Grolle 2001).
*** Plagiochila pinnilora Steph., Hedwigia 30 (5): 212, 1891 (Stephani 1891a).
*** Plagiochila praemorsa Steph., Bot. Jahrb. Syst. 8 (2): 92, 1886 (Stephani 1886d).
** Plagiochila propinqua Sande Lac., Plagiochila Sandei: 5, 1856 (Sande Lacoste 1856c).
** Plagiochila purpurascens Steph., Sp. Hepat. (Stephani) 6: 197, 1921 (Stephani 1921).
*** Plagiochila raddiana Lindenb., Sp. Hepat. (Lindenberg) 1: 9, 1839 (Lindenberg 1839).
* Plagiochila ragazzii Gola, Ann. Bot. (Rome) 13 (1): 67, 1914 (Gola 1914a).
*** Plagiochila repanda (Schwägr.) Lindenb., Sp. Hepat. (Lindenberg) 2-4: 62, 1840
(Lindenberg 1840). Bas.: Jungermannia repanda Schwägr., Hist. Musc. Hepat.
Prodr.: 26, 1814 (Schwägrichen 1814).
169 Plagiochila owaihiensis was considered conspeciic with Plagiochila remyana by Inoue (1976c), but So
(2000a) did not agree.
World checklist of hornworts and liverworts
237
** Plagiochila repanda var. perrotana (Steph.) Vanden Berghen, Bull. Jard. Bot. Natl.
Belg. 51 (1/2): 65, 1981 (Vanden Berghen 1981). Bas.: Plagiochila perrotana
Steph., Bull. Herb. Boissier (sér. 2) 4 (6): 586 (437), 1904 (Stephani 1904b).
*** Plagiochila rodriguezii Steph., Bot. Gaz. 15 (11): 290, 1890 (Stephani 1890c).
*** Plagiochila rudolii Pócs, Beih. Nova Hedwigia 90: 223, 1988 (Pócs 1988).
*** Plagiochila salacensis Gottsche, Natuurk. Tijdschr. Ned.-Indië 4: 576, 1853
(Gottsche 1853).
*** Plagiochila salvadorica Steph., Hedwigia 30 (6): 272, 1891 (Stephani 1891c).
** Plagiochila serrialata Herzog, Hedwigia 72 (6): 212, 1932 (Herzog 1932c).
*** Plagiochila shangaica Steph., Sp. Hepat. (Stephani) 6: 216, 1921 (Stephani 1921).
*** Plagiochila simplex (Sw.) Lindenb., Sp. Hepat. (Lindenberg) 2-4: 54, 1840 (Lindenberg 1840). Bas.: Jungermannia simplex Sw., Prodr. (Swartz): 143, 1788
(Swartz 1788).
*** Plagiochila squamulosa Mitt., J. Proc. Linn. Soc., Bot. 7 (27): 165, 1863 (Mitten 1863).
** Plagiochila squamulosa var. crispulocaudata (Gottsche) Vanden Berghen, Bull. Jard.
Bot. Natl. Belg. 51 (1/2): 74, 1981 (Vanden Berghen 1981). Bas.: Plagiochila crispulocaudata Gottsche, Abh. Naturwiss. Vereins Bremen 7: 340, 1882 (Gottsche 1882).
** Plagiochila squamulosa var. sinuosa (Mitt.) Vanden Berghen, Bull. Jard. Bot. Natl.
Belg. 51 (1/2): 75, 1981 (Vanden Berghen 1981). Bas.: Plagiochila sinuosa Mitt.,
J. Linn. Soc., Bot. 22 (146): 319, 1886 (Mitten 1886b).
** Plagiochila streimannii Inoue, J. Jap. Bot. 63 (11): 365, 1988 (Inoue 1988c).
*** Plagiochila strictifolia Steph., Hedwigia 30 (5): 210, 1891 (Stephani 1891a).
** Plagiochila sublabellata Colenso, Trans. & Proc. New Zealand Inst. 21: 51, 1889
(Colenso 1889).
** Plagiochila subjavanica M.L.So, Austral. Syst. Bot. 13 (5): 804, 2000 (So 2000a).
*** Plagiochila subtropica Steph., Bull. Soc. Roy. Bot. Belgique 38 (1): 46, 1899
(Stephani 1899h).
** Plagiochila tamariscina Steph., Bull. Herb. Boissier (sér. 2) 2 (8): 685 (222), 1902
(Stephani 1902a).
** Plagiochila tecta Inoue et Grolle, J. Hattori Bot. Lab. 33: 319, 1970 (Inoue 1970a).
*** Plagiochila terebrans Nees et Mont., Sp. Hepat. (Lindenberg) 2-4: 98, 1840 (Lindenberg 1840).
*** Plagiochila teysmannii Sande Lac., Plagiochila Sandei: 6, 1856 (Sande Lacoste 1856c).
** Plagiochila thyoides Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 498, 1885
(Spruce 1885).
** Plagiochila tocarema Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 106, 1864 (Gottsche
1864).
** Plagiochila ulata Inoue et Grolle, Bull. Natl. Sci. Mus. Tokyo, B 1 (3): 91, 1975
(Inoue 1975a).
** Plagiochila undata Sull., Amer. J. Sci. Arts (ser. 2) 1 (1): 73, 1846 (Gray 1846).
** Plagiochila undata subsp. crispata (Gottsche) R.M.Schust., Amer. Midl. Naturalist
63 (1): 122, 1960 (Schuster 1960c). Bas.: Plagiochila crispata Gottsche, Mexik.
Leverm.: 71, 1863 (Gottsche 1863).
238
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Plagiochila ungarangana Sande Lac., Syn. hepat. jav.: 10, 1856 [1857] (Sande Lacoste 1856b).
*** Plagiochila virginica A.Evans, Prelim. cat. l. W. Virginia: 497, 1892 (Evans 1892a).
** Plagiochila virginica var. caroliniana R.M.Schust., Amer. Midl. Naturalist 63 (1):
15, 1960 (Schuster 1960c).
** Plagiochila virginica var. euryphylla R.M.Schust., Amer. Midl. Naturalist 63 (1):
21, 1960 (Schuster 1960c).
*** Plagiochila wightii Nees, Sp. Hepat. (Lindenberg) 2-4: 43, 1840 (Lindenberg 1840).
** Plagiochila wilhelmina Inoue, J. Hattori Bot. Lab. 33: 317, 1970 (Inoue 1970a).
sect. Zanteniae (Inoue) Inoue, Gen. Plagiochila SE Asia: 45, 1984 (Inoue
1984b). Bas.: Plagiochila subsect. Zanteniae Inoue, J. Hattori Bot. Lab. 32: 109,
1969 (Inoue 1969).
** Plagiochila zantenii Inoue, J. Hattori Bot. Lab. 32: 107, 1969 (Inoue 1969).
*
*
**
**
*
*
**
*
*
**
**
*
*
**
**
**
Incertae sedis
Plagiochila abscendens Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 104, 1864 (Gottsche
1864).
Plagiochila aculeata (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat.
5: 627, 1847 (Gottsche et al. 1847). Bas.: Jungermannia aculeata Hook.f. et Taylor, London J. Bot. 3: 578, 1844 (Hooker and Taylor 1844c).
Plagiochila acuta Steph., Bull. Herb. Boissier (sér. 2) 3 (7): 607 (353), 1903
(Stephani 1903e).
Plagiochila albertii Steph., Biblioth. Bot. 87 (2): 188, 1916 (Stephani 1916a).
Plagiochila aliena Gottsche, Mexik. Leverm.: 22, 1863 (Gottsche 1863). 170
Plagiochila allionii Steph., Sp. Hepat. (Stephani) 6: 120, 1917 (Stephani 1917a).
Plagiochila ambigua Lindenb. et Hampe, Linnaea 24 (6): 640, 1851 [1852]
(Hampe 1851a).
Plagiochila ampliata Steph., Biblioth. Bot. 87 (2): 189, 1916 (Stephani 1916a).
Plagiochila andicola Mont. et Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 6: 187, 1856
(Montagne 1856a).
Plagiochila andina Steph., Sp. Hepat. (Stephani) 6: 121, 1917 (Stephani 1917a).
Plagiochila angolensis Steph., Sp. Hepat. (Stephani) 6: 122, 1917 (Stephani 1917a).
Plagiochila angusteoblonga Steph., Biblioth. Bot. 87 (2): 189, 1916 (Stephani 1916a).
Plagiochila angustisedens Steph., Bull. Herb. Boissier (sér. 2) 5 (8): 743 (542), 1905
(Stephani 1905i).
Plagiochila angustispina Steph., Bull. Herb. Boissier (sér. 2) 2 (8): 671 (208), 1902
(Stephani 1902a).
Plagiochila apicalis Gottsche, Mexik. Leverm.: 29, 1863 (Gottsche 1863).
170 Plagiochila aliena is conspeciic with Plagiochila ludoviciana in Schuster (1960c), but others (e.g.
Fulford and Sharp 1990, Heinrichs et al. 1998, Dauphin 2005) recognize it.
World checklist of hornworts and liverworts
239
** Plagiochila appalachiana Inoue, J. Hattori Bot. Lab. 40: 415, 1976 (Inoue 1976b).
Nom. nov. pro Plagiochila yokogurensis subsp. fragilifolia R.M.Schust., J. Hattori
Bot. Lab. 18: 18, 1957 (Schuster 1957c).
* Plagiochila arcta S.Winkl., Rev. Bryol. Lichénol. 42 (3): 818, 1976 (Winkler 1976).
** Plagiochila arcuata Lindenb., Sp. Hepat. (Lindenberg) 2-4: 91, 1840 (Lindenberg
1840).
** Plagiochila arenacea Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl., Denkschr. 111: 53, 1964 (Schifner and Arnell 1964).
* Plagiochila artsii Pócs, J. Hattori Bot. Lab. 100: 334, 2006 (Pócs 2006a).
** Plagiochila atrovirens Taylor, London J. Bot. 5: 266, 1846 (Taylor 1846a).
** Plagiochila balansae Gottsche, Contr. Étude Plagiochila: 149, 1928 (Dugas 1928).
** Plagiochila baldwinii Austin, Trans. Connecticut Acad. Arts 8 (15): 257, 1891
(Evans 1891).
* Plagiochila bamingensis Steph., Bull. Mus. Natl. Hist. Nat. 18 (2): 117, 1912
(Corbière 1912).
** Plagiochila bancroftii Steph., Sp. Hepat. (Stephani) 6: 127, 1917 (Stephani 1917a).
*** Plagiochila barbadensis Steph., Bull. Herb. Boissier (sér. 2) 5 (9): 897 (563), 1905
(Stephani 1905g).
* Plagiochila barbata Steph., Biblioth. Bot. 87 (2): 190, 1916 (Stephani 1916a).
* Plagiochila barbeyi Steph., Biblioth. Bot. 87 (2): 190, 1916 (Stephani 1916a).
* Plagiochila batava Steph., Sp. Hepat. (Stephani) 6: 128, 1917 (Stephani 1917a).
* Plagiochila berggrenii Steph., Biblioth. Bot. 87 (2): 191, 1916 (Stephani 1916a).
** Plagiochila beskeana Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 863 (232), 1902
(Stephani 1902g).
** Plagiochila bialata Mitt., Fl. vit.: 407, 1871 [1873] (Mitten 1871).
* Plagiochila biapiculata Steph., Bull. Herb. Boissier (sér. 2) 5 (9): 891 (557), 1905
(Stephani 1905g).
* Plagiochila bicaudata Steph., Sp. Hepat. (Stephani) 6: 130, 1918 (Stephani 1918).
** Plagiochila biciliata Steph., Bull. Herb. Boissier (sér. 2) 3 (6): 529 (333), 1903
(Stephani 1903d).
** Plagiochila bicornis Hampe et Gottsche, Linnaea 25 (3): 338, 1852 [1853] (Hampe and Gottsche 1852).
* Plagiochila bidentula Steph., Sp. Hepat. (Stephani) 6: 130, 1918 (Stephani 1918).
** Plagiochila binghamiae A.Evans, Trans. Connecticut Acad. Arts 18 (5): 304, 1914
(Evans 1914c).
** Plagiochila binominata Steph., Sp. Hepat. (Stephani) 6: 131, 1918 (Stephani 1918).
** Plagiochila bitexta Dugas, Contr. Étude Plagiochila: 58, 1928 (Dugas 1928).
** Plagiochila blepharobasis Herzog, Hedwigia 72 (6): 216, 1932 (Herzog 1932c).
** Plagiochila bogotensis Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 98, 1864 (Gottsche 1864).
* Plagiochila boliviana Spruce, Mem. Torrey Bot. Club 1 (3): 137, 1890 (Spruce 1890).
* Plagiochila borneensis Steph., Sp. Hepat. (Stephani) 6: 132, 1918 (Stephani 1918). 171
171 Plagiochila borneensis is a doubtful taxon. No specimen could be found by Inoue (1984b) or So and
Grolle (2000a).
240
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Plagiochila brassii Inoue et Grolle, J. Hattori Bot. Lab. 36: 492, 1972 [1973] (Inoue
1972b).
* Plagiochila brevicalycina Lindenb. et Gottsche, Linnaea 20 (3): 322, 1847 (Hampe
1847).
** Plagiochila brunneola Steph., Bull. Herb. Boissier (sér. 2) 4 (2): 164 (416), 1904
(Stephani 1904f).
*** Plagiochila bryhnii Steph., Biblioth. Bot. 87 (2): 192, 1916 (Stephani 1916a).
** Plagiochila bunburii Taylor, London J. Bot. 5: 269, 1846 (Taylor 1846a).
* Plagiochila byssacea Hampe, Linnaea 20 (3): 326, 1847 (Hampe 1847).
** Plagiochila caducidentata R.M.Schust., Phytologia 39 (4): 247, 1978 (Schuster 1978a).
** Plagiochila caldana Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 879 (248), 1902
(Stephani 1902g).
** Plagiochila callaensis Steph., Sp. Hepat. (Stephani) 6: 136, 1918 (Stephani 1918).
* Plagiochila camusii Steph., Biblioth. Bot. 87 (2): 193, 1916 (Stephani 1916a).
* Plagiochila capillicaulis Steph., Biblioth. Bot. 87 (2): 193, 1916 (Stephani 1916a).
** Plagiochila caribbeania R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 412, 1980
(Schuster 1980c).
* Plagiochila cava Steph., Bull. Herb. Boissier (sér. 2) 4 (12): 1213 (501), 1904 (Stephani 1904h).
** Plagiochila ceylanica Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 98, 1860 [1861] (Mitten
1860c).
** Plagiochila chacoënsis Herzog, Feddes Repert. Spec. Nov. Regni Veg. 55 (1): 7,
1952 (Herzog 1952h).
** Plagiochila chiloënsis Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26 (III,
6): 27, 1900 (Stephani 1900b).
** Plagiochila chinantlana Gottsche, Mexik. Leverm.: 12, 1863 (Gottsche 1863).
* Plagiochila chiovendae Gola, Ann. Bot. (Rome) 13 (1): 67, 1914 (Gola 1914a).
* Plagiochila cinchonae Steph., Bull. Herb. Boissier (sér. 2) 5 (10): 920 (570), 1905
(Stephani 1905f).
* Plagiochila circumvoluta Gerola, Lav. Bot. Ist. Bot. Univ. Padova 12: 475, 1947
(Gerola 1947).
*** Plagiochila cleeii Inoue, Stud. Cryptog. S. Peru: 95, 1987 (Inoue 1987c).
* Plagiochila concava Nees, Sp. Hepat. (Lindenberg) 2-4: 70, 1840 (Lindenberg 1840).
** Plagiochila conduplicata Steph., Sp. Hepat. (Stephani) 6: 139, 1918 (Stephani 1918).
* Plagiochila conferta Steph., Sp. Hepat. (Stephani) 6: 139, 1918 (Stephani 1918).
** Plagiochila confertissima Steph., Bull. Herb. Boissier (sér. 2) 5 (2): 182 (510), 1905
(Stephani 1905d).
** Plagiochila connata Lindenb. et Gottsche, Syn. Hepat. 5: 645, 1847 (Gottsche et
al. 1847).
** Plagiochila contorta Lindenb. et Hampe, Linnaea 24 (3): 301, 1851 [1852] (Hampe 1851b).
** Plagiochila convoluta Steph., Sp. Hepat. (Stephani) 6: 141, 1918 (Stephani 1918).
** Plagiochila convolutifolia Steph., Sp. Hepat. (Stephani) 6: 142, 1918 (Stephani 1918).
World checklist of hornworts and liverworts
241
*** Plagiochila corymbulosa Pearson, Forh. Vidensk.-Selsk. Kristiania 1887 (9): 14,
1887 (Pearson 1887b).
** Plagiochila costariensis Horik., Acta Phytotax. Geobot. 13: 213, 1943 (Horikawa
1943). Nom. nov. pro Plagiochila pinnata Steph., Bull. Herb. Boissier (sér. 2) 5 (8):
749 (548), 1905 (Stephani 1905i), nom. illeg.
** Plagiochila crispabilis Lindenb., Sp. Hepat. (Lindenberg) 1: 15, 1839 (Lindenberg
1839).
* Plagiochila crispabilis var. minima Lindenb., Sp. Hepat. (Lindenberg) 5: 155, 1843
(Lindenberg 1843).
** Plagiochila cristophylla Steph., Bull. Herb. Boissier (sér. 2) 3 (2): 117 (297), 1903
(Stephani 1903b).
*** Plagiochila cuatrecasii H.Rob., Bryologist 70 (1): 47, 1967 (Robinson 1967).
** Plagiochila cubensis Steph., Sp. Hepat. (Stephani) 6: 143, 1918 (Stephani 1918).
** Plagiochila cucullata Lindenb. et Gottsche, Syn. Hepat. 5: 642, 1847 (Gottsche et
al. 1847).
* Plagiochila cuervina Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 96, 1864 (Gottsche 1864).
** Plagiochila delapsa Inoue, Beih. Nova Hedwigia 90: 171, 1988 (Inoue 1988a).
* Plagiochila delognei Steph., Biblioth. Bot. 87 (2): 196, 1916 (Stephani 1916a).
** Plagiochila denigrata Inoue, Willdenowia 18 (2): 557, 1989 (Inoue 1989b).
** Plagiochila densa Herzog, Hedwigia 72 (6): 222, 1932 (Herzog 1932c).
** Plagiochila densilora Herzog, Hedwigia 72 (6): 226, 1932 (Herzog 1932c).
** Plagiochila densiramosa Steph., Biblioth. Bot. 87 (2): 196, 1916 (Stephani 1916a).
** Plagiochila deppeana Steph., Bull. Herb. Boissier (sér. 2) 5 (9): 886 (552), 1905
(Stephani 1905g).
** Plagiochila desciscens Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 867 (236), 1902
(Stephani 1902g).
* Plagiochila dilatata Steph., Bull. Herb. Boissier (sér. 2) 5 (9): 887 (553), 1905
(Stephani 1905g).
* Plagiochila distans Colenso, Trans. & Proc. New Zealand Inst. 19: 283, 1887 (Colenso 1887). 172
** Plagiochila divaricata Lindenb., Sp. Hepat. (Lindenberg) 5: 147, 1843 (Lindenberg 1843).
* Plagiochila doerleri Steph., Biblioth. Bot. 87 (2): 196, 1916 (Stephani 1916a).
** Plagiochila dolichoblasta Herzog, Trans. Brit. Bryol. Soc. 1 (4): 286, 1950 (Herzog
1950a).
** Plagiochila dussiana Steph., Symb. Antill. (Urban) 3 (2): 277, 1902 (Stephani 1902e).
** Plagiochila echinata R.M.Schust., Amer. Midl. Naturalist 62 (2): 341, 1959
(Schuster 1959b).
** Plagiochila echinella Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 8: 332, 1857 (Gottsche 1857).
* Plagiochila ecuadorensis Steph., Sp. Hepat. (Stephani) 6: 149, 1918 (Stephani 1918).
172 Plagiochila distans is a doubtful taxon. he type specimen could not be found (Hodgson 1944,
Hamlin 1972).
242
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
* Plagiochila efuseintricata Steph., Biblioth. Bot. 87 (2): 197, 1916 (Stephani 1916a).
** Plagiochila eggersii Inoue, Bull. Natl. Sci. Mus. Tokyo, B 14 (4): 138, 1988 (Inoue
1988b).
** Plagiochila ekmanii S.W.Arnell, Bryologist 59 (4): 274, 1956 (Arnell 1956c).
* Plagiochila electa Steph., Bull. Herb. Boissier (sér. 2) 5 (8): 739 (538), 1905 (Stephani 1905i).
* Plagiochila elegantula Herzog, Feddes Repert. Spec. Nov. Regni Veg. 57 (1/2):
161, 1955 (Herzog 1955).
* Plagiochila emarginatobidentula Steph., Biblioth. Bot. 87 (2): 197, 1916 (Stephani
1916a).
* Plagiochila erythraeae Herzog, Hedwigia 78 (3/4): 224, 1938 (Herzog 1938d).
** Plagiochila estrellensis Herzog, Repert. Spec. Nov. Regni Veg. 21 (1/7): 23, 1925
(Herzog 1925a).
** Plagiochila eurydictyon Herzog, Hedwigia 72 (6): 207, 1932 (Herzog 1932c).
* Plagiochila excisa Steph., Sp. Hepat. (Stephani) 6: 153, 1918 (Stephani 1918).
** Plagiochila exesa Lindenb. et Gottsche, Syn. Hepat. 5: 629, 1847 (Gottsche et al. 1847).
* Plagiochila exilis Colenso, Trans. & Proc. New Zealand Inst. 19: 282, 1887 (Colenso 1887). 173
** Plagiochila expansa Gottsche, Mexik. Leverm.: 18, 1863 (Gottsche 1863).
** Plagiochila facallonia Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 871 (240), 1902
(Stephani 1902g).
* Plagiochila falcato-oblonga Steph., Biblioth. Bot. 87 (2): 198, 1916 (Stephani 1916a).
** Plagiochila fallax Lindenb. et Hampe, Linnaea 24 (3): 300, 1851 [1852] (Hampe
1851b).
** Plagiochila faxinensis Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl., Denkschr. 111: 60, 1964 (Schifner and Arnell 1964).
** Plagiochila fendleri Mont., Ann. Sci. Nat. Bot. (sér. 4) 6: 198, 1856 (Montagne 1856a).
* Plagiochila ilicicola Steph., Bull. Herb. Boissier (sér. 2) 4 (4): 351 (427), 1904
(Stephani 1904e).
** Plagiochila issidentoides Taylor, London J. Bot. 5: 264, 1846 (Taylor 1846a).
* Plagiochila labellifrons Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 488, 1885
(Spruce 1885).
** Plagiochila lava Steph., Sp. Hepat. (Stephani) 6: 156, 1918 (Stephani 1918).
** Plagiochila lavescens (Gottsche, Lindenb. et Nees) Gottsche, Mexik. Leverm.: 52,
1863 (Gottsche 1863). Bas.: Plagiochila guilleminiana β lavescens Gottsche, Lindenb. et Nees, Syn. Hepat. 5: 644, 1847 (Gottsche et al. 1847).
* Plagiochila lavorufescens Steph., Biblioth. Bot. 87 (2): 199, 1916 (Stephani 1916a).
** Plagiochila footei A.Evans, Trans. Connecticut Acad. Arts 18 (5): 306, 1914 (Evans
1914c).
* Plagiochila formosa Nees, Contr. Étude Plagiochila: 147, 1928 (Dugas 1928).
173 Plagiochila exilis is a doubtful taxon. he type specimen could not be found (Hodgson 1944, Hamlin
1972).
World checklist of hornworts and liverworts
243
** Plagiochila fragmentata R.M.Schust., Phytologia 45 (5): 421, 1980 (Schuster 1980b).
* Plagiochila frausa Gottsche, Mexik. Leverm.: 66, 1863 (Gottsche 1863).
* Plagiochila frausa var. boliviana Spruce, Mem. Torrey Bot. Club 1 (3): 134, 1890
(Spruce 1890).
** Plagiochila frayjorgensis Hässel, Bol. Soc. Argent. Bot. 22 (1/4): 103, 1983 (Hässel
1983). Nom. nov. pro Plagiochila modesta Herzog, Rev. Bryol. Lichénol. 23 (1/2):
34, 1954 (Herzog 1954), nom. illeg.
* Plagiochila gafatensis Gottsche ex Schweinf., Beitr. Fl. Aethiop.: 227, 1866 [1867]
(Schweinfurth 1866).
** Plagiochila gaudichaudii Mont. et Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 6: 193,
1856 (Montagne 1856a).
** Plagiochila geniculata Lindenb., Sp. Hepat. (Lindenberg) 5: 131, 1843 (Lindenberg 1843).
* Plagiochila geppii Steph., Biblioth. Bot. 87 (2): 199, 1916 (Stephani 1916a).
** Plagiochila germana Gottsche, Mexik. Leverm.: 34, 1863 (Gottsche 1863).
** Plagiochila germanii Steph., Bull. Herb. Boissier (sér. 2) 5 (10): 938 (588), 1905
(Stephani 1905f).
* Plagiochila gibbilora Steph., Bull. Herb. Boissier (sér. 2) 4 (6): 590 (441), 1904
(Stephani 1904b).
** Plagiochila gittinsii Inoue, Bull. Natl. Sci. Mus. Tokyo, B 3 (4): 139, 1977 (Inoue
1977a).
* Plagiochila glauca Carl, Ann. Bryol., Suppl. 2: 129, 1931 (Carl 1931b). 174
* Plagiochila gracilicaulis Spruce, Mem. Torrey Bot. Club 1 (3): 132, 1890 (Spruce
1890).
** Plagiochila gracillima Austin, Trans. Connecticut Acad. Arts 8 (15): 256, 1891
(Evans 1891).
* Plagiochila granatensis Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 111, 1864
(Gottsche 1864).
** Plagiochila granditexta Steph., Bull. Herb. Boissier (sér. 2) 4 (2): 165 (417), 1904
(Stephani 1904f).
** Plagiochila grateloupii Mont., Ann. Sci. Nat. Bot. (sér. 4) 6: 188, 1856 (Montagne
1856a).
** Plagiochila guatemalensis Steph., Sp. Hepat. (Stephani) 6: 163, 1918 (Stephani 1918).
** Plagiochila guttisquama Inoue et Grolle, Bull. Natl. Sci. Mus. Tokyo, B 5 (1): 29,
1979 (Inoue 1979c).
** Plagiochila gymnocalyx Inoue, Trop. Bryol. 1: 34, 1989 (Gradstein and Florschützde Waard 1989).
** Plagiochila haeseliae Inoue, Stud. Cryptog. S. Chile: 97, 1984 (Inoue 1984a).
* Plagiochila hans-meyeri Steph., Sp. Hepat. (Stephani) 6: 164, 1918 (Stephani 1918).
** Plagiochila harlingii S.W.Arnell, Svensk Bot. Tidskr. 56 (2): 346, 1962 (Arnell
1962b).
174 Plagiochila glauca is a doubtful taxon. Hässel and Rubies (2009) could not ind any type material.
244
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Plagiochila haumanii Herzog, Repert. Spec. Nov. Regni Veg. 41 (14/25): 285,
1937 (Herzog 1937).
** Plagiochila hawaica Steph., Bull. Herb. Boissier (sér. 2) 3 (7): 598 (344), 1903
(Stephani 1903e).
** Plagiochila heteracantha Steph., Sp. Hepat. (Stephani) 6: 166, 1918 (Stephani 1918).
* Plagiochila heterofolia Steph., Biblioth. Bot. 87 (2): 200, 1916 (Stephani 1916a).
* Plagiochila hieronymii Steph., Biblioth. Bot. 87 (2): 201, 1916 (Stephani 1916a).
** Plagiochila hiroshiana Pócs, J. Hattori Bot. Lab. 100: 335, 2006 (Pócs 2006a).
** Plagiochila hoehnei Herzog, Hedwigia 72 (6): 220, 1932 (Herzog 1932c).
* Plagiochila holstii Steph., Hedwigia 45 (4): 214, 1906 (Stephani 1906b). Nom.
nov. pro Plagiochila prostrata Steph., Bull. Herb. Boissier (sér. 2) 4 (2): 168 (420),
1904 (Stephani 1904f), nom. illeg.
** Plagiochila horrida Gottsche, Mexik. Leverm.: 74, 1863 (Gottsche 1863).
* Plagiochila huatuscana Gottsche, Mexik. Leverm.: 24, 1863 (Gottsche 1863).
** Plagiochila huerlimannii Inoue, J. Hattori Bot. Lab. 33: 307, 1970 (Inoue 1970b).
* Plagiochila humboldtiana Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 112, 1864
(Gottsche 1864).
** Plagiochila hyalina Lindenb., Syn. Hepat. 5: 640, 1847 (Gottsche et al. 1847).
* Plagiochila incisa Dugas, Contr. Étude Plagiochila: 112, 1928 (Dugas 1928).
** Plagiochila inlata Steph., Bull. Herb. Boissier (sér. 2) 3 (11): 961 (376), 1903
(Stephani 1903a).
* Plagiochila informifolia Steph., Biblioth. Bot. 87 (2): 202, 1916 (Stephani 1916a).
** Plagiochila infuscata Mitt., J. Linn. Soc., Bot. 15 (82): 63, 1876 (Mitten 1876a).
* Plagiochila injasutiensis S.W.Arnell, Bot. Not. 110: 404, 1957 (Arnell 1957d).
** Plagiochila inouei Grolle, J. Bryol. 10 (3): 269, 1979 (Grolle 1979b). Nom. nov.
pro Plagiochila nudiuscula Inoue, Bull. Natl. Sci. Mus. Tokyo, B 3 (2): 45, 1977
(Inoue 1977c), nom. illeg.
** Plagiochila insularia Mitt., St. Helena: 366, 1875 (Mitten 1875).
** Plagiochila intertexta Hook.f. et Taylor, London J. Bot. 5: 267, 1846 (Taylor 1846a).
* Plagiochila inuensis Steph., Biblioth. Bot. 87 (2): 202, 1916 (Stephani 1916a).
** Plagiochila irregularis Gottsche, Mexik. Leverm.: 14, 1863 (Gottsche 1863).
** Plagiochila itatiajensis Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 874 (243), 1902
(Stephani 1902g).
** Plagiochila jamaicensis Lindenb. et Hampe, Linnaea 24 (3): 302, 1851 [1852]
(Hampe 1851b).
** Plagiochila jaramillii H.Rob., Bryologist 70 (1): 48, 1967 (Robinson 1967).
* Plagiochila karstenii Steph., Hedwigia 45 (4): 214, 1906 (Stephani 1906b). Nom.
nov. pro Plagiochila patentispina Steph., Bull. Herb. Boissier (sér. 2) 4 (1): 28
(400), 1904 (Stephani 1904g), nom. illeg. 175
* Plagiochila kaulfussiana Steph., Sp. Hepat. (Stephani) 6: 172, 1918 (Stephani 1918).
** Plagiochila keckiana Steph., Bull. Herb. Boissier (sér. 2) 5 (4): 358 (526), 1905
(Stephani 1905j).
175 Plagiochila karstenii is possibly conspeciic with Plagiochila bantamensis (So and Grolle 2000a).
World checklist of hornworts and liverworts
245
** Plagiochila kerneriana S.W.Arnell, Österr. Akad. Wiss., Math.-Naturwiss. Kl.,
Denkschr. 111: 60, 1964 (Schifner and Arnell 1964).
** Plagiochila koponenii Inoue et Piippo, Ann. Bot. Fenn. 26 (2): 216, 1989 (Piippo
1989b).
* Plagiochila lacerifolia Steph., Biblioth. Bot. 87 (2): 202, 1916 (Stephani 1916a).
* Plagiochila lachenaudii Steph., Biblioth. Bot. 87 (2): 202, 1916 (Stephani 1916a).
* Plagiochila laciniosa Dugas, Contr. Étude Plagiochila: 68, 1928 (Dugas 1928).
* Plagiochila laevifolia Gola, Ann. Bot. (Rome) 6 (2): 273, 1907 (Gola 1907).
** Plagiochila lansbergii Gottsche, Mexik. Leverm.: 13, 1863 (Gottsche 1863).
** Plagiochila latitrigona Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl., Denkschr. 111: 48, 1964 (Schifner and Arnell 1964).
* Plagiochila laxiramea Steph., Biblioth. Bot. 87 (2): 203, 1916 (Stephani 1916a).
* Plagiochila lecontei Steph., Bull. Herb. Boissier (sér. 2) 4 (2): 165 (417), 1904
(Stephani 1904f). 176
* Plagiochila ledermanniana Beauverd, Sp. Hepat. (Stephani) 6: 572, 1924 (Stephani 1924). Nom. nov. pro Plagiochila cucullifolia Steph., Sp. Hepat. (Stephani) 6:
243, 1922 (Stephani 1922), nom. illeg. 177
* Plagiochila ledieui Steph., Bull. Herb. Boissier (sér. 2) 4 (2): 161 (413), 1904
(Stephani 1904f). 178
** Plagiochila lehmanniana S.W.Arnell, Svensk Bot. Tidskr. 55 (1): 205, 1961 (Arnell
1961).
** Plagiochila liebmanniana Lehm. et Lindenb., Sp. Hepat. (Lindenberg) 2-4: 97,
1840 (Lindenberg 1840).
* Plagiochila lignicola Spruce, Mem. Torrey Bot. Club 1 (3): 135, 1890 (Spruce 1890).
* Plagiochila lindaui Steph., Biblioth. Bot. 87 (2): 204, 1916 (Stephani 1916a).
* Plagiochila lindigiana Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 105, 1864 (Gottsche
1864).
** Plagiochila lingua Steph., Bull. Herb. Boissier (sér. 2) 2 (8): 677 (214), 1902
(Stephani 1902a).
* Plagiochila longiissa Steph., Bull. Herb. Boissier (sér. 2) 5 (9): 891 (557), 1905
(Stephani 1905g).
* Plagiochila lotsyana Steph., Biblioth. Bot. 87 (2): 205, 1916 (Stephani 1916a).
* Plagiochila luteola Steph., Bull. Herb. Boissier (sér. 2) 5 (2): 175 (503), 1905
(Stephani 1905d).
** Plagiochila lutescens Steph., Sp. Hepat. (Stephani) 6: 179, 1921 (Stephani 1921).
** Plagiochila luzonensis Grolle et M.L.So, Bryologist 102 (1): 69, 1999 (Grolle and
So 1999a).
176 Plagiochila lecontei is allied to and perhaps conspeciic with Plagiochila ledieui and Plagiochila multilora
(Jones 1962).
177 Plagiochila ledermanniana is a doubtful taxon. he type specimen was burned in B (Piippo 1989b).
178 Plagiochila ledieui is allied to and perhaps conspeciic with Plagiochila lecontei and Plagiochila multilora
(Jones 1962).
246
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
* Plagiochila macra Taylor, London J. Bot. 7: 198, 1848 (Taylor 1848a).
** Plagiochila macrifolia Taylor, London J. Bot. 5: 270, 1846 (Taylor 1846a).
* Plagiochila macrifolia var. angustifolia Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 470, 1885 (Spruce 1885).
*** Plagiochila martiana (Nees) Lindenb., Sp. Hepat. (Lindenberg) 1: 12, 1839 (Lindenberg 1839). Bas.: Jungermannia martiana Nees, Linnaea 6 (4): 617, 1831
(Nees 1831).
* Plagiochila matanga Steph., Sp. Hepat. (Stephani) 6: 183, 1921 (Stephani 1921).
** Plagiochila maunakeana Steph., Sp. Hepat. (Stephani) 6: 186, 1921 (Stephani 1921).
** Plagiochila maximiliana Gottsche, Mexik. Leverm.: 18, 1863 (Gottsche 1863).
** Plagiochila meghalayensis K.K.Rawat et S.C.Srivast., Geophytology 35 (1/2): 49,
2005 (Rawat and Srivastava 2005).
** Plagiochila meridana Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 8: 328, 1857 (Gottsche
1857).
** Plagiochila microdonta Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 97, 1860 [1861]
(Mitten 1860c).
* Plagiochila mildbreadiana Beauverd, Sp. Hepat. (Stephani) 6: 571, 1924 (Stephani
1924). Nom. nov. pro Plagiochila andongensis Steph., Sp. Hepat. (Stephani) 6: 122,
1917 (Stephani 1917a), nom. illeg.
** Plagiochila minarum Herzog, Hedwigia 72 (6): 207, 1932 (Herzog 1932c).
** Plagiochila minutiretis Reimers, Repert. Spec. Nov. Regni Veg. 21 (8/20): 264,
1925 (Reimers 1925).
** Plagiochila miqueliana Lehm. et Lindenb., Sp. Hepat. (Lindenberg) 2-4: 95, 1840
(Lindenberg 1840).
** Plagiochila molliuscula Inoue, Ann. Bot. Fenn. 13 (3): 134, 1976 (Engel 1976a).
** Plagiochila mollusca Lehm., Nov. Stirp. Pug. 10: 4, 1857 (Lehmann 1857).
** Plagiochila moniliformis R.M.Schust., Phytologia 39 (4): 247, 1978 (Schuster 1978a).
** Plagiochila muelleriana Gottsche, Mexik. Leverm.: 38, 1863 (Gottsche 1863).
* Plagiochila multilora Steph., Planzenw. Ost-Afrikas C: 64, 1895 (Stephani
1895d). 179
** Plagiochila mutila Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 8: 327, 1857 (Gottsche 1857).
* Plagiochila naranjoënsis Steph., Bull. Herb. Boissier (sér. 2) 2 (8): 687 (224), 1902
(Stephani 1902a).
** Plagiochila neesiana Lindenb., Sp. Hepat. (Lindenberg) 2-4: 71, 1840 (Lindenberg
1840).
* Plagiochila neglecta Steph., Bull. Herb. Boissier (sér. 2) 5 (4): 351 (519), 1905
(Stephani 1905j).
* Plagiochila negrensis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 466, 1885
(Spruce 1885).
* Plagiochila negrii Gola, Ann. Bot. (Rome) 13 (1): 66, 1914 (Gola 1914a).
179 Plagiochila multilora is allied to and perhaps conspeciic with Plagiochila lecontei and Plagiochila
ledieui (Jones 1962).
World checklist of hornworts and liverworts
*
*
*
**
**
*
**
**
**
*
*
*
**
**
**
*
**
*
*
**
**
**
**
*
**
247
Plagiochila nigricaulis Steph., Biblioth. Bot. 87 (2): 206, 1916 (Stephani 1916a).
Plagiochila notha Steph., Bull. Herb. Boissier (sér. 2) 5 (8): 737 (536), 1905
(Stephani 1905i).
Plagiochila nova Steph., Sp. Hepat. (Stephani) 6: 189, 1921 (Stephani 1921).
Plagiochila nudifolia (Steph.) Grolle, Feddes Repert. 82 (1): 89, 1971 (Grolle
1971b). Bas.: Jamesoniella nudifolia Steph., Biblioth. Bot. 87 (2): 184, 1916
(Stephani 1916a).
Plagiochila nutans Steph., Bull. Herb. Boissier (sér. 2) 3 (11): 960 (375), 1903
(Stephani 1903a).
Plagiochila obliquetruncata Steph., Biblioth. Bot. 87 (2): 207, 1916 (Stephani 1916a).
Plagiochila oblonga Inoue, Bull. Natl. Sci. Mus. Tokyo (n.ser.) 8 (3): 398, 1965
(Inoue 1965a).
Plagiochila oerstediana Lindenb. et Hampe, Linnaea 24 (3): 301, 1851 [1852]
(Hampe 1851b).
Plagiochila olivacea Steph., Bull. Herb. Boissier (sér. 2) 5 (2): 190 (518), 1905
(Stephani 1905d).
Plagiochila ornata Wilson ex Lindenb., Sp. Hepat. (Lindenberg) 5: 164, 1843
(Lindenberg 1843).
Plagiochila ovato-obconica Steph., Sp. Hepat. (Stephani) 6: 191, 1921 (Stephani 1921).
Plagiochila ovifolia Steph., Sp. Hepat. (Stephani) 6: 191, 1921 (Stephani 1921).
Plagiochila palangiensis S.C.Srivast., K.K.Rawat et P.K.Verma, Natl. Acad. Sci.
Lett. 29 (7/8): 267, 2006 (Srivastava et al. 2006).
Plagiochila panamensis Inoue, Bull. Natl. Sci. Mus. Tokyo, B 15 (3): 94, 1989
(Inoue 1989a).
Plagiochila paramicola Herzog, Beih. Bot. Centralbl. 61B (3): 563, 1942 (Herzog
1942d).
Plagiochila parvitexta Steph., Bull. Herb. Boissier (sér. 2) 2 (8): 674 (211), 1902
(Stephani 1902a).
Plagiochila parvivittata Inoue, Bull. Natl. Sci. Mus. Tokyo, B 13 (2): 50, 1987
(Inoue 1987b).
Plagiochila parvula Steph., Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr.
88: 726, 1913 (Stephani 1913b).
Plagiochila pastasensis Steph., Sp. Hepat. (Stephani) 6: 197, 1921 (Stephani 1921).
Plagiochila patentispina Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 877 (246),
1902 (Stephani 1902g).
Plagiochila patuloides Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl., Denkschr. 111: 58, 1964 (Schifner and Arnell 1964).
Plagiochila paucidentata Mont. et Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 6: 197,
1856 (Montagne 1856a).
Plagiochila paupercula Gottsche, Mexik. Leverm.: 22, 1863 (Gottsche 1863).
Plagiochila pearceana Steph., Bull. Herb. Boissier (sér. 2) 5 (8): 742 (541), 1905
(Stephani 1905i).
Plagiochila pellucida Lindenb. et Gottsche, Linnaea 20 (3): 321, 1847 (Hampe 1847).
248
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Plagiochila perrottetiana Mont. et Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 6: 195,
1856 (Montagne 1856a).
** Plagiochila perrottetiana var. minor Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 101,
1864 (Gottsche 1864).
** Plagiochila pilifera Steph., Sp. Hepat. (Stephani) 6: 198, 1921 (Stephani 1921).
** Plagiochila pinnatidens Steph., Sp. Hepat. (Stephani) 6: 199, 1921 (Stephani 1921).
** Plagiochila pittieri Steph., Bull. Herb. Boissier (sér. 2) 2 (8): 673 (210), 1902
(Stephani 1902a).
** Plagiochila platyphylla Herzog, Hedwigia 72 (6): 223, 1932 (Herzog 1932c).
** Plagiochila polopolensis Herzog, Hedwigia 67 (6): 261, 1927 (Herzog 1927).
* Plagiochila prostrata Steph., Bull. Herb. Boissier (sér. 2) 2 (8): 668 (205), 1902
(Stephani 1902a).
* Plagiochila pseudopatula Herzog, Hedwigia 72 (6): 227, 1932 (Herzog 1932c).
** Plagiochila pseudoradicans Herzog, Repert. Spec. Nov. Regni Veg. 21 (1/7): 24,
1925 (Herzog 1925a).
** Plagiochila ptychanthoidea Steph., Bull. Herb. Boissier (sér. 2) 3 (2): 121 (301),
1903 (Stephani 1903b).
** Plagiochila pulchella Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 871 (240), 1902
(Stephani 1902g).
** Plagiochila pulvinata Steph., Bull. Herb. Boissier (sér. 2) 3 (6): 526 (330), 1903
(Stephani 1903d).
** Plagiochila ratkowskiana Inoue, Brunonia 3 (1): 141, 1980 (Inoue 1980).
** Plagiochila regeliana Steph., Bull. Herb. Boissier (sér. 2) 2 (8): 675 (212), 1902
(Stephani 1902a).
** Plagiochila remyana Steph., Bull. Herb. Boissier (sér. 2) 3 (11): 963 (378), 1903
(Stephani 1903a).
*** Plagiochila revolvens Mitt., Hooker’s J. Bot. Kew Gard. Misc. 3: 358, 1851 (Mitten
1851).
** Plagiochila rigidula Lindenb. et Gottsche, Linnaea 20 (3): 323, 1847 (Hampe 1847).
** Plagiochila rosana Steph., Bull. Herb. Boissier (sér. 2) 5 (9): 892 (558), 1905
(Stephani 1905g).
* Plagiochila rubricaulis Steph., Bot. Jahrb. Syst. 20 (3): 311, 1895 (Stephani 1895a).
* Plagiochila ruifolia Steph., Biblioth. Bot. 87 (2): 209, 1916 (Stephani 1916a).
** Plagiochila rufoviridis Spruce, Mem. Torrey Bot. Club 1 (3): 136, 1890 (Spruce
1890).
* Plagiochila rusbyi Spruce, Mem. Torrey Bot. Club 1 (3): 135, 1890 (Spruce 1890).
* Plagiochila sabensis Steph., Sp. Hepat. (Stephani) 6: 214, 1921 (Stephani 1921).
** Plagiochila sachapatensis Steph., Bull. Herb. Boissier (sér. 2) 2 (8): 679 (216), 1902
(Stephani 1902a).
** Plagiochila salazariae Inoue, Bull. Natl. Sci. Mus. Tokyo, B 15 (3): 91, 1989 (Inoue
1989a).
** Plagiochila saltuensis Spruce ex Steph., Bull. Herb. Boissier (sér. 2) 5 (10): 927
(577), 1905 (Stephani 1905f).
World checklist of hornworts and liverworts
249
** Plagiochila saltuensis var. spinosissima Herzog, Feddes Repert. Spec. Nov. Regni
Veg. 57 (1/2): 163, 1955 (Herzog 1955).
* Plagiochila samoana Steph., Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr.
85: 197, 1910 (Stephani 1910a).
** Plagiochila sarmentosa (Lehm. et Lindenb.) Lindenb., Sp. Hepat. (Lindenberg)
2-4: 86, 1840 (Lindenberg 1840). Bas.: Jungermannia sarmentosa Lehm. et Lindenb., Linnaea 9 (4): 427, 1835 (Lehmann 1835).
* Plagiochila schifneri Steph., Biblioth. Bot. 87 (2): 209, 1916 (Stephani 1916a).
* Plagiochila schinzei Steph., Biblioth. Bot. 87 (2): 209, 1916 (Stephani 1916a).
* Plagiochila schmidtii Steph., Biblioth. Bot. 87 (2): 209, 1916 (Stephani 1916a).
* Plagiochila schraderbergii Steph., Sp. Hepat. (Stephani) 6: 244, 1922 (Stephani
1922). 180
** Plagiochila schubertiana Steph., Sp. Hepat. (Stephani) 6: 208, 1921 (Stephani 1921).
** Plagiochila schusteri Inoue et Grolle, J. Hattori Bot. Lab. 33: 326, 1970 (Inoue 1970a).
** Plagiochila scissifolia Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 865 (234), 1902
(Stephani 1902g).
* Plagiochila scotica Macvicar ex Steph., Sp. Hepat. (Stephani) 6: 209, 1921 (Stephani
1921). 181
** Plagiochila semiamplexicaulis Steph., Bull. Herb. Boissier (sér. 2) 5 (10): 936 (586),
1905 (Stephani 1905f).
** Plagiochila semiermis Dugas, Contr. Étude Plagiochila: 66, 1928 (Dugas 1928).
** Plagiochila silvatica Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 108, 1864 (Gottsche
1864).
* Plagiochila similis Steph., Biblioth. Bot. 87 (2): 210, 1916 (Stephani 1916a).
** Plagiochila sisparensis Steph., Sp. Hepat. (Stephani) 6: 207, 1921 (Stephani 1921).
* Plagiochila slateri Steph., Biblioth. Bot. 87 (2): 211, 1916 (Stephani 1916a).
** Plagiochila solitaria Gottsche, Sp. Hepat. (Stephani) 6: 225, 1921 (Stephani 1921).
*** Plagiochila spinifera Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 30 (5): 113,
1873 (Ångström 1873).
* Plagiochila sprucei Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 860 (229), 1902
(Stephani 1902g).
* Plagiochila staudtiana Steph., Bull. Herb. Boissier (sér. 2) 4 (4): 352 (428), 1904
(Stephani 1904e).
** Plagiochila stipata Steph., Sp. Hepat. (Stephani) 6: 209, 1921 (Stephani 1921).
* Plagiochila stolzii Steph., Sp. Hepat. (Stephani) 6: 244, 1922 (Stephani 1922).
** Plagiochila subcontigua Herzog, Hedwigia 72 (6): 230, 1932 (Herzog 1932c).
* Plagiochila subconvoluta Gottsche, Mexik. Leverm.: 24, 1863 (Gottsche 1863).
* Plagiochila subdenudata Steph., Bull. Herb. Boissier (sér. 2) 2 (8): 673 (210), 1902
(Stephani 1902a).
180 Plagiochila schraderbergii is a doubtful taxon. he type specimen was burned in B (Piippo 1989b).
181 Plagiochila scotica was described from Europe, but it has neither been recognized in any recent European study nor synonymized.
250
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Plagiochila subedentata Steph., Bull. Herb. Boissier (sér. 2) 2 (8): 679 (216), 1902
(Stephani 1902a).
** Plagiochila subfragilis Inoue, Stud. Cryptog. S. Peru: 98, 1987 (Inoue 1987c).
* Plagiochila subhyalina Steph., Biblioth. Bot. 87 (2): 212, 1916 (Stephani 1916a).
** Plagiochila subligulata Steph., Sp. Hepat. (Stephani) 6: 224, 1921 (Stephani 1921).
** Plagiochila sublyallii M.L.So, New Zealand J. Bot. 39 (1): 109, 2001 (So 2001b).
* Plagiochila subrara Herzog, Hedwigia 74 (2): 87, 1934 (Herzog 1934a).
* Plagiochila subrotundifolia Steph., Bull. Herb. Boissier (sér. 2) 5 (10): 935 (585),
1905 (Stephani 1905f).
** Plagiochila subundulata Lindenb., Sp. Hepat. (Lindenberg) 5: 137, 1843 (Lindenberg
1843).
* Plagiochila suringarii Steph., Sp. Hepat. (Stephani) 6: 208, 1921 (Stephani 1921).
** Plagiochila sylvicultrix Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 486, 1885
(Spruce 1885).
* Plagiochila symmetrica Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 876 (245), 1902
(Stephani 1902g).
** Plagiochila tabinana Gottsche, Mexik. Leverm.: 46, 1863 (Gottsche 1863).
** Plagiochila talinayi S.W.Arnell, Svensk Bot. Tidskr. 50 (2): 312, 1956 (Arnell
1956d).
** Plagiochila tambillensis Loitl., Diagn. pl. nov.: 22, 1894 (Szyszylowicz 1894).
** Plagiochila tarapotensis Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 863 (232), 1902
(Stephani 1902g).
* Plagiochila tenera Steph., Sp. Hepat. (Stephani) 6: 231, 1921 (Stephani 1921).
** Plagiochila tenuis Lindenb., Sp. Hepat. (Lindenberg) 2-4: 50, 1840 (Lindenberg
1840).
* Plagiochila tenuispica Steph., Sp. Hepat. (Stephani) 6: 234, 1921 (Stephani 1921).
** Plagiochila thamniopsis Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: cc, 1889
[1890] (Spruce 1889).
* Plagiochila thollonii Steph., Bull. Herb. Boissier (sér. 2) 2 (12): 980 (270), 1902
(Stephani 1902h).
** Plagiochila thrausta Inoue et Grolle, Bull. Natl. Sci. Mus. Tokyo, B 5 (1): 34, 1979
(Inoue 1979c).
* Plagiochila tonduzana Steph., Sp. Hepat. (Stephani) 6: 227, 1921 (Stephani 1921).
** Plagiochila tortuosa Lindenb. et Gottsche, Mexik. Leverm.: 70, 1863 (Gottsche 1863).
* Plagiochila tovarina Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 102, 1864 (Gottsche
1864).
* Plagiochila trabeculatospinosa Herzog, Hedwigia 72 (6): 213, 1932 (Herzog 1932c).
* Plagiochila trabutii Steph., Biblioth. Bot. 87 (2): 213, 1916 (Stephani 1916a).
** Plagiochila trianae Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 114, 1864 (Gottsche 1864).
** Plagiochila trichomanes Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: cc, 1889
[1890] (Spruce 1889).
* Plagiochila tricuspis Steph., Sp. Hepat. (Stephani) 6: 229, 1921 (Stephani 1921).
World checklist of hornworts and liverworts
251
** Plagiochila tristaniana Váňa et J.J.Engel, Mem. New York Bot. Gard. 105: 89, 2013
(Váňa and Engel 2013). Nom. nov. pro Plagiochila fragilifolia S.W.Arnell, Results
Norweg. Sci. Exped. Tristan da Cunha 42: 25, 1958 (Arnell 1958b), nom. illeg.
** Plagiochila tristis Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 880 (249), 1902 (Stephani 1902g).
** Plagiochila trollii Herzog, Hedwigia 74 (2): 90, 1934 (Herzog 1934a).
** Plagiochila truncata Gottsche, Mexik. Leverm.: 25, 1863 (Gottsche 1863).
** Plagiochila uleana Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 868 (237), 1902
(Stephani 1902g).
** Plagiochila umbrosioides L.Söderstr., Phytotaxa 208 (1): 85, 2015 (Söderström et
al. 2015b). Nom. nov. pro Plagiochila umbrosa Steph., Sp. Hepat. (Stephani) 6:
235, 1921 (Stephani 1921), nom. illeg.
* Plagiochila unduavensis Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 882 (251),
1902 (Stephani 1902g).
* Plagiochila usambarana Steph., Bull. Herb. Boissier (sér. 2) 2 (12): 980 (270),
1902 (Stephani 1902h).
** Plagiochila variedentata Steph., Bull. Herb. Boissier (sér. 2) 2 (10): 887 (256),
1902 (Stephani 1902g).
** Plagiochila vastifolia Steph., Bull. Herb. Boissier (sér. 2) 2 (8): 680 (217), 1902
(Stephani 1902a).
** Plagiochila velata Inoue et Piippo, Ann. Bot. Fenn. 26 (2): 207, 1989 (Piippo 1989b).
* Plagiochila venezuelana Steph., Bull. Herb. Boissier (sér. 2) 5 (8): 749 (548), 1905
(Stephani 1905i).
* Plagiochila ventricosotrigona Steph., Biblioth. Bot. 87 (2): 214, 1916 (Stephani 1916a).
* Plagiochila verrucosa Steph., Bull. Herb. Boissier (sér. 2) 5 (9): 885 (551), 1905
(Stephani 1905g).
* Plagiochila vetustisilva Steph., Sp. Hepat. (Stephani) 6: 245, 1922 (Stephani 1922). 182
* Plagiochila viminea Spruce, Mem. Torrey Bot. Club 1 (3): 134, 1890 (Spruce 1890).
* Plagiochila viridis Steph., Sp. Hepat. (Stephani) 6: 239, 1921 (Stephani 1921).
** Plagiochila viridonigra (E.A.Hodgs.) Inoue, Bryologist 68 (2): 218, 1965 (Inoue
1965b). Bas.: Syzygiella viridonigra E.A.Hodgs., Rec. Domin. Mus. 4 (11): 120,
1962 (Hodgson 1962a).
** Plagiochila vittiana Inoue, Beih. Nova Hedwigia 90: 171, 1988 (Inoue 1988a).
** Plagiochila vittifolia Steph., Sp. Hepat. (Stephani) 6: 239, 1921 (Stephani 1921).
** Plagiochila vulcanica Steph., Bull. Herb. Boissier (sér. 2) 2 (8): 671 (208), 1902
(Stephani 1902a).
** Plagiochila wacei S.W.Arnell ex Váňa et J.J.Engel, Mem. New York Bot. Gard.
105: 89, 2013 (Váňa and Engel 2013).
* Plagiochila wallisiana Steph., Bull. Herb. Boissier (sér. 2) 5 (10): 936 (586), 1905
(Stephani 1905f).
182 Plagiochila vetustisilva is a doubtful taxon. he type specimen was burned in B (Piippo 1989b).
252
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Plagiochila wattsiana J.J.Engel et G.L.Merr., Nova Hedwigia 91 (3/4): 511, 2010
(Engel and Smith Merrill 2010). Nom. nov. pro Plagiochila wattsii Steph., Sp.
Hepat. (Stephani) 6: 240, 1921 (Stephani 1921), nom. illeg.
** Plagiochila wettsteiniana S.W.Arnell, Österr. Akad. Wiss., Math.-Naturwiss. Kl.,
Denkschr. 111: 53, 1964 (Schifner and Arnell 1964).
* Plagiochila weymouthiana Steph., Biblioth. Bot. 87 (2): 215, 1916 (Stephani 1916a).
** Plagiochila wiemanniana S.W.Arnell, Österr. Akad. Wiss., Math.-Naturwiss. Kl.,
Denkschr. 111: 54, 1964 (Schifner and Arnell 1964).
*** Plagiochila winteri Steph., Bull. Herb. Boissier (sér. 2) 2 (12): 981 (271), 1902
(Stephani 1902h).
** Plagiochila wrightii Steph., Bull. Herb. Boissier (sér. 2) 2 (8): 681 (218), 1902
(Stephani 1902a).
** Plagiochila xalapensis Gottsche, Mexik. Leverm.: 21, 1863 (Gottsche 1863).
* Plagiochila xanthochroma Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 489,
1885 (Spruce 1885).
* Plagiochila yoshinagana Steph., Sp. Hepat. (Stephani) 6: 242, 1922 (Stephani 1922).
** Plagiochila zacuapana Gottsche, Mexik. Leverm.: 20, 1863 (Gottsche 1863).
** Plagiochilidium Herzog, Mitt. Inst. Allg. Bot. Hamburg 7 (3): 186, 1931 (Herzog
1931a).
** Plagiochilidium bidentulum (Steph.) Grolle, J. Hattori Bot. Lab. 65: 408, 1988
(Grolle 1988c). Bas.: Tylimanthus bidentulus Steph., Bull. Herb. Boissier (sér. 2) 5
(12): 1134 (6), 1905 (Stephani 1905b).
** Plagiochilion S.Hatt., Biosphaera 1 (1): 7, 1947 (Hattori 1947a).
*** Plagiochilion braunianum (Nees) S.Hatt., Biosphaera 1 (1): 7, 1947 (Hattori
1947a). Bas.: Jungermannia brauniana Nees, Enum. Pl. Crypt. Javae: 80, 1830
(Nees 1830).
** Plagiochilion combinatum (Mitt.) Inoue, J. Hattori Bot. Lab. 27: 55, 1964 (Inoue
1964b). Bas.: Plagiochila combinata Mitt., Fl. vit.: 408, 1871 [1873] (Mitten 1871).
*** Plagiochilion conjugatum (Hook.) R.M.Schust., J. Hattori Bot. Lab. 26: 285, 1963
(Schuster 1963b). Bas.: Jungermannia conjugata Hook., Musci Exot. 1: tab. 91,
1818 (Hooker 1818).
** Plagiochilion imbriatum (Mitt.) Inoue, J. Hattori Bot. Lab. 27: 57, 1964 (Inoue
1964b). Bas.: Plagiochila imbriata Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 97, 1860
[1861] (Mitten 1860c).
** Plagiochilion giulianettii (Steph.) Inoue, J. Hattori Bot. Lab. 27: 57, 1964 (Inoue
1964b). Bas.: Plagiochila giulianettii Steph., Bull. Herb. Boissier (sér. 2) 4 (1): 30
(402), 1904 (Stephani 1904g).
** Plagiochilion herzogii Inoue, Bull. Natl. Sci. Mus. Tokyo (n.ser.) 14 (2): 270, 1971
(Inoue 1971c).
** Plagiochilion intermedium R.M.Schust., Phytologia 45 (5): 421, 1980 (Schuster
1980b).
World checklist of hornworts and liverworts
253
** Plagiochilion mayebarae S.Hatt., J. Hattori Bot. Lab. 3: 39, 1948 [1950] (Hattori
1948a).
*** Plagiochilion oppositum (Reinw., Blume et Nees) S.Hatt., Biosphaera 1 (1): 7,
1947 (Hattori 1947a). Bas.: Jungermannia opposita Reinw., Blume et Nees, Nova
Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 236, 1824 [1825]
(Reinwardt et al. 1824a).
** Plagiochilion pachycephalum (De Not.) Inoue, J. Hattori Bot. Lab. 27: 55, 1964
(Inoue 1964b). Bas.: Plagiochila pachycephala De Not., Epat. Borneo: 14, 1874
(De Notaris 1874).
*** Plagiochilion proliferum (Mitt.) R.M.Schust., J. Hattori Bot. Lab. 26: 285, 1963
(Schuster 1963b). Bas.: Plagiochila prolifera Mitt., Bot. antarct. voy. II (Fl. Nov.Zel. 2): 130, 1854 (Mitten 1854).
** Plagiochilion theriotanum (Steph.) Inoue, J. Hattori Bot. Lab. 27: 59, 1964 (Inoue
1964b). Bas.: Plagiochila theriotana Steph., Sp. Hepat. (Stephani) 6: 228, 1921
(Stephani 1921).
** Pseudolophocolea R.M.Schust. et J.J.Engel, Lindbergia 8 (2): 71, 1982 (Schuster and Engel 1982). Based on: Pseudolophocolea R.M.Schust. et J.J.Engel, Phytologia 47 (4): 310, 1981 (Schuster and Engel 1981).
** Pseudolophocolea denticulata R.M.Schust. et J.J.Engel, Lindbergia 8 (2): 73, 1982
(Schuster and Engel 1982). Based on: Pseudolophocolea denticulata R.M.Schust. et
J.J.Engel, Phytologia 47 (4): 311, 1981 (Schuster and Engel 1981), nom. inval.
** Xenochila R.M.Schust., Amer. Midl. Naturalist 62 (1): 15, 1959 (Schuster 1959a).
** Xenochila integrifolia (Mitt.) Inoue, Bull. Natl. Sci. Mus. Tokyo (n.ser.) 6 (4): 373,
1963 (Inoue 1963). Bas.: Plagiochila integrifolia Mitt., J. Proc. Linn. Soc., Bot. 5
(18): 96, 1860 [1861] (Mitten 1860c).
***
Pseudolepicoleaceae Fulford et J.Taylor
by M. von Konrat
Blepharostoma was recognized as an element within Pseudolepicoleaceae by CrandallStotler et al. (2009), but we follow the concept of Frey and Stech (2008) with Blepharostoma as the single genus in Blepharostomataceae.
*** Archeophylla R.M.Schust., J. Hattori Bot. Lab. 26: 263, 1963 (Schuster 1963b).
** Archeophylla paradoxa R.M.Schust., Trans. Brit. Bryol. Soc. 4 (5): 810, 1965
(Schuster 1965c).
** Archeophylla pungens (Herzog) R.M.Schust., Candollea 21 (1): 86, 1966 (Schuster
1966e). Bas.: Blepharostoma pungens Herzog, Rev. Bryol. Lichénol. 29 (3/4): 189,
1960 [1961] (Herzog 1960).
254
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Archeophylla schusteri (E.A.Hodgs. et Allison) R.M.Schust., J. Hattori Bot. Lab. 26:
263, 1963 (Schuster 1963b). Bas.: Temnoma schusteri E.A.Hodgs. et Allison, Trans.
Roy. Soc. New Zealand, Bot. 1 (12): 147, 1962 (Hodgson and Allison 1962).
*** Castanoclobos J.J.Engel et Glenny, Novon 17 (4): 424, 2007 (Engel and Glenny
2007).
*** Castanoclobos julaceus (Hatcher ex J.J.Engel) J.J.Engel et Glenny, Novon 17 (4):
425, 2007 (Engel and Glenny 2007). Bas.: Leiomitra julacea Hatcher ex J.J.Engel,
Novon 9 (1): 26, 1999 (Engel 1999a).
** Chaetocolea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 346, 1885 (Spruce
1885).
*** Chaetocolea palmata Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 346, 1885
(Spruce 1885).
*** Herzogiaria Fulford ex Hässel, Lindbergia 7 (1): 23, 1981 (Hässel 1981).
*** Herzogiaria teres (Steph.) Fulford ex Hässel, Lindbergia 7 (1): 24, 1981 (Hässel
1981). Bas.: Lepicolea teres Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26
(III, 17): 26, 1901 (Stephani 1901b).
** Isophyllaria E.A.Hodgs. et Allison, Trans. Roy. Soc. New Zealand, Bot. 3 (4):
68, 1965 (Hodgson 1965).
*** Isophyllaria attenuata (Rodway) E.A.Hodgs., J. Roy. Soc. New Zealand 2 (1): 111,
1972 (Hodgson 1972). Bas.: Isotachis attenuata Rodway, Pap. & Proc. Roy. Soc.
Tasmania 1916: 47, 1917 (Rodway 1917a).
** Isophyllaria fuegiana (Hässel) R.M.Schust., Beih. Nova Hedwigia 118: 141, 2000
(Schuster 2000a). Bas.: Fulfordiella fuegiana Hässel, Comun. Mus. Argent. Ci.
Nat. “Bernardino Rivadavia,” Ci. Bot. 2 (9): 48, 1974 (Hässel 1974).
*** Pseudolepicolea Fulford et J.Taylor, Nova Hedwigia 1 (3/4): 412, 1959 [1960]
(Fulford and Taylor 1959b).
** Pseudolepicolea andoi (R.M.Schust.) Inoue, Bull. Natl. Sci. Mus. Tokyo, B 4 (3):
94, 1978 (Inoue 1978a). Bas.: Lophochaete andoi R.M.Schust., J. Hattori Bot. Lab.
26: 261, 1963 (Schuster 1963b).
** Pseudolepicolea fryei (Perss.) Grolle et Ando, Hikobia 3 (3): 180, 1963 (Ando
1963). Bas.: Lepicolea fryei Perss., Bryologist 49 (2): 47, 1946 (Persson 1946).
*** Pseudolepicolea grolleana (R.M.Schust.) Grolle, Ann. Bot. Fenn. 21 (1): 30, 1984
(Piippo 1984a). Bas.: Archeochaete grolleana R.M.Schust., Nova Hedwigia 15:
441, 1968 (Schuster 1968b).
*** Pseudolepicolea kuehnemannii (R.M.Schust.) Hässel, Fl. Criptog. Tierra del Fuego 15: 125, 1975 (Hässel and Solari 1975). Bas.: Archeochaete kuehnemannii
R.M.Schust., J. Hattori Bot. Lab. 26: 262, 1963 (Schuster 1963b).
World checklist of hornworts and liverworts
255
*** Pseudolepicolea quadrilaciniata (Sull.) Fulford et J.Taylor, Nova Hedwigia 1 (3/4):
413, 1959 [1960] (Fulford and Taylor 1959b). Bas.: Sendtnera quadrilaciniata
Sull., Hooker’s J. Bot. Kew Gard. Misc. 2: 317, 1850 (Sullivant 1850).
*** Pseudolepicolea temnomoides (R.M.Schust.) Váňa et J.J.Engel, Mem. New York
Bot. Gard. 105: 92, 2013 (Váňa and Engel 2013). Bas.: Archeochaete temnomoides
R.M.Schust., Candollea 21 (1): 129, 1966 (Schuster 1966e).
** Pseudolepicolea trollii (Herzog) Grolle et Ando, Hikobia 3 (3): 177, 1963 (Ando
1963). Bas.: Blepharostoma trollii Herzog, Ann. Bryol. 12: 80, 1939 (Herzog
1939b).
* Pseudolepicolea trollii var. darjeelingensis S.Hatt. et Mizut., J. Hattori Bot. Lab. 31:
252, 1968 (Hattori and Mizutani 1968).
*** Temnoma Mitt., Handb. N. Zeal. l. 2: 750, 1867 (Hooker 1867).
*** Temnoma angustifolium R.M.Schust., Candollea 21 (2): 279, 1966 [1967] (Schuster
1966c).
*** Temnoma chaetophyllum R.M.Schust., Phytologia 39 (4): 239, 1978 (Schuster 1978a).
** Temnoma palmatum (Lindb. ex Pearson) R.M.Schust., Bryologist 62 (4): 240,
1959 [1960] (Schuster 1959c). Bas.: Blepharostoma palmatum Lindb. ex Pearson,
J. Bot. 25: 193, 1887 (Pearson 1887a).
** Temnoma palmatum var. cuneatum R.M.Schust., Candollea 21 (2): 347, 1966
[1967] (Schuster 1966c).
** Temnoma palmatum var. laxifolium R.M.Schust., Candollea 21 (2): 345, 1966
[1967] (Schuster 1966c).
** Temnoma palmatum var. pseudospiniferum R.M.Schust., Candollea 21 (2): 348,
1966 [1967] (Schuster 1966c).
** Temnoma patagonicum R.M.Schust., Candollea 21 (2): 313, 1966 [1967] (Schuster
1966c).
*** Temnoma paucisetigerum R.M.Schust., Candollea 21 (2): 266, 1966 [1967]
(Schuster 1966c).
*** Temnoma pilosum (A.Evans) R.M.Schust., Bryologist 62 (4): 240, 1959 [1960]
(Schuster 1959c). Bas.: Blepharostoma pilosum A.Evans, Bull. Torrey Bot. Club 25
(8): 413, 1898 (Evans 1898).
*** Temnoma pulchellum (Hook.) Mitt., Handb. N. Zeal. l. 2: 753, 1867 (Hooker 1867).
Bas.: Jungermannia pulchella Hook., Musci Exot. 1: tab. 94, 1818 (Hooker 1818).
*** Temnoma quadriidum (Mitt.) Mitt. ex E.A.Hodgs. et Allison, Trans. Roy. Soc. New
Zealand, Bot. 1 (12): 142, 1962 (Hodgson and Allison 1962). Bas.: Jungermannia
quadriida Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 128, 1854 (Mitten 1854).
*** Temnoma quadripartitum (Hook.) Mitt., J. Linn. Soc., Bot. 15 (82): 68, 1876
(Mitten 1876a). Bas.: Jungermannia quadripartita Hook., Musci Exot. 2: tab. 117,
1820 (Hooker 1820).
** Temnoma quadripartitum var. pseudopungens R.M.Schust., Candollea 21 (2): 312,
1966 [1967] (Schuster 1966c).
256
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Temnoma quadripartitum var. randii (S.W.Arnell) R.M.Schust., Candollea 21 (2):
307, 1966 [1967] (Schuster 1966c). Bas.: Lepidozia randii S.W.Arnell, Svensk
Bot. Tidskr. 47 (3): 417, 1953 (Arnell 1953c).
*** Temnoma setigerum (Lindenb.) R.M.Schust., Nova Hedwigia 5: 35, 1963 (Schuster 1963c). Bas.: Jungermannia setigera Lindenb., Syn. Hepat. 1: 131, 1844
(Gottsche et al. 1844).
** Temnoma setigerum var. hawaiicum Inoue, Bull. Natl. Sci. Mus. Tokyo (n.ser.) 17
(3): 228, 1974 (Inoue 1974b).
** Temnoma townrowii R.M.Schust., Candollea 21 (2): 351, 1966 [1967] (Schuster
1966c).
***
Trichocoleaceae Nakai
by T. Katagiri
he treatment of Trichocoleaceae follows what was outlined in Katagiri and Deguchi
(2012) and Katagiri et al. (2012, 2013). Recent nomenclatural and taxonomic notes
can also be found in Katagiri (2013).
*** Eotrichocolea R.M.Schust., J. Hattori Bot. Lab. 26: 252, 1963 (Schuster 1963b).
*** Eotrichocolea furukii T.Katag., Bryologist 115 (4): 519, 2012 (Katagiri et al. 2012).
*** Eotrichocolea polyacantha (Hook.f. et Taylor) R.M.Schust., J. Hattori Bot. Lab. 26:
264, 1963 (Schuster 1963b). Bas.: Jungermannia polyacantha Hook.f. et Taylor,
London J. Bot. 3: 290 [390], 1844 (Hooker and Taylor 1844a).
*** Leiomitra Lindb., Acta Soc. Sci. Fenn. 10: 515, 1875 (Lindberg 1875).
** subg. Brachygyna R.M.Schust., Nova Hedwigia 73 (3/4): 480, 2001 (Schuster
2001b).
** Leiomitra mastigophoroides R.M.Schust., Phytologia 45 (5): 416, 1980 (Schuster
1980b).
** subg. Leiomitra, Nova Hedwigia 73 (3/4): 480, 2001 (Schuster 2001b).
*** Leiomitra breviseta (Steph.) R.M.Schust., Beih. Nova Hedwigia 118: 152, 2000
(Schuster 2000a). Bas.: Trichocolea breviseta Steph., Sp. Hepat. (Stephani) 4: 60,
1909 (Stephani 1909d).
*** Leiomitra capillata Lindb., Acta Soc. Sci. Fenn. 10: 515, 1875 (Lindberg 1875).
*** Leiomitra elegans (Lehm.) Hässel, Novon 12 (4): 465, 2002 (Hässel 2002). Bas.:
Trichocolea elegans Lehm., Nov. Stirp. Pug. 10: 8, 1857 (Lehmann 1857).
** Leiomitra elliottii (Steph.) R.M.Schust., Nova Hedwigia 73 (3/4): 469, 2001
(Schuster 2001b). Bas.: Trichocolea elliottii Steph., Sp. Hepat. (Stephani) 4: 55,
1909 (Stephani 1909d).
World checklist of hornworts and liverworts
257
*** Leiomitra laccida Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 349, 1885
(Spruce 1885).
** Leiomitra hirticaulis R.M.Schust., Nova Hedwigia 73 (3/4): 472, 2001 (Schuster
2001b).
*** Leiomitra lanata (Hook.) R.M.Schust., Phytologia 45 (5): 417, 1980 (Schuster
1980b). Bas.: Jungermannia lanata Hook., Musci Exot. 2: tab. 116, 1820 (Hooker
1820).
*** Leiomitra merrillana (Steph.) T.Katag., Bryologist 115 (4): 488, 2012 (Katagiri
and Deguchi 2012). Bas.: Trichocolea merrillana Steph., Sp. Hepat. (Stephani) 6:
374, 1923 (Stephani 1923).
*** Leiomitra paraphyllina Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 350, 1885
(Spruce 1885).
** Leiomitra robusta (Steph.) R.M.Schust., Nova Hedwigia 73 (3/4): 469, 2001
(Schuster 2001b). Bas.: Trichocolea robusta Steph., Sp. Hepat. (Stephani) 4: 58,
1909 (Stephani 1909d).
** Leiomitra smaragdina Hässel, Novon 12 (4): 467, 2002 (Hässel 2002).
*** Leiomitra tomentosa (Sw.) Lindb., Acta Soc. Sci. Fenn. 10: 515, 1875 (Lindberg
1875). Bas.: Jungermannia tomentosa Sw., Prodr. (Swartz): 145, 1788 (Swartz 1788).
*** Trichocolea Dumort., Commentat. Bot. (Dumortier): 113, 1822 (Dumortier
1822) nom. conserv.
** Trichocolea argentea Herzog, Arch. Bot. São Paulo 1 (2): 40, 1925 (Herzog 1925b).
*** Trichocolea breviissa Steph., Sp. Hepat. (Stephani) 4: 54, 1909 (Stephani 1909d).
** Trichocolea comptonii Pearson, J. Linn. Soc., Bot. 46 (305): 27, 1922 (Pearson
1922b).
*** Trichocolea ilicaulis Steph., Sp. Hepat. (Stephani) 4: 59, 1909 (Stephani 1909d).
* Trichocolea loccosa Herzog et Hatcher, Lloydia 20 (3): 148, 1957 [1958] (Hatcher
1957). 183
** Trichocolea geniculata Pearson, J. Linn. Soc., Bot. 46 (305): 28, 1922 (Pearson 1922b).
** Trichocolea gracillima Austin, Bot. Gaz. 3 (1): 6, 1878 (Austin 1878).
*** Trichocolea hatcheri E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot. 3 (4): 69,
1965 (Hodgson 1965).
*** Trichocolea iriomotensis T.Katag., Hattoria 4: 6, 2013 (Katagiri et al. 2013).
*** Trichocolea japonica T.Katag., Bryologist 114 (4): 744, 2011 (Katagiri et al. 2011).
*** Trichocolea magna T.Katag., Hattoria 4: 7, 2013 (Katagiri et al. 2013).
** Trichocolea minutifolia Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2):
134, 1914 (Stephani and Watts 1914).
*** Trichocolea mollissima (Hook.f. et Taylor) Gottsche, Ann. Sci. Nat. Bot. (sér. 5)
1: 132, 1864 (Gottsche 1864). Bas.: Jungermannia mollissima Hook.f. et Taylor,
London J. Bot. 3: 290 [390], 1844 (Hooker and Taylor 1844a).
183 Trichocolea loccosa is treated as a variety of Trichocolea laccida (=Leiomitra laccida by Schultze-Motel
& Menzel (1987).
258
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Trichocolea pluma (Reinw., Blume et Nees) Mont., Voy. Bonite, Bot. 1: 238, 1846
(Montagne 1846). Bas.: Jungermannia pluma Reinw., Blume et Nees, Nova Acta
Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 209, 1824 [1825] (Reinwardt et al. 1824a).
*** Trichocolea rigida R.M.Schust., Nova Hedwigia 15: 447, 1968 (Schuster 1968b).
*** Trichocolea rudimentaris Steph., Sp. Hepat. (Stephani) 6: 376, 1923 (Stephani 1923).
* Trichocolea sprucei Steph., Sp. Hepat. (Stephani) 4: 59, 1909 (Stephani 1909d).
Nom. nov. pro Trichocolea gracillima Spruce, J. Linn. Soc., Bot. 30 (210): 353,
1895 (Gepp 1895b), nom. illeg. 184
*** Trichocolea tomentella (Ehrh.) Dumort., Syll. Jungerm. Europ.: 67, 1831 (Dumortier 1831). Bas.: Jungermannia tomentella Ehrh., Hannover. Mag. 21 (18): 277,
1783 (Ehrhart 1783).
** Trichocolea udarii D.K.Singh, Bull. Bot. Surv. India 25: 177, 1983 [1985] (Singh
1983a).
** Trichocolea wattsiana Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2):
135, 1914 (Stephani and Watts 1914).
Myliineae J.J.Engel et Braggins ex Crand.-Stotl.,Váňa, Stotler et J.J.Engel
***
Myliaceae Schljakov
by L. Söderström
Söderström et al. (2015c) outlines the controversy as to the placement of Mylia anomala
where they advocate recognition at the subgeneric level. It has previously been recognized
as a segregate genus, Leiomylia J.J.Engel and Braggins (cf. also Shaw et al. 2015).
*** Mylia Gray, Nat. Arr. Brit. Pl. 1: 693, 1821 (Gray 1821) nom. conserv.
** subg. Anomalae (R.M.Schust. ex Potemkin) L.Söderstr., Phytotaxa 202 (1):
70, 2015 (Söderström et al. 2015c). Bas.: Mylia sect. Anomalae R.M.Schust. ex
Potemkin, Arctoa 2: 1, 1993 (Potemkin and Kazanovsky 1993).
*** Mylia anomala (Hook.) Gray, Nat. Arr. Brit. Pl. 1: 693, 1821 (Gray 1821). Bas.:
Jungermannia anomala Hook., Brit. Jungermann.: tab. 34, 1813 (Hooker 1813).
** subg. Mylia
*** Mylia taylorii (Hook.) Gray, Nat. Arr. Brit. Pl. 1: 693, 1821 (Gray 1821). Bas.:
Jungermannia taylorii Hook., Brit. Jungermann.: tab. 57, 1813 (Hooker 1813).
*** Mylia verrucosa Lindb., Acta Soc. Sci. Fenn. 10: 236, 1872 [1873] (Lindberg
1872b).
184 Trichocolea sprucei is suggested to be a variety of Trichocolea laccida (=Leiomitra laccida) in SchultzeMotel & Menzel (1987)
World checklist of hornworts and liverworts
259
*** Mylia verrucosa subsp. nuda (Inoue et B.Y.Yang) Potemkin et Kazan., Arctoa 2:
5, 1993 (Potemkin and Kazanovsky 1993). Bas.: Mylia nuda Inoue et B.Y.Yang,
Taiwania 12 (1): 35, 1966 (Inoue and Yang 1966).
Perssoniellineae R.M.Schust.
***
Schistochilaceae H.Buch
by X. He and D. Glenny
He et al. (2014b) provided an historical account of Schistochilaceae summarizing
studies that have showed that the phylogenetic structure of the family does not match
units that have resulted from morphologically-based investigations. Further studies
are needed until a natural division of the family can be proposed. hus, we follow the
broad concept of He and Glenny (2010) and He and Sun (2013) here, treating Schistochilaceae as comprising a single genus as discussed by He et al. (2014b).
*** Schistochila Dumort., Recueil Observ. Jungerm.: 15, 1835 (Dumortier 1835).
** Schistochila acuminata Steph., Sp. Hepat. (Stephani) 4: 81, 1909 (Stephani
1909d).
*** Schistochila aequiloba Steph., Sp. Hepat. (Stephani) 4: 80, 1909 (Stephani 1909d).
*** Schistochila alata (Lehm.) Schifn., Hepat. (Engl.-Prantl): 111, 1893 (Schifner
1893b). Bas.: Jungermannia alata Lehm., Linnaea 4: 359, 1829 (Lehmann 1829).
*** Schistochila aligera (Nees et Blume) J.B.Jack et Steph., Hedwigia 31 (1): 12, 1892 (Jack
and Stephani 1892). Bas.: Jungermannia aligera Nees et Blume, Nova Acta Phys.-Med.
Acad. Caes. Leop.-Carol. Nat. Cur. 11: 135, 1823 (Blume and Nees 1823).
* Schistochila aligera var. laxa (Nees) Schifn., Consp. Hepat. Arch. Ind.: 212, 1898
(Schifner 1898b). Bas.: Jungermannia aligera γ laxa Nees, Enum. Pl. Crypt. Javae:
68, 1830 (Nees 1830).
*** Schistochila altissima E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot. 3 (4): 85,
1965 (Hodgson 1965).
** Schistochila altissima subsp. polystratosa R.M.Schust. et J.J.Engel, Phytologia 30
(4): 241, 1975 (Schuster and Engel 1975).
*** Schistochila antara Grolle, J. Hattori Bot. Lab. 29: 249, 1966 (Grolle 1966h).
*** Schistochila appendiculata (Hook.) Dumort. ex Trevis., Mem. Reale Ist. Lombardo
Sci. (Ser. 3), C. Sci. Mat. 4 (13): 392, 1877 (Trevisan 1877). Bas.: Jungermannia
appendiculata Hook., Musci Exot. 1: tab. 15, 1818 (Hooker 1818).
* Schistochila baileyana Steph., Sp. Hepat. (Stephani) 4: 85, 1909 (Stephani 1909d). 185
*** Schistochila balfouriana (Hook.f. et Taylor) Steph., Sp. Hepat. (Stephani) 4: 91,
1909 (Stephani 1909d). Bas.: Jungermannia balfouriana Hook.f. et Taylor, London J. Bot. 3: 556, 1844 (Hooker and Taylor 1844d).
185 Schistochila baileyana is possibly conspeciic with Schistochila beccariana (D. Meagher, pers. comm.).
260
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Schistochila beccariana (De Not.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser.
3), C. Sci. Mat. 4 (13): 392, 1877 (Trevisan 1877). Bas.: Gottschea beccariana De
Not., Epat. Borneo: 9, 1874 (De Notaris 1874).
*** Schistochila berggrenii (J.J.Engel et R.M.Schust.) Xiao L.He et Glenny, Austral.
Syst. Bot. 23 (4): 237, 2010 (He and Glenny 2010). Bas.: Pachyschistochila berggrenii J.J.Engel et R.M.Schust., J. Hattori Bot. Lab. 58: 476, 1985 (Schuster and
Engel 1985).
*** Schistochila berteroana (Hook.) Steph., Sp. Hepat. (Stephani) 4: 96, 1909 (Stephani
1909d). Bas.: Jungermannia berteroana Hook., Bot. Misc. 2: 148, 1831 (Hooker
1831).
*** Schistochila blumei (Nees) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 392, 1877 (Trevisan 1877). Bas.: Jungermannia blumei Nees, Nova
Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 11: 136, 1823 (Blume and
Nees 1823).
** Schistochila brassii Grolle, J. Hattori Bot. Lab. 31: 5, 1968 (Grolle 1968a).
** Schistochila caledonica Steph., Sp. Hepat. (Stephani) 4: 77, 1909 (Stephani 1909d).
*** Schistochila carnosa (Mitt.) Steph., Sp. Hepat. (Stephani) 4: 93, 1909 (Stephani
1909d). Bas.: Gottschea carnosa Mitt., J. Linn. Soc., Bot. 15 (82): 72, 1876 (Mitten
1876a).
*** Schistochila caudata R.M.Schust. et J.J.Engel, Phytologia 30 (4): 242, 1975 (Schuster and Engel 1975).
*** Schistochila childii (R.M.Schust. et J.J.Engel) Xiao L.He et Glenny, Austral. Syst. Bot.
23 (4): 237, 2010 (He and Glenny 2010). Bas.: Pachyschistochila childii R.M.Schust.
et J.J.Engel, J. Hattori Bot. Lab. 58: 471, 1985 (Schuster and Engel 1985).
*** Schistochila chlorophylla (Colenso) E.A.Hodgs., Trans. & Proc. Roy. Soc. New
Zealand 71 (3): 192, 1941 (Hodgson 1941). Bas.: Gottschea chlorophylla Colenso,
Trans. & Proc. New Zealand Inst. 18: 240, 1886 (Colenso 1886b).
*** Schistochila ciliata (Mitt.) Steph., Sp. Hepat. (Stephani) 4: 87, 1909 (Stephani
1909d). Bas.: Gottschea ciliata Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 151,
1854 (Mitten 1854).
*** Schistochila compacta (Colenso) E.A.Hodgs., Trans. & Proc. Roy. Soc. New Zealand 71 (3): 192, 1941 (Hodgson 1941). Bas.: Gottschea compacta Colenso, Trans.
& Proc. New Zealand Inst. 16: 349, 1884 (Colenso 1884).
*** Schistochila conchophylla Herzog ex E.A.Hodgs. et Allison, Trans. & Proc. Roy.
Soc. New Zealand 71 (3): 191, 1941 (Hodgson 1941).
** Schistochila conchophylla var. multidentata (J.J.Engel) Xiao L.He et Glenny, Phytotaxa 173 (1): 92, 2014 (He et al. 2014b). Bas.: Gottschea conchophylla var. multidentata J.J.Engel, Nova Hedwigia 93 (3/4): 407, 2011 (Engel 2011).
*** Schistochila congoana Steph., Sp. Hepat. (Stephani) 4: 71, 1909 (Stephani 1909d).
** Schistochila cookei (H.A.Mill.) R.M.Schust., J. Hattori Bot. Lab. 42: 274, 1977
(Schuster and Engel 1977). Bas.: Fulfordistria cookei H.A.Mill., Phytologia 20 (5):
320, 1970 (Miller 1970).
** Schistochila crinita Grolle, J. Hattori Bot. Lab. 31: 7, 1968 (Grolle 1968a).
World checklist of hornworts and liverworts
261
*** Schistochila cristata Steph., Hedwigia 28 (4): 274, 1889 (Stephani 1889c).
*** Schistochila cunninghamii Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26
(III, 17): 27, 1901 (Stephani 1901b).
** Schistochila doriae (De Not.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 392, 1877 (Trevisan 1877). Bas.: Gottschea doriae De Not., Epat.
Borneo: 10, 1874 (De Notaris 1874).
** Schistochila engleriana Steph., Sp. Hepat. (Stephani) 4: 69, 1909 (Stephani 1909d).
*** Schistochila exalata Herzog, Rev. Bryol. Lichénol. 29 (3/4): 191, 1960 [1961]
(Herzog 1960).
** Schistochila ijiensis H.Buch et Herzog, Memoranda Soc. Fauna Fl. Fennica 27: 94,
1952 (Herzog 1952c).
*** Schistochila glaucescens (Hook.) A.Evans, Rev. Bryol. 32 (4): 60, 1905 (Evans
1905c). Bas.: Jungermannia glaucescens Hook., Musci Exot. 1: tab. 39, 1818 (Hooker 1818).
** Schistochila hattorii Grolle, J. Hattori Bot. Lab. 29: 247, 1966 (Grolle 1966h).
** Schistochila integerrima Steph., Sp. Hepat. (Stephani) 6: 492, 1924 (Stephani 1924).
** Schistochila isotachyphylla (J.J.Engel et R.M.Schust.) Xiao L.He et Glenny, Austral.
Syst. Bot. 23 (4): 237, 2010 (He and Glenny 2010). Bas.: Paraschistochila isotachyphylla J.J.Engel et R.M.Schust., J. Hattori Bot. Lab. 58: 429, 1985 (Schuster
and Engel 1985).
*** Schistochila kirkiana Steph., Sp. Hepat. (Stephani) 4: 86, 1909 (Stephani 1909d).
*** Schistochila kunkelii S.W.Arnell, Ark. Bot. (n.ser.) 4 (1): 12, 1957 (Arnell 1957b).
** Schistochila lacerata Steph., Sp. Hepat. (Stephani) 6: 492, 1924 (Stephani 1924).
*** Schistochila lamellata (Hook.) Dumort. ex A.Evans, Contr. U.S. Natl. Herb. 1 (5):
141, 1892 (Evans 1892b). Bas.: Jungermannia lamellata Hook., Musci Exot. 1:
tab. 49, 1818 (Hooker 1818).
*** Schistochila laminigera (Hook.f. et Taylor) A.Evans, Contr. U.S. Natl. Herb. 1
(5): 141, 1892 (Evans 1892b). Bas.: Jungermannia laminigera Hook.f. et Taylor,
London J. Bot. 3: 456, 1844 (Hooker and Taylor 1844b).
*** Schistochila latiloba (R.M.Schust. et J.J.Engel) Xiao L.He et Glenny, Austral. Syst.
Bot. 23 (4): 237, 2010 (He and Glenny 2010). Bas.: Pachyschistochila latiloba
R.M.Schust. et J.J.Engel, J. Hattori Bot. Lab. 58: 493, 1985 (Schuster and Engel
1985).
*** Schistochila lehmanniana (Lindenb.) Steph., Sp. Hepat. (Stephani) 4: 86, 1909
(Stephani 1909d). Bas.: Jungermannia lehmanniana Lindenb., Nov. Stirp. Pug. 4:
60, 1832 (Lehmann 1832).
*** Schistochila leucophylla (Lehm. ex Gottsche, Lindenb. et Nees) Steph., Sp. Hepat.
(Stephani) 4: 98, 1910 (Stephani 1910b). Bas.: Gottschea leucophylla Lehm. ex
Gottsche, Lindenb. et Nees, Syn. Hepat. 1: 17, 1844 (Gottsche et al. 1844).
** Schistochila macrodonta W.E.Nicholson, Symb. Sin. 5: 29, 1930 (Nicholson et al.
1930).
** Schistochila minor C.Gao et Y.H.Wu, J. Hattori Bot. Lab. 95: 267, 2004 (Gao and
Wu 2004).
262
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Schistochila monticola R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 14 (4):
623, 1971 (Schuster 1971c).
*** Schistochila muricata E.A.Hodgs. et Allison, Trans. & Proc. Roy. Soc. New Zealand 71 (3): 186, 1941 (Hodgson 1941).
*** Schistochila nadeaudiana Steph., Sp. Hepat. (Stephani) 4: 76, 1909 (Stephani
1909d).
** Schistochila neesii (Mont.) Lindb., J. Linn. Soc., Bot. 13 (67): 194, 1872 [1873]
(Lindberg 1872a). Bas.: Gottschea neesii Mont., Ann. Sci. Nat. Bot. (sér. 2) 19:
244, 1843 (Montagne 1843).
*** Schistochila nitidissima R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 11 (1):
27, 1968 (Schuster 1968a).
*** Schistochila nivicola (R.M.Schust. et J.J.Engel) Xiao L.He et Glenny, Austral. Syst. Bot.
23 (4): 237, 2010 (He and Glenny 2010). Bas.: Pachyschistochila nivicola R.M.Schust.
et J.J.Engel, J. Hattori Bot. Lab. 58: 512, 1985 (Schuster and Engel 1985).
*** Schistochila nobilis (Hook.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 392, 1877 (Trevisan 1877). Bas.: Jungermannia nobilis Hook.,
Musci Exot. 1: tab. 11, 1818 (Hooker 1818).
*** Schistochila nuda Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 215, 1934
(Horikawa 1934).
*** Schistochila pachyphylla (Lehm.) Steph., Sp. Hepat. (Stephani) 4: 99, 1910
(Stephani 1910b). Bas.: Jungermannia pachyphylla Lehm., Nov. Stirp. Pug. 6: 61,
1834 (Lehmann 1834).
*** Schistochila papillifera R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 11 (1):
27, 1968 (Schuster 1968a).
*** Schistochila parvistipula Rodway, Pap. & Proc. Roy. Soc. Tasmania 1916: 47,
1917 (Rodway 1917a).
*** Schistochila pellucida R.M.Schust. et J.J.Engel, J. Hattori Bot. Lab. 58: 286, 1985
(Schuster and Engel 1985).
*** Schistochila piligera Steph., Bot. Gaz. 15 (11): 291, 1890 (Stephani 1890c).
*** Schistochila pinnatifolia (Hook.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 392, 1877 (Trevisan 1877). Bas.: Jungermannia pinnatifolia
Hook., Musci Exot. 2: tab. 114, 1820 (Hooker 1820).
*** Schistochila pluriciliata R.M.Schust. et J.J.Engel, J. Hattori Bot. Lab. 58: 395,
1985 (Schuster and Engel 1985).
*** Schistochila pseudociliata R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 14 (4):
623, 1971 (Schuster 1971c).
*** Schistochila quadriida A.Evans, Contr. U.S. Natl. Herb. 1 (5): 141, 1892 (Evans
1892b).
** Schistochila ramentacea Steph., Sp. Hepat. (Stephani) 6: 494, 1924 (Stephani 1924).
*** Schistochila relexa (Mont.) Steph., Sp. Hepat. (Stephani) 4: 97, 1910 (Stephani
1910b). Bas.: Gottschea relexa Mont., Ann. Sci. Nat. Bot. (sér. 3) 4: 347, 1845
(Montagne 1845b).
World checklist of hornworts and liverworts
263
*** Schistochila relexistipula J.J.Engel et R.M.Schust., Phytologia 30 (4): 245, 1975
(Schuster and Engel 1975).
*** Schistochila reinwardtii (Nees) Schifn., Consp. Hepat. Arch. Ind.: 218, 1898
(Schifner 1898b). Bas.: Jungermannia reinwardtii Nees, Enum. Pl. Crypt. Javae:
66, 1830 (Nees 1830).
*** Schistochila repleta (Hook.f. et Taylor) Steph., Sp. Hepat. (Stephani) 4: 90, 1909
(Stephani 1909d). Bas.: Jungermannia repleta Hook.f. et Taylor, London J. Bot. 3:
557, 1844 (Hooker and Taylor 1844d).
** Schistochila rubriseta Steph., Sp. Hepat. (Stephani) 4: 78, 1909 (Stephani 1909d).
** Schistochila schultzei Steph., Sp. Hepat. (Stephani) 6: 494, 1924 (Stephani 1924).
*** Schistochila sciophila R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 14 (4): 632,
1971 (Schuster 1971c).
*** Schistochila sciurea (Nees) Schifn., Nova Acta Acad. Caes. Leop.-Carol. German.
Nat. Cur. 60 (2): 251, 1893 (Schifner 1893a). Bas.: Jungermannia sciurea Nees,
Enum. Pl. Crypt. Javae: 34, 1830 (Nees 1830).
*** Schistochila simulans (C.Massal.) Xiao L.He et Yu Sun, Polish Bot. J. 58 (2): 473,
2013 (He and Sun 2013). Bas.: Cephalozia simulans C.Massal., Nuovo Giorn.
Bot. Ital. 17 (3): 236, 1885 (Massalongo 1885).
*** Schistochila spegazziniana (C.Massal.) Steph., Bih. Kongl. Svenska Vetensk.-Akad.
Handl. 26 (III, 6): 60, 1900 (Stephani 1900b). Bas.: Gottschea spegazziniana
C.Massal., Nuovo Giorn. Bot. Ital. 17 (3): 206, 1885 (Massalongo 1885).
*** Schistochila sphagnoides (Schwägr.) Lindb. ex Steph., Sp. Hepat. (Stephani) 4: 70,
1909 (Stephani 1909d). Bas.: Jungermannia sphagnoides Schwägr., Hist. Musc.
Hepat. Prodr.: 23, 1814 (Schwägrichen 1814).
*** Schistochila splachnophylla (Hook.f. et Taylor) Steph., Bih. Kongl. Svenska
Vetensk.-Akad. Handl. 26 (III, 17): 28, 1901 (Stephani 1901b). Bas.: Jungermannia
splachnophylla Hook.f. et Taylor, London J. Bot. 3: 455, 1844 (Hooker and Taylor
1844b).
** Schistochila stratosa (Mont.) A.Evans, Contr. U.S. Natl. Herb. 1 (5): 141, 1892
(Evans 1892b). Bas.: Gottschea stratosa Mont., Ann. Sci. Nat. Bot. (sér. 3) 4: 346,
1845 (Montagne 1845b).
*** Schistochila subhyalina R.M.Schust., Phytologia 30 (4): 246, 1975 (Schuster and
Engel 1975).
** Schistochila subhyalina var. grandidentata (J.J.Engel et R.M.Schust.) Xiao L.He
et Glenny, Austral. Syst. Bot. 23 (4): 237, 2010 (He and Glenny 2010). Bas.:
Pachyschistochila subhyalina var. grandidentata J.J.Engel et R.M.Schust., J. Hattori
Bot. Lab. 58: 507, 1985 (Schuster and Engel 1985).
*** Schistochila subimmersa J.J.Engel et R.M.Schust., Phytologia 30 (4): 247, 1975
(Schuster and Engel 1975).
*** Schistochila succulenta (J.J.Engel et R.M.Schust.) Xiao L.He et Glenny, Austral. Syst.
Bot. 23 (4): 237, 2010 (He and Glenny 2010). Bas.: Pachyschistochila succulenta J.J.Engel et R.M.Schust., J. Hattori Bot. Lab. 58: 517, 1985 (Schuster and Engel 1985).
264
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Schistochila tasmanica Steph., Sp. Hepat. (Stephani) 4: 86, 1909 (Stephani 1909d).
*** Schistochila trispiralis R.M.Schust., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 11 (1): 28,
1968 (Schuster 1968a).
*** Schistochila tuloides (Hook.f. et Taylor) Steph., Sp. Hepat. (Stephani) 4: 89, 1909
(Stephani 1909d). Bas.: Jungermannia tuloides Hook.f. et Taylor, London J. Bot.
3: 558, 1844 (Hooker and Taylor 1844d).
*** Schistochila undulatifolia Piippo, Ann. Bot. Fenn. 23 (1): 8, 1986 (Piippo 1986b).
*** Schistochila virescens R.M.Schust., Phytologia 30 (4): 248, 1975 (Schuster and Engel
1975).
*** Schistochila vitreocincta (Herzog) Xiao L.He et Glenny, Austral. Syst. Bot. 23 (4):
237, 2010 (He and Glenny 2010). Bas.: Perssoniella vitreocincta Herzog, Ark. Bot.
(n.ser.) 2 (4): 265, 1952 (Herzog 1952g).
** Schistochila volans Grolle, J. Hattori Bot. Lab. 29: 244, 1966 (Grolle 1966h).
*** Schistochila yakushimensis N.Ohnishi et Deguchi, Bryologist 106 (3): 451, 2003
(Ohnishi and Deguchi 2003).
** Schistochila zantenii Grolle, J. Hattori Bot. Lab. 29: 243, 1966 (Grolle 1966h).
Porellales Schljakov
Jubulineae Müll.Frib.
***
Frullaniaceae Lorch
by M. von Konrat, J. Hentschel, J. Uribe, P. Sukkharak, J. Heinrichs, J. Larraín, R.
Stotler and L. Zhang
he subgeneric treatment of Frullaniaceae follows Hentschel et al. (2015), which includes a slight departure from formal infrageneric ranks where they apply clades referred to as Diastaloba I, II, III, and IV that appear to represent distinct subgenera.
*** Frullania Raddi, Jungermanniogr. Etrusca: 9, 1818 (Raddi 1818a).
*** subg. Chonanthelia Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 8, 1884
(Spruce 1884).
* Frullania lammea Taylor, Trans. & Proc. Bot. Soc. Edinburgh 15: 29, 1884
(Spruce 1884).
*** Frullania lindmanii Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 23 (III, 2):
19, 1897 (Stephani 1897a).
** Frullania paranensis Steph., Sp. Hepat. (Stephani) 4: 607, 1911 (Stephani 1911e).
** Frullania spegazzinii M.E.Reiner, Bol. Soc. Argent. Bot. 25 (3/4): 310, 1988
(Reiner 1988).
*** sect. Chonanthelia (Spruce) Yuzawa ex Hentschel et von Konrat, Phytotaxa
220 (2): 129, 2015 (Hentschel et al. 2015). Bas.: Frullania subg. Chonanthelia
Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 8, 1884 (Spruce 1884).
World checklist of hornworts and liverworts
265
*** Frullania gibbosa Nees, Syn. Hepat. 3: 411, 1845 (Gottsche et al. 1845b).
*** sect. Cladocarpicae Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 43, 1884
(Spruce 1884).
*** Frullania beauverdii Steph., Biblioth. Bot. 87 (2): 241, 1916 (Stephani 1916a).
*** Frullania blepharozia Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 18, 1884
(Spruce 1884).
*** Frullania bogotensis Steph., Sp. Hepat. (Stephani) 4: 327, 1910 (Stephani 1910b).
* Frullania brachycarpa Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: cciv, 1889
[1890] (Spruce 1889).
*** Frullania confertiloba Steph., Sp. Hepat. (Stephani) 4: 326, 1910 (Stephani 1910b).
*** Frullania decidua Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 30, 1884
(Spruce 1884).
*** Frullania ecklonii (Spreng.) Gottsche, Lindenb. et Nees, Syn. Hepat. 3: 413, 1845
(Gottsche et al. 1845b). Bas.: Jungermannia ecklonii Spreng. Syst. Veg. (ed. 16)
[Sprengel] 4 (2): 324, 1827 (Sprengel 1827b).
* Frullania ecklonii var. robustior (Gottsche, Lindenb. et Nees) Sim, Trans. Roy.
Soc. South Africa 15 (1): 39, 1926 (Sim 1926). Bas.: Frullania ecklonii a robustior
Gottsche, Lindenb. et Nees, Syn. Hepat. 5: 771, 1847 (Gottsche et al. 1847).
* Frullania ecklonii var. rufescens Gottsche, Lindenb. et Nees, Syn. Hepat. 5: 771,
1847 (Gottsche et al. 1847).
* Frullania ecklonii var. tenerior (Gottsche, Lindenb. et Nees) Sim, Trans. Roy.
Soc. South Africa 15 (1): 39, 1926 (Sim 1926). Bas.: Frullania ecklonii b tenerior
Gottsche, Lindenb. et Nees, Syn. Hepat. 5: 771, 1847 (Gottsche et al. 1847).
*** Frullania holostipula S.Hatt. et D.G.Griin, Misc. Bryol. Lichenol. 8 (3): 47, 1978
(Hattori and Griin 1978).
*** Frullania megalostipa Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 15, 1884
(Spruce 1884).
*** Frullania obscura (Sw.) Mont., Ann. Sci. Nat. Bot. (sér. 2) 14: 333, 1840 (Montagne 1840a). Bas.: Jungermannia obscura Sw., Fl. Ind. Occid. 3: 1869, 1806
(Swartz 1806).
*** Frullania obscura var. spiniloba (Steph.) Hentschel et von Konrat, Phytotaxa 220
(2): 136, 2015 (Hentschel et al. 2015). Bas.: Frullania spiniloba Steph., Sp. Hepat.
(Stephani) 4: 336, 1910 (Stephani 1910b).
*** Frullania ringens Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 17, 1884 (Spruce 1884).
*** Frullania rio-janeirensis (Raddi) Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 33
(7): 88, 1876 [1877] (Ångström 1876). Bas.: Frullanoides rio-janeirensis Raddi,
Critt. Brasil.: 13, 1822 (Raddi 1822).
*** Frullania sphaerocephala Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 17, 1884
(Spruce 1884).
** Frullania tunguraguana L.Clark et Frye, Bryologist 55 (2): 133, 1952 (Clark and
Frye 1952). Nom. nov. pro Frullania brachyclada Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 15, 1884 (Spruce 1884), nom. illeg.
266
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** sect. Pluricarinatae (Yuzawa, Mues et S.Hatt.) Hentschel et von Konrat, Phytotaxa 220 (2): 129, 2015 (Hentschel et al. 2015). Bas.: Frullania ser. Pluricarinatae
Yuzawa, Mues et S.Hatt., J. Hattori Bot. Lab. 63: 428, 1987 (Yuzawa et al. 1987).
*** Frullania albertii Steph., Biblioth. Bot. 87 (2): 237, 1916 (Stephani 1916a).
*** Frullania arsenii Steph., Sp. Hepat. (Stephani) 6: 530, 1924 (Stephani 1924).
** Frullania bonariensis M.E.Reiner, Bol. Soc. Argent. Bot. 25 (3/4): 313, 1988
(Reiner 1988).
*** Frullania cuencensis Taylor, London J. Bot. 5: 406, 1846 (Taylor 1846b).
*** Frullania depressa Mitt., J. Proc. Linn. Soc., Bot. 7 (27): 168, 1863 (Mitten 1863).
*** Frullania dusenii Steph., Arch. Mus. Nac. Rio de Janeiro 13: 115 (9), 1905 (Stephani 1905c).
*** Frullania gradsteinii Yuzawa, Mues et S.Hatt., J. Hattori Bot. Lab. 63: 429, 1987
(Yuzawa et al. 1987).
*** Frullania haematocysta Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 54, 1884
(Spruce 1884).
*** Frullania jelskii Loitl., Diagn. pl. nov.: 18, 1894 (Loitlesberger 1894).
*** Frullania laxilora Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 26, 1884
(Spruce 1884).
* Frullania laxilora var. crossii Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 27,
1884 (Spruce 1884).
*** Frullania neurota Taylor, London J. Bot. 5: 400, 1846 (Taylor 1846b).
*** Frullania planifolia Steph., Sp. Hepat. (Stephani) 4: 337, 1910 (Stephani 1910b).
*** Frullania pluricarinata Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 168, 1864
(Gottsche 1864).
** Frullania sandvicensis Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 29 (4): 28,
1872 (Ångström 1872).
*** Frullania standaertii Steph., Sp. Hepat. (Stephani) 4: 342, 1910 (Stephani 1910b).
*** Frullania stenostipa Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 29, 1884
(Spruce 1884).
*** Frullania tetraptera Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 9: 47, 1838 (Montagne 1838).
** Frullania trinervis (Lehm.) Drège, Flora, Beig. 26: 186, 1843 (Drège 1843). Bas.:
Jungermannia trinervis Lehm., Linnaea 9 (4): 426, 1835 (Lehmann 1835). 186
*** Frullania winteri Steph., Sp. Hepat. (Stephani) 4: 338, 1910 (Stephani 1910b).
*** Frullania winteri var. vanderhammenii (Haarbrink) Yuzawa, J. Hattori Bot. Lab.
70: 233, 1991 (Yuzawa 1991). Bas.: Frullania vanderhammenii Haarbrink, Lindbergia 7 (1): 56, 1981 (Haarbrink 1981).
*** subg. Diastaloba Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 55, 1884
(Spruce 1884).
** Frullania antaresensis S.Hatt., J. Hattori Bot. Lab. 47: 92, 1980 (Hattori 1980d).
186 Frullania trinervis is possibly a species complex (Vanden Berghen 1976b).
World checklist of hornworts and liverworts
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267
Frullania armatifolia Verd., Bull. Jard. Bot. Buitenzorg (sér. 3) 12 (1): 61, 1932
(Verdoorn 1932a).
Frullania curvilobula Schäf.-Verw., D.F.Peralta et S.M.Siqueira, Phytotaxa 57 (4):
27, 2012 (Schäfer-Verwimp et al. 2012).
Frullania gracilicaulis S.Hatt., J. Hattori Bot. Lab. 43: 421, 1977 [1978] (Hattori
1977b).
Frullania humbertii Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 46 (1/2): 24,
1976 (Vanden Berghen 1976b).
Frullania hypoleucula S.Hatt., J. Hattori Bot. Lab. 57: 413, 1984 (Hattori 1984a).
Frullania incurva S.Hatt., J. Hattori Bot. Lab. 65: 431, 1988 (Hattori 1988a).
Frullania klotzschii Nees, Sp. Hepat. (Stephani) 4: 558, 1911 (Stephani 1911e).
Frullania letestui Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 46 (1/2): 38, 1976
(Vanden Berghen 1976b).
Frullania miradorensis Lindenb. et Gottsche, Syn. Hepat. 5: 781, 1847 (Gottsche
et al. 1847).
Frullania odontostipa Spruce, Mem. Torrey Bot. Club 1 (3): 120, 1890 (Spruce
1890).
Frullania pilibracteola S.Hatt., J. Hattori Bot. Lab. 43: 428, 1977 [1978] (Hattori
1977b).
Frullania pilistipula Steph., Sp. Hepat. (Stephani) 4: 648, 1911 (Stephani 1911e).
Frullania ramuligera (Nees) Mont., Ann. Sci. Nat. Bot. (sér. 2) 18: 14, 1842
(Montagne 1842b). Bas.: Jungermannia ramuligera Nees, Enum. Pl. Crypt. Javae:
52, 1830 (Nees 1830).
Frullania subpilibracteola S.Hatt., J. Hattori Bot. Lab. 43: 434, 1977 [1978] (Hattori 1977b).
Frullania subtilissima (Nees ex Mont.) Lindenb., Syn. Hepat. 3: 443, 1845
(Gottsche et al. 1845b). Bas.: Frullania atrata β subtilissima Nees ex Mont., Ann.
Sci. Nat. Bot. (sér. 2) 14: 333, 1840 (Montagne 1840a).
Frullania vandenberghenii Pócs, Acta Bot. Acad. Sci. Hung. 25 (3/4): 229, 1979
[1980] (Bizot and Pócs 1979). Nom. nov. pro Frullania epiphylla Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 46 (1/2): 30, 1976 (Vanden Berghen 1976b),
nom. illeg.
grp. Diastaloba I
** Frullania akiyamae S.Hatt., J. Hattori Bot. Lab. 60: 240, 1986 (Hattori 1986b).
** Frullania apiculata (Reinw., Blume et Nees) Nees, Syn. Hepat. 3: 452, 1845
(Gottsche et al. 1845b). Bas.: Jungermannia apiculata Reinw., Blume et Nees,
Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 222, 1824
[1825] (Reinwardt et al. 1824a).
** Frullania apiculata var. goebelii Schifn., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 60 (2): 222, 1893 (Schifner 1893a).
*** Frullania armitiana Steph., Sp. Hepat. (Stephani) 4: 538, 1911 (Stephani 1911e).
268
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Frullania armitiana var. inlexula S.Hatt., J. Hattori Bot. Lab. 65: 415, 1988 (Hattori 1988a).
** Frullania aterrima (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 3:
450, 1845 (Gottsche et al. 1845b). Bas.: Jungermannia aterrima Hook.f. et Taylor,
London J. Bot. 3: 395, 1844 (Hooker and Taylor 1844a).
*** Frullania attenuata Steph., Sp. Hepat. (Stephani) 4: 538, 1911 (Stephani 1911e).
*** Frullania bella Steph., Sp. Hepat. (Stephani) 4: 643, 1911 (Stephani 1911e).
** Frullania changii S.Hatt. et C.Gao, J. Jap. Bot. 60 (1): 1, 1985 (Hattori and Gao 1985).
* Frullania claviloba Steph., Sp. Hepat. (Stephani) 4: 651, 1911 (Stephani 1911e). 187
*** Frullania colliculosa von Konrat, Braggins, Hentschel et Heinrichs, Nova Hedwigia 91 (3/4): 494, 2010 (von Konrat et al. 2010b).
*** Frullania cordistipula (Reinw., Blume et Nees) Nees, Voy. Amér. Mérid. 7 (2): 68,
1839 (Montagne 1839a). Bas.: Jungermannia cordistipula Reinw., Blume et Nees,
Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 220, 1824
[1825] (Reinwardt et al. 1824a). 188
** Frullania cordistipula var. dentistipula S.Hatt., J. Hattori Bot. Lab. 60: 242, 1986
(Hattori 1986b).
* Frullania cordistipula var. mutica (Gottsche, Lindenb. et Nees) Schifn., Consp.
Hepat. Arch. Ind.: 323, 1898 (Schifner 1898b). Bas.: Frullania cordistipula β mutica Gottsche, Lindenb. et Nees, Syn. Hepat. 3: 454, 1845 (Gottsche et al. 1845b).
* Frullania crenatiloba Steph., Sp. Hepat. (Stephani) 6: 551, 1924 (Stephani 1924). 189
*** Frullania curvistipula Steph., Sp. Hepat. (Stephani) 4: 548, 1911 (Stephani 1911e).
** Frullania curvistipula var. falcatidentata S.Hatt., Misc. Bryol. Lichenol. 9 (6): 124,
1982 (Hattori 1982a).
** Frullania curvistipula var. lamii Verd., Ann. Bryol., Suppl. 1: 91, 1930 (Verdoorn
1930c).
** Frullania cuspidifolia Steph., Sp. Hepat. (Stephani) 4: 543, 1911 (Stephani 1911e).
* Frullania degelii S.W.Arnell, Svensk Bot. Tidskr. 53 (4): 503, 1959 (Arnell 1959).
** Frullania dentifera S.Hatt. et Streimann, J. Hattori Bot. Lab. 59: 102, 1985 (Hattori and Streimann 1985).
*** Frullania dentiloba S.Hatt., J. Jap. Bot. 50 (6): 161, 1975 (Hattori 1975a).
** Frullania durifolia Steph., Hedwigia 33 (3): 162, 1894 (Stephani 1894d).
** Frullania exilis Taylor, London J. Bot. 5: 405, 1846 (Taylor 1846b).
** Frullania gabonensis Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 46 (1/2): 81,
1976 (Vanden Berghen 1976b).
*** Frullania gracilis (Reinw., Blume et Nees) Nees, Syn. Hepat. 3: 452, 1845
(Gottsche et al. 1845b). Bas.: Jungermannia gracilis Reinw., Blume et Nees, Nova
Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 221, 1824 [1825]
(Reinwardt et al. 1824a).
187 Frullania claviloba is possibly conspeciic with Frullania gracilis (Söderström et al. 2010a).
188 Frullania cordistipula is a species complex also including Frullania serrata.
189 Frullania crenatiloba is conspeciic with Frullania apiculata in Verdoorn (1930c), but it was accepted
by So and Wang (2006).
World checklist of hornworts and liverworts
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269
Frullania gracilis var. brevior (Gottsche, Lindenb. et Nees) Schifn., Consp. Hepat.
Arch. Ind.: 327, 1898 (Schifner 1898b). Bas.: Frullania gracilis β brevior Gottsche,
Lindenb. et Nees, Syn. Hepat. 3: 453, 1845 (Gottsche et al. 1845b).
Frullania gracilis var. vittata S.Hatt., J. Hattori Bot. Lab. 60: 243, 1986 (Hattori
1986b).
Frullania gracilis subsp. zennoskei S.Hatt. et haithong, J. Hattori Bot. Lab. 44:
183, 1978 (Hattori and haithong 1978b).
Frullania hasskarliana Lindenb., Syn. Hepat. 3: 453, 1845 (Gottsche et al. 1845b).
Frullania hasskarliana var. gracilis S.Hatt., J. Hattori Bot. Lab. 60: 243, 1986
(Hattori 1986b).
Frullania hasskarliana var. parvidentata S.Hatt., J. Hattori Bot. Lab. 60: 245, 1986
(Hattori 1986b).
Frullania hattorii von Konrat et Braggins, New Zealand J. Bot. 41 (1): 56, 2003
(von Konrat and Braggins 2003).
Frullania hodgsoniae von Konrat, Braggins, Hentschel et Heinrichs, Nova Hedwigia
91 (3/4): 492, 2010 (von Konrat et al. 2010b). Nom. nov. pro Frullania aterrima
var. lepida E.A.Hodgs., Trans. & Proc. Roy. Soc. New Zealand 77 (3): 386, 1949
(Hodgson 1949).
Frullania hottana S.Hatt., J. Hattori Bot. Lab. 40: 479, 1976 (Hattori 1976d).
Frullania inconstans Verd., Ann. Bryol., Suppl. 1: 83, 1930 (Verdoorn 1930c).
Frullania inconstans var. grossedentata Kamim. et S.Hatt., J. Hattori Bot. Lab. 37:
530, 1973 (Hattori and Kamimura 1973).
Frullania johnsonii Steph., Hedwigia 33 (3): 163, 1894 (Stephani 1894d).
Frullania macgregorii Steph., Hedwigia 33 (3): 154, 1894 (Stephani 1894d).
Frullania macgregorii var. rostellula (S.Hatt.) S.Hatt., J. Hattori Bot. Lab. 51: 220,
1982 (Hattori 1982d). Bas.: Frullania reimersii var. rostellula S.Hatt., J. Hattori
Bot. Lab. 38: 258, 1974 (Hattori 1974c).
Frullania madens Steph., Sp. Hepat. (Stephani) 6: 553, 1924 (Stephani 1924).
Frullania mehrana S.Hatt., Recent Adv. Bot.: 66, 1976 (Hattori 1976e).
Frullania motoyana Steph., Sp. Hepat. (Stephani) 4: 646, 1911 (Stephani 1911e).
Frullania multilacera Steph., Sp. Hepat. (Stephani) 4: 650, 1911 (Stephani 1911e). 190
Frullania multilacera subsp. gracilior S.Hatt., Mem. New York Bot. Gard. 45: 547,
1987 (Hattori 1987a).
Frullania multilacera var. lacerissima S.Hatt., J. Hattori Bot. Lab. 39: 294, 1975
(Hattori 1975d).
Frullania multilaceroides S.Hatt., Mem. New York Bot. Gard. 45: 549, 1987 (Hattori 1987a).
Frullania neosheana S.Hatt., J. Hattori Bot. Lab. 45: 350, 1979 (Hattori 1979b).
Frullania papillata Steph., Sp. Hepat. (Stephani) 4: 615, 1911 (Stephani 1911e).
Frullania pulogensis Steph., Sp. Hepat. (Stephani) 4: 545, 1911 (Stephani 1911e).
Frullania purpurea Steph., Sp. Hepat. (Stephani) 4: 626, 1911 (Stephani 1911e).
190 Frullania multilacera is possibly conspeciic with Frullania vaga.
270
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Frullania reimersii Verd., Ann. Bryol., Suppl. 1: 84, 1930 (Verdoorn 1930c).
** Frullania saepisdentata S.Hatt. et Streimann, J. Hattori Bot. Lab. 59: 116, 1985
(Hattori and Streimann 1985).
** Frullania schifneri Verd., Ann. Bryol. 2: 150, 1929 (Verdoorn 1929a).
** Frullania schusterana S.Hatt., J. Hattori Bot. Lab. 36: 411, 1972 [1973] (Hattori
1972b).
** Frullania seriatifolia Steph., Hedwigia 33 (3): 167, 1894 (Stephani 1894d).
** Frullania serrata Gottsche, Syn. Hepat. 3: 453, 1845 (Gottsche et al. 1845b).
** Frullania serrata var. ceramensis S.Hatt., J. Hattori Bot. Lab. 60: 247, 1986 (Hattori
1986b).
** Frullania serrata subsp. grolleana (S.Hatt.) S.Hatt., J. Hattori Bot. Lab. 51: 225,
1982 (Hattori 1982d). Bas.: Frullania grolleana S.Hatt., J. Hattori Bot. Lab. 36:
416, 1972 [1973] (Hattori 1972b).
** Frullania serrata var. hamatispina (S.Hatt.) S.Hatt., J. Hattori Bot. Lab. 51: 225,
1982 (Hattori 1982d). Bas.: Frullania serrata subsp. hamatispina S.Hatt., J. Hattori Bot. Lab. 38: 262, 1974 (Hattori 1974c).
** Frullania serrata var. pertenuis (Nees) Schifn., Consp. Hepat. Arch. Ind.: 342,
1898 (Schifner 1898b). Bas.: Jungermannia cordistipula γ pertenuis Nees, Enum.
Pl. Crypt. Javae: 49, 1830 (Nees 1830).
** Frullania serrata subsp. spinistipula S.Hatt., J. Hattori Bot. Lab. 51: 225, 1982
(Hattori 1982d). Nom. nov. pro Frullania spinistipula S.Hatt., J. Hattori Bot. Lab.
36: 413, 1972 [1973] (Hattori 1972b), nom. illeg.
** Frullania setacea S.Hatt., J. Hattori Bot. Lab. 65: 447, 1988 (Hattori 1988a).
** Frullania sheana S.Hatt., J. Hattori Bot. Lab. 45: 356, 1979 (Hattori 1979b).
** Frullania simmondsii Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2): 109,
1914 (Stephani and Watts 1914).
*** Frullania sinuata Sande Lac., Ned. Kruidk. Arch. 3: 424, 1854 [1855] (Sande
Lacoste 1854).
** Frullania steereana S.Hatt., Mem. New York Bot. Gard. 45: 553, 1987 (Hattori
1987a).
** Frullania stipatiloba Steph., Hedwigia 33 (3): 168, 1894 (Stephani 1894d).
** Frullania subdentata Steph., Sp. Hepat. (Stephani) 4: 545, 1911 (Stephani 1911e).
** Frullania subdentata var. latistipula (S.Hatt.) S.Hatt., J. Hattori Bot. Lab. 51: 228,
1982 (Hattori 1982d). Bas.: Frullania curvistipula var. latistipula S.Hatt., J. Hattori
Bot. Lab. 44: 530, 1978 (Hattori 1978b).
** Frullania submultilacera S.Hatt. et Koike, J. Hattori Bot. Lab. 75: 190, 1994
(Koike 1994).
** Frullania subocellata S.Hatt., J. Hattori Bot. Lab. 60: 248, 1986 (Hattori 1986b).
*** Frullania taxodiocola R.M.Schust., Phytologia 53 (5): 364, 1983 (Schuster 1983b).
*** Frullania ternatensis Gottsche, Syn. Hepat. 4: 465, 1846 (Gottsche et al. 1846).
** Frullania ternatensis var. non-appendiculata S.Hatt., J. Hattori Bot. Lab. 38: 174,
1974 (Hattori 1974a).
World checklist of hornworts and liverworts
271
*** Frullania trichodes Mitt., Bonplandia 10 (2): 19, 1862 (Mitten 1862).
** Frullania vaga Mitt., Fl. vit.: 418, 1871 [1873] (Mitten 1871).
*** Frullania vaginata (Sw.) Nees, Syn. Hepat. 4: 465, 1846 (Gottsche et al. 1846).
Bas.: Jungermannia vaginata Sw., Meth. Musc.: 35, 1781 (Swartz 1781).
* Frullania vaginata var. nigricans (Gottsche, Lindenb. et Nees) Schifn., Consp.
Hepat. Arch. Ind.: 348, 1898 (Schifner 1898b). Bas.: Frullania vaginata β nigricans Gottsche, Lindenb. et Nees, Syn. Hepat. 4: 465, 1846 (Gottsche et al. 1846).
* Frullania van-zantenii Kamim. et S.Hatt., J. Hattori Bot. Lab. 37: 528, 1973
(Hattori and Kamimura 1973). 191
** Frullania venusta S.Hatt., J. Hattori Bot. Lab. 38: 217, 1974 (Hattori 1974d).
** Frullania verdoorniana S.Hatt., J. Hattori Bot. Lab. 37: 122, 1973 (Hattori 1973b).
*** Frullania vitalii Yuzawa et S.Hatt., J. Jap. Bot. 63 (1): 30, 1988 (Yuzawa and Hattori
1988a).
** Frullania vittata S.Hatt., J. Hattori Bot. Lab. 38: 270, 1974 (Hattori 1974c).
** Frullania vivipara Pócs, Fieldiana, Bot. (n.ser.) 47: 151, 2008 (Pócs 2008a).
** Frullania wairua von Konrat et Braggins, New Zealand J. Bot. 43 (4): 886, 2005
(von Konrat and Braggins 2005).
** Frullania warnckeana S.Hatt., J. Hattori Bot. Lab. 38: 213, 1974 (Hattori 1974d).
** Frullania warnckeana var. dentosa S.Hatt., Misc. Bryol. Lichenol. 7 (8): 162, 1977
(Hattori 1977a).
***
***
**
***
***
**
**
grp. Diastaloba II
Frullania baumannii S.Hatt., J. Hattori Bot. Lab. 43: 410, 1977 [1978] (Hattori
1977b).
Frullania congesta Gottsche, Lindenb. et Nees, Syn. Hepat. 3: 451, 1845 (Gottsche
et al. 1845b). Nom. nov. pro Jungermannia congesta Hook.f. et Taylor, London J.
Bot. 3: 396, 1844 (Hooker and Taylor 1844a), nom. illeg.
Frullania ocellata S.Hatt. et Kamim., J. Hattori Bot. Lab. 37: 531, 1973 (Hattori
and Kamimura 1973).
Frullania ptychantha Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 257, 1843 (Montagne
1843).
Frullania repandistipula Sande Lac., Ned. Kruidk. Arch. 3: 422, 1854 [1855]
(Sande Lacoste 1854).
Frullania repandistipula subsp. queenslandica S.Hatt., Mem. New York Bot. Gard.
45: 550, 1987 (Hattori 1987a).
Frullania repandistipula subsp. spinibractea S.Hatt., Bull. Natl. Sci. Mus. Tokyo,
B 1 (4): 150, 1975 (Hattori 1975f).
grp. Diastaloba III
** Frullania grossiclava Steph., Sp. Hepat. (Stephani) 4: 384, 1910 (Stephani 1910b).
191 Frullania van-zantenii is possibly conspeciic with Frullania serrata.
272
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Frullania loricata Pearson, Forh. Vidensk.-Selsk. Kristiania 1890 (2): 6, 1891
(Pearson 1891).
* Frullania loricata var. laxa Pearson, Forh. Vidensk.-Selsk. Kristiania 1890 (2): 8,
1891 (Pearson 1891).
*** Frullania usambarana Schifn., Hedwigia 33 (3): 160, 1894 (Stephani 1894d).
** Frullania usambarana var. reducta Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 46
(1/2): 91, 1976 (Vanden Berghen 1976b).
*
***
***
***
***
*
**
*
grp. Diastaloba IV
Frullania brunea (Spreng.) Drège, Flora, Beig. 26: 186, 1843 (Drège 1843). Bas.:
Jungermannia brunea Spreng. Syst. Veg. (ed. 16) [Sprengel] 4 (2): 325, 1827
(Sprengel 1827b). 192
Frullania caulisequa (Nees) Mont., Ann. Sci. Nat. Bot. (sér. 2) 12: 51, 1839 (Montagne 1839c). Bas.: Jungermannia caulisequa Nees, Fl. Bras. (Martius) 1 (1): 373,
1833 (Nees 1833a).
Frullania grossifolia Steph., Sp. Hepat. (Stephani) 4: 633, 1911 (Stephani 1911e).
Frullania hypoleuca Nees, Observ. bot.: 470, 1843 (Gottsche et al. 1843).
Frullania lindenbergii Lehm., Nov. Stirp. Pug. 8: 17, 1844 (Lehmann 1844).
Frullania lindenbergii var. fusca Gottsche, Lindenb. et Nees, Syn. Hepat. 5: 780,
1847 (Gottsche et al. 1847).
Frullania ponapensis S.Hatt. et Koike, J. Hattori Bot. Lab. 75: 186, 1994 (Koike 1994).
Frullania tricarinata Sande Lac., Plagiochila Sandei: 10, 1856 (Sande Lacoste 1856c). 193
subg. Diversitextae (Kamim.) S.Hatt., J. Hattori Bot. Lab. 59: 154, 1985 (Hattori and Lin 1985a). Bas.: Frullania subsect. Diversitextae Kamim., J. Hattori Bot.
Lab. 24: 80, 1961 (Kamimura 1961).
*** Frullania diversitexta Steph., Bull. Herb. Boissier 5 (2): 89, 1897 (Stephani 1897b).
*
*** subg. Frullania
* Frullania amamiensis Kamim., Bull. Kochi Gakuen Jun. Coll. 1: 51, 1970
(Kamimura 1970).
** Frullania amplicrania Steph., Sp. Hepat. (Stephani) 4: 404, 1910 (Stephani 1910b).
*** Frullania ampullifera J.B.Jack et Steph., Hedwigia 33 (3): 139, 1894 (Stephani
1894d).
*** Frullania anderssonii Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 30 (5): 144,
1873 (Ångström 1873).
** Frullania angustistipa Steph., Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr.
81: 297, 1907 (Stephani 1907a).
192 Frullania brunea belongs to the Frullania lindenbergii complex (Vanden Berghen 1976b).
193 Frullania tricarinata is possibly conspeciic with Frullania hypoleuca (Söderström et al. 2010a).
World checklist of hornworts and liverworts
273
** Frullania aposinensis S.Hatt. et P.J.Lin, J. Hattori Bot. Lab. 59: 131, 1985 (Hattori
and Lin 1985a). Nom. nov. pro Frullania chinensis Steph., Sp. Hepat. (Stephani) 4:
469, 1911 (Stephani 1911e), nom. illeg.
** Frullania appendistipula S.Hatt., J. Hattori Bot. Lab. 36: 424, 1972 [1973] (Hattori 1972b).
** Frullania appendistipula var. spinifera S.Hatt., J. Hattori Bot. Lab. 38: 226, 1974
(Hattori 1974c).
** Frullania auriculata S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 11 (1): 11, 1985
(Hattori 1985).
** Frullania benjaminiana Inoue, Bull. Natl. Sci. Mus. Tokyo, B 1 (3): 109, 1975
(Hattori 1975b).
** Frullania bergmanii S.Hatt., J. Hattori Bot. Lab. 38: 192, 1974 (Hattori 1974d).
*** Frullania berthoumieui Steph., Hedwigia 33 (3): 140, 1894 (Stephani 1894d).
** Frullania bhutanensis S.Hatt., Fl. E. Himalaya 2: 232, 1971 (Hattori 1971a).
** Frullania blastopetala S.Hatt., J. Hattori Bot. Lab. 57: 407, 1984 (Hattori 1984a).
*** Frullania bonincola S.Hatt., J. Hattori Bot. Lab. 44: 551, 1978 (Hattori 1978b).
Nom. nov. pro Frullania viridis Horik., Sci. Rep. Tôhoku Imp. Univ., Ser. 4, Biol.
5 (4): 646, 1929 [1930] (Horikawa 1929c), nom. illeg.
* Frullania brevicalycina Steph., Hedwigia 33 (3): 141, 1894 (Stephani 1894d).
** Frullania brittoniae A.Evans, Trans. Connecticut Acad. Arts 10 (1): 15, 1899
(Evans 1899).
*** Frullania bullata Steph., Sp. Hepat. (Stephani) 4: 371, 1910 (Stephani 1910b).
*** Frullania cafraria Steph., Hedwigia 33 (3): 141, 1894 (Stephani 1894d).
*** Frullania calcarata Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 30 (5): 137,
1873 (Ångström 1873).
** Frullania carrii Kamim. et S.Hatt., J. Hattori Bot. Lab. 37: 520, 1973 (Hattori
and Kamimura 1973).
** Frullania chenii S.Hatt. et P.J.Lin, J. Jap. Bot. 60 (4): 106, 1985 (Hattori and Lin
1985b).
** Frullania chilensis Steph., Hedwigia 33 (3): 142, 1894 (Stephani 1894d).
*** Frullania chodatii Beauverd, Sp. Hepat. (Stephani) 6: 533, 1924 (Stephani 1924).
Based on: Frullania grandistipula Steph., Sp. Hepat. (Stephani) 6: 533, 1924
(Stephani 1924), nom. inval.
*** Frullania cobrensis Gottsche, Hedwigia 33 (3): 142, 1894 (Stephani 1894d).
** Frullania consociata Steph., Sp. Hepat. (Stephani) 4: 461, 1911 (Stephani 1911e).
* Frullania contracta Steph., Sp. Hepat. (Stephani) 4: 469, 1911 (Stephani 1911e). 194
* Frullania cornuta Steph., Sp. Hepat. (Stephani) 4: 467, 1911 (Stephani 1911e). 195
*** Frullania crassitexta Steph., Sp. Hepat. (Stephani) 4: 423, 1910 (Stephani 1910b).
*** Frullania crispiplicata Yuzawa et S.Hatt., J. Jap. Bot. 58 (2): 43, 1983 (Yuzawa and
Hattori 1983).
194 Frullania contracta is conspeciic with Frullania squarrosa in Verdoorn (1930c), but it was accepted
by Hattori (1986e).
195 Frullania cornuta is closely related to the very variable Frullania ornithocephala.
274
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Frullania cristata S.Hatt., J. Hattori Bot. Lab. 49: 165, 1981 (Hattori 1981a).
* Frullania cuneiloba Nees, Syn. Hepat. 3: 427, 1845 (Gottsche et al. 1845b).
** Frullania cyparioides (Schwägr.) Nees, Naturgesch. Eur. Leberm. 3: 210, 1838
(Nees 1838b). Bas.: Jungermannia cyparioides Schwägr., Hist. Musc. Hepat.
Prodr.: 14, 1814 (Schwägrichen 1814).
*** Frullania davurica Hampe ex Gottsche, Lindenb. et Nees, Syn. Hepat. 3: 422,
1845 (Gottsche et al. 1845b).
** Frullania debilis Steph. ex S.Hatt., J. Hattori Bot. Lab. 38: 190, 1974 (Hattori
1974d).
** Frullania deppii Lehm., Nov. Stirp. Pug. 8: 15, 1844 (Lehmann 1844).
** Frullania diptera (Lehm.) Drège, Flora, Beig. 26: 186, 1843 (Drège 1843). Bas.:
Jungermannia diptera Lehm., Linnaea 9 (4): 425, 1835 (Lehmann 1835).
*** Frullania duthiana Steph., Sp. Hepat. (Stephani) 4: 351, 1910 (Stephani 1910b).
** Frullania duthiana var. appendiculata S.Hatt., Fl. E. Himalaya 2: 234, 1971 (Hattori 1971a).
** Frullania duthiana var. laevis S.Hatt., Bull. Univ. Mus. Univ. Tokyo 8: 233, 1975
(Hattori 1975e).
** Frullania duthiana var. szechuanensis S.Hatt. et C.Gao, J. Jap. Bot. 60 (1): 2, 1985
(Hattori and Gao 1985).
** Frullania echinantha S.Hatt., J. Hattori Bot. Lab. 38: 233, 1974 (Hattori 1974c).
** Frullania echinatella S.Hatt., J. Hattori Bot. Lab. 65: 423, 1988 (Hattori 1988a).
* Frullania elegans Lehm., Nov. Stirp. Pug. 10: 16, 1857 (Lehmann 1857).
*** Frullania elephantum S.Hatt., J. Hattori Bot. Lab. 43: 416, 1977 [1978] (Hattori
1977b).
** Frullania epiphylla S.Hatt., J. Hattori Bot. Lab. 38: 235, 1974 (Hattori 1974c).
* Frullania epiphylla subsp. ijiensis S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 11 (1):
14, 1985 (Hattori 1985).
** Frullania erostrata S.Hatt., J. Hattori Bot. Lab. 38: 237, 1974 (Hattori 1974c).
* Frullania esenbeckiana Beauverd, Sp. Hepat. (Stephani) 6: 570, 1924 (Stephani
1924). Nom. nov. pro Frullania grossiloba Steph., Sp. Hepat. (Stephani) 4: 531,
1911 (Stephani 1911e), nom. illeg.
** Frullania evelynae S.Hatt. et haithong, J. Hattori Bot. Lab. 44: 179, 1978 (Hattori and haithong 1978b).
* Frullania evelynae var. devendrae Sushil K.Singh et Barbhuiya, Taiwania 57 (2):
109, 2012 (Singh and Barbhuiya 2012).
* Frullania evelynae var. srivastavae Sushil K.Singh et Barbhuiya, Taiwania 57 (2):
112, 2012 (Singh and Barbhuiya 2012).
*** Frullania eymae S.Hatt., J. Hattori Bot. Lab. 39: 284, 1975 (Hattori 1975d).
** Frullania eymae var. crispidentata S.Hatt. et Streimann, J. Hattori Bot. Lab. 59:
105, 1985 (Hattori and Streimann 1985).
*** Frullania falciloba Lehm., Nov. Stirp. Pug. 8: 20, 1844 (Lehmann 1844).
** Frullania falsicornuta S.Hatt., J. Hattori Bot. Lab. 60: 212, 1986 (Hattori 1986e).
** Frullania fauriana Steph., Hedwigia 33 (3): 144, 1894 (Stephani 1894d).
World checklist of hornworts and liverworts
275
** Frullania fengyangshanensis R.L.Zhu et M.L.So, Bryologist 100 (3): 356, 1997
(Zhu and So 1997b).
** Frullania ferdinandi-muelleri Steph., Sp. Hepat. (Stephani) 4: 417, 1910 (Stephani
1910b).
* Frullania lexuosa S.Hatt., J. Hattori Bot. Lab. 54: 146, 1983 (Hattori 1983). 196
** Frullania fuscovirens Steph., Sp. Hepat. (Stephani) 4: 401, 1910 (Stephani 1910b).
** Frullania fuscovirens var. gemmipara (R.M.Schust. et S.Hatt.) S.Hatt. et P.J.Lin, J.
Hattori Bot. Lab. 59: 135, 1985 (Hattori and Lin 1985a). Bas.: Frullania gemmipara R.M.Schust. et S.Hatt., J. Hattori Bot. Lab. 44: 547, 1978 (Hattori 1978b).
** Frullania gaoligongensis X.L.Bai et C.Gao, Hikobia 13 (1): 87, 1999 (Bai and Gao
1999).
** Frullania gemmulosa S.Hatt. et haithong, J. Hattori Bot. Lab. 43: 449, 1977
[1978] (Hattori et al. 1977).
*** Frullania gigantea Steph., Sp. Hepat. (Stephani) 4: 467, 1911 (Stephani 1911e).
** Frullania giraldiana C.Massal., Hepat. Shen-si: 41, 1897 (Massalongo 1897). 197
** Frullania giraldiana var. handelii (Verd.) S.Hatt., J. Hattori Bot. Lab. 36: 123,
1972 [1973] (Hattori 1972c). Bas.: Frullania nepalensis var. handelii Verd., Symb.
Sin. 5: 41, 1930 (Nicholson et al. 1930).
** Frullania globosa S.Hatt. et Streimann, J. Hattori Bot. Lab. 59: 107, 1985 (Hattori
and Streimann 1985).
** Frullania grandilobula S.Hatt. et Piippo, Acta Bot. Fenn. 133: 45, 1986 (Hattori
and Piippo 1986).
*** Frullania grandistipula Lindenb., Syn. Hepat. 3: 430, 1845 (Gottsche et al. 1845b).
** Frullania hainanensis S.Hatt. et P.J.Lin, J. Jap. Bot. 61 (10): 307, 1986 (Hattori
and Lin 1986).
** Frullania hamatiloba Steph., Sp. Hepat. (Stephani) 4: 400, 1910 (Stephani 1910b).
** Frullania handelii Verd., Symb. Sin. 5: 36, 1930 (Nicholson et al. 1930).
** Frullania handel-mazzettii S.Hatt., J. Hattori Bot. Lab. 49: 150, 1981 (Hattori 1981a).
** Frullania hattoriantha Udar et V.Nath, Misc. Bryol. Lichenol. 9 (2): 44, 1981
(Udar and Nath 1981).
* Frullania hebridensis Steph., Sp. Hepat. (Stephani) 4: 469, 1911 (Stephani 1911e). 198
** Frullania hicksiae S.Hatt., Cryptog. Bryol. Lichénol. 5 (1/2): 182, 1984 (Hattori
1984c).
** Frullania higuchii Yuzawa, Koike et S.Hatt., J. Hattori Bot. Lab. 75: 194, 1994
(Yuzawa and Koike 1994).
** Frullania hiroshii S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 6 (1): 34, 1980 (Hattori
1980b).
* Frullania hirtilora Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 35, 1884
(Spruce 1884).
196 Frullania lexuosa is possibly conspeciic with Frullania rubella.
197 Frullania giraldiana belongs to the Frullania nepalensis complex (Hattori 1973b).
198 Frullania hebridensis belongs to the Frullania ericoides species complex (Verdoorn 1930c, 1930a).
276
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*** Frullania howeana Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2): 107,
1914 (Stephani and Watts 1914).
** Frullania ignatovii Sofronova, Mamontov et Potemkin, Novosti Sist. Nizš. Rast.
47: 335, 2013 (Sofronova et al. 2013).
** Frullania incisoduthiana S.Hatt., J. Hattori Bot. Lab. 46: 391, 1979 (Mizutani 1979c).
* Frullania incisoduthiana var. parva S.Hatt., J. Hattori Bot. Lab. 46: 391, 1979
(Mizutani 1979c).
* Frullania incisostipula Steph., Sp. Hepat. (Stephani) 6: 541, 1924 (Stephani 1924).
** Frullania inlexiloba S.Hatt., J. Hattori Bot. Lab. 57: 415, 1984 (Hattori 1984a).
** Frullania inouei S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 6 (1): 36, 1980 (Hattori
1980b).
*** Frullania irregularis S.Hatt. et Piippo, Acta Bot. Fenn. 133: 46, 1986 (Hattori and
Piippo 1986).
*** Frullania jackii Gottsche, Hepat. Eur., Leberm. 29-30: no. 294, 1863 (Gottsche
and Rabenhorst 1863).
** Frullania jacobsii S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 12 (4): 129, 1986 (Hattori
1986c).
* Frullania jacquinotii Gottsche, J. Bot. (Morot) 12: 138, 1898 (Bescherelle 1898).
** Frullania jovetiana von Konrat et Hentschel, Phytotaxa 220 (2): 135, 2015
(Hentschel et al. 2015). Based on: Frullania pseudericoides S.Hatt., Bull. Natl. Sci.
Mus. Tokyo, B 12 (4): 132, 1986 (Hattori 1986c), nom. illeg.
** Frullania kagoshimensis Steph., Sp. Hepat. (Stephani) 4: 353, 1910 (Stephani 1910b).
** Frullania kagoshimensis subsp. hunanensis (S.Hatt.) S.Hatt. et P.J.Lin, J. Hattori Bot. Lab. 59: 137, 1985 (Hattori and Lin 1985a). Bas.: Frullania hunanensis
S.Hatt., J. Hattori Bot. Lab. 49: 152, 1981 (Hattori 1981a).
** Frullania kalimantanensis S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 12 (4): 127,
1986 (Hattori 1986c).
** Frullania kashyapii Verd., Ann. Bryol. 5: 162, 1932 (Dixon et al. 1932).
** Frullania kitagawana S.Hatt., J. Hattori Bot. Lab. 57: 417, 1984 (Hattori 1984a).
** Frullania laeviperiantha X.L.Bai et C.Gao, Phytotaxa 220 (2): 136, 2015 (Hentschel
et al. 2015). Based on: Frullania laeviperiantha X.L.Bai et C.Gao, Nova Hedwigia
70 (1/2): 135, 2000 (Bai and Gao 2000), nom. inval.
* Frullania lancistyla Steph., Sp. Hepat. (Stephani) 4: 389, 1910 (Stephani 1910b).
* Frullania latilora Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 34, 1884
(Spruce 1884).
** Frullania latogaleata Herzog, Trans. Brit. Bryol. Soc. 1 (3): 189, 1949 (Herzog 1949b).
** Frullania lepida S.Hatt. et Piippo, Acta Bot. Fenn. 133: 48, 1986 (Hattori and
Piippo 1986).
* Frullania levieri Steph., Sp. Hepat. (Stephani) 4: 388, 1910 (Stephani 1910b).
** Frullania linii S.Hatt., J. Hattori Bot. Lab. 49: 155, 1981 (Hattori 1981a).
* Frullania longistyla Yuzawa et S.Hatt., J. Jap. Bot. 63 (11): 361, 1988 (Yuzawa and
Hattori 1988b). 199
199 Frullania longistyla is closely related to Frullania brevicalycina (Yuzawa and Hattori 1988b).
World checklist of hornworts and liverworts
277
* Frullania ludoviciae Steph., Rev. Bryol. 35 (2): 29, 1908 (Stephani 1908l). 200
** Frullania lushanensis S.Hatt. et P.J.Lin, J. Hattori Bot. Lab. 59: 137, 1985 (Hattori and Lin 1985a).
* Frullania macularis Taylor, London J. Bot. 5: 403, 1846 (Taylor 1846b).
** Frullania maymyoensis Svihla, Bryologist 61 (4): 376, 1958 [1959] (Svihla 1958).
** Frullania microauriculata Verd., Ann. Bryol. 2: 126, 1929 (Verdoorn 1929a).
** Frullania microauriculata var. rotundior Verd., Ann. Jard. Bot. Buitenzorg 40:
141, 1929 (Verdoorn 1929b).
** Frullania microrhyncha L.Clark et Svihla, Bryologist 53 (1): 63, 1950 (Clark and
Svihla 1950).
** Frullania mizoramensis Sushil K.Singh et Barbhuiya, Taiwania 57 (2): 106, 2012
(Singh and Barbhuiya 2012).
* Frullania montana Steph., Sp. Hepat. (Stephani) 4: 455, 1911 (Stephani 1911e). 201
** Frullania multituberculata Hentschel et von Konrat, Phytotaxa 220 (2): 136, 2015
(Hentschel et al. 2015). Nom. nov. pro Frullania kalimantanensis Piippo et S.Hatt.,
J. Hattori Bot. Lab. 72: 117, 1992 (Piippo and Tan 1992), nom. illeg.
** Frullania mutilata Steph., Sp. Hepat. (Stephani) 4: 673, 1911 (Stephani 1911e).
** Frullania nadeaudii Steph., Sp. Hepat. (Stephani) 4: 465, 1911 (Stephani 1911e).
*** Frullania nepalensis (Spreng.) Lehm. et Lindenb., Syn. Hepat. 3: 422, 1845
(Gottsche et al. 1845b). Bas.: Jungermannia nepalensis Spreng. Syst. Veg. (ed. 16)
[Sprengel] 4 (2): 324, 1827 (Sprengel 1827b). 202
*** Frullania nicholsonii E.A.Hodgs., Trans. & Proc. Roy. Soc. New Zealand 77 (3):
368, 1949 (Hodgson 1949). Nom. nov. pro Frullania berggrenii W.E.Nicholson,
Bryologist 28 (2): 17, 1925 (Nicholson 1925), nom. illeg.
** Frullania nigricaulis (Reinw., Blume et Nees) Nees, Syn. Hepat. 3: 457, 1845
(Gottsche et al. 1845b). Bas.: Jungermannia nigricaulis Reinw., Blume et Nees,
Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 225, 1824
[1825] (Reinwardt et al. 1824a).
* Frullania nigricaulis var. elongata Verd., Ann. Bryol., Suppl. 1: 155, 1930 (Verdoorn 1930c).
** Frullania nivimontana S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 8 (3): 95, 1982
(Hattori 1982c).
*** Frullania nobilis Steph., Hedwigia 33 (3): 154, 1894 (Stephani 1894d).
** Frullania nobilis var. cochleata (Steph.) S.Hatt., J. Hattori Bot. Lab. 59: 111, 1985
(Hattori and Streimann 1985). Bas.: Frullania cochleata Steph., Sp. Hepat. (Stephani) 4: 681, 1911 (Stephani 1911e).
** Frullania novocurvirostris S.Hatt., J. Hattori Bot. Lab. 49: 370, 1981 (Hattori
1981b). Nom. nov. pro Frullania curvirostris J.B.Jack et Steph., Hedwigia 33 (3):
143, 1894 (Stephani 1894d), nom. illeg.
200 Frullania ludoviciae seems to be close to Frullania pusilla (Hattori 1986e).
201 Frullania montana is possibly conspeciic with Frullania relexistipula.
202 Frullania nepalensis is a species complex (Hattori 1973b).
278
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Frullania oahuensis Hampe ex Gottsche, Lindenb. et Nees, Observ. bot.: 471,
1843 (Gottsche et al. 1843).
** Frullania obovata S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 8 (3): 97, 1982 (Hattori
1982c).
** Frullania obtusangula Hentschel et von Konrat, Phytotaxa 220 (2): 137, 2015
(Hentschel et al. 2015).
** Frullania okinawensis Kamim., Misc. Bryol. Lichenol. 9 (4): 90, 1982 (Kamimura
1982).
** Frullania orbicularis Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 227, 1869
(Austin 1869).
*** Frullania orientalis Sande Lac., Plagiochila Sandei: 10, 1856 (Sande Lacoste 1856c).
** Frullania orinocensis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 30, 1884
(Spruce 1884).
*** Frullania ornithocephala (Reinw., Blume et Nees) Nees, Syn. Hepat. 3: 425, 1845
(Gottsche et al. 1845b). Bas.: Jungermannia ornithocephala Reinw., Blume et Nees,
Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 216, 1824
[1825] (Reinwardt et al. 1824a).
* Frullania ornithocephala var. major (Nees) Schifn., Consp. Hepat. Arch. Ind.:
336, 1898 (Schifner 1898b). Bas.: Jungermannia ornithocephala β major Nees,
Enum. Pl. Crypt. Javae: 47, 1830 (Nees 1830).
** Frullania ornithocephala var. pilosa Verd., Ann. Bryol. 2: 129, 1929 (Verdoorn 1929a).
** Frullania ornithocephala var. tuberculosa S.Hatt., J. Hattori Bot. Lab. 38: 179,
1974 (Hattori 1974a).
** Frullania pachyderma S.Hatt., J. Hattori Bot. Lab. 44: 525, 1978 (Hattori 1978b).
** Frullania pallidevirens Steph., Sp. Hepat. (Stephani) 4: 454, 1911 (Stephani 1911e).
** Frullania pariharii S.Hatt. et haithong, J. Jap. Bot. 53 (5): 130, 1978 (Hattori
and haithong 1978a).
* Frullania parvifolia Steph., Sp. Hepat. (Stephani) 4: 354, 1910 (Stephani 1910b).
** Frullania pauciramea Steph., Sp. Hepat. (Stephani) 4: 458, 1911 (Stephani 1911e).
** Frullania pauciramea var. pauciramella S.Hatt. et Piippo, Acta Bot. Fenn. 133: 50,
1986 (Hattori and Piippo 1986).
** Frullania paucirameoides S.Hatt. et Piippo, Acta Bot. Fenn. 133: 51, 1986 (Hattori and Piippo 1986).
** Frullania pedicellata Steph., Bull. Herb. Boissier 5 (2): 90, 1897 (Stephani 1897b).
*** Frullania physantha Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 121, 1860 [1861] (Mitten 1860c).
** Frullania piptophylla S.Hatt., J. Hattori Bot. Lab. 47: 87, 1980 (Hattori 1980d).
** Frullania piptophylla var. minor S.Hatt., J. Hattori Bot. Lab. 60: 247, 1986 (Hattori 1986b).
** Frullania piptophylloides S.Hatt., J. Hattori Bot. Lab. 47: 90, 1980 (Hattori 1980d).
* Frullania plicata Hentschel et von Konrat, Phytotaxa 220 (2): 137, 2015 (Hentschel
et al. 2015). Nom. nov. pro Frullania acutiloba Gerola, Lav. Bot. Ist. Bot. Univ.
Padova 12: 477, 1947 (Gerola 1947), nom. illeg.
World checklist of hornworts and liverworts
279
** Frullania pocsantha haithong et S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 3 (4):
149, 1977 (haithong and Hattori 1977).
** Frullania pran-nathii M.Dey et D.K.Singh, J. Jap. Bot. 83 (5): 281, 2008 (Dey
and Singh 2008).
** Frullania pringlei Fulford et Sharp, Mem. New York Bot. Gard. 63: 45, 1990
(Fulford and Sharp 1990). Nom. nov. pro Frullania spicata Steph., Sp. Hepat.
(Stephani) 4: 392, 1910 (Stephani 1910b), nom. illeg.
** Frullania prominula S.Hatt. et Streimann, J. Hattori Bot. Lab. 59: 112, 1985
(Hattori and Streimann 1985).
** Frullania propaginea S.Hatt. et Streimann, J. Hattori Bot. Lab. 59: 114, 1985
(Hattori and Streimann 1985).
* Frullania pseudericoides S.Hatt., J. Hattori Bot. Lab. 51: 256, 1982 (Hattori
1982d). Nom. nov. pro Frullania sharpii subsp. subrostrata S.Hatt., J. Hattori Bot.
Lab. 38: 265, 1974 (Hattori 1974c).
** Frullania pseudoschensiana S.Hatt., J. Hattori Bot. Lab. 47: 101, 1980 (Hattori
1980d).
** Frullania pseudoschensiana var. darjeelingensis S.Hatt., J. Hattori Bot. Lab. 49: 149,
1981 (Hattori 1981a).
** Frullania pullei Verd., Nova Guinea 14: 542, 1930 (Verdoorn 1930b).
** Frullania pusilla Mitt., Fl. vit.: 417, 1871 [1873] (Mitten 1871).
*** Frullania pycnantha (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat.
3: 411, 1845 (Gottsche et al. 1845b). Bas.: Jungermannia pycnantha Hook.f. et
Taylor, London J. Bot. 3: 566, 1844 (Hooker and Taylor 1844d).
** Frullania queenslandica Steph., Sp. Hepat. (Stephani) 4: 424, 1910 (Stephani
1910b).
** Frullania recurvistipula S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 1 (3): 114, 1975
(Hattori 1975b).
*** Frullania relexistipula Sande Lac., Ned. Kruidk. Arch. 3: 422, 1854 [1855] (Sande Lacoste 1854).
** Frullania relexistipula var. squarrosa S.Hatt. et Piippo, Acta Bot. Fenn. 133: 54,
1986 (Hattori and Piippo 1986).
** Frullania remotidens S.Hatt., J. Hattori Bot. Lab. 51: 257, 1982 (Hattori 1982d).
* Frullania reptans Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 161, 1855 (Mitten 1855).
*** Frullania retusa Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 119, 1860 [1861] (Mitten
1860c).
** Frullania retusa var. gymnantha S.Hatt. et haithong, J. Jap. Bot. 53 (5): 131,
1978 (Hattori and haithong 1978a).
** Frullania retusa var. hirsuta S.Hatt. et haithong, J. Hattori Bot. Lab. 44: 191,
1978 (Hattori and haithong 1978b).
*** Frullania rhystocolea Herzog, Symb. Sin. 5: 39, 1930 (Nicholson et al. 1930).
** Frullania rhytidantha S.Hatt., J. Hattori Bot. Lab. 47: 97, 1980 (Hattori 1980d).
** Frullania rigida Steph., Sp. Hepat. (Stephani) 4: 371, 1910 (Stephani 1910b).
*** Frullania riparia Hampe, Nov. Stirp. Pug. 7: 14, 1838 (Lehmann 1838).
280
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Frullania rizalii Piippo et S.Hatt., J. Hattori Bot. Lab. 72: 119, 1992 (Piippo and
Tan 1992).
*** Frullania rostellata Mitt., Handb. N. Zeal. l. 2: 755, 1867 (Hooker 1867).
** Frullania rubella Gottsche, Hedwigia 28 (3): 159, 1889 (Stephani 1889d).
** Frullania rubella var. elongata (Steph.) S.Hatt., J. Hattori Bot. Lab. 54: 166, 1983
(Hattori 1983). Bas.: Frullania elongata Steph., Sp. Hepat. (Stephani) 4: 423,
1910 (Stephani 1910b).
** Frullania rupicola Steph., Sp. Hepat. (Stephani) 6: 534, 1924 (Stephani 1924).
** Frullania saipanensis S.Hatt. et Koike, J. Hattori Bot. Lab. 75: 188, 1994 (Koike 1994).
*** Frullania schensiana C.Massal., Hepat. Shen-si: 40, 1897 (Massalongo 1897).
** Frullania schusteri S.Hatt., Beih. Nova Hedwigia 90: 154, 1988 (Hattori 1988b).
** Frullania scottiana S.Hatt., Mem. New York Bot. Gard. 45: 551, 1987 (Hattori
1987a).
** Frullania setchellii Pearson, Univ. Calif. Publ. Bot. 10 (4): 326, 1923 (Pearson 1923).
** Frullania shanensis Svihla, Bryologist 60 (4): 359, 1957 (Svihla 1957).
** Frullania sharpantha Udar et Ad.Kumar, Misc. Bryol. Lichenol. 9 (9): 192, 1983
(Udar and Kumar 1983a).
** Frullania sharpii S.Hatt., J. Hattori Bot. Lab. 38: 180, 1974 (Hattori 1974a).
** Frullania sinensis Steph., Nuovo Giorn. Bot. Ital. (n.ser.) 13 (4): 349, 1906 (Levier
1906).
** Frullania sinosphaerantha S.Hatt. et P.J.Lin, J. Hattori Bot. Lab. 59: 144, 1985
(Hattori and Lin 1985a).
** Frullania sphaerantha S.Hatt., J. Hattori Bot. Lab. 47: 99, 1980 (Hattori 1980d).
** Frullania sphaerolobulata S.H.Lin, Tunghai Journal 38: 104, 1997 (Lin and Chen
1997).
*** Frullania spinifera Taylor, London J. Bot. 5: 407, 1846 (Taylor 1846b).
** Frullania spinigastria S.Hatt., J. Hattori Bot. Lab. 45: 358, 1979 (Hattori 1979b).
** Frullania spiniplica S.Hatt., J. Hattori Bot. Lab. 36: 428, 1972 [1973] (Hattori
1972b).
*** Frullania spongiosa Steph., Hedwigia 33 (3): 147, 1894 (Stephani 1894d).
** Frullania squamuligera Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 33, 1884
(Spruce 1884).
*** Frullania squarrosula (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat.
3: 412, 1845 (Gottsche et al. 1845b). Bas.: Jungermannia squarrosula Hook.f. et
Taylor, London J. Bot. 4: 88, 1845 (Hooker and Taylor 1845).
** Frullania subcaduca S.Hatt., J. Hattori Bot. Lab. 38: 267, 1974 (Hattori 1974c).
** Frullania subclavata Steph., Sp. Hepat. (Stephani) 4: 354, 1910 (Stephani 1910b).
** Frullania subnigricaulis S.Hatt., J. Hattori Bot. Lab. 37: 89, 1973 (Hattori 1973c).
** Frullania subnigricaulis var. subtruncata S.Hatt., J. Hattori Bot. Lab. 39: 308,
1975 (Hattori 1975d).
* Frullania subpedicellata S.Hatt., J. Hattori Bot. Lab. 47: 93, 1980 (Hattori 1980d). 203
** Frullania subsquarrosa S.Hatt., J. Hattori Bot. Lab. 36: 429, 1972 [1973] (Hattori 1972b).
203 Frullania subpedicellata is possibly conspeciic with Frullania pedicellata.
World checklist of hornworts and liverworts
281
** Frullania subvalida S.Hatt. et haithong, J. Jap. Bot. 53 (6): 173, 1978 (Hattori
and haithong 1978c).
*** Frullania svihlana S.Hatt., J. Hattori Bot. Lab. 54: 180, 1983 (Hattori 1983).
* Frullania taiheizana Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 241,
1934 (Horikawa 1934).
** Frullania tamsuina Steph., Sp. Hepat. (Stephani) 4: 444, 1910 (Stephani 1910b).
*** Frullania taradakensis Steph., Sp. Hepat. (Stephani) 4: 352, 1910 (Stephani
1910b).
** Frullania tenuirostris Steph., Sp. Hepat. (Stephani) 4: 462, 1911 (Stephani 1911e).
** Frullania togashiana S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 1 (3): 118, 1975
(Hattori 1975b).
** Frullania tubercularis S.Hatt. et P.J.Lin, J. Jap. Bot. 60 (4): 107, 1985 (Hattori
and Lin 1985b).
*** Frullania usamiensis Steph., Bull. Herb. Boissier 5 (2): 91, 1897 (Stephani 1897b).
** Frullania valdiviensis J.B.Jack et Steph., Hedwigia 33 (3): 149, 1894 (Stephani 1894d).
** Frullania valida Steph., Sp. Hepat. (Stephani) 4: 402, 1910 (Stephani 1910b).
** Frullania variegata Steph., Hedwigia 33 (3): 149, 1894 (Stephani 1894d).
* Frullania victoriensis Steph., Sp. Hepat. (Stephani) 4: 418, 1910 (Stephani 1910b).
*** Frullania vittiana S.Hatt., Bryologist 90 (4): 368, 1987 [1988] (Hattori 1987b).
** Frullania wangii S.Hatt. et P.J.Lin, J. Hattori Bot. Lab. 59: 146, 1985 (Hattori
and Lin 1985a).
*** Frullania yuennanensis Steph., Hedwigia 33 (3): 161, 1894 (Stephani 1894d).
** Frullania yuennanensis var. siamensis (N.Kitag., haithong et S.Hatt.) S.Hatt. et
P.J.Lin, J. Hattori Bot. Lab. 59: 148, 1985 (Hattori and Lin 1985a). Bas.: Frullania siamensis N.Kitag., haithong et S.Hatt., J. Hattori Bot. Lab. 43: 452, 1977
[1978] (Hattori et al. 1977).
** Frullania yuzawana S.Hatt., J. Hattori Bot. Lab. 49: 157, 1981 (Hattori 1981a).
** Frullania zangii S.Hatt. et P.J.Lin, J. Hattori Bot. Lab. 59: 149, 1985 (Hattori and
Lin 1985a).
** Frullania zennoskeana S.Hatt., J. Jap. Bot. 59 (10): 308, 1984 (Hattori 1984b).
** sect. Acutilobae Verd., Ann. Bryol., Suppl. 1: 44, 1930 (Verdoorn 1930c).
** Frullania allanii E.A.Hodgs., Trans. & Proc. Roy. Soc. New Zealand 77 (3): 371,
1949 (Hodgson 1949).
*** Frullania clavata (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 3:
428, 1845 (Gottsche et al. 1845b). Bas.: Jungermannia clavata Hook.f. et Taylor,
London J. Bot. 4: 88, 1845 (Hooker and Taylor 1845).
** Frullania hamaticoma Steph., Hedwigia 28 (3): 158, 1889 (Stephani 1889d).
*** Frullania monocera (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat.
3: 418, 1845 (Gottsche et al. 1845b). Bas.: Jungermannia monocera Hook.f. et
Taylor, London J. Bot. 4: 89, 1845 (Hooker and Taylor 1845).
** Frullania monocera var. acutiloba (Mitt.) Hentschel et von Konrat, Phytotaxa 220
(2): 136, 2015 (Hentschel et al. 2015). Bas.: Frullania acutiloba Mitt., J. Proc.
Linn. Soc., Bot. 5 (18): 120, 1860 [1861] (Mitten 1860c).
282
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Frullania monocera var. depauperata S.Hatt., J. Hattori Bot. Lab. 57: 419, 1984
(Hattori 1984a).
** Frullania monocera var. schifneri (Verd.) S.Hatt., J. Hattori Bot. Lab. 46: 120,
1979 (Hattori 1979a). Bas.: Frullania acutiloba var. schifneri Verd., Ann. Bryol.
2: 123, 1929 (Verdoorn 1929a).
*** Frullania monocera var. subhampeana (E.A.Hodgs.) Hentschel et von Konrat, Phytotaxa
220 (2): 136, 2015 (Hentschel et al. 2015). Bas.: Frullania subhampeana E.A.Hodgs.,
Trans. & Proc. Roy. Soc. New Zealand 77 (3): 370, 1949 (Hodgson 1949).
*** Frullania monocera var. undulata (Kamim.) Hentschel et von Konrat, Phytotaxa
220 (2): 136, 2015 (Hentschel et al. 2015). Bas.: Frullania undulata Kamim., J.
Hattori Bot. Lab. 24: 50, 1961 (Kamimura 1961).
** Frullania osumiensis (S.Hatt.) S.Hatt., J. Hattori Bot. Lab. 16: 87, 1956 (Iwatsuki
and Hattori 1956). Bas.: Frullania hampeana var. osumiensis S.Hatt., Bull. Tokyo
Sci. Mus. 11: 144, 1944 (Hattori 1944d).
* Frullania pallidula S.Hatt., Beih. Nova Hedwigia 90: 152, 1988 (Hattori 1988b).
** Frullania pseudomonocera S.Hatt., J. Hattori Bot. Lab. 60: 216, 1986 (Hattori
1986e).
** Frullania seriata Gottsche, Hedwigia 28 (3): 160, 1889 (Stephani 1889d).
** Frullania spinistipula Steph., Sp. Hepat. (Stephani) 4: 463, 1911 (Stephani 1911e).
** Frullania streimannii S.Hatt., J. Hattori Bot. Lab. 54: 176, 1983 (Hattori 1983).
*** sect. Australes Verd., Ann. Bryol., Suppl. 1: 58, 1930 (Verdoorn 1930c).
*** Frullania anomala E.A.Hodgs., Trans. & Proc. Roy. Soc. New Zealand 77 (3):
374, 1949 (Hodgson 1949).
* Frullania baileyana Steph., Sp. Hepat. (Stephani) 4: 417, 1910 (Stephani 1910b).
*** Frullania baladina Gottsche, Hedwigia 33 (3): 140, 1894 (Stephani 1894d). 204
* Frullania belmorensis Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2): 106,
1914 (Stephani and Watts 1914).
*** Frullania campanulata Sande Lac., Ned. Kruidk. Arch. 3: 422, 1854 [1855] (Sande Lacoste 1854).
** Frullania campanulata var. caduca Verd., Ann. Bryol., Suppl. 1: 41, 1930 (Verdoorn 1930c).
** Frullania campanulata var. malesiaca (Verd.) S.Hatt., Bull. Natl. Sci. Mus. Tokyo,
B 1 (4): 158, 1975 (Hattori 1975f). Bas.: Frullania malesiaca Verd., Ann. Bryol.,
Suppl. 1: 59, 1930 (Verdoorn 1930c).
* Frullania cataractarum Steph., Sp. Hepat. (Stephani) 4: 657, 1911 (Stephani 1911e).
* Frullania crawfordii Steph., Hedwigia 33 (3): 143, 1894 (Stephani 1894d).
*** Frullania dentata S.Hatt., J. Hattori Bot. Lab. 38: 231, 1974 (Hattori 1974c).
** Frullania dentata var. secernens S.Hatt., J. Hattori Bot. Lab. 65: 422, 1988 (Hattori
1988a).
*** Frullania errans Verd., Ann. Bryol., Suppl. 1: 59, 1930 (Verdoorn 1930c).
204 Frullania baladina is a species complex also including Frullania fulfordiae.
World checklist of hornworts and liverworts
283
** Frullania errans var. angulistipula S.Hatt., J. Hattori Bot. Lab. 36: 431, 1972
[1973] (Hattori 1972b).
*** Frullania fugax (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 3:
445, 1845 (Gottsche et al. 1845b). Bas.: Jungermannia fugax Hook.f. et Taylor,
London J. Bot. 4: 87, 1845 (Hooker and Taylor 1845). 205
** Frullania fulfordiae S.Hatt., Bryologist 90 (4): 365, 1987 [1988] (Hattori 1987b).
*** Frullania glomerata (Lehm. et Lindenb.) Nees et Mont., Ann. Sci. Nat. Bot. (sér.
2) 9: 46, 1838 (Montagne 1838). Bas.: Jungermannia glomerata Lehm. et Lindenb., Nov. Stirp. Pug. 5: 21, 1833 (Lehmann 1833).
*** Frullania incumbens Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 162, 1855 (Mitten
1855).
*** Frullania inlexa Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 120, 1860 [1861] (Mitten
1860c).
** Frullania media (E.A.Hodgs.) S.Hatt., J. Hattori Bot. Lab. 54: 153, 1983 (Hattori
1983). Bas.: Frullania fugax var. media E.A.Hodgs., Trans. & Proc. Roy. Soc. New
Zealand 77 (3): 375, 1949 (Hodgson 1949).
** Frullania mizutanii Kamim. et S.Hatt., J. Hattori Bot. Lab. 37: 524, 1973 (Hattori and Kamimura 1973).
*** Frullania obscurifolia Mitt., Philos. Trans. 168: 400, 1879 (Mitten 1879).
*** Frullania patagonica Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46
(9): 88, 1911 (Stephani 1911b).
** Frullania pentapleura Taylor, London J. Bot. 5: 402, 1846 (Taylor 1846b).
** Frullania polyptera Taylor, London J. Bot. 5: 401, 1846 (Taylor 1846b).
** Frullania polyptera var. angustata (Mitt.) S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 1
(2): 74, 1975 (Hattori 1975g). Bas.: Frullania angustata Mitt., J. Proc. Linn. Soc.,
Bot. 5 (18): 122, 1860 [1861] (Mitten 1860c).
** Frullania probosciphora Taylor, London J. Bot. 5: 402, 1846 (Taylor 1846b).
** Frullania pulchella Herzog, Rev. Bryol. Lichénol. 23 (1/2): 60, 1954 (Herzog 1954).
*** Frullania sinskeana J.J.Engel et B.C.Tan, J. Hattori Bot. Lab. 60: 335, 1986 (Tan
and Engel 1986). Nom. nov. pro Frullania spathulistipa Steph., Sp. Hepat. (Stephani) 4: 415, 1910 (Stephani 1910b), nom. illeg.
** Frullania socotrana Mitt., Trans. Roy Soc. Edinburgh 31: 335, 1888 (Mitten 1888).
* Frullania solanderiana Colenso, Trans. & Proc. New Zealand Inst. 21: 75, 1889
(Colenso 1889). 206
** Frullania subincumbens S.Hatt., Bryologist 90 (4): 367, 1987 [1988] (Hattori
1987b).
* Frullania subtropica Steph., Sp. Hepat. (Stephani) 4: 416, 1910 (Stephani 1910b).
* Frullania tjibodensis S.Hatt. et haithong, J. Jap. Bot. 52 (10): 289, 1977 (Hattori
and haithong 1977). 207
205 Frullania fugax is a species complex possibly including Frullania baileyana, Frullania belmorensis,
Frullania cataractarum, Frullania media and Frullania subtropica (Hattori 1979a).
206 Frullania solanderiana is morphologically similar to Frullania pentapleura and possibly conspeciic.
207 Frullania tjibodensis is possibly conspeciic with Frullania campanulata (Söderström et al. 2010a).
284
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Frullania tuyamae S.Hatt. et haithong, J. Jap. Bot. 53 (6): 175, 1978 (Hattori
and haithong 1978c).
*** sect. Frullania
*** Frullania appalachiana R.M.Schust., Phytologia 53 (5): 366, 1983 (Schuster
1983b).
*** Frullania azorica Sim-Sim, Sérgio, Mues et Kraut, Cryptog. Bryol. Lichénol. 16
(2): 112, 1995 (Sim-Sim et al. 1995).
*** Frullania catalinae A.Evans, Trans. Connecticut Acad. Arts 10 (1): 11, 1899
(Evans 1899).
*** Frullania dilatata (L.) Dumort., Recueil Observ. Jungerm.: 13, 1835 (Dumortier
1835). Bas.: Jungermannia dilatata L., Sp. Pl. 1: 1133, 1753 (Linnaeus 1753).
** Frullania dilatata subsp. asiatica S.Hatt., J. Jap. Bot. 57 (9): 258, 1982 (Hattori
1982b).
*** Frullania eboracensis Lehm., Nov. Stirp. Pug. 8: 14, 1844 (Lehmann 1844). 208
*** Frullania ericoides (Nees) Mont., Ann. Sci. Nat. Bot. (sér. 2) 12: 51, 1839 (Montagne 1839c). Bas.: Jungermannia ericoides Nees, Fl. Bras. (Martius) 1 (1): 346,
1833 (Nees 1833a). 209
* Frullania ericoides var. laxa (Gottsche, Lindenb. et Nees) Schifn., Consp. Hepat.
Arch. Ind.: 324, 1898 (Schifner 1898b). Bas.: Frullania squarrosa γ laxa Gottsche,
Lindenb. et Nees, Syn. Hepat. 5: 772, 1847 (Gottsche et al. 1847).
* Frullania ericoides var. minor Kamim., Bull. Kochi Gakuen Jun. Coll. 2: 22, 1971
(Kamimura 1971).
** Frullania ericoides var. verrucosa (Kamim.) Hentschel et von Konrat, Phytotaxa
220 (2): 134, 2015 (Hentschel et al. 2015). Bas.: Frullania squarrosa var. verrucosa
Kamim., J. Hattori Bot. Lab. 24: 19, 1961 (Kamimura 1961).
*** Frullania fragilifolia (Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 3: 437,
1845 (Gottsche et al. 1845b). Bas.: Jungermannia fragilifolia Taylor, Ann. Mag.
Nat. Hist. 12 (76): 172, 1843 (Taylor 1843).
** Frullania fuegiana Steph., Sp. Hepat. (Stephani) 4: 428, 1910 (Stephani 1910b).
** Frullania hattoriana J.D.Godfrey et G.Godfrey, J. Hattori Bot. Lab. 48: 321, 1980
(Godfrey and Godfrey 1980).
** Frullania koponenii S.Hatt., Ann. Bot. Fenn. 15 (2): 111, 1978 (Koponen et al. 1978).
*** Frullania muscicola Steph., Hedwigia 33 (3): 146, 1894 (Stephani 1894d).
*** Frullania oakesiana Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 225, 1869 (Austin 1869).
** Frullania oakesiana subsp. takayuensis (Steph.) R.M.Schust., Hepat. Anthocerotae
N. Amer. 5: 195, 1992 (Schuster 1992b). Bas.: Frullania takayuensis Steph., Sp.
Hepat. (Stephani) 4: 399, 1910 (Stephani 1910b).
208 Frullania eboracensis is a species complex also including Frullania appalachiana, Frullania parvistipula
and Frullania virginica.
209 Frullania ericoides is enormously variable (Schuster 1992b) and is a species complex with at least two
lineages (Hentschel et al. 2009).
World checklist of hornworts and liverworts
285
*** Frullania parvistipula Steph., Sp. Hepat. (Stephani) 4: 397, 1910 (Stephani 1910b).
** Frullania sabaliana R.M.Schust., Phytologia 53 (5): 365, 1983 (Schuster 1983b).
** Frullania semivillosa Lindenb. et Gottsche, Syn. Hepat. 5: 774, 1847 (Gottsche et
al. 1847).
** Frullania stylifera (R.M.Schust.) R.M.Schust., Hepat. Anthocerotae N. Amer. 5:
210, 1992 (Schuster 1992b). Bas.: Frullania inlata var. stylifera R.M.Schust., Phytologia 53 (5): 366, 1983 (Schuster 1983b).
* Frullania subdilatata C.Massal., Nuovo Giorn. Bot. Ital. (n.ser.) 13 (4): 349, 1906
(Levier 1906).
*** Frullania virginica Lehm., Nov. Stirp. Pug. 8: 19, 1844 (Lehmann 1844).
** sect. Irregulares E.A.Hodgs. ex S.Hatt., J. Hattori Bot. Lab. 54: 143, 1983 (Hattori 1983).
* Frullania astrolabea Steph., Sp. Hepat. (Stephani) 4: 460, 1910 (Stephani 1910b). 210
*** Frullania deplanata Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 161, 1855 (Mitten
1855).
*** Frullania morobensis S.Hatt. et Streimann, J. Hattori Bot. Lab. 59: 109, 1985
(Hattori and Streimann 1985).
*** Frullania patula Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 159, 1854 (Mitten 1854).
*** Frullania scandens Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 258, 1843 (Montagne 1843).
** sect. Planae R.M.Schust., Phytologia 57 (5): 372, 1985 (Schuster 1985a).
*** Frullania plana Sull., Mem. Amer. Acad. Arts (n.ser.) 4: 175, 1849 (Sullivant 1849).
*** subg. Homotropantha Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 35, 1884
(Spruce 1884).
*** Frullania delexa Mitt., Bonplandia 10 (2): 19, 1862 (Mitten 1862).
*** Frullania integristipula (Nees) Nees, Syn. Hepat. 3: 431, 1845 (Gottsche et al.
1845b). Bas.: Jungermannia integristipula Nees, Enum. Pl. Crypt. Javae: 54, 1830
(Nees 1830).
** Frullania integristipula var. emarginata Verd., Ann. Bryol. 2: 153, 1929 (Verdoorn
1929a).
** Frullania macrophylla S.Hatt., J. Hattori Bot. Lab. 47: 220, 1980 (Hattori 1980a).
** Frullania sabahana S.Hatt., J. Hattori Bot. Lab. 40: 493, 1976 (Hattori 1976d).
** Frullania sackawana Steph., Bull. Herb. Boissier 5 (2): 91, 1897 (Stephani 1897b).
** Frullania sarawakensis S.Hatt., J. Hattori Bot. Lab. 40: 496, 1976 (Hattori 1976d).
** Frullania umbonata Mitt., Sp. Hepat. (Stephani) 4: 579, 1911 (Stephani 1911e).
*** Frullania utriculata Steph., Hedwigia 33 (3): 152, 1894 (Stephani 1894d).
*** sect. Fallaces Verd., Rev. Bryol. Lichénol. 1: 112, 1928 (Verdoorn 1928a).
* Frullania fallax Gottsche, Syn. Hepat. 3: 432, 1845 (Gottsche et al. 1845b).
210 Frullania astrolabea is possibly conspeciic with Frullania scandens (Hattori 1981b).
286
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Frullania intermedia (Reinw., Blume et Nees) Nees, Syn. Hepat. 3: 434, 1845
(Gottsche et al. 1845b). Bas.: Jungermannia intermedia Reinw., Blume et Nees,
Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 218, 1824
[1825] (Reinwardt et al. 1824a). 211
** Frullania intermedia subsp. morokensis (Steph.) S.Hatt., J. Hattori Bot. Lab. 47:
194, 1980 (Hattori 1980a). Bas.: Frullania morokensis Steph., Sp. Hepat. (Stephani) 4: 578, 1911 (Stephani 1911e).
* Frullania intermedia var. non-apiculata S.Hatt., J. Hattori Bot. Lab. 39: 291, 1975
(Hattori 1975d). 212
** Frullania novoguineensis Schifn., Leberm., Forschungsr. Gazelle 4 (4): 37, 1890
(Schifner 1890).
** Frullania regularis Schifn., Leberm., Forschungsr. Gazelle 4 (4): 38, 1890
(Schifner 1890).
***
*
**
**
***
sect. Nodulosae Verd., Rev. Bryol. Lichénol. 1: 116, 1928 (Verdoorn 1928a).
Frullania brotheri Steph., Hedwigia 33 (3): 150, 1894 (Stephani 1894d). 213
Frullania hamata Steph., Sp. Hepat. (Stephani) 4: 582, 1911 (Stephani 1911e).
Frullania leeuwenii Verd., Nova Guinea 14: 545, 1930 (Verdoorn 1930b).
Frullania nodulosa (Reinw., Blume et Nees) Nees, Syn. Hepat. 3: 433, 1845
(Gottsche et al. 1845b). Bas.: Jungermannia nodulosa Reinw., Blume et Nees,
Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 217, 1824
[1825] (Reinwardt et al. 1824a).
** sect. Remotilobae Verd., Rev. Bryol. Lichénol. 1: 119, 1928 (Verdoorn 1928a).
*** Frullania heteromorpha Schifn., Leberm., Forschungsr. Gazelle 4 (4): 38, 1890
(Schifner 1890).
** Frullania remotiloba Steph., Hedwigia 33 (3): 152, 1894 (Stephani 1894d).
* subg. Mammillosae S.Hatt., J. Hattori Bot. Lab. 60: 226, 1986 (Hattori 1986e).
** Frullania huerlimannii S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 2 (3): 84, 1976
(Hattori 1976b).
* Frullania huerlimannii var. dioica S.Hatt., J. Hattori Bot. Lab. 57: 413, 1984
(Hattori 1984a).
** Frullania involvens S.Hatt. et Kamim., J. Hattori Bot. Lab. 37: 526, 1973 (Hattori
and Kamimura 1973).
** Frullania iriomotensis S.Hatt., J. Jap. Bot. 55 (5): 133, 1980 (Hattori 1980c).
** Frullania mammillosa S.Hatt., J. Hattori Bot. Lab. 43: 424, 1977 [1978] (Hattori
1977b).
211 Frullania intermedia is a species complex (Hattori 1980a).
212 Frullania intermedia var. non-apiculata was reduced to a synonym of subsp. intermedia by Hattori
(1980a), but it was re-instated by Hattori (1985).
213 Frullania brotheri is probably just a high-elevation form of Frullania nodulosa (Söderström et al. 2010a).
World checklist of hornworts and liverworts
287
*** Frullania meijeri S.Hatt., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 17 (4): 307, 1974
(Hattori 1974b).
** Frullania notarisii Steph., Sp. Hepat. (Stephani) 4: 651, 1911 (Stephani 1911e).
** Frullania papulosa Steph., Sp. Hepat. (Stephani) 4: 654, 1911 (Stephani 1911e).
** Frullania rudoliana S.Hatt., J. Hattori Bot. Lab. 36: 437, 1972 [1973] (Hattori
1972b).
*** Frullania thiersiae S.Hatt., Beih. Nova Hedwigia 90: 156, 1988 (Hattori 1988b).
** Frullania tixieri S.Hatt., J. Jap. Bot. 51 (7): 193, 1976 (Hattori 1976a).
*** subg. Meteoriopsis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 37, 1884
(Spruce 1884).
* Frullania caldensis Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 33 (7): 89, 1876
[1877] (Ångström 1876).
** Frullania evoluta Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 122, 1860 [1861] (Mitten
1860c).
*** Frullania tagawana (S.Hatt. et haithong) S.Hatt., J. Hattori Bot. Lab. 59: 160,
1985 (Hattori and Lin 1985a). Bas.: Frullania evoluta var. tagawana S.Hatt. et
haithong, J. Hattori Bot. Lab. 43: 441, 1977 [1978] (Hattori et al. 1977).
***
***
**
***
***
***
***
**
*
*
*
sect. Intumescentes R.M.Schust., Phytologia 57 (5): 370, 1985 (Schuster 1985a).
Frullania aculeata Taylor, London J. Bot. 5: 407, 1846 (Taylor 1846b).
Frullania ambronnii Steph., Biblioth. Bot. 87 (2): 242, 1916 (Stephani 1916a).
Frullania atrata (Sw.) Nees ex Mont., Ann. Sci. Nat. Bot. (sér. 2) 12: 51, 1839
(Montagne 1839c). Bas.: Jungermannia atrata Sw., Prodr. (Swartz): 144, 1788
(Swartz 1788).
Frullania beyrichiana (Lehm. et Lindenb.) Lehm. et Lindenb., Syn. Hepat. 3: 460,
1845 (Gottsche et al. 1845b). Bas.: Jungermannia beyrichiana Lehm. et Lindenb.,
Nov. Stirp. Pug. 5: 25, 1833 (Lehmann 1833).
Frullania bicornistipula Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 46, 1884
(Spruce 1884).
Frullania brasiliensis Raddi, Critt. Brasil.: 12, 1822 (Raddi 1822). 214
Frullania brasiliensis var. elegantula Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 50, 1884 (Spruce 1884).
Frullania breuteliana Gottsche, Syn. Hepat. 3: 461, 1845 (Gottsche et al. 1845b). 215
Frullania compacta Gottsche, Sp. Hepat. (Stephani) 4: 493, 1911 (Stephani
1911e). 216
Frullania crenulifolia J.B.Jack et Steph., Hedwigia 31 (1): 14, 1892 (Jack and Stephani 1892).
214 Frullania brasiliensis is a species complex with controversial synonymy (Schuster 1992b, Hentschel et
al. 2009).
215 Frullania breuteliana is possibly conspeciic with Frullania beyrichiana.
216 Frullania compacta is conspeciic with Frullania brasiliensis in Stotler (1969), but Schuster (1992b)
doubts the correctness of this.
288
**
**
***
**
*
***
*
*
**
*
***
*
***
*
***
**
**
*
***
**
**
***
**
***
**
***
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Frullania crispiloba Steph., Hedwigia 33 (3): 156, 1894 (Stephani 1894d).
Frullania curviramea Steph., Sp. Hepat. (Stephani) 4: 684, 1911 (Stephani 1911e).
Frullania ecuadorensis Steph., Sp. Hepat. (Stephani) 4: 526, 1911 (Stephani 1911e).
Frullania formosa Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 46, 1884
(Spruce 1884).
Frullania granatensis Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 173 (79), 1864
(Gottsche 1864). 217
Frullania griithsiana Gottsche, Syn. Hepat. 4: 466, 1846 (Gottsche et al. 1846).
Frullania guadalupensis Gottsche, Sp. Hepat. (Stephani) 4: 496, 1911 (Stephani 1911e).
Frullania gualaquizana Steph., Sp. Hepat. (Stephani) 4: 531, 1911 (Stephani 1911e).
Frullania hamilora Herzog et L.Clark, Bryologist 56 (3): 180, 1953 (Clark and
Schultz 1953).
Frullania humilis Spruce, Mem. Torrey Bot. Club 1 (3): 119, 1890 (Spruce 1890).
Frullania intumescens (Lehm. et Lindenb.) Lehm. et Lindenb., Syn. Hepat. 3: 460,
1845 (Gottsche et al. 1845b). Bas.: Jungermannia intumescens Lehm. et Lindenb.,
Nov. Stirp. Pug. 6: 52, 1834 (Lehmann 1834).
Frullania laticaulis Spruce, Mem. Torrey Bot. Club 1 (3): 120, 1890 (Spruce
1890).
Frullania lobatohastata Steph., Sp. Hepat. (Stephani) 4: 499, 1911 (Stephani 1911e).
Frullania longistipula var. apiculata Demaret et Vanden Berghen, Bull. Jard. Bot.
État Bruxelles 20 (1): 4, 1950 (Demaret and Vanden Berghen 1950).
Frullania macrocephala (Lehm. et Lindenb.) Lehm. et Lindenb., Syn. Hepat. 3:
460, 1845 (Gottsche et al. 1845b). Bas.: Jungermannia macrocephala Lehm. et
Lindenb., Nov. Stirp. Pug. 5: 20, 1833 (Lehmann 1833).
Frullania madothecoides Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 47, 1884
(Spruce 1884).
Frullania meridana Steph., Sp. Hepat. (Stephani) 4: 500, 1911 (Stephani 1911e).
Frullania microcephala Gottsche, Mexik. Leverm.: 251, 1863 (Gottsche 1863).
Frullania montagnei Gottsche, Syn. Hepat. 3: 456, 1845 (Gottsche et al. 1845b).
Frullania moritziana Lindenb. et Gottsche, Syn. Hepat. 5: 782, 1847 (Gottsche et
al. 1847).
Frullania osculatiana De Not., Mem. Reale Accad. Sci. Torino (ser. 2) 16: 236,
1857 (De Notaris 1857).
Frullania paradoxa Lehm. et Lindenb., Syn. Hepat. 3: 462, 1845 (Gottsche et al.
1845b).
Frullania pearceana Steph., Sp. Hepat. (Stephani) 4: 515, 1911 (Stephani 1911e).
Frullania pittieri Steph., Primit. l. costar.: 113, 1892 [1893] (Stephani 1892e).
Frullania rigescens Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 52, 1884
(Spruce 1884).
Frullania setigera Steph., Hedwigia 33 (3): 159, 1894 (Stephani 1894d).
217 Frullania granatensis is possibly conspeciic with Frullania beyrichiana, but the type specimen was lost
in B (Stotler 1969).
World checklist of hornworts and liverworts
289
** Frullania speciosa Herzog, Memoranda Soc. Fauna Fl. Fennica 25: 59, 1950 (Herzog
1950c).
** Frullania supradecomposita (Lehm. et Lindenb.) Lehm. et Lindenb., Syn. Hepat.
3: 431, 1845 (Gottsche et al. 1845b). Bas.: Jungermannia supradecomposita Lehm.
et Lindenb., Nov. Stirp. Pug. 5: 23, 1833 (Lehmann 1833).
* Frullania trianae Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 173, 1864 (Gottsche
1864).
** Frullania triquetra Lindenb. et Gottsche, Syn. Hepat. 5: 780, 1847 (Gottsche et
al. 1847).
* Frullania trollii Herzog, Beih. Bot. Centralbl. 61B (3): 575, 1942 (Herzog 1942d).
*** Frullania uleana Steph., Hedwigia 33 (3): 155, 1894 (Stephani 1894d).
** sect. Meteoriopsis Uribe, von Konrat et Hentschel, Phytotaxa 220 (2): 132,
2015 (Hentschel et al. 2015).
*** Frullania convoluta Lindenb. et Hampe, Linnaea 24 (3): 303, 1851 [1852] (Hampe 1851b).
* Frullania convoluta var. ampliata Herzog, Rev. Bryol. Lichénol. 20 (1/2): 129,
1951 [1952] (Herzog 1951a).
*** Frullania darwinii Gradst. et Uribe, Cryptog. Bryol. 25 (4): 296, 2004 (Uribe
2004).
*** Frullania dulimensis Uribe, Cryptog. Bryol. 27 (3): 309, 2006 (Uribe 2006).
*** Frullania grandifolia Steph., Sp. Hepat. (Stephani) 4: 684, 1911 (Stephani 1911e).
*** Frullania peruviana Gottsche, Syn. Hepat. 4: 465, 1846 (Gottsche et al. 1846).
*** Frullania phalangilora Steph., Biblioth. Bot. 87 (2): 247, 1916 (Stephani 1916a).
*** Frullania weberbaueri Steph., Sp. Hepat. (Stephani) 4: 510, 1911 (Stephani 1911e).
*** sect. Obtusilobae Verd., Ann. Bryol., Suppl. 1: 81, 1930 (Verdoorn 1930c).
** Frullania angulata Mitt., J. Proc. Linn. Soc., Bot. 7 (27): 169, 1863 (Mitten
1863).
** Frullania angulata var. laciniata Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 46
(1/2): 72, 1976 (Vanden Berghen 1976b).
** Frullania apicalis Mitt., Philos. Trans. 168: 401, 1879 (Mitten 1879).
** Frullania apicalis var. camerunensis Vanden Berghen, Bull. Jard. Bot. Natl. Belg.
46 (1/2): 53, 1976 (Vanden Berghen 1976b).
* Frullania borbonica Lindenb., Syn. Hepat. 3: 455, 1845 (Gottsche et al. 1845b). 218
*** Frullania capensis Gottsche, Syn. Hepat. 3: 449, 1845 (Gottsche et al. 1845b).
** Frullania donnellii Austin, Bull. Torrey Bot. Club 6 (52): 301, 1879 (Austin 1879).
** Frullania eplicata Steph., Sp. Hepat. (Stephani) 4: 679, 1911 (Stephani 1911e).
** Frullania imerinensis Steph., Sp. Hepat. (Stephani) 4: 484, 1911 (Stephani 1911e).
*** Frullania kunzei (Lehm. et Lindenb.) Lehm. et Lindenb., Syn. Hepat. 3: 449,
1845 (Gottsche et al. 1845b). Bas.: Jungermannia kunzei Lehm. et Lindenb., Nov.
Stirp. Pug. 6: 50, 1834 (Lehmann 1834).
218 Frullania borbonica is possibly conspeciic with Frullania apicalis (Vanden Berghen 1976b).
290
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Frullania kunzei var. maritima R.M.Schust., Phytotaxa 220 (2): 135, 2015
(Hentschel et al. 2015). Based on: Frullania kunzei var. maritima R.M.Schust., J.
Hattori Bot. Lab. 70: 145, 1991 (Schuster 1991b), nom. inval.
** Frullania longistipula Steph., Bull. Soc. Roy. Bot. Belgique, Compt. Rend. 30 (2):
199, 1891 [1892] (Stephani 1891b).
*** Frullania meyeniana Lindenb., Syn. Hepat. 3: 455, 1845 (Gottsche et al. 1845b).
** Frullania meyeniana var. dioica S.Hatt., J. Hattori Bot. Lab. 43: 426, 1977 [1978]
(Hattori 1977b).
** Frullania onraedtii Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 46 (1/2): 60,
1976 (Vanden Berghen 1976b).
** Frullania papuana Verd., Ann. Bryol., Suppl. 1: 82, 1930 (Verdoorn 1930c).
*** Frullania schimperi Nees, Syn. Hepat. 3: 454, 1845 (Gottsche et al. 1845b). 219
** Frullania schimperi var. laciniata Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 46
(1/2): 59, 1976 (Vanden Berghen 1976b).
*** subg. Microfrullania (R.M.Schust.) R.M.Schust., Hepat. Anthocerotae N.
Amer. 5: 34, 1992 (Schuster 1992b). Bas.: Neohattoria subg. Microfrullania
R.M.Schust., J. Hattori Bot. Lab. 33: 280, 1970 (Schuster 1970a).
*** Frullania fertilis De Not., Mem. Reale Accad. Sci. Torino (ser. 2) 16: 235, 1857
(De Notaris 1857).
*** Frullania knightbridgei von Konrat et de Lange, PhytoKeys 8: 28, 2012 (von Konrat et al. 2012b).
*** Frullania magellanica F.Weber et Nees, Syn. Hepat. 3: 446, 1845 (Gottsche et al.
1845b). Nom. nov. pro Jungermannia magellanica Spreng. Ann. Wetterauischen
Ges. Gesammte Naturk. 1: 25, 1809 (Sprengel 1809), nom. illeg.
** Frullania magellanica subsp. tristaniana (S.W.Arnell) Váňa et J.J.Engel, Mem.
New York Bot. Gard. 105: 59, 2013 (Váňa and Engel 2013). Bas.: Frullania
tristaniana S.W.Arnell, Results Norweg. Sci. Exped. Tristan da Cunha 42: 9, 1958
(Arnell 1958b).
* Frullania matafaoica H.A.Mill., Phytologia 47 (4): 322, 1981 (Miller 1981). Nom.
nov. pro Frullania minutissima Pearson, Amer. Samoa: 140, 1924 (Pearson 1924a),
nom. illeg.
*** Frullania toropuku von Konrat, de Lange et Larraín, Polish Bot. J. 58 (2): 439,
2013 (von Konrat et al. 2013).
*** sect. Amphijubula (R.M.Schust.) von Konrat, Hentschel, Heinrichs et Braggins, Bryologist 114 (1): 53, 2011 (von Konrat et al. 2011). Bas.: Amphijubula
R.M.Schust., J. Hattori Bot. Lab. 33: 298, 1970 (Schuster 1970a).
*** Frullania lobulata (Hook.) Hook. et Nees, Syn. Hepat. 3: 445, 1845 (Gottsche
et al. 1845b). Bas.: Jungermannia lobulata Hook., Musci Exot. 2: tab. 119, 1820
(Hooker 1820).
219 Frullania schimperi belongs to a species complex also including Frullania onraedtii, Frullania capensis,
Frullania apicalis, Frullania imerinensis and Frullania meyeniana (Vanden Berghen 1976b).
World checklist of hornworts and liverworts
291
*** Frullania microcaulis Gola, Nuovo Giorn. Bot. Ital. (n.ser.) 29 (1/4): 172, 1922
[1923] (Gola 1922).
*** Frullania truncatistyla von Konrat, Hentschel, Heinrichs et Braggins, Bryologist
114 (1): 63, 2011 (von Konrat et al. 2011).
*** sect. Microfrullania (R.M.Schust.) von Konrat et Hentschel, Phytotaxa 220
(2): 133, 2015 (Hentschel et al. 2015). Bas.: Neohattoria sect. Microfrullania
R.M.Schust., J. Hattori Bot. Lab. 33: 288, 1970 (Schuster 1970a).
*** Frullania chevalieri (R.M.Schust.) R.M.Schust., Hepat. Anthocerotae N. Amer.
5: 34, 1992 (Schuster 1992b). Bas.: Neohattoria chevalieri R.M.Schust., J. Hattori
Bot. Lab. 33: 289, 1970 (Schuster 1970a). 220
*** Frullania microscopica Pearson, J. Linn. Soc., Bot. 46 (305): 33, 1922 (Pearson 1922b).
** Frullania neocaledonica J.J.Engel, Novon 9 (3): 344, 1999 (Engel and Smith Merrill 1999a). Nom. nov. pro Neohattoria caledonica R.M.Schust., J. Hattori Bot. Lab.
33: 291, 1970 (Schuster 1970a).
*** Frullania parhamii (R.M.Schust.) R.M.Schust. ex von Konrat, L.Söderstr. et
A.Hagborg, Telopea 13 (3): 407, 2011 (Söderström et al. 2011a). Bas.: Neohattoria parhamii R.M.Schust., J. Hattori Bot. Lab. 26: 243, 1963 (Schuster 1963b).
*** sect. Regulares Verd., Ann. Bryol., Suppl. 1: 133, 1930 (Verdoorn 1930c).
*** Frullania junghuhniana Gottsche, Syn. Hepat. 3: 444, 1845 (Gottsche et al.
1845b). 221
** Frullania junghuhniana var. bisexualis S.Hatt., J. Hattori Bot. Lab. 40: 485, 1976
(Hattori 1976d).
** Frullania junghuhniana var. tenella (Sande Lac.) Grolle et S.Hatt., Misc. Bryol.
Lichenol. 9 (6): 123, 1982 (Hattori 1982a). Bas.: Frullania tenella Sande Lac.,
Ned. Kruidk. Arch. 3: 423, 1854 [1855] (Sande Lacoste 1854).
** Frullania mcveanii S.Hatt., J. Hattori Bot. Lab. 37: 55, 1973 (Hattori 1973a).
** Frullania pseudomeyeniana S.Hatt., J. Hattori Bot. Lab. 60: 231, 1986 (Hattori
1986e).
*** Frullania rostrata (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 3:
445, 1845 (Gottsche et al. 1845b). Bas.: Jungermannia rostrata Hook.f. et Taylor,
London J. Bot. 4: 87, 1845 (Hooker and Taylor 1845). 222
** Frullania scalaris S.Hatt., J. Hattori Bot. Lab. 43: 432, 1977 [1978] (Hattori 1977b).
** subg. Saccophora Verd., Ann. Bryol. 2: 121, 1929 (Verdoorn 1929a).
*** Frullania gaudichaudii (Nees et Mont.) Nees et Mont., Syn. Hepat. 3: 435, 1845
(Gottsche et al. 1845b). Bas.: Jubula gaudichaudii Nees et Mont., Ann. Sci. Nat.
Bot. (sér. 2) 5: 64, 1836 (Nees and Montagne 1836).
220 Frullania chevalieri is a species complex (Hattori 1984a).
221 Frullania junghuhniana is a species complex (Hattori 1976b).
222 Frullania rostrata is a species complex with several lineages, some of which have been recently described (e.g. von Konrat et al. 2013).
292
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Frullania gaudichaudii var. ceylanica (Nees) S.Hatt., J. Hattori Bot. Lab. 47: 104,
1980 (Hattori 1980d). Bas.: Frullania ceylanica Nees, Syn. Hepat. 3: 436, 1845
(Gottsche et al. 1845b).
*** Frullania hedrantha S.Hatt. et Kamim., J. Hattori Bot. Lab. 37: 519, 1973 (Hattori and Kamimura 1973).
** Frullania immersa Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 315, 1896 (Stephani 1896a).
** Frullania pancheri Gottsche, Hedwigia 33 (3): 159, 1894 (Stephani 1894d).
** Frullania papillilobula S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 1 (4): 141, 1975
(Hattori 1975f).
** Frullania sublignosa Steph., Hedwigia 33 (3): 148, 1894 (Stephani 1894d).
subg. Steerea (S.Hatt. et Kamim.) R.M.Schust., Hepat. Anthocerotae N. Amer.
5: 32, 1992 (Schuster 1992b). Bas.: Steerea S.Hatt. et Kamim., J. Hattori Bot.
Lab. 34: 429, 1971 (Hattori and Kamimura 1971).
*** Frullania clemensiana Verd., Ned. Kruidk. Arch. (ser. 3) 42 (2): 493, 1932 (Verdoorn 1932b).
*
*** subg. hyopsiella Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 41, 1884
(Spruce 1884).
*** Frullania acicularis Hentschel et von Konrat, Phytotaxa 220 (2): 134, 2015
(Hentschel et al. 2015). Nom. nov. pro Frullania tamarisci var. azorica J.-P.Frahm,
Trop. Bryol. 27: 102, 2006 (Frahm 2006). 223
** Frullania alstonii Verd., Ann. Bryol., Suppl. 1: 76, 1930 (Verdoorn 1930c).
** Frullania aoshimensis Horik., Sci. Rep. Tôhoku Imp. Univ., Ser. 4, Biol. 4 (1): 64,
1929 (Horikawa 1929b).
*** Frullania appendiculata Steph., Bull. Herb. Boissier 5 (2): 88, 1897 (Stephani
1897b). 224
*** Frullania asagrayana Mont., Ann. Sci. Nat. Bot. (sér. 2) 18: 14, 1842 (Montagne
1842b). 225
*** Frullania calcarifera Steph., Bol. Soc. Brot. 4: 241, 1886 [1887] (Henriques 1886). 226
*** Frullania californica (M.Howe) A.Evans, Trans. Connecticut Acad. Arts 10 (1):
25, 1899 (Evans 1899). Bas.: Frullania asagrayana var. californica M.Howe, Erythea 2 (6): 98, 1894 (Howe 1894).
*** Frullania densiloba Steph. ex A.Evans, Proc. Wash. Acad. Sci. 8: 157, 1906 (Evans
1906b).
*** Frullania franciscana M.Howe, Erythea 2 (6): 99, 1894 (Howe 1894).
223 Frullania acicularis was shown to be a species separate from Frullania tamarisci (Heinrichs et al. 2010,
Vilnet et al. (2014). It may, however, not be the earliest available name.
224 Frullania appendiculata was shown by Vilnet et al. (2014) to be a species separate from Frullania
tamarisci. It may, however, not be the earliest available name.
225 Frullania asagrayana consists of two geographically separated clades (Ramaiya et al. 2010).
226 Frullania calcarifera was shown to be a species separate from Frullania tamarisci by Heinrichs et al.
(2010) and Vilnet et al. (2014). It may, however, not be the earliest available name.
World checklist of hornworts and liverworts
293
** Frullania iwatsukii S.Hatt., J. Hattori Bot. Lab. 35: 240, 1972 (Hattori 1972a).
*** Frullania microphylla (Gottsche) Pearson, J. Bot. 32: 328, 1894 (Pearson 1894).
Bas.: Frullania tamarisci var. microphylla Gottsche, Hepat. Eur., Leberm. 21-22:
no. 109 [209], 1862 (Rabenhorst 1862).
*** Frullania moniliata (Reinw., Blume et Nees) Mont., Ann. Sci. Nat. Bot. (sér. 2)
18: 13, 1842 (Montagne 1842b). Bas.: Jungermannia moniliata Reinw., Blume
et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 224,
1824 [1825] (Reinwardt et al. 1824a). 227
*** Frullania nisquallensis Sull., Mem. Amer. Acad. Arts (n.ser.) 4: 175, 1849 (Sullivant 1849).
*** Frullania polysticta Lindenb., Syn. Hepat. 3: 440, 1845 (Gottsche et al. 1845b).
** Frullania pseudoalstonii Tsudo et J.Haseg., Bryol. Res. 9 (3): 44, 2006 (Tsudo and
Hasegawa 2006).
** Frullania punctata Reimers, Hedwigia 71 (1/2): 36, 1931 (Reimers 1931).
** Frullania schaefer-verwimpii Yuzawa et S.Hatt., J. Jap. Bot. 64 (2): 37, 1989 (Yuzawa and Hattori 1989).
** Frullania selwyniana Pearson, List. Canad. Hepat.: 1, 1890 (Pearson 1890).
*** Frullania sergiae Sim-Sim, Fontinha, Mues et Lion, Nova Hedwigia 71 (1/2): 186,
2000 (Sim-Sim et al. 2000).
*** Frullania subarctica Vilnet, Borovich. et Bakalin, Phytotaxa 173 (1): 67, 2014
(Vilnet et al. 2014). 228
*** Frullania tamarisci (L.) Dumort., Recueil Observ. Jungerm.: 13, 1835 (Dumortier
1835). Bas.: Jungermannia tamarisci L., Sp. Pl. 1: 1134, 1753 (Linnaeus 1753).
*** Frullania tenerifae (F.Weber) Nees, Naturgesch. Eur. Leberm. 3: 239, 1838 (Nees
1838b). Bas.: Jungermannia tenerifae F.Weber, Hist. Musc. Hepat. Prodr.: 23,
1815 (Weber 1815).
** Frullania trigona L.Clark, Jovet-Ast et Frye, Bryologist 50 (1): 52, 1947 (Clark et
al. 1947).
Incertae sedis
Frullania ainis Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 257, 1843 (Montagne 1843). 229
* Frullania allionii Steph., Sp. Hepat. (Stephani) 4: 394, 1910 (Stephani 1910b).
* Frullania alpina Steph., Sp. Hepat. (Stephani) 4: 533, 1911 (Stephani 1911e).
** Frullania alternans Nees, Syn. Hepat. 3: 430, 1845 (Gottsche et al. 1845b).
* Frullania apertilobula Gerola, Lav. Bot. Ist. Bot. Univ. Padova 12: 478, 1947
(Gerola 1947).
** Frullania armata Herzog et L.Clark, Bryologist 57 (1): 36, 1954 (Clark 1954).
*
227 Frullania moniliata is a species complex (Vilnet et al. 2014).
228 Frullania subarctica was shown to be a species separate from the Frullania tamarisci complex (Vilnet
et al. 2014), but it is maybe not the oldest name.
229 Frullania ainis is possibly conspeciic with Frullania ecklonii (Arnell 1963b).
294
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Frullania bolanderi Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 226, 1869 (Austin
1869).
*** Frullania boveana C.Massal., Nuovo Giorn. Bot. Ital. 17 (3): 244, 1885 (Massalongo 1885).
* Frullania caespitans Beauverd, Sp. Hepat. (Stephani) 6: 537, 1924 (Stephani
1924). Nom. nov. pro Frullania campanulata Steph., Biblioth. Bot. 87 (2): 242,
1916 (Stephani 1916a), nom. illeg.
* Frullania canaliculata Gottsche, Sp. Hepat. (Stephani) 4: 391, 1910 (Stephani 1910b).
** Frullania capillaris Steph., Sp. Hepat. (Stephani) 4: 616, 1911 (Stephani 1911e).
* Frullania cavallii Gola, Ann. Bot. (Rome) 6 (2): 275, 1907 (Gola 1907).
* Frullania chiapasana Steph., Sp. Hepat. (Stephani) 4: 392, 1910 (Stephani 1910b).
* Frullania chilcootiensis Steph., Bot. Jahrb. Syst. 8 (2): 98, 1886 (Stephani 1886a).
* Frullania chiovendae Gola, Ann. Bot. (Rome) 13 (1): 71, 1914 (Gola 1914a).
** Frullania ciliata Lindenb. et Gottsche, Syn. Hepat. 5: 775, 1847 (Gottsche et al.
1847).
** Frullania cinchonae Gottsche, Syn. Hepat. 3: 455, 1845 (Gottsche et al. 1845b).
* Frullania complicata Steph., Sp. Hepat. (Stephani) 4: 671, 1911 (Stephani 1911e).
* Frullania cordaeana Lindenb., Syn. Hepat. 3: 463, 1845 (Gottsche et al. 1845b).
** Frullania cuneatistipula Steph., Sp. Hepat. (Stephani) 6: 538, 1924 (Stephani 1924).
* Frullania cuspiloba Steph., Sp. Hepat. (Stephani) 4: 394, 1910 (Stephani 1910b).
* Frullania dispar Nees, Syn. Hepat. 3: 429, 1845 (Gottsche et al. 1845b).
* Frullania guatemalensis Steph., Sp. Hepat. (Stephani) 4: 497, 1911 (Stephani 1911e).
*** Frullania inlata Gottsche, Syn. Hepat. 3: 424, 1845 (Gottsche et al. 1845b).
** Frullania inlata var. communis R.M.Schust., Phytologia 57 (5): 372, 1985 (Schuster
1985a).
** Frullania inlata var. dioica S.Hatt. et haithong, J. Hattori Bot. Lab. 44: 185,
1978 (Hattori and haithong 1978b).
** Frullania inlata var. mayebarae (S.Hatt.) K.Yamada, Misc. Bryol. Lichenol. 6 (9):
163, 1974 (Yamada 1974). Bas.: Frullania mayebarae S.Hatt., Bot. Mag. (Tokyo)
65 (763/764): 13, 1952 (Hattori 1952a).
* Frullania laetevirens Hampe ex Gottsche, Lindenb. et Nees, Syn. Hepat. 3: 420,
1845 (Gottsche et al. 1845b).
* Frullania larjiana Sushil K.Singh et D.K.Singh, J. Bryol. 27 (2): 105, 2005 (Singh
and Singh 2005).
* Frullania larjiana var. didyhatii S.N.Srivast. et M.Rai, Geophytology 41 (1/2):
109, 2011 (Srivastava and Rai 2011).
* Frullania leana Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 227, 1869 (Austin 1869).
* Frullania madagascariensis Gottsche, Abh. Naturwiss. Vereins Bremen 7: 364,
1882 (Gottsche 1882).
* Frullania mauritiana Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 227, 1869
(Austin 1869).
*** Frullania mirabilis J.B.Jack et Steph., Hedwigia 31 (1): 15, 1892 (Jack and Stephani
1892).
World checklist of hornworts and liverworts
295
** Frullania monoica Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26 (III, 6):
67, 1900 (Stephani 1900b).
* Frullania ocanniensis Steph., Sp. Hepat. (Stephani) 6: 542, 1924 (Stephani 1924).
*** Frullania platycalyx Herzog, Feddes Repert. Spec. Nov. Regni Veg. 55 (1): 10,
1952 (Herzog 1952h).
* Frullania pyricalycina Steph., Sp. Hepat. (Stephani) 4: 394, 1910 (Stephani 1910b).
** Frullania quillotensis (Nees et Mont.) Nees et Mont., Syn. Hepat. 3: 427, 1845
(Gottsche et al. 1845b). Bas.: Jubula quillotensis Nees et Mont., Ann. Sci. Nat.
Bot. (sér. 2) 5: 64, 1836 (Nees and Montagne 1836).
** Frullania reicheana Steph., Sp. Hepat. (Stephani) 4: 427, 1910 (Stephani 1910b).
* Frullania sabanetica Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 170, 1864 (Gottsche
1864).
* Frullania semienana Gola, Ann. Bot. (Rome) 13 (1): 71, 1914 (Gola 1914a).
** Frullania serrifolia Steph., Sp. Hepat. (Stephani) 4: 526, 1911 (Stephani 1911e).
** Frullania subpyricalycina Herzog, Rev. Bryol. Lichénol. 23 (1/2): 57, 1954 (Herzog
1954).
** Frullania subtruncata Steph., Sp. Hepat. (Stephani) 4: 389, 1910 (Stephani 1910b).
** Frullania turfosa Lindenb. et Gottsche, Syn. Hepat. 5: 779, 1847 (Gottsche et al.
1847).
** Frullania udarii V.Nath et Ajit P.Singh, Curr. Sci. 91 (6): 744, 2006 (Nath and
Singh 2006).
** Frullania valparaisiana Lehm., Nov. Stirp. Pug. 10: 17, 1857 (Lehmann 1857).
** Frullania wagneri Steph., Sp. Hepat. (Stephani) 4: 392, 1910 (Stephani 1910b).
***
Jubulaceae H.Klinggr.
by J. Hentschel and M. von Konrat
he systematic placement of Neohattoria herzogii has been contentious since its description six decades ago. It has been interpreted as either a member of the genus Frullania or segregated into its own genus, Neohattoria, due to morphological similarities
with both Frullania and Jubula. We follow Larrain et al. (2015) that provided molecular evidence supporting the recognition of the genus Neohattoria and its inclusion
within the Jubulaceae, together with Jubula and Nipponolejeunea.
*** Jubula Dumort., Commentat. Bot. (Dumortier): 112, 1822 (Dumortier 1822)
nom. conserv.
*** Jubula blepharophylla Grolle, J. Hattori Bot. Lab. 28: 47, 1965 (Grolle 1965b).
** Jubula hattorii Udar et V.Nath, Misc. Bryol. Lichenol. 8 (3): 49, 1978 (Udar and
Nath 1978).
** Jubula hattorii var. muthukuzhiana A.E.D.Daniels et P.Daniel, Bryol. Southernm. W Ghats: 181, 2013 (Daniels and Daniel 2013).
296
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Jubula himalayensis S.C.Srivast. et D.Sharma, Proc. Indian Acad. Sci. Pl. Sci. 100
(2): 85, 1990 (Srivastava and Sharma 1990).
*** Jubula hutchinsiae (Hook.) Dumort., Syll. Jungerm. Europ.: 36, 1831 (Dumortier
1831). Bas.: Jungermannia hutchinsiae Hook., Brit. Jungermann.: tab. 1, 1812
(Hooker 1812). 230
** Jubula hutchinsiae subsp. australiae Pócs et A.Cairns, Nova Hedwigia 86 (1/2):
232, 2008 (Pócs and Cairns 2008).
** Jubula hutchinsiae subsp. bogotensis (Steph.) Verd., Ann. Cryptog. Exot. 1 (2): 215,
1928 (Verdoorn 1928b). Bas.: Jubula bogotensis Steph., Sp. Hepat. (Stephani) 4:
687, 1911 (Stephani 1911e). 231
*** Jubula hutchinsiae subsp. caucasica Konstant. et Vilnet, Arctoa 20: 234, 2011
(Konstantinova and Vilnet 2011).
*** Jubula hutchinsiae subsp. japonica (Steph.) Horik. et Ando, J. Sci. Hiroshima
Univ., Ser. B, Div. 2, Bot. 6: 300, 1954 (Horikawa and Ando 1954). Bas.: Jubula
japonica Steph., Bull. Herb. Boissier 5 (2): 92, 1897 (Stephani 1897b).
*** Jubula hutchinsiae subsp. javanica (Steph.) Verd., Ann. Cryptog. Exot. 1 (2): 216,
1928 (Verdoorn 1928b). Bas.: Jubula javanica Steph., Sp. Hepat. (Stephani) 4:
688, 1911 (Stephani 1911e). 232
*** Jubula hutchinsiae subsp. pennsylvanica (Steph.) Verd., Ann. Cryptog. Exot. 1 (2):
215, 1928 (Verdoorn 1928b). Bas.: Frullania pennsylvanica Steph., Hedwigia 22
(10): 147, 1883 (Stephani 1883).
** Jubula kwangsiensis C.Gao et K.C.Chang, Bull. Bot. Res., Harbin 4 (3): 89, 1984
(Chang and Gao 1984).
*** Neohattoria Kamim., J. Jap. Bot. 37 (7): 218, 1962 (Kamimura 1962). Nom.
nov. pro Hattoria Kamim., J. Hattori Bot. Lab. 24: 93, 1961 (Kamimura 1961).
*** Neohattoria herzogii (S.Hatt.) Kamim., J. Jap. Bot. 37 (7): 218, 1962 (Kamimura
1962). Bas.: Frullania herzogii S.Hatt., Feddes Repert. Spec. Nov. Regni Veg. 58:
53, 1955 (Hattori 1955).
*** Nipponolejeunea S.Hatt., Bull. Tokyo Sci. Mus. 11: 124, 1944 (Hattori 1944d).
*** Nipponolejeunea pilifera (Steph.) S.Hatt., Bull. Tokyo Sci. Mus. 11: 125, 1944
(Hattori 1944d). Bas.: Pycnolejeunea pilifera Steph., Sp. Hepat. (Stephani) 5: 624,
1914 (Stephani 1914b).
*** Nipponolejeunea subalpina (Horik.) S.Hatt., Bull. Tokyo Sci. Mus. 11: 125, 1944
(Hattori 1944d). Bas.: Pycnolejeunea subalpina Horik., J. Jap. Bot. 15: 360, 1939
(Horikawa 1939).
230 Jubula hutchinsiae has subspecies that may be better recognized at the species level.
231 Jubula hutchinsiae subsp. bogotensis is closely related to subsp. pennsylvanica (Pätsch et al. 2010).
232 Jubula hutchinsiae subsp. javanica includes a cryptic taxon from China (Pätsch et al. 2010).
World checklist of hornworts and liverworts
***
297
Lejeuneaceae Cavers
by S.R. Gradstein, T. Pócs and R.-L. Zhu with contributions by G. Dauphin (Ceratolejeunea), X. He (Pycnolejeunea), A.-L. Ilkiu-Borges (Prionolejeunea), E. ReinerDrehwald (Rectolejeunea, Lejeunea), A. Sass-Gyarmati (Lopholejeunea), A. Schäfer-Verwimp (Diplasiolejeunea) and P. Sukkharak (hysananthus)
he subdivision of Lejeuneaceae follows Gradstein (2013c) with updates from Heinrichs et al. (2013, 2014) and Schäfer-Verwimp et al. (2014). Some nomenclatural and
taxonomic notes can also be found in Gradstein (2013b), Shu and Zhu (2014), Pócs
et al. (2015a, 2015c), Qui et al. (2014) and Söderström et al. (2015a).
***
***
***
Lejeuneoideae
trib. Brachiolejeuneeae
subtrib. Brachiolejeuneinae Gradst.
** Acanthocoleus R.M.Schust., Bull. Torrey Bot. Club 97 (6): 339, 1970 [1971]
(Schuster 1970b).
*** Acanthocoleus aberrans (Lindenb. et Gottsche) Kruijt, Bryophyt. Biblioth. 36: 62,
1988 (Kruijt 1988). Bas.: Lejeunea aberrans Lindenb. et Gottsche, Syn. Hepat. 5:
751, 1847 (Gottsche et al. 1847).
*** Acanthocoleus aberrans var. laevis Gradst., Fl. Neotrop. Monogr. 62: 193, 1994
(Gradstein 1994).
*** Acanthocoleus chrysophyllus (Lehm.) Kruijt, Bryophyt. Biblioth. 36: 72, 1988
(Kruijt 1988). Bas.: Jungermannia chrysophylla Lehm., Linnaea 9 (4): 423, 1835
(Lehmann 1835).
** Acanthocoleus elgonensis Gyarmati et Pócs, Cryptog. Bryol. 35 (2): 120, 2014 (SassGyarmati and Pócs 2014).
*** Acanthocoleus gilvus (Gottsche) Kruijt, Bryophyt. Biblioth. 36: 79, 1988 (Kruijt
1988). Bas.: Lejeunea gilva Gottsche, Abh. Naturwiss. Vereins Bremen 7: 353,
1882 (Gottsche 1882).
*** Acanthocoleus javanicus (Steph.) Kruijt, Bryophyt. Biblioth. 36: 85, 1988 (Kruijt
1988). Bas.: Dicranolejeunea javanica Steph., Sp. Hepat. (Stephani) 5: 169, 1912
(Stephani 1912c).
*** Acanthocoleus juddii Kruijt, Bryophyt. Biblioth. 36: 93, 1988 (Kruijt 1988).
** Acanthocoleus madagascariensis (Steph.) Kruijt, Bryophyt. Biblioth. 36: 98, 1988
(Kruijt 1988). Bas.: Dicranolejeunea madagascariensis Steph., Sp. Hepat. (Stephani) 5: 158, 1912 (Stephani 1912c).
*** Acanthocoleus trigonus (Nees et Mont.) Gradst., Contr. Univ. Michigan Herb. 18:
101, 1992 (Gradstein 1992b). Bas.: Lejeunea trigona Nees et Mont., Ann. Sci.
Nat. Bot. (sér. 2) 5: 61, 1836 (Nees and Montagne 1836).
298
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Acanthocoleus yoshinaganus (S.Hatt.) Kruijt, Bryophyt. Biblioth. 36: 105, 1988
(Kruijt 1988). Bas.: Lopholejeunea subfusca var. yoshinagana S.Hatt., Bot. Mag.
(Tokyo) 58 (686): 38, 1944 (Hattori 1944a).
*** Blepharolejeunea S.W.Arnell, Svensk Bot. Tidskr. 56 (2): 335, 1962 (Arnell 1962b).
*** Blepharolejeunea chimantaensis van Slageren et Kruijt, Beih. Nova Hedwigia 80:
126, 1985 (van Slageren and Kruijt 1985).
*** Blepharolejeunea fuegiana (Besch. et C.Massal.) Gradst., Beih. Nova Hedwigia 80:
130, 1985 (van Slageren and Kruijt 1985). Bas.: Lejeunea fuegiana Besch. et C.
Massal., Bull. Mens. Soc. Linn. Paris 1 (79): 638, 1886 (Bescherelle and Massalongo 1886).
*** Blepharolejeunea incongrua (Lindenb. et Gottsche) van Slageren et Kruijt, Beih.
Nova Hedwigia 80: 133, 1985 (van Slageren and Kruijt 1985). Bas.: Lejeunea
incongrua Lindenb. et Gottsche, Syn. Hepat. 5: 750, 1847 (Gottsche et al. 1847).
*** Blepharolejeunea saccata (Steph.) van Slageren et Kruijt, Beih. Nova Hedwigia 80:
138, 1985 (van Slageren and Kruijt 1985). Bas.: Dicranolejeunea saccata Steph.,
Hedwigia 35 (3): 78, 1896 (Stephani 1896b).
*** Blepharolejeunea securifolia (Steph.) R.M.Schust., Phytologia 45 (5): 424, 1980
(Schuster 1980b). Bas.: Brachiolejeunea securifolia Steph., Sp. Hepat. (Stephani) 5:
128, 1912 (Stephani 1912c).
*** Brachiolejeunea (Spruce) Schifn., Hepat. (Engl.-Prantl): 128, 1893 (Schifner
1893b). Bas.: Lejeunea subg. Brachiolejeunea Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 129, 1884 (Spruce 1884).
*** Brachiolejeunea conduplicata (Steph.) Gradst., Fl. Neotrop. Monogr. 62: 175,
1994 (Gradstein 1994). Bas.: Archilejeunea conduplicata Steph., Sp. Hepat. (Stephani) 4: 712, 1911 (Stephani 1911e).
*** Brachiolejeunea fernandeziana S.W.Arnell, Ark. Bot. (n.ser.) 4 (1): 18, 1957 (Arnell 1957b).
*** Brachiolejeunea laxifolia (Taylor) Schifn., Hepat. (Engl.-Prantl): 128, 1893
(Schifner 1893b). Bas.: Phragmicoma laxifolia Taylor, London J. Bot. 6: 341,
1847 (Taylor 1847b).
*** Brachiolejeunea leiboldiana (Gottsche et Lindenb.) Schifn., Hedwigia 33 (4): 182,
1894 (Schifner 1894). Bas.: Phragmicoma leiboldiana Gottsche et Lindenb., Syn.
Hepat. 2: 296, 1845 (Gottsche et al. 1845a).
*** Brachiolejeunea phyllorhiza (Nees) Kruijt et Gradst., Nova Hedwigia 43 (3/4):
299, 1986 (Kruijt and Gradstein 1986). Bas.: Jungermannia phyllorhiza Nees, Fl.
Bras. (Martius) 1 (1): 348, 1833 (Nees 1833a).
*** Brachiolejeunea spruceana (C.Massal.) Schifn., Hepat. (Engl.-Prantl): 128, 1893
(Schifner 1893b). Bas.: Lejeunea spruceana C.Massal., Nuovo Giorn. Bot. Ital. 17
(3): 246, 1885 (Massalongo 1885).
World checklist of hornworts and liverworts
299
*** Dicranolejeunea (Spruce) Schifn., Hepat. (Engl.-Prantl): 128, 1893 (Schifner
1893b). Bas.: Lejeunea subg. Dicranolejeunea Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 138, 1884 (Spruce 1884).
*** Dicranolejeunea axillaris (Nees et Mont.) Schifn., Hepat. (Engl.-Prantl): 128,
1893 (Schifner 1893b). Bas.: Lejeunea axillaris Nees et Mont., Ann. Sci. Nat.
Bot. (sér. 2) 5: 59, 1836 (Nees and Montagne 1836).
* Dicranolejeunea bovonei Gola, Nuovo Giorn. Bot. Ital. (n.ser.) 27 (2/4): 248, 1920
(Gola 1920).
*** Lindigianthus Kruijt et Gradst., Beih. Nova Hedwigia 80: 165, 1985 (Kruijt and
Gradstein 1985).
*** Lindigianthus cipaconeus (Gottsche) Kruijt et Gradst., Beih. Nova Hedwigia 80:
166, 1985 (Kruijt and Gradstein 1985). Bas.: Lejeunea cipaconea Gottsche, Ann.
Sci. Nat. Bot. (sér. 5) 1: 150, 1864 (Gottsche 1864).
*** Odontolejeunea (Spruce) Schifn., Hepat. (Engl.-Prantl): 127, 1893 (Schifner
1893b). Bas.: Lejeunea subg. Odontolejeunea Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 142, 1884 (Spruce 1884).
*** Odontolejeunea decemdentata (Spruce) Steph., Sp. Hepat. (Stephani) 5: 175, 1912
(Stephani 1912c). Bas.: Lejeunea decemdentata Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 148, 1884 (Spruce 1884).
*** Odontolejeunea lunulata (F.Weber) Schifn., Hepat. (Engl.-Prantl): 128, 1893
(Schifner 1893b). Bas.: Jungermannia lunulata F.Weber, Hist. Musc. Hepat.
Prodr.: 33, 1815 (Weber 1815).
*** Odontolejeunea rhomalea (Spruce) Steph., Sp. Hepat. (Stephani) 5: 176, 1912
(Stephani 1912c). Bas.: Lejeunea rhomalea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 147, 1884 (Spruce 1884).
***
subtrib. Stictolejeuneinae Gradst.
*** Neurolejeunea (Spruce) Schifn., Hepat. (Engl.-Prantl): 131, 1893 (Schifner
1893b). Bas.: Lejeunea subg. Neurolejeunea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 84, 1884 (Spruce 1884).
*** Neurolejeunea breutelii (Gottsche) A.Evans, Bull. Torrey Bot. Club 34 (1): 13,
1907 (Evans 1907b). Bas.: Lejeunea breutelii Gottsche, Syn. Hepat. 3: 324, 1845
(Gottsche et al. 1845b).
** Neurolejeunea breutelii var. africana Pócs, Herzogia 28 (1): 63, 2015 (Pócs et al. 2015b).
*** Neurolejeunea catenulata (Nees) Schifn., Hepat. (Engl.-Prantl): 131, 1893
(Schifner 1893b). Bas.: Lejeunea catenulata Nees, Syn. Hepat. 3: 323, 1845
(Gottsche et al. 1845b).
*** Neurolejeunea sastreana Gradst., Bryologist 92 (3): 345, 1989 (Gradstein 1989).
300
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Neurolejeunea seminervis (Spruce) Schifn., Hepat. (Engl.-Prantl): 131, 1893
(Schifner 1893b). Bas.: Lejeunea seminervis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 84, 1884 (Spruce 1884).
*** Stictolejeunea (Spruce) Schifn., Hepat. (Engl.-Prantl): 131, 1893 (Schifner
1893b). Bas.: Lejeunea subg. Stictolejeunea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 81, 1884 (Spruce 1884).
*** subg. Leptostictolejeunea R.M.Schust., Phytologia 56 (2): 70, 1984 (Schuster
1984).
*** Stictolejeunea balfourii (Mitt.) E.W.Jones, J. Bryol. 9 (1): 50, 1976 (Jones 1976).
Bas.: Lejeunea balfourii Mitt., Philos. Trans. 168: 398, 1879 (Mitten 1879).
*** Stictolejeunea balfourii var. bekkeri Gradst., Beih. Nova Hedwigia 80: 214, 1985
(Gradstein 1985b).
*** Stictolejeunea iwatsukii Mizut., J. Hattori Bot. Lab. 44: 134, 1978 (Mizutani 1978).
*** subg. Stictolejeunea
*** Stictolejeunea squamata (Willd.) Schifn., Hepat. (Engl.-Prantl): 131, 1893
(Schifner 1893b). Bas.: Jungermannia squamata Willd. ex F.Weber, Hist. Musc.
Hepat. Prodr.: 33, 1815 (Weber 1815).
*** trib. Lejeuneeae Dumort.
** Dactylophorella R.M.Schust., Phytologia 45 (5): 427, 1980 (Schuster 1980b).
*** Dactylophorella muricata (Gottsche) R.M.Schust., Phytologia 45 (5): 427, 1980
(Schuster 1980b). Bas.: Lejeunea muricata Gottsche, Syn. Hepat. 3: 348, 1845
(Gottsche et al. 1845b).
*** Metalejeunea Grolle, Bryophyt. Biblioth. 48: 17, 1995 (Grolle 1995).
** Metalejeunea crassitexta (J.B.Jack et Steph.) Pócs, Telopea 13 (3): 456, 2011 (Pócs
et al. 2011). Bas.: Microlejeunea crassitexta J.B.Jack et Steph., Bot. Centralbl. 60
(4): 106, 1894 (Jack and Stephani 1894).
*** Metalejeunea cucullata (Reinw., Blume et Nees) Grolle, Bryophyt. Biblioth. 48:
100, 1995 (Grolle 1995). Bas.: Jungermannia cucullata Reinw., Blume et Nees,
Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 227, 1824
[1825] (Reinwardt et al. 1824a).
*** Metalejeunea winkleri R.L.Zhu et Grolle, Nova Hedwigia 74 (3/4): 498, 2002
(Zhu and Grolle 2002).
*** Pictolejeunea Grolle, Feddes Repert. 88 (4): 248, 1977 (Grolle 1977b).
*** Pictolejeunea levis Grolle et M.E.Reiner, J. Bryol. 27 (3): 281, 2005 (Grolle and
Reiner-Drehwald 2005).
World checklist of hornworts and liverworts
301
*** Pictolejeunea mizutanii Grolle, Feddes Repert. 88 (4): 255, 1977 (Grolle 1977b).
Nom. nov. pro Cheilolejeunea picta Mizut., J. Hattori Bot. Lab. 33: 226, 1970 (Mizutani 1970).
** Pictolejeunea piconii Pócs, Acta Bot. Hung. 49 (1/2): 110, 2007 (Pócs 2007).
*** Pictolejeunea picta (Steph.) Grolle, Feddes Repert. 88 (4): 252, 1977 (Grolle
1977b). Bas.: Prionolejeunea picta Steph., Sp. Hepat. (Stephani) 5: 223, 1913
(Stephani 1913a).
*** Pictolejeunea reginae Ilk.-Borg., Brittonia 54 (4): 318, 2002 [2003] (Ilkiu-Borges
2002).
*** Pictolejeunea sprucei Grolle, Feddes Repert. 88 (4): 249, 1977 (Grolle 1977b).
**
subtrib. Ceratolejeuneinae Gradst.
*** Ceratolejeunea (Spruce) J.B.Jack et Steph., Hedwigia 31 (1): 16, 1892 (Jack and
Stephani 1892). Bas.: Lejeunea subg. Ceratolejeunea Spruce, Trans. & Proc. Bot.
Soc. Edinburgh 15: 198, 1884 (Spruce 1884). 233
*** subg. Ceratolejeunea
*** Ceratolejeunea andringitrae Pócs, Polish Bot. J. 56 (2): 144, 2011 (Pócs 2011c).
** Ceratolejeunea atlantica Alvarenga et Ilk.-Borg., Nova Hedwigia 86 (1/2): 238,
2008 (Ilkiu-Borges and Alvarenga 2008).
** Ceratolejeunea belangeriana (Gottsche) Steph., Sp. Hepat. (Stephani) 5: 396, 1913
(Stephani 1913a). Bas.: Lejeunea belangeriana Gottsche, Syn. Hepat. 3: 398, 1845
(Gottsche et al. 1845b).
** Ceratolejeunea beninensis E.W.Jones et Vanden Berghen, Bull. Jard. Bot. État
Bruxelles 21 (1/2): 63, 1951 (Vanden Berghen 1951b).
*** Ceratolejeunea brevinervis (Spruce) A.Evans, Bull. Torrey Bot. Club 32 (6): 282,
1905 (Evans 1905a). Bas.: Lejeunea brevinervis Spruce, J. Linn. Soc., Bot. 30
(210): 342, 1895 (Gepp 1895b).
*** Ceratolejeunea ceratantha (Nees et Mont.) Schifn., Bot. Jahrb. Syst. 23 (5): 582,
1897 (Schifner 1897). Bas.: Lejeunea ceratantha Nees et Mont., Ann. Sci. Nat.
Bot. (sér. 2) 14: 335, 1840 (Montagne 1840a).
** Ceratolejeunea coalita (Ångstr.) Steph., Sp. Hepat. (Stephani) 5: 402, 1913
(Stephani 1913a). Bas.: Lejeunea coalita Ångstr., Öfvers. Kongl. Vetensk.-Akad.
Förh. 30 (5): 135, 1873 (Ångström 1873).
*** Ceratolejeunea coarina (Gottsche) Schifn., Hepat. (Engl.-Prantl): 125, 1893
(Schifner 1893b). Bas.: Lejeunea coarina Gottsche, Syn. Hepat. 3: 395, 1845
(Gottsche et al. 1845b).
233 he treatment of Ceratolejeunea follows Dauphin (2003) except that his subg. Caduciloba becomes
subg. Ceratolejeunea and his subg. Ceratolejeunea becomes subg. Ceratophora as a consequence of the
introduction of ICN Art. 41.4 (cf. Söderström et al. 2015a).
302
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Ceratolejeunea confusa R.M.Schust., J. Elisha Mitchell Sci. Soc. 72 (2): 313, 1956
(Schuster 1956a).
*** Ceratolejeunea cornuta (Lindenb.) Steph., Planzenw. Ost-Afrikas C: 65, 1895
(Stephani 1895d). Bas.: Jungermannia cornuta Lindenb., Syn. hepat. eur: 23, 1829
(Lindenberg 1829).
*** Ceratolejeunea cubensis (Mont.) Schifn., Hepat. (Engl.-Prantl): 125, 1893
(Schifner 1893b). Bas.: Lejeunea cubensis Mont., Hist. Phys. Cuba, Bot., Pl. Cell.:
481, 1842 (Montagne 1842a).
*** Ceratolejeunea dentistipula Steph., Sp. Hepat. (Stephani) 5: 407, 1913 (Stephani 1913a).
*** Ceratolejeunea fallax (Lehm. et Lindenb.) Bonner, Candollea 14: 189, 1953 (Bonner 1953a). Bas.: Jungermannia fallax Lehm. et Lindenb., Nov. Stirp. Pug. 5: 17,
1833 (Lehmann 1833).
*** Ceratolejeunea ilaria (Taylor) Steph., Sp. Hepat. (Stephani) 5: 412, 1913 (Stephani
1913a). Bas.: Lejeunea ilaria Taylor, Nov. Stirp. Pug. 8: 28, 1844 (Lehmann 1844).
** Ceratolejeunea loribunda Steph., Sp. Hepat. (Stephani) 5: 412, 1913 (Stephani
1913a).
*** Ceratolejeunea guianensis (Nees et Mont.) Steph., Sp. Hepat. (Stephani) 5: 416,
1913 (Stephani 1913a). Bas.: Lejeunea guianensis Nees et Mont., Ann. Sci. Nat.
Bot. (sér. 2) 14: 335, 1840 (Montagne 1840a).
* Ceratolejeunea karstenii Steph., Sp. Hepat. (Stephani) 5: 420, 1913 (Stephani 1913a).
* Ceratolejeunea kuerschneri Eb.Fisch. et Vanderp., Beih. Nova Hedwigia 138: 87,
2010 (Fischer and Vanderpoorten 2010). 234
*** Ceratolejeunea laetefusca (Austin) R.M.Schust., J. Elisha Mitchell Sci. Soc. 72 (2):
306, 1956 (Schuster 1956a). Bas.: Lejeunea laetefusca Austin, Bot. Bull. (Hanover)
1 (8): 36, 1876 (Austin 1876a).
* Ceratolejeunea ledermannii Steph., Sp. Hepat. (Stephani) 6: 399, 1923 (Stephani 1923).
*** Ceratolejeunea malleigera (Spruce) Steph., Sp. Hepat. (Stephani) 5: 422, 1913
(Stephani 1913a). Bas.: Lejeunea malleigera Spruce, Mem. Torrey Bot. Club 1 (3):
123, 1890 (Spruce 1890).
** Ceratolejeunea maranhensis Silva Brito et Ilk.-Borg., Nova Hedwigia 95 (3/4): 424,
2012 (Brito and Ilkiu-Borges 2012).
** Ceratolejeunea minor Mizut., J. Hattori Bot. Lab. 49: 311, 1981 (Mizutani 1981).
*** Ceratolejeunea minuta G.Dauphin, Fl. Neotrop. Monogr. 90: 66, 2003 (Dauphin
2003).
** Ceratolejeunea moniliata Herzog, Mitt. Inst. Allg. Bot. Hamburg 7 (3): 205, 1931
(Herzog 1931a).
** Ceratolejeunea oculata (Gottsche) Steph., Bull. Herb. Boissier 5 (10): 842, 1897
(Stephani 1897c). Bas.: Lejeunea oculata Gottsche, Syn. Hepat. 3: 357, 1845
(Gottsche et al. 1845b).
*** Ceratolejeunea oxygonia Steph., Sp. Hepat. (Stephani) 5: 429, 1913 (Stephani
1913a).
234 Ceratolejeunea kuerschneri is very similar to Ceratolejeunea papulilora and may be conspeciic with it.
World checklist of hornworts and liverworts
303
*** Ceratolejeunea papulilora Steph., Sp. Hepat. (Stephani) 5: 430, 1913 (Stephani
1913a).
*** Ceratolejeunea patentissima (Hampe et Gottsche) A.Evans, Bull. Torrey Bot. Club
32 (6): 286, 1905 (Evans 1905a). Bas.: Lejeunea patentissima Hampe et Gottsche,
Linnaea 25 (3): 355, 1852 [1853] (Hampe and Gottsche 1852).
*** Ceratolejeunea pungens Steph., Sp. Hepat. (Stephani) 5: 434, 1913 (Stephani
1913a).
*** Ceratolejeunea rubiginosa Steph., Hedwigia 34 (5): 237, 1895 (Stephani 1895b).
*** Ceratolejeunea saroltae Pócs, Polish Bot. J. 56 (2): 150, 2011 (Pócs 2011c).
* Ceratolejeunea sinensis P.C.Chen et P.C.Wu, Acta Phytotax. Sin. 9 (3): 232, 1964
(Chen and Wu 1964). 235
** Ceratolejeunea singapurensis (Lindenb.) Schifn., Consp. Hepat. Arch. Ind.: 273,
1898 (Schifner 1898b). Bas.: Lejeunea singapurensis Lindenb., Syn. Hepat. 3: 397,
1845 (Gottsche et al. 1845b).
*** Ceratolejeunea spinosa (Gottsche) Steph., Hedwigia 34 (5): 238, 1895 (Stephani 1895b).
Bas.: Lejeunea spinosa Gottsche, Syn. Hepat. 3: 402, 1845 (Gottsche et al. 1845b).
*** Ceratolejeunea temnantha (Spruce) M.E.Reiner, Cryptog. Bryol. 32 (2): 95, 2011
(Reiner-Drehwald 2011). Bas.: Lejeunea temnantha Spruce, Trans. & Proc. Bot.
Soc. Edinburgh 15: 250, 1884 (Spruce 1884).
*** Ceratolejeunea umbonata Steph., Sp. Hepat. (Stephani) 5: 446, 1913 (Stephani 1913a).
** Ceratolejeunea vitiensis Steph., Sp. Hepat. (Stephani) 5: 448, 1913 (Stephani 1913a).
** Ceratolejeunea zenkeri Steph., Sp. Hepat. (Stephani) 5: 449, 1914 (Stephani 1914b).
*** subg. Ceratophora R.M.Schust., J. Elisha Mitchell Sci. Soc. 72 (2): 294, 1956
(Schuster 1956a).
*** Ceratolejeunea desciscens (Sande Lac.) Schifn., Hepat. (Engl.-Prantl): 126, 1893
(Schifner 1893b). Bas.: Lejeunea desciscens Sande Lac., Syn. hepat. jav.: 107, 1856
[1857] (Sande Lacoste 1856b).
*** Ceratolejeunea globulifera Herzog, Rev. Bryol. Lichénol. 13: 23, 1942 (Herzog 1942e).
*** Ceratolejeunea grandiloba J.B.Jack et Steph., Hedwigia 31 (1): 16, 1892 (Jack and
Stephani 1892).
** Ceratolejeunea grandiloba subsp. inlata (Mizut.) Gradst., Phytotaxa 81 (1): 5,
2013 (Gradstein 2013d). Bas.: Ceratolejeunea inlata Mizut., J. Hattori Bot. Lab.
49: 313, 1981 (Mizutani 1981).
*** Ceratolejeunea szyszylowiczii (Loitl.) Steph., Sp. Hepat. (Stephani) 5: 443, 1913
(Stephani 1913a). Bas.: Lejeunea szyszylowiczii Loitl., Diagn. pl. nov.: 19, 1894
(Loitlesberger 1894).
Incertae sedis
** Ceratolejeunea aliena Herzog, Trans. Brit. Bryol. Soc. 2 (1): 71, 1952 (Herzog 1952a).
235 Ceratolejeunea sinensis may be conspeciic with Drepanolejeunea erecta (Zhu et al. 2005).
304
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Luteolejeunea Piippo, Acta Bot. Fenn. 132: 56, 1986 (Piippo 1986a).
*** Luteolejeunea herzogii (Buchloh) Piippo, Acta Bot. Fenn. 132: 57, 1986 (Piippo
1986a). Bas.: Stictolejeunea herzogii Buchloh, Nova Hedwigia 3 (4): 515, 1961
(Buchloh 1961).
*** Otigoniolejeunea (Spruce) Schifn., Hepat. (Engl.-Prantl): 125, 1893 (Schifner
1893b). Bas.: Lejeunea subg. Otigoniolejeunea Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 226, 1884 (Spruce 1884).
* Otigoniolejeunea crenulata Steph., Sp. Hepat. (Stephani) 6: 408, 1923 (Stephani 1923). 236
*** Otigoniolejeunea huctumalcensis (Lindenb. et Gottsche) Y.M.Wei, R.L.Zhu et
Gradst., Phytotaxa 162 (4): 237, 2014 (Wei et al. 2014). Bas.: Lejeunea huctumalcensis Lindenb. et Gottsche, Syn. Hepat. 5: 762, 1847 (Gottsche et al. 1847).
* Otigoniolejeunea ledermannii Steph., Sp. Hepat. (Stephani) 6: 409, 1923 (Stephani
1923). 237
*** Otigoniolejeunea portoricensis (Hampe et Gottsche) Y.M.Wei, R.L.Zhu et Gradst.,
Phytotaxa 162 (4): 237, 2014 (Wei et al. 2014). Bas.: Lejeunea portoricensis Hampe et Gottsche, Linnaea 25 (3): 352, 1852 [1853] (Hampe and Gottsche 1852).
***
subtrib. Cheilolejeuneinae Gradst.
* Aureolejeunea R.M.Schust., Phytologia 39 (6): 428, 1978 (Schuster 1978b). 238
*** Aureolejeunea aurifera R.M.Schust., Phytologia 39 (6): 429, 1978 (Schuster 1978b).
*** Aureolejeunea lumae (Herzog) van Slageren, Meded. Bot. Mus. Herb. Rijks Univ.
Utrecht 544: 121, 1985 (van Slageren 1985). Bas.: Brachiolejeunea lumae Herzog,
Beih. Bot. Centralbl. 60B (1/2): 15, 1939 (Herzog 1939c).
*** Aureolejeunea paramicola (Herzog) R.M.Schust., Phytologia 61 (7): 446, 1987
(Schuster 1987c). Bas.: Brachiolejeunea paramicola Herzog, Hedwigia 74 (2): 95,
1934 (Herzog 1934a).
*** Aureolejeunea quinquecarinata R.M.Schust., Phytologia 39 (6): 429, 1978 (Schuster
1978b).
*** Aureolejeunea tonduzana (Steph.) Gradst., Phytotaxa 76 (3): 46, 2013 (Gradstein
2013a). Bas.: Archilejeunea tonduzana Steph., Sp. Hepat. (Stephani) 4: 721, 1911
(Stephani 1911e).
236 Otigoniolejeunea crenulata is a doubtful taxon. he type specimen was burned in B and it is unclear
where the taxon belongs (Grolle and Piippo (1984).
237 Otigoniolejeunea ledermannii is a doubtful taxon. he type specimen was burned in B and it is unclear
where the taxon belongs (Grolle and Piippo (1984).
238 Aureolejeunea is nested in Cheilolejeunea (Ye et al. 2011, Gradstein 2013a, Ye et al. 2015). A formal
transfer of the genus and its species to Cheilolejeunea is found in Ye et al. (2015).
World checklist of hornworts and liverworts
305
*** Cheilolejeunea (Spruce) Steph., Bot. Gaz. 15 (11): 284, 1890 (Stephani 1890c).
Bas.: Lejeunea subg. Cheilolejeunea Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 251, 1884 (Spruce 1884). 239
** Cheilolejeunea sandvicensis (Prantl) Steph., Bull. Herb. Boissier 5 (10): 842, 1897
(Stephani 1897c). Bas.: Lejeunea sandvicensis Prantl, Hedwigia 29: xvii, 1890
(Prantl 1890).
** subg. Cheilolejeunea
*** Cheilolejeunea acanthina (Spruce) Gradst. et Ilk.-Borg., Mem. New York Bot.
Gard. 76 (4): 62, 2009 (Gradstein and Ilkiu-Borges 2009). Bas.: Lejeunea acanthina Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 182, 1884 (Spruce 1884).
*** Cheilolejeunea adnata (Kunze ex Lehm.) Grolle, J. Bryol. 9 (4): 529, 1977 [1978]
(Grolle 1977a). Bas.: Jungermannia adnata Kunze ex Lehm., Nov. Stirp. Pug. 6:
46, 1834 (Lehmann 1834).
** Cheilolejeunea adnata var. autoica Gradst. et Ilk.-Borg., Mem. New York Bot.
Gard. 76 (4): 64, 2009 (Gradstein and Ilkiu-Borges 2009).
** Cheilolejeunea albovirens (Hook.f. et Taylor) E.A.Hodgs., Rec. Domin. Mus. 4
(11): 127, 1962 (Hodgson 1962a). Bas.: Jungermannia albovirens Hook.f. et Taylor, London J. Bot. 3: 397, 1844 (Hooker and Taylor 1844a).
*** Cheilolejeunea aneogyna (Spruce) A.Evans, Trans. Connecticut Acad. Arts 10 (8):
440, 1900 (Evans 1900a). Bas.: Lejeunea aneogyna Spruce, Trans. & Proc. Bot.
Soc. Edinburgh 15: 254, 1884 (Spruce 1884).
** Cheilolejeunea ascensionis (Hook.f. et Taylor) Grolle, Haussknechtia 4: 45, 1988
(Grolle 1988a). Bas.: Jungermannia ascensionis Hook.f. et Taylor, London J. Bot.
4: 91, 1845 (Hooker and Taylor 1845).
*** Cheilolejeunea asperilora (Spruce) Gradst. et Ilk.-Borg., Mem. New York Bot.
Gard. 76 (4): 62, 2009 (Gradstein and Ilkiu-Borges 2009). Bas.: Lejeunea asperilora Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 183, 1884 (Spruce 1884).
** Cheilolejeunea australis Solari, Comun. Mus. Argent. Ci. Nat. “Bernardino Rivadavia,” Ci. Bot. 2 (11): 70, 1981 (Solari 1981).
** Cheilolejeunea baumannii Hürl., Bauhinia 11 (3): 160, 1995 (Hürlimann 1995).
*** Cheilolejeunea beyrichii (Lindenb.) M.E.Reiner, Nova Hedwigia 83 (3/4): 474,
2006 (Reiner-Drehwald 2006). Bas.: Lejeunea beyrichii Lindenb., Syn. Hepat. 3:
371, 1845 (Gottsche et al. 1845b).
** Cheilolejeunea boninensis Mizut., J. Hattori Bot. Lab. 51: 153, 1982 (Mizutani 1982).
239 Cheilolejeunea is here organized into subgenera that are maintained for practical purposes. hey may
not be natural lineages. An improved infrageneric classiication based on molecular analysis is in
preparation (Ye et al. 2015). he genus includes Euosmolejeunea, Strepsilejeunea and Trachylejeunea,
but several taxa have neither been transferred nor synonymized. hey are listed under “Names in
genera not currently accepted” below.
306
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Cheilolejeunea caducifolia (Gradst. et Schäf.-Verw.) W.Ye et R.L.Zhu, J. Bryol.
32 (4): 280, 2010 (Ye and Zhu 2010). Bas.: Leucolejeunea caducifolia Gradst. et
Schäf.-Verw., J. Hattori Bot. Lab. 74: 64, 1993 (Gradstein et al. 1993).
** Cheilolejeunea camerunensis S.W.Arnell, Svensk Bot. Tidskr. 52 (1): 63, 1958 (Arnell
1958a).
** Cheilolejeunea celebensis (Steph.) Mizut., J. Hattori Bot. Lab. 36: 157, 1972 [1973]
(Mizutani 1972a). Bas.: Trachylejeunea celebensis Steph., Sp. Hepat. (Stephani) 5:
312, 1913 (Stephani 1913a).
** Cheilolejeunea chenii R.L.Zhu et M.L.So, Taxon 48 (4): 663, 1999 (Zhu et al.
1999). Nom. nov. pro Neurolejeunea fukiensis P.C.Chen et P.C.Wu, Acta Phytotax.
Sin. 9 (3): 227, 1964 (Chen and Wu 1964).
*** Cheilolejeunea clypeata (Schwein.) W.Ye et R.L.Zhu, J. Bryol. 32 (4): 280, 2010
(Ye and Zhu 2010). Bas.: Jungermannia clypeata Schwein., Spec. Fl. Amer. Crypt.:
12, 1821 (Schweinitz 1821).
** Cheilolejeunea compressa (Herzog) Grolle, Bryophyt. Biblioth. 48: 38, 1995
(Grolle 1995). Bas.: Strepsilejeunea compressa Herzog, Bot. Not. 100 (4): 325,
1947 (Herzog 1947).
*** Cheilolejeunea conchifolia (A.Evans) W.Ye et R.L.Zhu, J. Bryol. 32 (4): 280, 2010
(Ye and Zhu 2010). Bas.: Archilejeunea conchifolia A.Evans, Mem. Torrey Bot.
Club 8 (2): 128, 1902 (Evans 1902a).
*** Cheilolejeunea cordigera (Steph.) Grolle, J. Bryol. 9 (4): 530, 1977 [1978] (Grolle 1977a).
Bas.: Hygrolejeunea cordigera Steph., Hedwigia 35 (3): 100, 1896 (Stephani 1896b).
** Cheilolejeunea coronalis (Gottsche) R.M.Schust., Phytologia 45 (5): 431, 1980
(Schuster 1980b). Bas.: Lejeunea coronalis Gottsche, Syn. Hepat. 3: 361, 1845
(Gottsche et al. 1845b).
* Cheilolejeunea curvatilobula (Herzog) Grolle, J. Bryol. 21 (1): 41, 1999 (Grolle
and Reiner-Drehwald 1999). Bas.: Harpalejeunea curvatilobula Herzog, Feddes
Repert. Spec. Nov. Regni Veg. 57 (1/2): 184, 1955 (Herzog 1955).
*** Cheilolejeunea decursiva (Sande Lac.) R.M.Schust., Beih. Nova Hedwigia 9: 112,
1963 (Schuster 1963a). Bas.: Lejeunea decursiva Sande Lac., Ned. Kruidk. Arch. 3:
522, 1855 (Sande Lacoste 1855).
*** Cheilolejeunea discoidea (Lehm. et Lindenb.) R.M.Schust. et Kachroo, J. Linn.
Soc., Bot. 56 (368): 509, 1961 (Kachroo and Schuster 1961). Bas.: Jungermannia
discoidea Lehm. et Lindenb., Nov. Stirp. Pug. 6: 47, 1834 (Lehmann 1834).
** Cheilolejeunea diversifolia Augier, Ann. Fac. Sci. Univ. Féd. Cameroun 11: 66,
1972 (Augier 1972).
** Cheilolejeunea ecarinata Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 54 (1/2): 11,
1984 (Vanden Berghen 1984b).
** Cheilolejeunea erostrata R.M.Schust., Phytologia 39 (6): 427, 1978 (Schuster 1978b).
*** Cheilolejeunea exinnovata E.W.Jones, J. Bryol. 12 (1): 37, 1982 (Jones 1982).
*** Cheilolejeunea fragrantissima (Spruce) R.M.Schust., Phytologia 45 (5): 431, 1980
(Schuster 1980b). Bas.: Lejeunea fragrantissima Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 243, 1884 (Spruce 1884).
World checklist of hornworts and liverworts
*
*
**
**
***
**
**
**
*
**
***
**
**
**
**
**
**
***
307
Cheilolejeunea fukiensis (P.C.Chen et P.C.Wu) Piippo, J. Hattori Bot. Lab. 68:
133, 1990 (Piippo 1990). Bas.: Euosmolejeunea fukiensis P.C.Chen et P.C.Wu,
Acta Phytotax. Sin. 9 (3): 232, 1964 (Chen and Wu 1964). 240
Cheilolejeunea galliotii Steph., Sp. Hepat. (Stephani) 5: 656, 1914 (Stephani 1914b).
Cheilolejeunea germanii (Besch. et Spruce) Grolle, Acta Bot. Fenn. 125: 64, 1984
(Grolle and Piippo 1984). Bas.: Lejeunea germanii Besch. et Spruce, Bull. Soc. Bot.
France (Congr. Bot.) 36: clxxxvii, 1889 [1890] (Bescherelle and Spruce 1889).
Cheilolejeunea ghatensis G.Asthana, S.C.Srivast. et A.K.Asthana, Lindbergia 20
(2/3): 132, 1995 [1996] (Asthana et al. 1995).
Cheilolejeunea gradsteinii (Grolle et Piippo) W.Ye et R.L.Zhu, J. Bryol. 32 (4):
280, 2010 (Ye and Zhu 2010). Bas.: Leucolejeunea gradsteinii Grolle et Piippo,
Ann. Bot. Fenn. 27 (2): 122, 1990 (Grolle and Piippo 1990).
Cheilolejeunea grandibracteata Steph., Sp. Hepat. (Stephani) 5: 657, 1914 (Stephani
1914b).
Cheilolejeunea hamlinii Grolle, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Reihe 31 (2): 212, 1982 (Grolle 1982). Nom. nov. pro Strepsilejeunea
curnowii Steph., Hedwigia 35 (3): 129, 1896 (Stephani 1896b).
Cheilolejeunea hawaica Steph., Bull. Herb. Boissier 5 (10): 847, 1897 (Stephani 1897c).
Cheilolejeunea herzogiana Steph., Biblioth. Bot. 87 (2): 267, 1916 (Stephani 1916a).
Cheilolejeunea implexicaulis (Hook.f. et Taylor) R.M.Schust., J. Hattori Bot. Lab.
26: 245, 1963 (Schuster 1963b). Bas.: Jungermannia implexicaulis Hook.f. et Taylor, London J. Bot. 3: 397, 1844 (Hooker and Taylor 1844a).
Cheilolejeunea insecta Grolle et Gradst., Taxon 50 (4): 1071, 2001 [2002] (Grolle
et al. 2001).
Cheilolejeunea insignis Jovet-Ast et Tixier, Rev. Bryol. Lichénol. 31 (1/2): 25, 1962
(Jovet-Ast and Tixier 1962).
Cheilolejeunea intricata (Steph.) J.J.Engel, Bryologist 79 (4): 514, 1976 [1977]
(Engel 1976b). Bas.: Harpalejeunea intricata Steph., Sp. Hepat. (Stephani) 5: 269,
1913 (Stephani 1913a).
Cheilolejeunea invaginata R.M.Schust., Phytologia 39 (6): 427, 1978 (Schuster 1978b).
Cheilolejeunea jamaicensis Steph., Hedwigia 34 (5): 241, 1895 (Stephani 1895b).
Cheilolejeunea japonica (Horik.) W.Ye et R.L.Zhu, J. Bryol. 32 (4): 280, 2010 (Ye
and Zhu 2010). Bas.: Archilejeunea japonica Horik., J. Sci. Hiroshima Univ., Ser.
B, Div. 2, Bot. 1: 84, 1932 (Horikawa 1932a).
Cheilolejeunea kitagawae W.Ye et R.L.Zhu, J. Bryol. 32 (4): 280, 2010 (Ye and
Zhu 2010). Nom. nov. pro Leucolejeunea paroica N.Kitag., Acta Phytotax. Geobot.
18 (7): 190, 1960 (Kitagawa 1960a).
Cheilolejeunea lacerata C.J.Bastos et Gradst., J. Bryol. 28 (2): 133, 2006 (Bastos
and Gradstein 2006).
240 Cheilolejeunea fukiensis may be conspeciic with Lejeunea lava (Zhu and So 2001), but they did not
see the type specimen.
308
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Cheilolejeunea larsenii Mizut., Dansk Bot. Ark. 27 (1): 95, 1969 (Hattori and Mizutani 1969).
* Cheilolejeunea laurentii Steph., Sp. Hepat. (Stephani) 5: 647, 1914 (Stephani
1914b). 241
** Cheilolejeunea leptophylla (Ångstr.) Steph., Sp. Hepat. (Stephani) 5: 657, 1914
(Stephani 1914b). Bas.: Lejeunea leptophylla Ångstr., Öfvers. Kongl. Vetensk.-Akad.
Förh. 33 (7): 86, 1876 [1877] (Ångström 1876).
* Cheilolejeunea longilora (Taylor) R.M.Schust., Phytologia 45 (5): 431, 1980
(Schuster 1980b). Bas.: Lejeunea longilora Taylor, London J. Bot. 5: 396, 1846
(Taylor 1846b). 242
** Cheilolejeunea longispina (Herzog) R.M.Schust., Beih. Nova Hedwigia 9: 111,
1963 (Schuster 1963a). Bas.: Harpalejeunea longispina Herzog, Rev. Bryol. Lichénol. 23 (1/2): 61, 1954 (Herzog 1954).
** Cheilolejeunea loriana (Steph.) W.Ye et R.L.Zhu, J. Bryol. 32 (4): 280, 2010 (Ye
and Zhu 2010). Bas.: Symbiezidium lorianum Steph., Sp. Hepat. (Stephani) 5:
106, 1912 (Stephani 1912c).
** Cheilolejeunea ludoviciae Steph., Sp. Hepat. (Stephani) 5: 668, 1914 (Stephani 1914b).
** Cheilolejeunea lurida (Lindenb.) Steph., Sp. Hepat. (Stephani) 5: 658, 1914 (Stephani 1914b). Bas.: Lejeunea lurida Lindenb., Syn. Hepat. 3: 379, 1845 (Gottsche
et al. 1845b).
** Cheilolejeunea macroloba (Herzog) Grolle, Wiss. Z. Friedrich-Schiller-Univ. Jena,
Math.-Naturwiss. Reihe 31 (2): 212, 1982 (Grolle 1982). Bas.: Strepsilejeunea macroloba Herzog, Nat. Hist. Juan Fernandez (Botany) 2 (5): 742, 1942 (Herzog 1942a).
*** Cheilolejeunea malaccensis (G.Hofm.) Xiao L.He, Ann. Bot. Fenn. 33 (1): 59,
1996 (He 1996b). Bas.: Pycnolejeunea malaccensis G.Hofm., Ann. Bryol. 8: 118,
1935 (Hofman 1935).
** Cheilolejeunea mammifera R.M.Schust., Phytologia 45 (5): 429, 1980 (Schuster 1980b).
*** Cheilolejeunea mariana (Gottsche) B.M.hiers et Gradst., Mem. New York Bot.
Gard. 52: 75, 1989 (hiers and Gradstein 1989). Bas.: Lejeunea mariana Gottsche,
Syn. Hepat. 3: 337, 1845 (Gottsche et al. 1845b).
** Cheilolejeunea mexicana Steph., Sp. Hepat. (Stephani) 6: 417, 1923 (Stephani 1923).
* Cheilolejeunea micholitzii (Steph.) R.M.Schust. et Kachroo, J. Linn. Soc., Bot. 56
(368): 509, 1961 (Kachroo and Schuster 1961). Bas.: Pycnolejeunea micholitzii
Steph., Sp. Hepat. (Stephani) 5: 627, 1914 (Stephani 1914b). 243
241 Cheilolejeunea laurentii is not a Cheilolejeunea species (Jones 1954b, Wigginton and Grolle 1996).
We do not know where to refer it.
242 Cheilolejeunea longilora is possibly conspeciic with Cheilolejeunea trifaria (Gradstein and Costa
2003).
243 Cheilolejeunea micholitzii is conspeciic with Cheilolejeunea longidens in hiers (1992b), but accepted
by Hürlimann (1995).
World checklist of hornworts and liverworts
309
** Cheilolejeunea microscypha (Hook.f. et Taylor) M.Wigginton, J. Bryol. 34 (4):
269, 2012 (Wigginton 2012). Bas.: Jungermannia microscypha Hook.f. et Taylor,
London J. Bot. 4: 90, 1845 (Hooker and Taylor 1845).
*** Cheilolejeunea mizutanii W.Ye et R.L.Zhu, J. Bryol. 32 (4): 281, 2010 (Ye and
Zhu 2010). Nom. nov. pro Hygrolejeunea decurrens Steph., Hedwigia 35 (3): 101,
1896 (Stephani 1896b).
*** Cheilolejeunea neblinensis Ilk.-Borg. et Gradst., Nova Hedwigia 87 (3/4): 522,
2008 (Ilkiu-Borges and Gradstein 2008).
** Cheilolejeunea nietneri (Steph.) Mizut., J. Hattori Bot. Lab. 37: 191, 1973 (Mizutani 1973). Bas.: Harpalejeunea nietneri Steph., Sp. Hepat. (Stephani) 6: 392,
1923 (Stephani 1923).
*** Cheilolejeunea norisiae G.Dauphin et Gradst., J. Bryol. 25 (4): 259, 2003 (Dauphin
and Gradstein 2003).
** Cheilolejeunea novaezelandiae R.M.Schust., Phytologia 56 (7): 459, 1985 (Schuster
1985c).
*** Cheilolejeunea obcordata Herzog, Mitt. Inst. Allg. Bot. Hamburg 7 (3): 209, 1931
(Herzog 1931a).
** Cheilolejeunea obtruncata (Mont.) Solari, J. Hattori Bot. Lab. 54: 539, 1983 (Solari 1983a). Bas.: Lejeunea obtruncata Mont., Ann. Sci. Nat. Bot. (sér. 3) 4: 354,
1845 (Montagne 1845b).
** Cheilolejeunea obtusa (Herzog) Solari, J. Hattori Bot. Lab. 54: 537, 1983 (Solari
1983a). Bas.: Harpalejeunea obtusa Herzog, Rev. Bryol. Lichénol. 29 (3/4): 193,
1960 [1961] (Herzog 1960).
** Cheilolejeunea obtusifolia (Steph.) S.Hatt., J. Hattori Bot. Lab. 18: 116, 1957
(Hattori 1957a). Bas.: Harpalejeunea obtusifolia Steph., Sp. Hepat. (Stephani) 5:
265, 1913 (Stephani 1913a).
** Cheilolejeunea occlusa (Herzog) T.Kodama et N.Kitag., Bull. Osaka Mus. Nat.
Hist. 28: 40, 1974 (Kitagawa and Kodama 1974). Bas.: Strepsilejeunea occlusa Herzog, Trans. Brit. Bryol. Soc. 1 (4): 320, 1950 (Herzog 1950a).
** Cheilolejeunea orientalis (Gottsche) Mizut., J. Hattori Bot. Lab. 35: 399, 1972
(Mizutani 1972b). Bas.: Lejeunea orientalis Gottsche, Syn. Hepat. 3: 371, 1845
(Gottsche et al. 1845b).
** Cheilolejeunea ovistipula Steph., Hedwigia 34 (5): 244, 1895 (Stephani 1895b).
** Cheilolejeunea panurensis (Spruce) Steph., Sp. Hepat. (Stephani) 5: 660, 1914
(Stephani 1914b). Bas.: Lejeunea panurensis Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 255, 1884 (Spruce 1884).
** Cheilolejeunea papillata Solari, Comun. Mus. Argent. Ci. Nat. “Bernardino Rivadavia,” Ci. Bot. 2 (11): 72, 1981 (Solari 1981).
** Cheilolejeunea paroica Mizut., J. Hattori Bot. Lab. 46: 364, 1979 (Mizutani 1979b).
** Cheilolejeunea pirilora Schifn., Bot. Jahrb. Syst. 23 (5): 592, 1897 (Schifner
1897). Based on: Lejeunea pirilora Gottsche ex Pol., J. Bot. 15: 227, 1877 (Polakowski 1877), nom. inval.
310
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Cheilolejeunea polystachya (Spruce) Gradst. et Ilk.-Borg., Mem. New York Bot.
Gard. 76 (4): 62, 2009 (Gradstein and Ilkiu-Borges 2009). Bas.: Lejeunea polystachya Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 250, 1884 (Spruce 1884).
** Cheilolejeunea renneri (G.Hofm.) Xiao L.He, Ann. Bot. Fenn. 33 (1): 62, 1996
(He 1996b). Bas.: Pycnolejeunea renneri G.Hofm., Ann. Bryol. 8: 117, 1935
(Hofman 1935).
*** Cheilolejeunea rotundistipula (Lindenb. ex Lehm.) Malombe, Acta Bot. Hung. 51
(3/4): 322, 2009 (Malombe 2009). Bas.: Jungermannia rotundistipula Lindenb. ex
Lehm., Linnaea 4: 360, 1829 (Lehmann 1829).
** Cheilolejeunea rufescens (Lindenb.) Grolle, Wiss. Z. Friedrich-Schiller-Univ. Jena,
Math.-Naturwiss. Reihe 31 (2): 212, 1982 (Grolle 1982). Bas.: Lejeunea rufescens
Lindenb., Syn. Hepat. 3: 366, 1845 (Gottsche et al. 1845b).
** Cheilolejeunea stenoschiza (Ångstr.) A.Evans, Trans. Connecticut Acad. Arts 10
(8): 436, 1900 (Evans 1900a). Bas.: Lejeunea stenoschiza Ångstr., Öfvers. Kongl.
Vetensk.-Akad. Förh. 29 (4): 26, 1872 (Ångström 1872).
** Cheilolejeunea subcrenulata (Spruce) R.M.Schust., Phytologia 45 (5): 431, 1980
(Schuster 1980b). Bas.: Lejeunea subcrenulata Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 245, 1884 (Spruce 1884).
* Cheilolejeunea suborbicularis (Herzog) H.A.Mill., Bonner et Bischl., Nova Hedwigia 4: 559, 1962 [1963] (Miller et al. 1962). Bas.: Microlejeunea suborbicularis Herzog, Memoranda Soc. Fauna Fl. Fennica 26: 65, 1950 [1951] (Herzog 1950b).
*** Cheilolejeunea suprema (Grolle et Piippo) W.Ye et R.L.Zhu, J. Bryol. 32 (4): 281,
2010 (Ye and Zhu 2010). Bas.: Leucolejeunea suprema Grolle et Piippo, Ann. Bot.
Fenn. 27 (2): 123, 1990 (Grolle and Piippo 1990).
** Cheilolejeunea surrepens (Mitt.) E.W.Jones, J. Bryol. 9 (1): 49, 1976 (Jones 1976).
Bas.: Lejeunea surrepens Mitt., Philos. Trans. 168: 399, 1879 (Mitten 1879).
*** Cheilolejeunea turgida (Mitt.) W.Ye et R.L.Zhu, J. Bryol. 32 (4): 281, 2010 (Ye
and Zhu 2010). Bas.: Lejeunea turgida Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 110,
1860 [1861] (Mitten 1860c).
* Cheilolejeunea ulugurica Malombe, Eb.Fisch. et Pócs, Acta Biol. Pl. Agr. 1: 24,
2010 [2011] (Malombe et al. 2010). 244
*** Cheilolejeunea unciloba (Lindenb.) Malombe, Acta Bot. Hung. 51 (3/4): 325,
2009 (Malombe 2009). Bas.: Lejeunea unciloba Lindenb., Syn. Hepat. 3: 331,
1845 (Gottsche et al. 1845b).
** Cheilolejeunea upoluensis S.W.Arnell, Svensk Bot. Tidskr. 50 (3): 516, 1956 (Arnell
1956a).
** Cheilolejeunea valenciae (Steph.) Xiao L.He, Ann. Bot. Fenn. 33 (1): 55, 1996 (He
1996a). Bas.: Pycnolejeunea valenciae Steph., Sp. Hepat. (Stephani) 5: 605, 1914
(Stephani 1914b).
244 Cheilolejeunea ulugurica is possibly conspeciic with Cheilolejeunea chenii (Ye et al. 2013a).
World checklist of hornworts and liverworts
311
** Cheilolejeunea virescens (Lehm. et Lindenb.) Steph., Sp. Hepat. (Stephani) 5: 662,
1914 (Stephani 1914b). Bas.: Lejeunea virescens Lehm. et Lindenb., Nov. Stirp.
Pug. 7: 21, 1838 (Lehmann 1838).
* Cheilolejeunea viridis Steph., Sp. Hepat. (Stephani) 5: 673, 1914 (Stephani 1914b).
** Cheilolejeunea warnstorii (Steph.) Solari, Darwiniana 20 (3/4): 387, 1976 (Solari 1976). Bas.: Strepsilejeunea warnstorii Steph., Hedwigia 35 (3): 131, 1896
(Stephani 1896b).
* Cheilolejeunea wrightii Steph., Sp. Hepat. (Stephani) 5: 662, 1914 (Stephani 1914b).
*** Cheilolejeunea xanthocarpa (Lehm. et Lindenb.) Malombe, Acta Bot. Hung. 51
(3/4): 326, 2009 (Malombe 2009). Bas.: Jungermannia xanthocarpa Lehm. et Lindenb., Nov. Stirp. Pug. 5: 8, 1833 (Lehmann 1833).
** Cheilolejeunea xanthophylla (Lindenb.) Steph., Sp. Hepat. (Stephani) 5: 663, 1914
(Stephani 1914b). Bas.: Lejeunea xanthophylla Lindenb., Syn. Hepat. 3: 370, 1845
(Gottsche et al. 1845b).
** subg. Euosmolejeunea (Spruce) Kachroo, Ceylon J. Sci., Biol. Sci. 8 (1): 6, 1968
(Kachroo 1968). Bas.: Lejeunea subg. Euosmolejeunea Spruce, Trans. & Proc. Bot.
Soc. Edinburgh 15: 241, 1884 (Spruce 1884).
** Cheilolejeunea aciculifera R.M.Schust., Phytologia 45 (5): 428, 1980 (Schuster 1980b).
** Cheilolejeunea acutangula (Nees) Grolle, J. Hattori Bot. Lab. 45: 173, 1979 (Grolle
1979c). Bas.: Jungermannia acutangula Nees, Fl. Bras. (Martius) 1 (1): 357, 1833
(Nees 1833a).
*** Cheilolejeunea asperrima (Steph.) Grolle, J. Hattori Bot. Lab. 58: 197, 1985 (Grolle 1985b). Bas.: Taxilejeunea asperrima Steph., Biblioth. Bot. 87 (2): 259, 1916
(Stephani 1916a).
** Cheilolejeunea birmensis (Steph.) Mizut., J. Hattori Bot. Lab. 27: 139, 1964 (Mizutani 1964a). Bas.: Strepsilejeunea birmensis Steph., Sp. Hepat. (Stephani) 5: 286,
1913 (Stephani 1913a).
** Cheilolejeunea campbelliensis (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 245,
1963 (Schuster 1963b). Bas.: Strepsilejeunea campbelliensis Steph., Hedwigia 35
(3): 128, 1896 (Stephani 1896b).
** Cheilolejeunea cedercreutzii (H.Buch et Perss.) Grolle, Feddes Repert. 87 (3/4):
188, 1976 (Grolle 1976a). Bas.: Euosmolejeunea cedercreutzii H.Buch et Perss.,
Bryophyt. Azoren Madeira: 9, 1941 (Buch and Persson 1941).
** Cheilolejeunea choachina (Gottsche) Gradst., Mem. New York Bot. Gard. 84: 69,
1999 (Gradstein 1999). Bas.: Lejeunea choachina Gottsche, Ann. Sci. Nat. Bot.
(sér. 5) 1: 156, 1864 (Gottsche 1864).
*** Cheilolejeunea clausa (Nees et Mont.) R.M.Schust., Hepat. Anthocerotae N. Amer.
4: 863, 1980 (Schuster 1980c). Bas.: Lejeunea clausa Nees et Mont., Ann. Sci. Nat.
Bot. (sér. 2) 14: 337, 1840 (Montagne 1840a).
** Cheilolejeunea clavata Mizut., J. Hattori Bot. Lab. 46: 361, 1979 (Mizutani 1979b).
312
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Cheilolejeunea comans (Spruce) R.M.Schust., Phytologia 45 (5): 431, 1980 (Schuster 1980b). Bas.: Lejeunea comans Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:
246, 1884 (Spruce 1884).
** Cheilolejeunea convexa (S.W.Arnell) S.W.Arnell, Hepat. South Africa: 209, 1963 (Arnell
1963b). Bas.: Lejeunea convexa S.W.Arnell, Bot. Not. 106: 272, 1953 (Arnell 1953b).
** Cheilolejeunea cookiensis (Steph.) R.M.Schust. et Kachroo, J. Linn. Soc., Bot. 56
(368): 509, 1961 (Kachroo and Schuster 1961). Bas.: Pycnolejeunea cookiensis
Steph., Sp. Hepat. (Stephani) 5: 617, 1914 (Stephani 1914b).
** Cheilolejeunea cordistipula (Steph.) Grolle ex E.W.Jones, J. Bryol. 13 (3): 395,
1985 (Jones 1985). Bas.: Strepsilejeunea cordistipula Steph., Sp. Hepat. (Stephani)
5: 276, 1913 (Stephani 1913a).
** Cheilolejeunea ischeri Malombe, Acta Bot. Hung. 51 (3/4): 319, 2009 (Malombe 2009).
** Cheilolejeunea gaoi R.L.Zhu, M.L.So et Grolle, Bryologist 103 (3): 499, 2000
(Zhu et al. 2000).
** Cheilolejeunea hallingii B.M.hiers, Brittonia 44 (2): 160, 1992 (hiers 1992a).
*** Cheilolejeunea inlexa (Hampe) Grolle, J. Hattori Bot. Lab. 45: 174, 1979 (Grolle
1979c). Bas.: Lejeunea inlexa Hampe, Nov. Stirp. Pug. 7: 22, 1838 (Lehmann 1838).
*** Cheilolejeunea intertexta (Lindenb.) Steph., Bull. Herb. Boissier 5 (2): 79, 1897
(Stephani 1897b). Bas.: Lejeunea intertexta Lindenb., Syn. Hepat. 3: 379, 1845
(Gottsche et al. 1845b).
*** Cheilolejeunea krakakammae (Lindenb.) R.M.Schust., Beih. Nova Hedwigia 9:
112, 1963 (Schuster 1963a). Bas.: Lejeunea krakakammae Lindenb., Syn. Hepat.
3: 353, 1845 (Gottsche et al. 1845b).
*** Cheilolejeunea laciniata D.F.Peralta et M.E.Reiner, Bryologist 116 (1): 54, 2013
(Peralta and Reiner-Drehwald 2013).
*** Cheilolejeunea laevicalyx (J.B.Jack et Steph.) Grolle, J. Hattori Bot. Lab. 65: 403,
1988 (Grolle 1988c). Bas.: Strepsilejeunea laevicalyx J.B.Jack et Steph., Sp. Hepat.
(Stephani) 5: 283, 1913 (Stephani 1913a).
** Cheilolejeunea laeviuscula (Mitt.) Steph., Sp. Hepat. (Stephani) 5: 668, 1914
(Stephani 1914b). Bas.: Lejeunea laeviuscula Mitt., J. Proc. Linn. Soc., Bot. 5 (18):
114, 1860 [1861] (Mitten 1860c).
*** Cheilolejeunea lindenbergii (Gottsche) Mizut., J. Hattori Bot. Lab. 33: 226, 1970
(Mizutani 1970). Bas.: Lejeunea lindenbergii Gottsche, Syn. Hepat. 3: 336, 1845
(Gottsche et al. 1845b).
** Cheilolejeunea mimosa (Hook.f. et Taylor) R.M.Schust., J. Hattori Bot. Lab. 26:
245, 1963 (Schuster 1963b). Bas.: Jungermannia mimosa Hook.f. et Taylor, London
J. Bot. 3: 398, 1844 (Hooker and Taylor 1844a).
** Cheilolejeunea nana R.M.Schust., Phytologia 39 (6): 426, 1978 (Schuster 1978b).
** Cheilolejeunea ngongensis Malombe et Pócs, Acta Bot. Hung. 51 (3/4): 317, 2009
(Malombe 2009).
** Cheilolejeunea nipponica (S.Hatt.) S.Hatt., Misc. Bryol. Lichenol. 1 (14): 1, 1957
(Hattori 1957c). Bas.: Strepsilejeunea nipponica S.Hatt., Bull. Tokyo Sci. Mus. 11:
134, 1944 (Hattori 1944d).
World checklist of hornworts and liverworts
313
*** Cheilolejeunea omphalogastria Pócs, Trop. Bryol. 9: 131, 1994 (Pócs 1994b).
*** Cheilolejeunea oncophylla (Ångstr.) Grolle et M.E.Reiner, J. Bryol. 19 (4): 781,
1997 (Grolle and Reiner-Drehwald 1997). Bas.: Lejeunea oncophylla Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 33 (7): 86, 1876 [1877] (Ångström 1876).
** Cheilolejeunea ornata C.J.Bastos, J. Bryol. 33 (1): 86, 2011 (Bastos 2011).
** Cheilolejeunea osumiensis (S.Hatt.) Mizut., Misc. Bryol. Lichenol. 8 (7): 148, 1980
(Mizutani 1980). Bas.: Euosmolejeunea osumiensis S.Hatt., Bull. Tokyo Sci. Mus.
11: 105, 1944 (Hattori 1944d).
** Cheilolejeunea pluriplicata (Pearson) R.M.Schust., Phytologia 45 (5): 430, 1980
(Schuster 1980b). Bas.: Lejeunea pluriplicata Pearson, Forh. Vidensk.-Selsk. Kristiania 1887 (9): 5, 1887 (Pearson 1887b).
** Cheilolejeunea pocsii E.W.Jones, J. Bryol. 15 (1): 156, 1988 (Jones 1988).
** Cheilolejeunea polyantha A.Evans, Mem. Torrey Bot. Club 8 (2): 141, 1902 (Evans
1902a).
** Cheilolejeunea polyantha var. caduciloba R.M.Schust., Hepat. Anthocerotae N.
Amer. 4: 887, 1980 (Schuster 1980c).
*** Cheilolejeunea revoluta (Herzog) Gradst. et Grolle, J. Hattori Bot. Lab. 74: 59,
1993 (Gradstein et al. 1993). Bas.: Pycnolejeunea revoluta Herzog, Feddes Repert.
Spec. Nov. Regni Veg. 57 (1/2): 193, 1955 (Herzog 1955).
*** Cheilolejeunea rigidula (Nees ex Mont.) R.M.Schust., Castanea 36 (2): 102, 1971
(Schuster 1971a). Bas.: Lejeunea rigidula Nees ex Mont., Ann. Sci. Nat. Bot. (sér.
2) 14: 336, 1840 (Montagne 1840a).
** Cheilolejeunea ruwenzorensis (S.W.Arnell) R.M.Schust., Beih. Nova Hedwigia 9:
112, 1963 (Schuster 1963a). Bas.: Euosmolejeunea ruwenzorensis S.W.Arnell, Ark.
Bot. (n.ser.) 3 (16): 529, 1956 (Arnell 1956e).
** Cheilolejeunea ryukyuensis Mizut., J. Hattori Bot. Lab. 51: 162, 1982 (Mizutani 1982).
** Cheilolejeunea subopaca (Mitt.) Mizut., J. Hattori Bot. Lab. 26: 183, 1963 (Mizutani 1963). Bas.: Lejeunea subopaca Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 116, 1860
[1861] (Mitten 1860c).
*** Cheilolejeunea trifaria (Reinw., Blume et Nees) Mizut., J. Hattori Bot. Lab. 27:
132, 1964 (Mizutani 1964b). Bas.: Jungermannia trifaria Reinw., Blume et Nees,
Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 226, 1824
[1825] (Reinwardt et al. 1824a).
* Cheilolejeunea udarii G.Asthana, S.C.Srivast. et A.K.Asthana, Lindbergia 20 (2/3):
142, 1995 [1996] (Asthana et al. 1995). 245
** Cheilolejeunea usambarana (Steph.) Grolle, J. Hattori Bot. Lab. 46: 344, 1979
(Grolle 1979d). Bas.: Strepsilejeunea usambarana Steph., Akad. Wiss. Wien,
Math.-Naturwiss. Kl., Denkschr. 88: 731, 1913 (Stephani 1913b).
** Cheilolejeunea verrucosa Steph., Sp. Hepat. (Stephani) 5: 673, 1914 (Stephani 1914b).
245 Cheilolejeunea udarii is possibly conspeciic with Cheilolejeunea krakakammae (Zhu 2006a).
314
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** subg. Renilejeunea R.M.Schust., Beih. Nova Hedwigia 9: 112, 1963 (Schuster 1963a).
*** Cheilolejeunea montagnei (Gottsche ex Mont.) R.M.Schust., Beih. Nova Hedwigia
9: 112, 1963 (Schuster 1963a). Bas.: Lejeunea montagnei Gottsche ex Mont., Ann.
Sci. Nat. Bot. (sér. 2) 19: 261, 1843 (Montagne 1843).
** subg. Xenolejeunea Kachroo et R.M.Schust., J. Linn. Soc., Bot. 56 (368): 496,
1961 (Kachroo and Schuster 1961).
** Cheilolejeunea ceylanica (Gottsche) R.M.Schust. et Kachroo, J. Linn. Soc., Bot. 56
(368): 509, 1961 (Kachroo and Schuster 1961). Bas.: Lejeunea ceylanica Gottsche,
Syn. Hepat. 3: 359, 1845 (Gottsche et al. 1845b).
** Cheilolejeunea eximia (Jovet-Ast et Tixier) R.L.Zhu et M.L.So, Beih. Nova Hedwigia 121: 114, 2001 (Zhu and So 2001). Bas.: Pycnolejeunea eximia Jovet-Ast et
Tixier, Rev. Bryol. Lichénol. 31 (1/2): 31, 1962 (Jovet-Ast and Tixier 1962).
*** Cheilolejeunea falsinervis (Sande Lac.) R.M.Schust. et Kachroo, J. Linn. Soc., Bot.
56 (368): 509, 1961 (Kachroo and Schuster 1961). Bas.: Lejeunea falsinervis Sande
Lac., Ned. Kruidk. Arch. 3: 421, 1854 [1855] (Sande Lacoste 1854).
** Cheilolejeunea gardneri (Mitt.) Mizut., J. Hattori Bot. Lab. 26: 181, 1963 (Mizutani 1963). Bas.: Lejeunea gardneri Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 115, 1860
[1861] (Mitten 1860c).
** Cheilolejeunea gigantea (Steph.) R.M.Schust. et Kachroo, J. Linn. Soc., Bot. 56
(368): 509, 1961 (Kachroo and Schuster 1961). Bas.: Pycnolejeunea gigantea Steph., Hedwigia 35 (3): 125, 1896 (Stephani 1896b).
** Cheilolejeunea huerlimannii Tixier, Misc. Bryol. Lichenol. 9 (9): 184, 1983 (Tixier
1983b).
** Cheilolejeunea hyalomarginata R.L.Zhu et Frank Müll., Bryologist 115 (2): 218,
2012 (Zhu and Müller 2012).
*** Cheilolejeunea incisa (Gottsche) R.M.Schust. et Kachroo, J. Linn. Soc., Bot. 56
(368): 509, 1961 (Kachroo and Schuster 1961). Bas.: Lejeunea incisa Gottsche,
Syn. Hepat. 3: 360, 1845 (Gottsche et al. 1845b).
** Cheilolejeunea incisa var. teretilora B.M.hiers, J. Hattori Bot. Lab. 82: 321, 1997
(hiers 1997a).
** Cheilolejeunea longidens (Steph.) R.M.Schust. et Kachroo, J. Linn. Soc., Bot. 56
(368): 509, 1961 (Kachroo and Schuster 1961). Bas.: Pycnolejeunea longidens
Steph., Sp. Hepat. (Stephani) 5: 634, 1914 (Stephani 1914b).
** Cheilolejeunea meyeniana (Nees, Lindenb. et Gottsche) R.M.Schust. et Kachroo,
J. Linn. Soc., Bot. 56 (368): 509, 1961 (Kachroo and Schuster 1961). Bas.:
Lejeunea meyeniana Nees, Lindenb. et Gottsche, Observ. bot.: 472, 1843
(Gottsche et al. 1843).
** Cheilolejeunea obtusilobula (S.Hatt.) S.Hatt., Misc. Bryol. Lichenol. 1 (14): 2,
1957 (Hattori 1957c). Bas.: Pycnolejeunea obtusilobula S.Hatt., J. Hattori Bot.
Lab. 3: 44, 1948 [1950] (Hattori 1948a).
** Cheilolejeunea parvidens B.M.hiers, J. Hattori Bot. Lab. 82: 323, 1997 (hiers 1997a).
World checklist of hornworts and liverworts
315
** Cheilolejeunea streimannii Pócs et Ninh, Acta Bot. Hung. 47 (1/2): 162, 2005
(Pócs and Ninh 2005).
*** Cheilolejeunea trapezia (Nees) Kachroo et R.M.Schust., J. Linn. Soc., Bot. 56
(368): 509, 1961 (Kachroo and Schuster 1961). Bas.: Jungermannia trapezia Nees,
Enum. Pl. Crypt. Javae: 41, 1830 (Nees 1830).
** Cheilolejeunea ventricosa (Schifn. ex P.Syd.) Xiao L.He, Acta Bot. Fenn. 163: 60,
1999 (He 1999). Bas.: Pycnolejeunea ventricosa Schifn. ex P.Syd., Just’s Bot. Jahresber. 19: 246, 1894 (Sydow 1894).
*** Cheilolejeunea vittata (Steph. ex G.Hofm.) R.M.Schust. et Kachroo, J. Linn. Soc.,
Bot. 56 (368): 509, 1961 (Kachroo and Schuster 1961). Bas.: Pycnolejeunea vittata
Steph. ex G.Hofm., Ann. Bryol. 8: 115, 1935 (Hofman 1935).
Incertae sedis
** Cheilolejeunea gottscheana C.J.Bastos, J. Bryol. 34 (4): 316, 2012 (Bastos 2012b).
Nom. nov. pro Strepsilejeunea lindenbergii Steph., Hedwigia 35 (3): 130, 1896
(Stephani 1896b).
* Cheilolejeunea heteroclada (Spruce) Schifn., Hepat. (Engl.-Prantl): 124, 1893
(Schifner 1893b). Bas.: Lejeunea heteroclada Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 256, 1884 (Spruce 1884). 246
** Cheilolejeunea minutilobula Amakawa, J. Jap. Bot. 35 (12): 365, 1960 (Amakawa
1960a).
** Cheilolejeunea norrisii (Grolle) M.A.M.Renner, Bryologist 115 (4): 550, 2012
(Renner 2012). Bas.: Lejeunea norrisii Grolle, Wiss. Z. Friedrich-Schiller-Univ.
Jena, Math.-Naturwiss. Reihe 31 (2): 215, 1982 (Grolle 1982).
** Cheilolejeunea oscilla M.A.M.Renner, Bryologist 115 (4): 551, 2012 (Renner 2012).
** Cheilolejeunea papulosa Schifn., Hepat. (Engl.-Prantl): 124, 1893 (Schifner
1893b). Nom. nov. pro Lejeunea papulosa Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 258, 1884 (Spruce 1884), nom. illeg.
*** Cheilolejeunea riparia (Steph.) M.E.Reiner, Nova Hedwigia 95 (3/4): 467, 2012
(Reiner-Drehwald and Grolle 2012). Bas.: Potamolejeunea riparia Steph., Sp.
Hepat. (Stephani) 5: 639, 1914 (Stephani 1914b).
*** Cheilolejeunea rotalis (Hook.f. et Taylor) M.Wigginton, J. Bryol. 34 (4): 270,
2012 (Wigginton 2012). Bas.: Jungermannia rotalis Hook.f. et Taylor, London J.
Bot. 4: 89, 1845 (Hooker and Taylor 1845).
** Cheilolejeunea tenerrima (Steph.) C.J.Bastos, J. Bryol. 34 (4): 317, 2012 (Bastos
2012b). Bas.: Strepsilejeunea tenerrima Steph., Sp. Hepat. (Stephani) 5: 286, 1913
(Stephani 1913a).
246 Cheilolejeunea heteroclada was considered conspeciic with Cheilolejeunea aneogyna by X.-L. He (in Gradstein and Costa 2003), but it was tentatively accepted by Reiner-Drehwald and Grolle (2012). Lejeunea
heteroclada var. subandina Spruce may be a diferent taxon (cf. Reiner-Drehwald & Grolle 2012).
316
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Cheilolejeunea urubuensis (Zartman et I.L.Ackerman) R.L.Zhu et Y.M.Wei, Phytotaxa 152 (1): 50, 2013 (Wei et al. 2013). Bas.: Vitalianthus urubuensis Zartman
et I.L.Ackerman, Bryologist 105 (2): 267, 2002 (Zartman and Ackerman 2002).
* Cyrtolejeunea A.Evans, Bull. Torrey Bot. Club 30 (10): 553, 1903 (Evans 1903c).
*** Cyrtolejeunea holostipa (Spruce) A.Evans, Bull. Torrey Bot. Club. 30 (10): 553,
1903 (Evans 1903c). Bas.: Lejeunea holostipa Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 171 (Spruce 1884).
** Cystolejeunea A.Evans, Bull. Torrey Bot. Club 33 (1): 16, 1906) (Evans 1906a)
*** Cystolejeunea lineata (Lehm. et Lindenb.) A.Evans, Bull. Torrey Bot. Club. 33
(1): 17, 1906 (Evans 1906a). Bas.: Jungermannia lineata Lehm. et Lindenb., Nov.
Stirp. Pug. 4: 53, 1832 (Lehmann 1832).
*
**
***
**
**
**
***
***
***
Omphalanthus Lindenb. et Nees, Syn. Hepat. 2: 303, 1845 (Gottsche et al.
1845a). 247
Omphalanthus baracoensis Mustelier, M.E.Reiner et Gradst., J. Bryol. 29 (2): 95,
2007 (Reiner-Drehwald et al. 2007).
Omphalanthus iliformis (Sw.) Nees, Syn. Hepat. 2: 304, 1845 (Gottsche et al. 1845a).
Bas.: Jungermannia iliformis Sw., Prodr. (Swartz): 144, 1788 (Swartz 1788).
Omphalanthus iliformis var. platycoleus (Herzog) Gradst., Phytotaxa 76 (3): 46,
2013 (Gradstein 2013a). Bas.: Omphalanthus platycoleus Herzog, Feddes Repert.
Spec. Nov. Regni Veg. 57 (1/2): 171, 1955 (Herzog 1955).
Omphalanthus iliformis var. wallisii (Prantl) Gradst., Phytotaxa 76 (3): 46, 2013 (Gradstein 2013a). Bas.: Lejeunea wallisii Prantl, Hedwigia 31: xvii, 1892 (Prantl 1892).
Omphalanthus huanucensis (Gottsche) Gradst., Beih. Nova Hedwigia 80: 109,
1985 (Gradstein and Buskes 1985). Bas.: Lejeunea huanucensis Gottsche, Syn. Hepat. 3: 335, 1845 (Gottsche et al. 1845b).
Omphalanthus jackii (Prantl) Gradst., Proc. Kon. Ned. Akad. Wetensch. C 80:
410, 1977 (Gradstein et al. 1977). Bas.: Lejeunea jackii Prantl, Hedwigia 31: xvii,
1892 (Prantl 1892).
Omphalanthus ovalis (Lindenb. et Gottsche) Gradst., Proc. Kon. Ned. Akad.
Wetensch. C 80: 411, 1977 (Gradstein et al. 1977). Bas.: Lejeunea ovalis Lindenb.
et Gottsche, Syn. Hepat. 5: 754, 1847 (Gottsche et al. 1847).
Omphalanthus roccatii (Gola) R.M.Schust., Beih. Nova Hedwigia 9: 96, 1963
(Schuster 1963a). Bas.: Acrolejeunea roccatii Gola, Ann. Bot. (Rome) 6 (2): 275,
1907 (Gola 1907).
247 Omphalanthus is nested in Cheilolejeunea (Wilson et al. 2007, Ye et al. 2011, Gradstein 2013a). A
formal transfer of the genus and its species to Cheilolejeunea is found in Ye et al. (2015).
World checklist of hornworts and liverworts
***
317
subtrib. Cololejeuneinae Gradst.
** Aphanotropis Herzog, Trans. Brit. Bryol. Soc. 2 (1): 63, 1952 (Herzog 1952b).
*** Aphanotropis saxicola Herzog, Trans. Brit. Bryol. Soc. 2 (1): 63, 1952 (Herzog 1952b).
** Calatholejeunea K.I.Goebel, Ann. Jard. Bot. Buitenzorg 39: 8, 1928 (Goebel 1928).
*** Calatholejeunea lamii Mizut., J. Hattori Bot. Lab. 56: 334, 1984 (Mizutani 1984a).
*** Calatholejeunea paradoxa (Schifn.) K.I.Goebel, Ann. Jard. Bot. Buitenzorg 39:
8, 1928 (Goebel 1928). Bas.: Lejeunea paradoxa Schifn., Nova Acta Acad. Caes.
Leop.-Carol. German. Nat. Cur. 60 (2): 243, 1893 (Schifner 1893a).
*** Cololejeunea (Spruce) Steph., Hedwigia 30 (5): 208, 1891 (Stephani 1891a).
Bas.: Lejeunea subg. Cololejeunea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:
291, 1884 (Spruce 1884). 248
** Cololejeunea micrandroecia (Spruce) M.Menzel, Willdenowia 14: 492, 1984
[1985] (Menzel 1984). Bas.: Lejeunea micrandroecia Spruce, Trans. & Proc. Bot.
Soc. Edinburgh 15: 298, 1884 (Spruce 1884).
*
***
***
***
***
***
subg. Aphanolejeunea (A.Evans) Pócs, Phytotaxa 202 (1): 64, 2015 (Pócs et al.
2015c). Bas.: Aphanolejeunea A.Evans, Bull. Torrey Bot. Club 38 (6): 272, 1911
(Evans 1911).
Cololejeunea berneckerae Pócs, Polish Bot. J. 54 (1): 4, 2009 (Pócs and Bernecker
2009). Nom. nov. pro Aphanolejeunea pocsii Bern.-Lück., Nova Hedwigia 66 (1/2):
168, 1998 (Bernecker-Lücking 1998).
Cololejeunea cingens (Herzog) Bernecker et Pócs, Polish Bot. J. 54 (1): 4, 2009
(Pócs and Bernecker 2009). Bas.: Aphanolejeunea cingens Herzog, Svensk Bot.
Tidskr. 46 (1): 104, 1952 (Herzog 1952e).
Cololejeunea cornutissima (R.M.Schust.) Stotler et Crand.-Stotl., Bryologist 80 (3):
411, 1977 (Stotler and Crandall-Stotler 1977). Bas.: Aphanolejeunea cornutissima
R.M.Schust., Bryologist 59 (3): 217, 1956 (Schuster 1956c).
Cololejeunea costaricensis (Bern.-Lück.) Bernecker et Pócs, Polish Bot. J. 54 (1): 5,
2009 (Pócs and Bernecker 2009). Bas.: Aphanolejeunea costaricensis Bern.-Lück.,
Nova Hedwigia 66 (1/2): 164, 1998 (Bernecker-Lücking 1998).
Cololejeunea cubensis Pócs, Polish Bot. J. 54 (1): 5, 2009 (Pócs and Bernecker
2009). Nom. nov. pro Aphanolejeunea evansii Herzog, Beih. Bot. Centralbl. 61B
(3): 583, 1942 (Herzog 1942d).
248 Cololejeunea is here organized into subgenera that are not supported by the molecular study of Yu et
al. (2013). hey are referred to following the traditional use (e.g. as in Pócs et al. 2014). Leptocolea
and Physocolea also belong here, but a few taxa have neither been transferred nor synonymized. hey
are listed in the “Names in genera not currently accepted” section below.
318
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*** Cololejeunea gracilis (Jovet-Ast) Pócs, Cryptog. Bryol. 29 (3): 233, 2008 (Dauphin
et al. 2008). Bas.: Aphanolejeunea gracilis Jovet-Ast, Rev. Bryol. Lichénol. 16 (1/2):
21, 1947 [1948] (Jovet-Ast 1947b).
** Cololejeunea gracilis var. linearifolia (R.M.Schust.) Pócs, Acta Bot. Hung. 56
(1/2): 189, 2014 (Pócs et al. 2014). Bas.: Aphanolejeunea gracilis var. linearifolia
R.M.Schust., Phytologia 45 (5): 434, 1980 (Schuster 1980b).
*** Cololejeunea grossepapillosa (Horik.) N.Kitag., Hikobia, Suppl. 1: 68, 1981 (Kitagawa 1981a). Bas.: Aphanolejeunea grossepapillosa Horik., J. Sci. Hiroshima Univ.,
Ser. B, Div. 2, Bot. 1: 92, 1932 (Horikawa 1932a).
** Cololejeunea iwatsukiana (Pócs) Pócs, Polish Bot. J. 54 (1): 6, 2009 (Pócs and
Bernecker 2009). Bas.: Aphanolejeunea iwatsukiana Pócs, Hikobia 11: 457, 1994
(Pócs 1994d).
*** Cololejeunea jovetastiana (Pócs) Pócs, Polish Bot. J. 54 (1): 6, 2009 (Pócs and
Bernecker 2009). Bas.: Aphanolejeunea jovetastiana Pócs, Cryptog. Bryol. Lichénol. 5 (3): 251, 1984 (Pócs 1984a).
** Cololejeunea lisowskii (Pócs) Pócs, Polish Bot. J. 54 (1): 6, 2009 (Pócs and Bernecker 2009). Bas.: Aphanolejeunea lisowskii Pócs, Cryptog. Bryol. Lichénol. 5 (3):
259, 1984 (Pócs 1984a).
*** Cololejeunea madeirensis Schifn., Hedwigia 41 (5): 279, 1902 (Schifner 1902).
*** Cololejeunea microscopica (Taylor) Schifn., Hepat. (Engl.-Prantl): 122, 1893
(Schifner 1893b). Bas.: Jungermannia microscopica Taylor, Mackay, Fl. Hibern.
2: 59, 1836 (Taylor 1836a).
*** Cololejeunea microscopica var. africana (Pócs) Pócs et Bernecker, Cryptog. Bryol.
29 (3): 234, 2008 (Dauphin et al. 2008). Bas.: Aphanolejeunea exigua var. africana
Pócs, Cryptog. Bryol. Lichénol. 5 (3): 247, 1984 (Pócs 1984a).
*** Cololejeunea microscopica var. exigua (A.Evans) Pócs, Mem. New York Bot. Gard.
76 (4): 73, 2009 (Gradstein and Ilkiu-Borges 2009). Bas.: Aphanolejeunea exigua
A.Evans, Bull. Torrey Bot. Club 38 (6): 273, 1911 (Evans 1911).
*** Cololejeunea minuscula Pócs, Polish Bot. J. 54 (1): 7, 2009 (Pócs and Bernecker
2009). Nom. nov. pro Aphanolejeunea minuta R.M.Schust., Hepat. Anthocerotae
N. Amer. 4: 1310, 1980 (Schuster 1980c).
** Cololejeunea moramangae Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 241, 1977
[1979] (Tixier 1977a).
*** Cololejeunea norrisii (Pócs) Pócs, Polish Bot. J. 54 (1): 7, 2009 (Pócs and Bernecker 2009). Bas.: Aphanolejeunea norrisii Pócs, Acta Bot. Fenn. 165: 95, 1999
(Pócs and Piippo 1999).
*** Cololejeunea papillosa (K.I.Goebel) Mizut., J. Hattori Bot. Lab. 29: 156, 1966
(Mizutani 1966). Bas.: Physocolea papillosa K.I.Goebel, Ann. Jard. Bot. Buitenzorg
39: 41, 1928 (Goebel 1928).
*** Cololejeunea sicifolia (Gottsche ex A.Evans) Pócs et Bernecker, Polish Bot. J. 54
(1): 8, 2009 (Pócs and Bernecker 2009). Bas.: Aphanolejeunea sicifolia Gottsche ex
A.Evans, Bull. Torrey Bot. Club 38 (6): 277, 1911 (Evans 1911).
World checklist of hornworts and liverworts
319
*** Cololejeunea sicifolia subsp. jamaicensis (R.M.Schust.) Bernecker et Pócs, Polish
Bot. J. 54 (1): 8, 2009 (Pócs and Bernecker 2009). Bas.: Aphanolejeunea jamaicensis R.M.Schust., Phytologia 45 (5): 434, 1980 (Schuster 1980b).
*** Cololejeunea sintenisii (Steph.) Pócs, Cryptog. Bryol. 29 (3): 235, 2008 (Dauphin
et al. 2008). Bas.: Aphanolejeunea sintenisii Steph., Sp. Hepat. (Stephani) 5: 861,
1916 (Stephani 1916b).
*** Cololejeunea subsphaeroidea (R.M.Schust.) Pócs, Cryptog. Bryol. 29 (3): 235, 2008
(Dauphin et al. 2008). Bas.: Aphanolejeunea subsphaeroidea R.M.Schust., Phytologia 39 (6): 431, 1978 (Schuster 1978b).
** Cololejeunea thiersiae (Pócs) Pócs, Polish Bot. J. 54 (1): 9, 2009 (Pócs and Bernecker 2009). Bas.: Aphanolejeunea thiersiae Pócs, Hikobia 11: 459, 1994 (Pócs
1994d).
** Cololejeunea veillonii Tixier, Nova Hedwigia 31: 757, 1979 (Tixier 1979b).
*** Cololejeunea winkleri (M.I.Morales et A.Lücking) Pócs, Mem. New York Bot.
Gard. 76 (4): 78, 2009 (Gradstein and Ilkiu-Borges 2009). Bas.: Aphanolejeunea
winkleri M.I.Morales et A.Lücking, Nova Hedwigia 60 (1/2): 120, 1995 (Morales
and Lücking 1995).
*
**
**
**
**
subg. Austrocololejeunea Tixier, Nova Hedwigia 31: 776, 1979 (Tixier 1979b).
Cololejeunea australis Tixier, Nova Hedwigia 31: 781, 1979 (Tixier 1979b).
Cololejeunea caledonica Gottsche, Hedwigia 34 (5): 246, 1895 (Stephani 1895b).
Cololejeunea sophiana Tixier, Bot. Not. 128: 428, 1975 [1976] (Tixier 1975a).
Cololejeunea virotana Tixier, Nova Hedwigia 31: 777, 1979 (Tixier 1979b).
*
subg. Chlorocolea R.M.Schust., Beih. Nova Hedwigia 9: 178, 1963 (Schuster
1963a).
Cololejeunea ceratilobula (P.C.Chen) R.M.Schust., Beih. Nova Hedwigia 9: 179,
1963 (Schuster 1963a). Bas.: Leptocolea ceratilobula P.C.Chen, Feddes Repert.
Spec. Nov. Regni Veg. 58: 49, 1955 (Chen 1955).
Cololejeunea desciscens Steph., Hedwigia 34 (5): 248, 1895 (Stephani 1895b).
Cololejeunea linopteroides H.Rob., Bryologist 67 (4): 457, 1964 (Robinson 1964).
Cololejeunea rotundilobula (P.C.Wu et P.J.Lin) Piippo, J. Hattori Bot. Lab. 68:
134, 1990 (Piippo 1990). Bas.: Pedinolejeunea rotundilobula P.C.Wu et P.J.Lin,
Acta Phytotax. Sin. 16 (2): 69, 1978 (Wu and Lin 1978). 249
Cololejeunea sigmoidea Jovet-Ast et Tixier, Rev. Bryol. Lichénol. 31 (1/2): 27,
1962 (Jovet-Ast and Tixier 1962).
Cololejeunea sigmoidea var. dubia Tixier, Bryophyt. Biblioth. 27: 114, 1985 (Tixier
1985a).
Cololejeunea standleyi Herzog, Rev. Bryol. Lichénol. 20 (1/2): 172, 1951 [1952]
(Herzog 1951a).
***
***
***
*
***
**
***
249 Cololejeunea rotundilobula is possibly conspeciic with Cololejeunea sigmoidea (Pócs et al. 2013).
320
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Cololejeunea stylilobula Tixier, Phytotaxa 202 (1): 65, 2015 (Pócs et al. 2015c).
Based on: Cololejeunea stylilobula Tixier, Bryophyt. Biblioth. 27: 119, 1985 (Tixier 1985a), nom. inval.
*** Cololejeunea zangii R.L.Zhu et M.L.So, Syst. Bot. 24 (4): 501, 1999 [2000] (Zhu
and So 1999b).
*** subg. Chlorolejeunea Benedix, Feddes Repert. Spec. Nov. Regni Veg. Beih. 134:
81, 1953 (Benedix 1953). 250
** Cololejeunea lacinulata Benedix, Feddes Repert. Spec. Nov. Regni Veg. Beih. 134:
82, 1953 (Benedix 1953).
*** Cololejeunea madothecoides (Steph.) Benedix, Feddes Repert. Spec. Nov. Regni
Veg. Beih. 134: 81, 1953 (Benedix 1953). Bas.: Physocolea madothecoides Steph.,
Sp. Hepat. (Stephani) 5: 898, 1916 (Stephani 1916b).
** Cololejeunea ombrophila Tixier, Nat. Hist. Bull. Siam Soc. 23 (4): 550, 1970 (Tixier 1970a).
*** Cololejeunea stotleriana Gradst., Ilk.-Borg. et Vanderp., Bryologist 114 (1): 13,
2011 (Gradstein et al. 2011).
*** subg. Chondriolejeunea Benedix, Feddes Repert. Spec. Nov. Regni Veg. Beih.
134: 75, 1953 (Benedix 1953). 251
*** Cololejeunea chinii Tixier, Nat. Hist. Bull. Siam Soc. 24 (3/4): 445, 1973 (Tixier
1973b).
*** Cololejeunea pseudostipulata Schifn. ex P.Syd., Just’s Bot. Jahresber. 19: 246, 1894
(Sydow 1894). Based on: Cololejeunea pseudostipulata Schifn., Leberm., Forschungsr.
Gazelle 4 (4): 33, 1890 (Schifner 1890), nom. inval.
*** Cololejeunea shimizui N.Kitag., Acta Phytotax. Geobot. 23 (5/6): 185, 1969 (Kitagawa 1969b).
*** Cololejeunea shimizui var. phangngana N.Kitag., Acta Phytotax. Geobot. 23 (5/6):
187, 1969 (Kitagawa 1969b).
*** subg. Cololejeunea, 252
** Cololejeunea albodentata P.C.Chen et P.C.Wu, Acta Phytotax. Sin. 9 (3): 252,
1964 (Chen and Wu 1964).
** Cololejeunea armata Tixier, Gard. Bull. Singapore 26 (1): 149, 1972 (Tixier
1972b).
*** Cololejeunea bhutanica Grolle et Mizut., J. Bryol. 15 (2): 281, 1989 (Grolle 1989e).
250 Cololejeunea subg. Chlorolejeunea forms a separate clade in Yu et al. (2013) with three sequenced species
(Cololejeunea ceratilobula, Cololejeunea madothecoides (the type) and Cololejeunea raduliloba).
251 Cololejeunea subg. Chondriolejeunea is supported by Yu et al. (2013) who included Cololejeunea chinii,
Cololejeunea pseudostipulata (the type, not sequenced) and Cololejeunea shimizui.
252 Subg. Cololejeunea includes subg. Aphanolejeunea in the phylogeny by Yu et al. (2013).
World checklist of hornworts and liverworts
321
** Cololejeunea biddlecomiae (Austin) A.Evans, Mem. Torrey Bot. Club 8 (2): 168,
1902 (Evans 1902a). Bas.: Lejeunea biddlecomiae Austin, List. Canad. Hepat.: 5,
1890 (Pearson 1890).
* Cololejeunea caihuaella But et P.C.Wu, Hepat. Fl. Hong Kong: 132, 2009 (Wu
and But 2009).
*** Cololejeunea calcarea (Lib.) Steph., Bot. Gaz. 17 (6): 171, 1892 (Stephani 1892f).
Bas.: Lejeunea calcarea Lib., Ann. Gen. Sci. Phys. 6: 373, 1820 (Libert 1820).
** Cololejeunea capuronii Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 241, 1977
[1979] (Tixier 1977a).
** Cololejeunea dinghuiana R.L.Zhu et Y.F.Wang, J. E. China Norm. Univ., Nat.
Sci. Ed. 2: 91, 1992 (Zhu and Wang 1992).
** Cololejeunea dolichodonta Tixier, Bryophyt. Biblioth. 27: 215, 1985 (Tixier 1985a).
*** Cololejeunea dozyana (Sande Lac.) Schifn., Hedwigia 39 (4): 199, 1900 (Schifner
1900b). Bas.: Lejeunea dozyana Sande Lac., Ned. Kruidk. Arch. 3: 522, 1855
(Sande Lacoste 1855).
*** Cololejeunea elegans Steph., Hedwigia 30 (5): 208, 1891 (Stephani 1891a).
** Cololejeunea falcidentata R.M.Schust., Nova Hedwigia 15: 507, 1968 (Schuster 1968b).
** Cololejeunea ilidens Benedix, Feddes Repert. Spec. Nov. Regni Veg. Beih. 134: 49,
1953 (Benedix 1953).
* Cololejeunea frahmii Tixier, Trop. Bryol. 11: 63, 1995 (Tixier 1995b). 253
*** Cololejeunea grolleana Pócs, J. Hattori Bot. Lab. 48: 312, 1980 (Pócs 1980b).
*** Cololejeunea haskarliana (Lehm.) Schifn., Consp. Hepat. Arch. Ind.: 244, 1898
(Schifner 1898b). Bas.: Lejeunea haskarliana Lehm., Nov. Stirp. Pug. 8: 26, 1844
(Lehmann 1844).
*** Cololejeunea hyalina G.Asthana et S.C.Srivast., Bryophyt. Biblioth. 60: 25, 2003
(Asthana and Srivastava 2003).
* Cololejeunea kahuziensis Tixier, Trop. Bryol. 11: 63, 1995 (Tixier 1995b). 254
*** Cololejeunea karnatakensis G.Asthana et S.C.Srivast., Bryophyt. Biblioth. 60: 26,
2003 (Asthana and Srivastava 2003).
** Cololejeunea kodamae Kamim., Feddes Repert. Spec. Nov. Regni Veg. 58: 55,
1955 (Kamimura 1955).
*** Cololejeunea kolombangarae Pócs, Acta Bryolichenol. Asiat. 4: 67, 2011 (Pócs and
Piippo 2011).
** Cololejeunea kolombangarae subsp. sepikensis Pócs, Acta Bryolichenol. Asiat. 4: 68,
2011 (Pócs and Piippo 2011).
*** Cololejeunea konratii Pócs, Acta Bot. Hung. 54 (1/2): 156, 2012 (Pócs 2012b).
*** Cololejeunea kuciana Pócs et Schäf.-Verw., Polish Bot. J. 57 (1): 51, 2012 (Pócs
and Schäfer-Verwimp 2012).
*** Cololejeunea longiana Grolle et Mizut., J. Bryol. 15 (2): 284, 1989 (Grolle 1989e).
253 Cololejeunea frahmii is probably only a depauperate form of Cololejeunea elegans or Cololejeunea zenkeri.
254 Cololejeunea kahuziensis is probably only a depauperate form of Cololejeunea elegans or Cololejeunea
zenkeri.
322
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Cololejeunea macounii (Spruce) A.Evans, Mem. Torrey Bot. Club 8 (2): 171, 1902
(Evans 1902a). Bas.: Lejeunea macounii Spruce, Bull. Torrey Bot. Club 17 (10):
259, 1890 (Underwood 1890).
** Cololejeunea magillii Pócs, J. Hattori Bot. Lab. 74: 49, 1993 (Pócs 1993).
*** Cololejeunea malanjae Steph., Bot. Jahrb. Syst. 30 (2): 261, 1901 (Stephani 1901g).
** Cololejeunea mamillata (Ångstr.) E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot.
3 (11): 184, 1967 (Hodgson 1967). Bas.: Lejeunea mamillata Ångstr., Öfvers.
Kongl. Vetensk.-Akad. Förh. 29 (4): 14, 1872 (Ångström 1872).
** Cololejeunea mizutaniana Udar et G.Srivast., Misc. Bryol. Lichenol. 9 (7): 138,
1983 (Udar and Srivastava 1983).
*** Cololejeunea mocambiquensis S.W.Arnell, Mitt. hüring. Bot. Ges. 1 (1): 7, 1955
[1956] (Arnell 1955a).
** Cololejeunea nanhutashanensis J.D.Yang et S.H.Lin, Phytotaxa 177 (1): 56, 2014
(Yang and Lin 2014).
*** Cololejeunea nilgiriensis G.Asthana et S.C.Srivast., Bryophyt. Biblioth. 60: 27,
2003 (Asthana and Srivastava 2003).
*** Cololejeunea ornata A.Evans, Bryologist 41 (4): 73, 1938 (Evans 1938c).
** Cololejeunea parva Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 47 (1/2): 239,
1977 (Vanden Berghen 1977).
*** Cololejeunea planilora Benedix, Feddes Repert. Spec. Nov. Regni Veg. Beih. 134:
60, 1953 (Benedix 1953).
*** Cololejeunea platyneura (Spruce) A.Evans, Mem. Torrey Bot. Club 8 (2): 172,
1902 (Evans 1902a). Bas.: Lejeunea platyneura Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 299, 1884 (Spruce 1884).
*** Cololejeunea pluridentata P.C.Wu et J.S.Lou, Acta Phytotax. Sin. 16 (4): 105,
1978 (Wu and Lou 1978).
** Cololejeunea pretiosa Benedix, Feddes Repert. Spec. Nov. Regni Veg. Beih. 134:
62, 1953 (Benedix 1953).
*** Cololejeunea pseudocristallina P.C.Chen et P.C.Wu, Acta Phytotax. Sin. 9 (3): 257,
1964 (Chen and Wu 1964).
** Cololejeunea pseudoplagiophylla P.C.Wu et J.S.Lou, Acta Phytotax. Sin. 16 (4):
106, 1978 (Wu and Lou 1978).
** Cololejeunea pseudoschmidtii Tixier, Gard. Bull. Singapore 26 (1): 145, 1972 (Tixier
1972b).
** Cololejeunea ramromensis Pócs, Biodiv., biogeogr., nat. conserv. Wallacea N.
Guinea II: 113, 2014 (Chantanaorrapint and Pócs 2014).
*** Cololejeunea rosellata Mizut., J. Hattori Bot. Lab. 29: 159, 1966 (Mizutani 1966).
*** Cololejeunea rossettiana (C.Massal.) Schifn., Hepat. (Engl.-Prantl): 122, 1893
(Schifner 1893b). Bas.: Lejeunea rossettiana C.Massal., Nuovo Giorn. Bot. Ital.
21 (3): 487, 1889 (Massalongo 1889).
*** Cololejeunea runssorensis (Steph.) Pócs, Acta Bot. Acad. Sci. Hung. 21 (3/4): 371,
1975 (Pócs 1975). Bas.: Aphanolejeunea runssorensis Steph., Sp. Hepat. (Stephani)
5: 858, 1916 (Stephani 1916b).
World checklist of hornworts and liverworts
323
** Cololejeunea schaeferi Grolle, J. Bryol. 13 (4): 488, 1986 (Grolle 1986b).
*** Cololejeunea schmidtii Steph., Bot. Tidsskr. 24 (3): 278, 1902 (Stephani 1902b).
** Cololejeunea schmidtii var. acutepapillosa Pócs, Acta Bot. Hung. 54 (1/2): 165,
2012 (Pócs 2012b).
** Cololejeunea selaginellicola Tixier, Gard. Bull. Singapore 26 (1): 149, 1972 (Tixier
1972b).
*** Cololejeunea serrata (Steph.) Benedix, Feddes Repert. Spec. Nov. Regni Veg. Beih.
134: 53, 1953 (Benedix 1953). Bas.: Physocolea serrata Steph., Sp. Hepat. (Stephani)
5: 905, 1916 (Stephani 1916b).
** Cololejeunea shikokiana (Horik.) S.Hatt., Bull. Tokyo Sci. Mus. 11: 101, 1944
(Hattori 1944d). Bas.: Physocolea shikokiana Horik., Bot. Mag. (Tokyo) 46 (544):
182, 1932 (Horikawa 1932b).
** Cololejeunea spinosa (Horik.) Pandé et R.N.Misra, J. Indian Bot. Soc. 22 (2/4):
166, 1943 (Pandé and Misra 1943). Bas.: Physocolea spinosa Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 1: 70, 1931 (Horikawa 1931a).
*** Cololejeunea stellaris Pócs, Acta Bryolichenol. Asiat. 4: 70, 2011 (Pócs and Piippo
2011).
** Cololejeunea subkodamae Mizut., J. Hattori Bot. Lab. 60: 448, 1986 (Mizutani 1986a).
** Cololejeunea tamasii Schäf.-Verw., Phytotaxa 60: 9, 2012 (Schäfer-Verwimp
2012).
** Cololejeunea tanneri Pócs, Acta Bot. Hung. 31 (1/4): 126, 1985 (Pócs 1985).
*** Cololejeunea tenella Benedix, Feddes Repert. Spec. Nov. Regni Veg. Beih. 134: 55,
1953 (Benedix 1953).
** Cololejeunea tenella var. dentiloba Onr., Bull. Jard. Bot. Natl. Belg. 59 (3/4): 436,
1989 (Onraedt 1989).
** Cololejeunea thiersiana Tixier, Cryptog. Bryol. Lichénol. 16 (3): 229, 1995 (Tixier
1995a). Based on: Cololejeunea thiersiana Tixier, Candollea 46 (2): 291, 1991
(Tixier 1991), nom. inval.
** Cololejeunea tuiwawana Pócs, Acta Bot. Hung. 54 (1/2): 165, 2012 (Pócs 2012b).
*** Cololejeunea verrucosa Steph., Hedwigia 34 (5): 253, 1895 (Stephani 1895b).
** Cololejeunea verrucosa var. rectispina (Herzog) Benedix, Feddes Repert. Spec.
Nov. Regni Veg. Beih. 134: 55, 1953 (Benedix 1953). Bas.: Physocolea verrucosa
var. rectispina Herzog, Mitt. Inst. Allg. Bot. Hamburg 7 (3): 214, 1931 (Herzog
1931a).
*** Cololejeunea zenkeri (Steph.) E.W.Jones, Trans. Brit. Bryol. Soc. 2 (3): 420, 1954
(Jones 1954c). Bas.: Aphanolejeunea zenkeri Steph., Sp. Hepat. (Stephani) 5: 858,
1916 (Stephani 1916b).
subg. Cryptolejeunea Benedix, Feddes Repert. Spec. Nov. Regni Veg. Beih. 134:
77, 1953 (Benedix 1953).
*** Cololejeunea angustilora (Steph.) Mizut., J. Hattori Bot. Lab. 28: 113, 1965 (Mizutani 1965). Bas.: Leptocolea angustilora Steph., Sp. Hepat. (Stephani) 5: 848,
1916 (Stephani 1916b).
*
324
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Cololejeunea drepanolejeuneoides (Horik.) R.M.Schust., Beih. Nova Hedwigia 9:
174, 1963 (Schuster 1963a). Bas.: Boninoleptocolea drepanolejeuneoides Horik.,
Bot. Mag. (Tokyo) 50 (598): 558, 1936 (Horikawa 1936). 255
** Cololejeunea hattoriana Mizut. et Pócs, Ann. Bot. Fenn. 31 (3): 188, 1994 (Pócs et
al. 1994). Nom. nov. pro Campylolejeunea pusilla Mizut., J. Hattori Bot. Lab. 29:
154, 1966 (Mizutani 1966).
*** Cololejeunea inlectens (Mitt.) Benedix, Feddes Repert. Spec. Nov. Regni Veg.
Beih. 134: 79, 1953 (Benedix 1953). Bas.: Lejeunea inlectens Mitt., J. Proc. Linn.
Soc., Bot. 5 (18): 117, 1860 [1861] (Mitten 1860c).
** Cololejeunea mouensis (Tixier) H.A.Mill., Phytologia 47 (4): 322, 1981 (Miller 1981).
Bas.: Campylolejeunea mouensis Tixier, Nova Hedwigia 31: 727, 1979 (Tixier 1979b).
*** Cololejeunea vesicaria (Sande Lac.) Schifn., Consp. Hepat. Arch. Ind.: 247, 1898
(Schifner 1898b). Bas.: Lejeunea vesicaria Sande Lac., Syn. hepat. jav.: 74, 1856
[1857] (Sande Lacoste 1856b).
*
***
**
*
***
***
***
***
**
***
***
subg. Diaphanae R.M.Schust., J. Elisha Mitchell Sci. Soc. 72 (1): 103, 1956
(Schuster 1956b).
Cololejeunea amphibola B.M.hiers, Beih. Nova Hedwigia 90: 130, 1988 (hiers
1988).
Cololejeunea antillana Pócs, Polish Bot. J. 54 (1): 3, 2009 (Pócs and Bernecker
2009). Nom. nov. pro Aphanolejeunea longifolia Jovet-Ast, Rev. Bryol. Lichénol. 16
(1/2): 23, 1947 [1948] (Jovet-Ast 1947b).
Cololejeunea augieri Tixier, Trop. Bryol. 11: 48, 1995 (Tixier 1995b).
Cololejeunea azorica V.Allorge et Jovet-Ast, Mitt. hüring. Bot. Ges. 1 (2/3): 17,
1955 (Allorge and Jovet-Ast 1955).
Cololejeunea camillii (Lehm.) A.Evans, Bryologist 15 (4): 59, 1912 (Evans 1912a).
Bas.: Lejeunea camillii Lehm., Nov. Stirp. Pug. 10: 15, 1857 (Lehmann 1857).
Cololejeunea contractiloba A.Evans, Amer. J. Bot. 5 (3): 131, 1918 (Evans 1918).
Cololejeunea cornuta E.W.Jones, Trans. Brit. Bryol. Soc. 2 (3): 436, 1954 (Jones
1954c).
Cololejeunea crenata (A.Evans) Pócs, Polish Bot. J. 54 (1): 5, 2009 (Pócs and Bernecker 2009). Bas.: Aphanolejeunea crenata A.Evans, Bull. Torrey Bot. Club 38
(6): 276, 1911 (Evans 1911).
Cololejeunea diaphana A.Evans, Bull. Torrey Bot. Club 32 (4): 184, 1905 (Evans
1905b).
Cololejeunea erostrata (Herzog) Bernecker et Pócs, Polish Bot. J. 54 (1): 5, 2009
(Pócs and Bernecker 2009). Bas.: Physocolea erostrata Herzog, Beih. Bot. Centralbl. 61B (3): 582, 1942 (Herzog 1942d).
255 Cololejeunea drepanolejeuneoides may based on its description be close to Cololejeunea angustilora and
especially to Cololejeunea inlectens, forming a natural group.
World checklist of hornworts and liverworts
325
** Cololejeunea gradsteinii M.J.Lai et R.L.Zhu, Ann. Bot. Fenn. 45 (5): 334, 2008
(Lai et al. 2008). Nom. nov. pro Cololejeunea pusilla Tixier, Nat. Hist. Bull. Siam
Soc. 23 (4): 552, 1970 (Tixier 1970a), nom. illeg.
** Cololejeunea koponenii (Pócs) Pócs, Polish Bot. J. 54 (1): 6, 2009 (Pócs and Bernecker 2009). Bas.: Aphanolejeunea koponenii Pócs, Acta Bot. Fenn. 165: 91, 1999
(Pócs and Piippo 1999).
*** Cololejeunea lanceolata E.W.Jones, Trans. Brit. Bryol. Soc. 2 (3): 428, 1954 (Jones
1954c).
*** Cololejeunea moralesiae (Bern.-Lück.) Bernecker et Pócs, Polish Bot. J. 54 (1): 7,
2009 (Pócs and Bernecker 2009). Bas.: Aphanolejeunea moralesiae Bern.-Lück.,
Nova Hedwigia 66 (1/2): 166, 1998 (Bernecker-Lücking 1998).
*** Cololejeunea morobensis (Pócs) Pócs, Polish Bot. J. 54 (1): 7, 2009 (Pócs and Bernecker 2009). Bas.: Aphanolejeunea morobensis Pócs, Ann. Bot. Fenn. 31 (3): 180,
1994 (Pócs et al. 1994).
*** Cololejeunea obtusifolia (E.W.Jones) Tixier, Trop. Bryol. 11: 42, 1995 (Tixier
1995b). Bas.: Cololejeunea pusilla var. obtusifolia E.W.Jones, Trans. Brit. Bryol.
Soc. 2 (3): 427, 1954 (Jones 1954c).
* Cololejeunea obtusifolia var. madecassa (Tixier) Pócs, Phytotaxa 202 (1): 64, 2015
(Pócs et al. 2015c). Bas.: Cololejeunea androphylla var. madecassa Tixier, Bull.
Acad. Malgache (n.ser) 55 (1/2): 216, 1977 [1979] (Tixier 1977a).
*** Cololejeunea papilliloba (Steph.) Steph., Hedwigia 34 (5): 250, 1895 (Stephani 1895b).
Bas.: Lejeunea papilliloba Steph., Hedwigia 29 (2): 73, 1890 (Stephani 1890b).
** Cololejeunea papulosa R.M.Schust., Phytologia 45 (5): 433, 1980 (Schuster 1980b).
*** Cololejeunea paucifolia (Spruce) Bernecker et Pócs, Polish Bot. J. 54 (1): 8, 2009
(Pócs and Bernecker 2009). Bas.: Lejeunea paucifolia Spruce, Bull. Soc. Bot. France
(Congr. Bot.) 36: cxciv, 1889 [1890] (Spruce 1889).
*** Cololejeunea peponiformis Mizut., J. Hattori Bot. Lab. 33: 258, 1970 (Mizutani 1970).
** Cololejeunea pseudocuspidata Tixier, Bauhinia 8 (4): 228, 1987 (Hürlimann 1987).
*** Cololejeunea pterocolea Herzog, Svensk Bot. Tidskr. 46 (1): 103, 1952 (Herzog 1952e).
*** Cololejeunea pusilla Steph., Hedwigia 34 (5): 251, 1895 (Stephani 1895b).
** Cololejeunea spathulata Jovet-Ast, Rev. Bryol. Lichénol. 29 (1/2): 36, 1960 (Jovet-Ast 1960).
** Cololejeunea taurifolia Inoue et H.A.Mill., Bull. Natl. Sci. Mus. Tokyo (n.ser.) 8
(2): 152, 1965 (Inoue and Miller 1965).
*** Cololejeunea wightii Steph., Hedwigia 34 (5): 253, 1895 (Stephani 1895b).
*** Cololejeunea yelitzae Pócs et Bernecker, Acta Bot. Hung. 55 (3/4): 386, 2013 (Pócs
and Bernecker 2013).
subg. Leptocolea (Spruce) Schifn., Hepat. (Engl.-Prantl): 122, 1893 (Schifner
1893b). Bas.: Lejeunea sect. Leptocolea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 294, 1884 (Spruce 1884).
** Cololejeunea acuminata Mizut., J. Hattori Bot. Lab. 33: 261, 1970 (Mizutani 1970).
*
326
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Cololejeunea aequabilis (Sande Lac.) Schifn., Consp. Hepat. Arch. Ind.: 242, 1898
(Schifner 1898b). Bas.: Lejeunea aequabilis Sande Lac., Ann. Mus. Bot. LugdunoBatavi 1: 310, 1864 (Sande Lacoste 1864).
** Cololejeunea altimontana Pócs, Acta Bryolichenol. Asiat. 4: 78, 2011 (Pócs and
Piippo 2011).
*** Cololejeunea amaniensis Pócs, Acta Bot. Hung. 31 (1/4): 120, 1985 (Pócs 1985).
*** Cololejeunea angustibracteata Schifn. ex P.Syd., Just’s Bot. Jahresber. 19: 246,
1894 (Sydow 1894). Based on: Cololejeunea angustibracteata Schifn., Leberm.,
Forschungsr. Gazelle 4 (4): 34, 1890 (Schifner 1890), nom. inval.
*** Cololejeunea apiculata (E.W.Jones) R.M.Schust., Beih. Nova Hedwigia 9: 174,
1963 (Schuster 1963a). Bas.: Leptocolea apiculata E.W.Jones, Trans. Brit. Bryol.
Soc. 2 (3): 415, 1954 (Jones 1954c).
* Cololejeunea arrectifolia (Mitt.) Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 309, 1896
(Stephani 1896a). Bas.: Lejeunea arrectifolia Mitt., Fl. vit.: 415, 1871 [1873] (Mitten
1871).
** Cololejeunea aurantia (Tixier) houvenot, Cryptog. Bryol. 32 (4): 290, 2011 (houvenot et al. 2011). Bas.: Jovetastella aurantia Tixier, Cryptog. Bryol. Lichénol. 3 (1):
29, 1982 (Tixier 1982).
** Cololejeunea bandamiae Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 220, 1977
[1979] (Tixier 1977a).
** Cololejeunea bebourensis Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 229, 1977
[1979] (Tixier 1977a).
** Cololejeunea bergmansiana Tixier, Bull. Jard. Bot. Natl. Belg. 59 (3/4): 440, 1989
(Tixier 1989).
** Cololejeunea bidentula (Steph.) E.W.Jones, Trans. Brit. Bryol. Soc. 2 (3): 423,
1954 (Jones 1954c). Bas.: Physocolea bidentula Steph., Sp. Hepat. (Stephani) 5:
868, 1916 (Stephani 1916b).
** Cololejeunea bifalcata Pócs, Acta Bot. Hung. 54 (1/2): 146, 2012 (Pócs 2012b).
*** Cololejeunea blepharophylla Pócs, Acta Bot. Hung. 54 (1/2): 149, 2012 (Pócs 2012b).
** Cololejeunea borhidiana Pócs, J. Hattori Bot. Lab. 48: 305, 1980 (Pócs 1980b).
** Cololejeunea bosseriana Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 235, 1977
[1979] (Tixier 1977a).
** Cololejeunea calcarata E.W.Jones, Bull. Brit. Mus. (Nat. Hist.), Bot. 11 (3): 235,
1983 (Jones and Harrington 1983).
* Cololejeunea camusii Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 227, 1977
[1979] (Tixier 1977a).
*** Cololejeunea ceatocarpa (Ångstr.) Steph., Bull. Herb. Boissier 5 (10): 842, 1897
(Stephani 1897c). Bas.: Lejeunea ceatocarpa Ångstr., Öfvers. Kongl. Vetensk.-Akad.
Förh. 29 (4): 27, 1872 (Ångström 1872).
*** Cololejeunea ceylanica Onr., Acta Bot. Acad. Sci. Hung. 25 (1/2): 107, 1979 (Onraedt
1979).
World checklist of hornworts and liverworts
*
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***
**
**
**
**
*
**
**
*
**
**
**
**
*
***
327
Cololejeunea chamlongiana Tixier, Ann. Fac. Sci. Univ. Phnom Penh 3: 181, 1970
(Tixier 1970b). 256
Cololejeunea chenii Tixier, Bryophyt. Biblioth. 27: 219, 1985 (Tixier 1985a).
Nom. nov. pro Cololejeunea plagiophylla var. grossipapillosa P.C.Chen et P.C.Wu,
Acta Phytotax. Sin. 9 (3): 254, 1964 (Chen and Wu 1964).
Cololejeunea ciliata Pócs, Ann. Bot. Fenn. 31 (3): 182, 1994 (Pócs et al. 1994).
Cololejeunea comptonii (Pearson) H.A.Mill., Phytologia 47 (4): 321, 1981 (Miller
1981). Bas.: Leptocolea comptonii Pearson, J. Linn. Soc., Bot. 46 (305): 40, 1922
(Pearson 1922b).
Cololejeunea cookei A.Evans, Trans. Connecticut Acad. Arts 10 (8): 447, 1900
(Evans 1900a).
Cololejeunea cordilora Steph., Hedwigia 34 (5): 246, 1895 (Stephani 1895b).
Cololejeunea crateris Pócs, Acta Bot. Hung. 54 (1/2): 153, 2012 (Pócs 2012b).
Cololejeunea crenulata (Pearson) H.A.Mill., Phytologia 47 (4): 321, 1981 (Miller
1981). Bas.: Leptocolea crenulata Pearson, J. Linn. Soc., Bot. 46 (305): 41, 1922
(Pearson 1922b). 257
Cololejeunea cucullifolia (Herzog) E.A.Hodgs., Rec. Domin. Mus. 4 (11): 127,
1962 (Hodgson 1962a). Bas.: Physocolea cucullifolia Herzog, Trans. & Proc. Roy.
Soc. New Zealand 68 (1): 45, 1938 (Herzog 1938c).
Cololejeunea cuneifolia Steph., Hedwigia 31 (4): 166, 1892 (Stephani 1892g).
Cololejeunea dadeuniana Tixier, Ann. Fac. Sci. Univ. Phnom Penh 3: 186, 1970
(Tixier 1970b).
Cololejeunea decemplicata (Steph.) Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2):
213, 1977 [1979] (Tixier 1977a). Bas.: Physocolea decemplicata Steph., Sp. Hepat.
(Stephani) 5: 869, 1916 (Stephani 1916b).
Cololejeunea decliviloba Steph., Hedwigia 34 (5): 247, 1895 (Stephani 1895b).
Cololejeunea dentata (E.W.Jones) R.M.Schust., Beih. Nova Hedwigia 9: 175,
1963 (Schuster 1963a). Bas.: Leptocolea dentata E.W.Jones, Trans. Brit. Bryol.
Soc. 2 (2): 161, 1953 (Jones 1953a).
Cololejeunea denticulata (Horik.) S.Hatt., Bull. Tokyo Sci. Mus. 11: 99, 1944
(Hattori 1944d). Bas.: Physocolea denticulata Horik., J. Sci. Hiroshima Univ., Ser.
B, Div. 2, Bot. 2: 287, 1934 (Horikawa 1934).
Cololejeunea dentilobula (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 241, 1963
(Schuster 1963b). Bas.: Physocolea dentilobula Steph., Sp. Hepat. (Stephani) 5:
892, 1916 (Stephani 1916b). 258
Cololejeunea dianae M.Wigginton, J. Bryol. 28 (4): 364, 2006 (Wigginton 2006).
256 Cololejeunea chamlongiana is possibly conspeciic with Cololejeunea papulilora.
257 Cololejeunea crenulata is possibly conspeciic with Cololejeunea angustilora judging from the protologue.
258 Cololejeunea dentilobula is possibly conspeciic with Cololejeunea decliviloba judging from Stephani’s
Icones (unpublished).
328
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Cololejeunea dilatata (Steph.) Mizut., J. Hattori Bot. Lab. 28: 113, 1965 (Mizutani 1965). Bas.: Leptocolea dilatata Steph., Sp. Hepat. (Stephani) 5: 850, 1916
(Stephani 1916b).
*** Cololejeunea diplasiolejeunoides Tixier, Bryologist 82 (4): 602, 1979 (Tixier 1979d).
*** Cololejeunea distalopapillata (E.W.Jones) R.M.Schust., Beih. Nova Hedwigia 9:
173, 1963 (Schuster 1963a). Bas.: Leptocolea distalopapillata E.W.Jones, Trans.
Brit. Bryol. Soc. 3 (2): 202, 1957 (Jones 1957b).
* Cololejeunea efusa (Mitt.) Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 309, 1896 (Stephani
1896a). Bas.: Lejeunea efusa Mitt., Fl. vit.: 415, 1871 [1873] (Mitten 1871). 259
*** Cololejeunea elephantorum Tixier, Ann. Fac. Sci. Univ. Phnom Penh 3: 185, 1970
(Tixier 1970b).
** Cololejeunea ellipsoidea R.M.Schust., Nova Hedwigia 15: 508, 1968 (Schuster 1968b).
*** Cololejeunea equialbi Tixier, Ann. Fac. Sci. Univ. Phnom Penh 3: 178, 1970 (Tixier
1970b).
*** Cololejeunea ilicis (Herzog) Piippo, J. Hattori Bot. Lab. 68: 133, 1990 (Piippo
1990). Bas.: Leptocolea ilicis Herzog, Symb. Sin. 5: 52, 1930 (Nicholson et al. 1930).
* Cololejeunea ischeri Tixier, Trop. Bryol. 11: 59, 1995 (Tixier 1995b). 260
** Cololejeunea lavicans (Steph.) Mizut., J. Hattori Bot. Lab. 28: 115, 1965 (Mizutani 1965). Bas.: Physocolea lavicans Steph., Sp. Hepat. (Stephani) 5: 893, 1916
(Stephani 1916b).
* Cololejeunea fredericii Onr., Rev. Bryol. Lichénol. 44 (1): 80, 1978 (Onraedt 1978). 261
** Cololejeunea fusca (Steph.) Mizut., J. Hattori Bot. Lab. 28: 120, 1965 (Mizutani 1965).
Bas.: Physocolea fusca Steph., Sp. Hepat. (Stephani) 5: 893, 1916 (Stephani 1916b).
** Cololejeunea gottschei (Steph.) Pandé, K.P.Srivast. et Ahmad, J. Indian Bot. Soc.
36 (3): 345, 1957 (Pandé et al. 1957). Bas.: Physocolea gottschei Steph., Sp. Hepat.
(Stephani) 5: 894, 1916 (Stephani 1916b).
*** Cololejeunea grossestyla M.Wigginton, J. Bryol. 28 (4): 369, 2006 (Wigginton 2006).
*** Cololejeunea grushvitzkiana Pócs, Bot. Zhurn. (Moscow & Leningrad) 56 (5): 674,
1971 (Pócs 1971).
*** Cololejeunea hainanensis R.L.Zhu, J. Hattori Bot. Lab. 78: 87, 1995 (Zhu 1995).
*** Cololejeunea harrisii Pócs, Acta Bot. Acad. Sci. Hung. 21 (3/4): 357, 1975 (Pócs 1975).
*** Cololejeunea hildebrandii (Austin) Steph., Bull. Herb. Boissier 5 (10): 842, 1897
(Stephani 1897c). Bas.: Lejeunea hildebrandii Austin, Bot. Bull. (Hanover) 1 (8):
35, 1876 (Austin 1876a).
*** Cololejeunea hirta Steph., Bull. Misc. Inform. Kew 1899 (151/152): 125, 1899
(MacGregor 1899).
** Cololejeunea hodgsoniae (Herzog) E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot.
3 (11): 184, 1967 (Hodgson 1967). Bas.: Physocolea hodgsoniae Herzog, Trans. &
Proc. Roy. Soc. New Zealand 68 (1): 46, 1938 (Herzog 1938c).
259 Cololejeunea efusa may be conspeciic with Cololejeunea trichomanis.
260 Cololejeunea leischeri is near to or conspeciic with Cololejeunea bidentula.
261 Cololejeunea fredericii is possibly conspeciic with Cololejeunea polyantha.
World checklist of hornworts and liverworts
329
*** Cololejeunea horikawana (S.Hatt.) Mizut., J. Hattori Bot. Lab. 24: 254, 1961 (Mizutani 1961). Bas.: Leptocolea horikawana S.Hatt., J. Jap. Bot. 18 (11): 653, 1942
(Hattori 1942).
*** Cololejeunea huerlimannii Tixier, Nova Hedwigia 31: 773, 1979 (Tixier 1979b).
** Cololejeunea inlexifolia R.M.Schust., Phytologia 56 (7): 458, 1985 (Schuster 1985c).
*** Cololejeunea iradieri M.Infante et Heras, Trop. Bryol. 17: 14, 1999 (Infante and
Heras 1999).
** Cololejeunea irianensis Tixier, Bull. Jard. Bot. Natl. Belg. 59 (3/4): 440, 1989 (Tixier
1989).
*** Cololejeunea johannis-winkleri (Herzog) R.L.Zhu, Nova Hedwigia 79 (3/4): 528,
2004 (Zhu et al. 2004). Bas.: Leptocolea johannis-winkleri Herzog, Mitt. Inst. Allg.
Bot. Hamburg 7 (3): 214, 1931 (Herzog 1931a).
** Cololejeunea kegelii Steph., Hedwigia 34 (5): 249, 1895 (Stephani 1895b).
** Cololejeunea khanii Tixier, Dacca Univ. Stud., B 15: 9, 1967 (Tixier 1967).
* Cololejeunea kohkongensis Tixier, Bryophyt. Biblioth. 27: 302, 1985 (Tixier 1985a). 262
*** Cololejeunea lichenyae R.D.Porley, N.G.Hodgetts et M.Wigginton, J. Bryol. 29
(1): 7, 2007 (Wigginton et al. 2007).
* Cololejeunea lobulilineata Tixier, Trop. Bryol. 11: 42, 1995 (Tixier 1995b). 263
*** Cololejeunea longifolia (Mitt.) Benedix ex Mizut., J. Hattori Bot. Lab. 26: 184,
1963 (Mizutani 1963). Bas.: Lejeunea longifolia Mitt., J. Proc. Linn. Soc., Bot. 5
(18): 117, 1860 [1861] (Mitten 1860c).
*** Cololejeunea magna (Tixier) M.Infante et Heras, Trop. Bryol. 17: 17, 1999 (Infante and Heras 1999). Bas.: Cololejeunea harrisii var. magna Tixier, Trop. Bryol. 11:
56, 1995 (Tixier 1995b).
** Cololejeunea magniica Pócs, Acta Bryolichenol. Asiat. 4: 88, 2011 (Pócs and Piippo
2011).
** Cololejeunea magnilobula (Horik.) S.Hatt., Bull. Tokyo Sci. Mus. 11: 99, 1944
(Hattori 1944d). Bas.: Physocolea magnilobula Horik., J. Sci. Hiroshima Univ.,
Ser. B, Div. 2, Bot. 2: 288, 1934 (Horikawa 1934).
* Cololejeunea micronesica H.A.Mill. et Bonner, Beih. Nova Hedwigia 11: 63, 1963
(Miller et al. 1963). 264
** Cololejeunea mooreaensis Tixier, Bauhinia 8 (4): 226, 1987 (Hürlimann 1987).
* Cololejeunea ninguana Tixier, Nova Hedwigia 31: 776, 1979 (Tixier 1979b). 265
** Cololejeunea obcordata (Austin) A.Evans, Trans. Connecticut Acad. Arts 10 (8):
448, 1900 (Evans 1900a). Bas.: Lejeunea obcordata Austin, Bot. Bull. (Hanover) 1
(8): 36, 1876 (Austin 1876a).
262 Cololejeunea kohkongensis may be conspeciic with Cololejeunea plagiophylla.
263 Cololejeunea lobulilineata seems to be a depauperate form of some other species, maybe Cololejeunea
obtusifolia.
264 Cololejeunea micronesica is possibly conspeciic with Cololejeunea cookei ide Schuster (1980c, 1983a).
265 Cololejeunea ninguana is possibly conspeciic with Cololejeunea decliviloba (Yu et al. 2013).
330
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Cololejeunea obliqua (Nees et Mont.) Schifn., Bot. Jahrb. Syst. 23 (5): 586, 1897
(Schifner 1897). Bas.: Lejeunea obliqua Nees et Mont., Ann. Sci. Nat. Bot. (sér.
2) 19: 264, 1843 (Montagne 1843).
* Cololejeunea oblongiperianthia (P.C.Wu et J.S.Lou) Piippo, J. Hattori Bot. Lab. 68:
134, 1990 (Piippo 1990). Bas.: Leptocolea oblongiperianthia P.C.Wu et J.S.Lou,
Acta Phytotax. Sin. 16 (4): 109, 1978 (Wu and Lou 1978). 266
** Cololejeunea oleana Sim, Trans. Roy. Soc. South Africa 15 (1): 49, 1926 (Sim 1926).
** Cololejeunea ovalifolia A.Evans, Trans. Connecticut Acad. Arts 10 (8): 450, 1900
(Evans 1900a).
** Cololejeunea panchoana Tixier, Ann. Fac. Sci. Univ. Phnom Penh 3: 183, 1970
(Tixier 1970b).
*** Cololejeunea paniensis (Tixier) Grolle, J. Bryol. 8 (4): 485, 1975 (Grolle 1975d).
Bas.: Jovetastella paniensis Tixier, Rev. Bryol. Lichénol. 39 (4): 662, 1973 [1974]
(Tixier 1973c).
** Cololejeunea papulilora Steph., Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr. 85: 199, 1910 (Stephani 1910a).
** Cololejeunea pentagona (Mitt.) Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 309, 1896 (Stephani 1896a). Bas.: Lejeunea pentagona Mitt., Fl. vit.: 416, 1871 [1873] (Mitten
1871).
*** Cololejeunea plagiochiliana Tixier, Bot. Not. 128: 428, 1975 [1976] (Tixier 1975a).
** Cololejeunea planiuscula Tixier, Cryptog. Bryol. Lichénol. 16 (3): 229, 1995 (Tixier 1995a). Based on: Cololejeunea planiuscula Tixier, Candollea 46 (2): 289, 1991
(Tixier 1991), nom. inval.
*** Cololejeunea pseudoserrata Tixier, Nova Hedwigia 31: 770, 1979 (Tixier 1979b).
* Cololejeunea pteroporum Tixier, Bryophyt. Biblioth. 27: 271, 1985 (Tixier 1985a). 267
** Cololejeunea pulchella (Mitt.) R.M.Schust., J. Hattori Bot. Lab. 26: 241, 1963
(Schuster 1963b). Bas.: Lejeunea pulchella Mitt., Bot. antarct. voy. II (Fl. Nov.Zel. 2): 157, 1854 (Mitten 1854).
** Cololejeunea pulchella var. stylifera R.M.Schust., Phytologia 56 (7): 458, 1985
(Schuster 1985c).
*** Cololejeunea quadridentata (S.Hatt.) Grolle, Acta Bot. Fenn. 125: 65, 1984 (Grolle and Piippo 1984). Bas.: Leptocolea quadridentata S.Hatt., Bot. Mag. (Tokyo) 64
(755/756): 117, 1951 (Hattori 1951c).
** Cololejeunea retusula (Mitt.) H.A.Mill., Phytologia 47 (4): 322, 1981 (Miller
1981). Bas.: Lejeunea retusula Mitt., Fl. vit.: 416, 1871 [1873] (Mitten 1871).
** Cololejeunea salgadoi Onr., Bull. Jard. Bot. Natl. Belg. 59 (3/4): 433, 1989 (Onraedt
1989).
*** Cololejeunea sanctae-helenae M.Wigginton, J. Bryol. 28 (4): 366, 2006 (Wigginton
2006).
** Cololejeunea serrulata Steph., Hedwigia 34 (5): 252, 1895 (Stephani 1895b).
266 Cololejeunea oblongiperianthia is possibly conspeciic with Cololejeunea trichomanis (Zhu and So 2001).
267 Cololejeunea pteroporum is possibly conspeciic with Cololejeunea spathulifolia.
World checklist of hornworts and liverworts
331
*** Cololejeunea setiloba A.Evans, Bryologist 16 (4): 51, 1913 (Evans 1913).
*** Cololejeunea siangensis G.Asthana et S.C.Srivast., Bryophyt. Biblioth. 60: 57, 2003
(Asthana and Srivastava 2003).
*** Cololejeunea skottsbergii Herzog, Nat. Hist. Juan Fernandez (Botany) 2 (5): 749,
1942 (Herzog 1942a).
*** Cololejeunea societatis Tixier, Bauhinia 8 (4): 230, 1987 (Hürlimann 1987).
* Cololejeunea spathulifolia (Steph.) H.A.Mill., Phytologia 47 (4): 322, 1981 (Miller
1981). Bas.: Leptocolea spathulifolia Steph., Sp. Hepat. (Stephani) 5: 855, 1916
(Stephani 1916b). 268
** Cololejeunea spruceana Tixier, Cryptog. Bryol. Lichénol. 16 (3): 229, 1995 (Tixier 1995a). Based on: Cololejeunea spruceana Tixier, Candollea 46 (2): 286, 1991
(Tixier 1991), nom. inval.
** Cololejeunea stenophylla Herzog, Bot. Not. 100 (4): 330, 1947 (Herzog 1947).
*** Cololejeunea streimannii Pócs, Acta Bryolichenol. Asiat. 4: 96, 2011 (Pócs and
Piippo 2011).
** Cololejeunea streimannii subsp. solomonensis Pócs, Acta Bryolichenol. Asiat. 4: 96,
2011 (Pócs and Piippo 2011).
*** Cololejeunea subalpina Pócs, Acta Bryolichenol. Asiat. 4: 98, 2011 (Pócs and Piippo
2011).
** Cololejeunea subcristata A.Evans, Bryologist 20 (2): 24, 1917 (Evans 1917b).
* Cololejeunea takamakae Tixier, Bryophyt. Biblioth. 27: 319, 1985 (Tixier 1985a). 269
*** Cololejeunea tanzaniae Pócs, J. Hattori Bot. Lab. 48: 312, 1980 (Pócs 1980b).
** Cololejeunea tenuiparietata Tixier, Trop. Bryol. 11: 56, 1995 (Tixier 1995b).
** Cololejeunea teurnoumensis Tixier, Ann. Fac. Sci. Univ. Phnom Penh 3: 183, 1970
(Tixier 1970b).
*** Cololejeunea timoi Pócs, Acta Bryolichenol. Asiat. 4: 98, 2011 (Pócs and Piippo 2011).
** Cololejeunea touwii Pócs, Acta Bryolichenol. Asiat. 4: 101, 2011 (Pócs and Piippo
2011).
*** Cololejeunea tranninhiana Tixier, Ann. Hist.-Nat. Mus. Natl. Hung. 66: 97, 1974
(Tixier 1974).
*** Cololejeunea trichomanis (Gottsche) Besch., Rev. Bryol. 19 (1): 14, 1892 (Bescherelle 1892). Bas.: Lejeunea trichomanis Gottsche, Abh. Naturwiss. Vereins
Bremen 7: 362, 1882 (Gottsche 1882).
* Cololejeunea tuksapiana Tixier, Ann. Fac. Sci. Univ. Phnom Penh 3: 186, 1970
(Tixier 1970b). 270
** Cololejeunea vulcania Tixier, Ann. Fac. Sci. Univ. Phnom Penh 3: 181, 1970 (Tixier
1970b).
*** Cololejeunea yoshinagana (S.Hatt.) Mizut., J. Hattori Bot. Lab. 24: 250, 1961
(Mizutani 1961). Bas.: Leptocolea yoshinagana S.Hatt., Bull. Tokyo Sci. Mus. 11:
115, 1944 (Hattori 1944d).
268 Cololejeunea spathulifolia is possibly conspeciic with Cololejeunea obliqua (Pócs and Piippo 2011).
269 Cololejeunea takamakae is possibly conspeciic with Cololejeunea angustilora.
270 Cololejeunea tuksapiana is possibly conspeciic with Cololejeunea aequalis.
332
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subg. Metzgeriopsis (K.I.Goebel) Pócs, Acta Bryolichenol. Asiat. 4: 106, 2011
(Pócs and Piippo 2011). Bas.: Metzgeriopsis K.I.Goebel, Ann. Jard. Bot. Buitenzorg 7 (1): 54, 1888 (Goebel 1888).
*** Cololejeunea metzgeriopsis (K.I.Goebel) Gradst., R.Wilson, Ilk.-Borg. et Heinrichs, Bot. J. Linn. Soc. 151 (3): 306, 2006 (Gradstein et al. 2006). Bas.: Lejeunea
metzgeriopsis K.I.Goebel, Flora 72 (1): 2, 1889 (Goebel 1889).
*
*** subg. Pedinolejeunea Benedix ex Mizut., J. Hattori Bot. Lab. 24: 240, 1961
(Mizutani 1961).
** Cololejeunea abnormis Mizut., J. Hattori Bot. Lab. 33: 260, 1970 (Mizutani 1970).
*** Cololejeunea adhaesiva (Mitt.) R.M.Schust., Beih. Nova Hedwigia 9: 177, 1963
(Schuster 1963a). Bas.: Lejeunea adhaesiva Mitt., J. Linn. Soc., Bot. 22 (146): 325,
1886 (Mitten 1886b).
** Cololejeunea adnata Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 194, 1977
[1979] (Tixier 1977a).
*** Cololejeunea africana (Steph.) R.M.Schust., Beih. Nova Hedwigia 9: 173, 1963
(Schuster 1963a). Bas.: Physocolea africana Steph., Sp. Hepat. (Stephani) 5: 867,
1916 (Stephani 1916b).
** Cololejeunea ambeliensis Tixier, Bryophyt. Biblioth. 27: 142, 1985 (Tixier 1985a).
** Cololejeunea amieuensis Tixier, Nova Hedwigia 31: 748, 1979 (Tixier 1979b).
** Cololejeunea andapania Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 178, 1977
[1979] (Tixier 1977a).
*** Cololejeunea angulata (Steph.) Mizut., J. Hattori Bot. Lab. 28: 108, 1965 (Mizutani 1965). Bas.: Leptocolea angulata Steph., Sp. Hepat. (Stephani) 5: 847, 1916
(Stephani 1916b).
*** Cololejeunea ankaiana Tixier, Bryophyt. Biblioth. 27: 62, 1985 (Tixier 1985a).
** Cololejeunea attilana Pócs, Magyar Bot. Kut. Ezredf. Tanul. Borhidi: 186, 2002
(Pócs 2002a).
*** Cololejeunea auriculata (E.W.Jones) R.M.Schust., Beih. Nova Hedwigia 9: 177,
1963 (Schuster 1963a). Bas.: Leptocolea auriculata E.W.Jones, Trans. Brit. Bryol.
Soc. 2 (2): 152, 1953 (Jones 1953b).
** Cololejeunea autoica (Steph.) Grolle, Bryophyt. Biblioth. 48: 43, 1995 (Grolle
1995). Bas.: Physocolea autoica Steph., Sp. Hepat. (Stephani) 5: 867, 1916 (Stephani 1916b).
*** Cololejeunea bekkeri Tixier, Cryptog. Bryol. Lichénol. 16 (3): 229, 1995 (Tixier
1995a). Based on: Cololejeunea bekkeri Tixier, Candollea 46 (2): 269, 1991 (Tixier
1991), nom. inval.
*** Cololejeunea bischleriana Tixier, Bradea 3 (6): 36, 1980 (Tixier 1980a).
** Cololejeunea bolovenensis Tixier, Nat. Hist. Bull. Siam Soc. 24 (3/4): 442, 1973
(Tixier 1973b).
*** Cololejeunea borbonica Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 188, 1977
[1979] (Tixier 1977a).
World checklist of hornworts and liverworts
333
* Cololejeunea brunelii Tixier, Dacca Univ. Stud., B 15: 10, 1967 (Tixier 1967). 271
*** Cololejeunea cardiocarpa (Mont.) A.Evans, Mem. Torrey Bot. Club 8 (2): 172,
1902 (Evans 1902a). Bas.: Lejeunea cardiocarpa Mont., Hist. Phys. Cuba, Bot., Pl.
Cell.: 476, 1842 (Montagne 1842a).
* Cololejeunea chittagongensis Tixier, Bryophyt. Biblioth. 27: 97, 1985 (Tixier 1985a). 272
*** Cololejeunea cocoscola Tixier, Cryptog. Bryol. Lichénol. 14 (3): 353, 1993 (Tixier
1993).
** Cololejeunea cremersii Tixier, Cryptog. Bryol. Lichénol. 16 (3): 229, 1995 (Tixier
1995a). Based on: Cololejeunea cremersii Tixier, Candollea 46 (2): 271, 1991 (Tixier
1991), nom. inval.
** Cololejeunea cristata (Steph.) R.M.Schust., Beih. Nova Hedwigia 9: 173, 1963
(Schuster 1963a). Bas.: Physocolea cristata Steph., Sp. Hepat. (Stephani) 5: 869,
1916 (Stephani 1916b).
** Cololejeunea cuneata (Lehm. et Lindenb.) Herzog, Bot. Not. 100 (4): 320, 1947
(Herzog 1947). Bas.: Jungermannia cuneata Lehm. et Lindenb., Nov. Stirp. Pug.
4: 56, 1832 (Lehmann 1832).
* Cololejeunea deroinii Tixier, Cryptog. Bryol. Lichénol. 14 (3): 355, 1993 (Tixier 1993).
* Cololejeunea deslooveri Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 47 (1/2): 227,
1977 (Vanden Berghen 1977). 273
* Cololejeunea dzumacensis Tixier, Nova Hedwigia 31: 754, 1979 (Tixier 1979b). 274
*** Cololejeunea ecuadoriensis Pócs, Acta Bot. Hung. 44 (3/4): 372, 2002 (Pócs 2002b).
* Cololejeunea epiphylla G.Asthana et A.Shukla, Cryptog. Bryol. 31 (3): 218, 2010
(Asthana and Shukla 2010). 275
** Cololejeunea issilobula Herzog, Trans. Brit. Bryol. Soc. 1 (4): 323, 1950 (Herzog
1950a).
** Cololejeunea lorencei Tixier, Cryptog. Bryol. Lichénol. 14 (3): 355, 1993 (Tixier
1993).
** Cololejeunea foliicola S.C.Srivast. et G.Srivast., Proc. Indian Acad. Sci. Pl. Sci. 99
(2): 86, 1989 (Srivastava and Srivastava 1989a).
** Cololejeunea fructumarginata Tixier, Bryophyt. Biblioth. 27: 58, 1985 (Tixier
1985a).
*** Cololejeunea furcilobulata (Berrie et E.W.Jones) R.M.Schust., Beih. Nova Hedwigia 9: 178, 1963 (Schuster 1963a). Bas.: Leptocolea furcilobulata Berrie et
E.W.Jones, Trans. Brit. Bryol. Soc. 2 (3): 417, 1954 (Jones 1954c).
*** Cololejeunea geissleriana Tixier, Bradea 3 (6): 37, 1980 (Tixier 1980a).
271 Cololejeunea brunelii is conspeciic with Cololejeunea madothecoides in Tixier (1985a), but it belongs to
the Cololejeunea raduliloba, Cololejeunea furcilobulata, Cololejeunea paucimarginata species complex.
272 Cololejeunea chittagongensis is probably conspeciic with Cololejeunea schwabei.
273 Cololejeunea deslooveri is possibly conspeciic with Cololejeunea cristata.
274 Cololejeunea dzumacensis is possibly conspeciic with Cololejeunea lanciloba, but the type specimen
has not been localized.
275 Cololejeunea epiphylla is closely related to Cololejeunea chittagongensis and Cololejeunea schwabei.
334
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Cololejeunea georgiana Tixier, Bryophyt. Biblioth. 27: 145, 1985 (Tixier 1985a).
*** Cololejeunea guadelupensis Tixier, Cryptog. Bryol. Lichénol. 16 (3): 229, 1995
(Tixier 1995a). Based on: Cololejeunea guadelupensis Tixier, Candollea 46 (2): 276,
1991 (Tixier 1991), nom. inval.
*** Cololejeunea hebridensis Tixier, Bot. Not. 128: 425, 1975 [1976] (Tixier 1975a).
*** Cololejeunea hinidumae Onr., Acta Bot. Acad. Sci. Hung. 25 (1/2): 109, 1979
(Onraedt 1979).
* Cololejeunea hoabinhiana Tixier, Ann. Hist.-Nat. Mus. Natl. Hung. 66: 91, 1974
(Tixier 1974). 276
** Cololejeunea hoeana Tixier, Bryophyt. Biblioth. 27: 56, 1985 (Tixier 1985a).
** Cololejeunea hungii Tixier, Ann. Hist.-Nat. Mus. Natl. Hung. 66: 91, 1974 (Tixier 1974).
*** Cololejeunea indosinica Tixier, Bryophyt. Biblioth. 27: 63, 1985 (Tixier 1985a).
** Cololejeunea inoueana Mizut., J. Hattori Bot. Lab. 57: 440, 1984 (Mizutani
1984c).
** Cololejeunea japonica (Schifn.) Mizut., J. Hattori Bot. Lab. 24: 241, 1961 (Mizutani 1961). Bas.: Leptocolea japonica Schifn., Ann. Bryol. 2: 92, 1929 (Schifner
1929).
*** Cololejeunea jonesii Pócs, Acta Bot. Acad. Sci. Hung. 21 (3/4): 361, 1975 (Pócs 1975).
*** Cololejeunea kapingaensis H.A.Mill., Bryologist 59 (3): 170, 1956 (Miller 1956).
** Cololejeunea kiriromensis Tixier, Bryophyt. Biblioth. 27: 147, 1985 (Tixier 1985a).
*** Cololejeunea kulenensis Tixier, Bryophyt. Biblioth. 27: 71, 1985 (Tixier 1985a). Nom.
nov. pro Leptocolea verdoornii Herzog, Ann. Bryol. 5: 97, 1932 (Herzog 1932a).
** Cololejeunea laevigata (Mitt.) R.M.Schust., J. Hattori Bot. Lab. 26: 241, 1963
(Schuster 1963b). Bas.: Lejeunea laevigata Mitt., Bot. antarct. voy. II (Fl. Nov.Zel. 2): 157, 1854 (Mitten 1854).
*** Cololejeunea lanciloba Steph., Hedwigia 34 (5): 250, 1895 (Stephani 1895b).
*** Cololejeunea latilobula (Herzog) Tixier, Bryophyt. Biblioth. 27: 156, 1985 (Tixier
1985a). Bas.: Leptocolea latilobula Herzog, Symb. Sin. 5: 54, 1930 (Nicholson et
al. 1930).
*** Cololejeunea latistyla R.L.Zhu, Hikobia 11: 544, 1994 (Zhu et al. 1994).
*** Cololejeunea leloutrei (E.W.Jones) R.M.Schust., Beih. Nova Hedwigia 9: 173,
1963 (Schuster 1963a). Bas.: Leptocolea leloutrei E.W.Jones, Trans. Brit. Bryol.
Soc. 2 (2): 146, 1953 (Jones 1953b).
** Cololejeunea leloutrei var. microlobulata Tixier, Bryophyt. Biblioth. 27: 78, 1985
(Tixier 1985a).
** Cololejeunea leloutrei var. ulugurica Pócs ex Tixier, Bryophyt. Biblioth. 27: 74,
1985 (Tixier 1985a).
*** Cololejeunea lemuriana Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 191, 1977
[1979] (Tixier 1977a).
** Cololejeunea littoralis Tixier, Bryophyt. Biblioth. 27: 163, 1985 (Tixier 1985a).
276 Cololejeunea hoabinhiana is possibly conspeciic with Cololejeunea lanciloba.
World checklist of hornworts and liverworts
335
** Cololejeunea longistylis A.Evans, Trans. Connecticut Acad. Arts 10 (8): 453, 1900
(Evans 1900a).
*** Cololejeunea magnistyla (Horik.) Mizut., J. Hattori Bot. Lab. 24: 243, 1961 (Mizutani 1961). Bas.: Leptocolea magnistyla Horik., J. Sci. Hiroshima Univ., Ser. B,
Div. 2, Bot. 1: 131, 1932 (Horikawa 1932c).
* Cololejeunea malaccensis Tixier, Bryophyt. Biblioth. 27: 42, 1985 (Tixier 1985a). 277
** Cololejeunea malayana Tixier, Bryophyt. Biblioth. 27: 154, 1985 (Tixier 1985a).
*** Cololejeunea marginata (Lehm. et Lindenb.) Pearson, Forh. Vidensk.-Selsk. Kristiania 1892 (8): 9, 1892 (Pearson 1892). Bas.: Jungermannia marginata Lehm. et
Lindenb., Nov. Stirp. Pug. 5: 11, 1833 (Lehmann 1833).
* Cololejeunea maritima Tixier, Nova Hedwigia 31: 752, 1979 (Tixier 1979b).
*** Cololejeunea minutilobula Herzog, Rev. Bryol. Lichénol. 20 (1/2): 171, 1951
[1952] (Herzog 1951a).
*** Cololejeunea nigerica (E.W.Jones) R.M.Schust., Beih. Nova Hedwigia 9: 177,
1963 (Schuster 1963a). Bas.: Leptocolea nigerica E.W.Jones, Trans. Brit. Bryol.
Soc. 2 (2): 150, 1953 (Jones 1953b).
** Cololejeunea occidentalis (E.W.Jones) Vanden Berghen, Rev. Bryol. Lichénol. 44
(4): 449, 1978 (Vanden Berghen 1978). Bas.: Leptocolea cristata var. occidentalis
E.W.Jones, Trans. Brit. Bryol. Soc. 2 (2): 149, 1953 (Jones 1953b).
** Cololejeunea onraedtii Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 202, 1977
[1979] (Tixier 1977a).
*** Cololejeunea paciica Pócs, Acta Bot. Hung. 54 (1/2): 158, 2012 (Pócs 2012b).
** Cololejeunea panamensis G.Dauphin et Pócs, Trop. Bryol. 27: 76, 2006 (Dauphin
et al. 2006).
*** Cololejeunea paucimarginata Tixier, Bryophyt. Biblioth. 27: 100, 1985 (Tixier 1985a).
** Cololejeunea perakensis Tixier, Bryophyt. Biblioth. 27: 95, 1985 (Tixier 1985a).
*** Cololejeunea plagiophylla Benedix, Feddes Repert. Spec. Nov. Regni Veg. Beih.
134: 73, 1953 (Benedix 1953).
*** Cololejeunea planissima (Mitt.) Abeyw., Ceylon J. Sci., Biol. Sci. 2 (1): 73, 1959
(Abeywickrama 1959). Bas.: Lejeunea planissima Mitt., J. Proc. Linn. Soc., Bot. 5
(18): 117, 1860 [1861] (Mitten 1860c).
** Cololejeunea planissima var. chagosensis Pócs, J. Bryol. 28 (1): 14, 2006 (Seaward et
al. 2006).
** Cololejeunea praeruptorum Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 205,
1977 [1979] (Tixier 1977a).
** Cololejeunea producta (Mitt.) S.Hatt., Fl. E. Himalaya: 533, 1966 (Hattori 1966c).
Bas.: Lejeunea producta Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 117, 1860 [1861]
(Mitten 1860c).
277 Cololejeunea malaccensis may be conspeciic with Cololejeunea stylosa.
336
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Cololejeunea punctata (Gottsche) Pearson, Forh. Vidensk.-Selsk. Kristiania 1892
(14): 9, 1893 (Pearson 1893). Bas.: Lejeunea punctata Gottsche, Abh. Naturwiss.
Vereins Bremen 7: 361, 1882 (Gottsche 1882). 278
Cololejeunea raduliloba Steph., Hedwigia 34 (5): 251, 1895 (Stephani 1895b).
Cololejeunea reineckeana Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 309, 1896 (Stephani
1896a).
Cololejeunea saltuum Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 188, 1977
[1979] (Tixier 1977a).
Cololejeunea sambiroana Tixier, Bryologist 82 (4): 608, 1979 (Tixier 1979d).
Cololejeunea saroltae Pócs, Acta Bot. Hung. 54 (1/2): 160, 2012 (Pócs 2012b).
Cololejeunea schusteri Pócs, Acta Bot. Hung. 44 (3/4): 376, 2002 (Pócs 2002b).
Cololejeunea schwabei Herzog, J. Hattori Bot. Lab. 14: 54, 1955 (Herzog and
Noguchi 1955).
Cololejeunea selangorensis Tixier, Bryophyt. Biblioth. 27: 166, 1985 (Tixier 1985a).
Cololejeunea shibiensis Mizut., J. Hattori Bot. Lab. 57: 437, 1984 (Mizutani 1984c).
Cololejeunea smitinandii Tixier, Bryophyt. Biblioth. 27: 131, 1985 (Tixier 1985a).
Based on: Cololejeunea smitinandii Tixier, Nat. Hist. Bull. Siam Soc. 24 (3/4):
439, 1973 (Tixier 1973b), nom. inval.
Cololejeunea stoniana Tixier, Bot. Not. 128: 429, 1975 [1976] (Tixier 1975a).
Cololejeunea stylosa Steph., Trans. Connecticut Acad. Arts 10 (8): 454, 1900 (Evans 1900a). Based on: Cololejeunea stylosa Steph., Hedwigia 27 (11/12): 289, 1888
(Stephani 1888c), nom. inval.
Cololejeunea subcardiocarpa Tixier, Bradea 3 (6): 39, 1980 (Tixier 1980a).
Cololejeunea subinlata Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 182, 1977
[1979] (Tixier 1977a).
Cololejeunea submarginata Tixier, Bradea 3 (6): 40, 1980 (Tixier 1980a).
Cololejeunea subminutilobula Mizut., J. Hattori Bot. Lab. 24: 282, 1961 (Mizutani
1961). Nom. nov. pro Leptocolea minutilobula Horik., J. Sci. Hiroshima Univ., Ser.
B, Div. 2, Bot. 1: 16, 1931 (Horikawa 1931b).
Cololejeunea subscariosa (Spruce) Pócs, Acta Bot. Hung. 56 (1/2): 197, 2014 (Pócs
et al. 2014). Bas.: Lejeunea subscariosa Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 300, 1884 (Spruce 1884).
Cololejeunea subtriapiculata Tixier, Nova Hedwigia 31: 744, 1979 (Tixier 1979b).
Cololejeunea succinea Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 194, 1977
[1979] (Tixier 1977a).
Cololejeunea surinamensis Tixier, Bradea 3 (6): 42, 1980 (Tixier 1980a).
Cololejeunea tahitensis Tixier, Cryptog. Bryol. Lichénol. 14 (3): 359, 1993 (Tixier
1993).
Cololejeunea tamatavensis Tixier, Bull. Acad. Malgache (n.ser) 55 (1/2): 191, 1977
[1979] (Tixier 1977a).
278 Cololejeunea punctata is a doubtful taxon. he type specimen has been destroyed and the identity is
unclear (Grolle 1995).
World checklist of hornworts and liverworts
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Cololejeunea taprobanea Tixier, Bryophyt. Biblioth. 27: 158, 1985 (Tixier 1985a).
Cololejeunea thailandensis Tixier, Nat. Hist. Bull. Siam Soc. 24 (3/4): 441, 1973
(Tixier 1973b).
Cololejeunea triapiculata (Herzog) Tixier, Gard. Bull. Singapore 25 (3): 344, 1971
(Tixier 1971). Bas.: Leptocolea triapiculata Herzog, Ann. Bryol. 5: 95, 1932 (Herzog 1932a).
Cololejeunea tribracteata Tixier, Trop. Bryol. 11: 46, 1995 (Tixier 1995b). 279
Cololejeunea tridentata Tixier, Bryophyt. Biblioth. 27: 83, 1985 (Tixier 1985a).
Cololejeunea uchimae Amakawa, J. Jap. Bot. 33 (5): 142, 1958 (Amakawa 1958a). 280
Cololejeunea verwimpii Tixier, Cryptog. Bryol. Lichénol. 16 (3): 230, 1995 (Tixier
1995a).
Cololejeunea vidaliana Tixier, Nat. Hist. Bull. Siam Soc. 24 (3/4): 444, 1973 (Tixier
1973b).
Cololejeunea vietnamensis Tixier, Bryophyt. Biblioth. 27: 127, 1985 (Tixier 1985a).
Cololejeunea vitaliana Tixier, Cryptog. Bryol. Lichénol. 16 (3): 230, 1995 (Tixier
1995a).
Cololejeunea yakusimensis (S.Hatt.) Mizut., J. Hattori Bot. Lab. 57: 430, 1984
(Mizutani 1984c). Bas.: Leptocolea lanciloba var. yakusimensis S.Hatt., J. Jap. Bot.
18 (11): 655, 1942 (Hattori 1942).
subg. Protocolea R.M.Schust., Beih. Nova Hedwigia 9: 171, 1963 (Schuster
1963a).
** Cololejeunea chuahiana Pócs, Polish Bot. J. 47 (1): 11, 2002 (Pócs 2002c). 281
** Cololejeunea dauphinii R.L.Zhu, J. Bryol. 28 (3): 277, 2006 (Zhu 2006b). Nom.
nov. pro Cololejeunea tixieri M.I.Morales et G.Dauphin, Trop. Bryol. 14: 133,
1998 (Morales and Dauphin 1998), nom. illeg. 282
** Cololejeunea discilora Tixier, Bryologist 82 (4): 604, 1979 (Tixier 1979d). 283
*
*** subg. Taeniolejeunea (Zwickel) Benedix, Feddes Repert. Spec. Nov. Regni Veg.
Beih. 134: 21, 1953 (Benedix 1953). Bas.: Taeniolejeunea Zwickel, Ann. Bryol. 6:
106, 1933 (Zwickel 1933).
* Cololejeunea amoena Benedix, Feddes Repert. Spec. Nov. Regni Veg. Beih. 134:
25, 1953 (Benedix 1953). 284
*** Cololejeunea appressa (A.Evans) Benedix, Feddes Repert. Spec. Nov. Regni Veg.
Beih. 134: 31, 1953 (Benedix 1953). Bas.: Leptocolea appressa A.Evans, Bull. Torrey Bot. Club 39 (12): 606, 1912 [1913] (Evans 1912d).
279 Cololejeunea tribracteata is possibly conspeciic with Cololejeunea africana.
280 Cololejeunea uchimae is conspeciic with Cololejeunea raduliloba in Mizutani (1961), but it is accepted
in recent Japanese checklists.
281 Cololejeunea chuahiana is probably a Myriocoleopsis species (Yu et al. 2014).
282 Cololejeunea dauphinii is probably a Myriocoleopsis species (Yu et al. 2014).
283 Cololejeunea discilora is probably a Myriocoleopsis species (Yu et al. 2014).
284 Cololejeunea amoena is possibly conspeciic with Cololejeunea loccosa (Söderström et al. 2010a).
338
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Cololejeunea bachmaensis Tixier, Rev. Bryol. Lichénol. 36 (3/4): 560, 1969 [1970]
(Tixier 1969).
Cololejeunea bontocensis Tixier, Rev. Bryol. Lichénol. 36 (3/4): 583, 1969 [1970]
(Tixier 1969). 285
Cololejeunea crassipapillata Tixier, Rev. Bryol. Lichénol. 36 (3/4): 565, 1969
[1970] (Tixier 1969).
Cololejeunea eustacei Pócs, J. Bryol. 29 (2): 83, 2007 (Müller and Pócs 2007).
Cololejeunea falcata (Horik.) Benedix, Feddes Repert. Spec. Nov. Regni Veg. Beih.
134: 29, 1953 (Benedix 1953). Bas.: Physocolea falcata Horik., J. Sci. Hiroshima
Univ., Ser. B, Div. 2, Bot. 1: 22, 1931 (Horikawa 1931b).
Cololejeunea falcata var. madecassa Tixier, Bryologist 82 (4): 606, 1979 (Tixier 1979d).
Cololejeunea lavida P.C.Wu et J.S.Lou, Acta Phytotax. Sin. 16 (4): 103, 1978
(Wu and Lou 1978). 286
Cololejeunea lavovittata Pócs, Acta Bryolichenol. Asiat. 4: 120, 2011 (Pócs and
Piippo 2011).
Cololejeunea loccosa (Lehm. et Lindenb.) Schifn., Consp. Hepat. Arch. Ind.: 243,
1898 (Schifner 1898b). Bas.: Jungermannia loccosa Lehm. et Lindenb., Nov.
Stirp. Pug. 5: 26, 1833 (Lehmann 1833).
Cololejeunea loccosa var. amoenoides Tixier, Cryptog. Bryol. Lichénol. 2 (1): 59,
1981 (Tixier 1981).
Cololejeunea loccosa var. angustibracteata Tixier, Cryptog. Bryol. Lichénol. 2 (1):
65, 1981 (Tixier 1981).
Cololejeunea loccosa var. aurita Benedix, Feddes Repert. Spec. Nov. Regni Veg.
Beih. 134: 24, 1953 (Benedix 1953).
Cololejeunea loccosa var. conivens Benedix, Feddes Repert. Spec. Nov. Regni Veg.
Beih. 134: 24, 1953 (Benedix 1953).
Cololejeunea loccosa var. ocellata Tixier, Cryptog. Bryol. Lichénol. 2 (1): 69, 1981
(Tixier 1981).
Cololejeunea loccosa var. plicata Tixier, Cryptog. Bryol. Lichénol. 2 (1): 65, 1981
(Tixier 1981).
Cololejeunea loccosa var. trivittata Tixier, Cryptog. Bryol. Lichénol. 2 (1): 62,
1981 (Tixier 1981).
Cololejeunea gresicola Tixier, Rev. Bryol. Lichénol. 36 (3/4): 569, 1969 [1970]
(Tixier 1969).
Cololejeunea gynophthalma Benedix, Feddes Repert. Spec. Nov. Regni Veg. Beih.
134: 32, 1953 (Benedix 1953).
Cololejeunea inlata Steph., Hedwigia 34 (5): 249, 1895 (Stephani 1895b).
Cololejeunea khiavensis Tixier, Rev. Bryol. Lichénol. 36 (3/4): 557, 1969 [1970]
(Tixier 1969).
285 Cololejeunea bontocensis is possibly conspeciic with Cololejeunea pseudostephanii.
286 Cololejeunea lavida may be conspeciic with Cololejeunea perainis (Zhu and So 2001).
World checklist of hornworts and liverworts
339
** Cololejeunea koratensis Tixier, Rev. Bryol. Lichénol. 36 (3/4): 585, 1969 [1970]
(Tixier 1969).
* Cololejeunea manlinensis Tixier, Rev. Bryol. Lichénol. 36 (3/4): 563, 1969 [1970]
(Tixier 1969).
** Cololejeunea maquilingensis Tixier, Rev. Bryol. Lichénol. 36 (3/4): 579, 1969
[1970] (Tixier 1969).
** Cololejeunea mutabilis Benedix, Feddes Repert. Spec. Nov. Regni Veg. Beih. 134:
27, 1953 (Benedix 1953).
** Cololejeunea nakajimae S.Hatt., J. Hattori Bot. Lab. 10: 57, 1953 (Hattori and
Kodama 1953).
*** Cololejeunea ocellata (Horik.) Benedix, Feddes Repert. Spec. Nov. Regni Veg.
Beih. 134: 38, 1953 (Benedix 1953). Bas.: Leptocolea ocellata Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 1: 86, 1932 (Horikawa 1932a).
*** Cololejeunea ocelloides (Horik.) Mizut., J. Hattori Bot. Lab. 24: 277, 1961 (Mizutani 1961). Bas.: Leptocolea ocelloides Horik., J. Sci. Hiroshima Univ., Ser. B, Div.
2, Bot. 2: 280, 1934 (Horikawa 1934).
*** Cololejeunea perainis (Schifn.) Schifn., Consp. Hepat. Arch. Ind.: 245, 1898
(Schifner 1898b). Bas.: Lejeunea perainis Schifn., Nova Acta Acad. Caes. Leop.Carol. German. Nat. Cur. 60 (2): 242, 1893 (Schifner 1893a).
* Cololejeunea perainis var. ciconiae Tixier, Bull. Jard. Bot. Natl. Belg. 59 (3/4):
444, 1989 (Tixier 1989).
** Cololejeunea perainis var. elegans Benedix, Feddes Repert. Spec. Nov. Regni Veg.
Beih. 134: 35, 1953 (Benedix 1953).
** Cololejeunea perainis var. serrulata Schifn. ex Benedix, Feddes Repert. Spec. Nov.
Regni Veg. Beih. 134: 35, 1953 (Benedix 1953).
* Cololejeunea polisiana Tixier, Rev. Bryol. Lichénol. 36 (3/4): 550, 1969 [1970]
(Tixier 1969). 287
*** Cololejeunea pseudoloccosa (Horik.) Benedix, Feddes Repert. Spec. Nov. Regni
Veg. Beih. 134: 36, 1953 (Benedix 1953). Bas.: Leptocolea pseudoloccosa Horik., J.
Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 1: 87, 1932 (Horikawa 1932a).
*** Cololejeunea pseudostephanii Tixier, Rev. Bryol. Lichénol. 36 (3/4): 571, 1969
[1970] (Tixier 1969).
** Cololejeunea setosa Mizut., J. Hattori Bot. Lab. 29: 163, 1966 (Mizutani 1966).
*** Cololejeunea sharpii Mizut., J. Hattori Bot. Lab. 39: 258, 1975 (Mizutani 1975).
*** Cololejeunea siamensis Steph., Bot. Tidsskr. 24 (3): 279, 1902 (Stephani 1902b).
*** Cololejeunea sphaerodonta Mizut., J. Hattori Bot. Lab. 29: 165, 1966 (Mizutani 1966).
*** Cololejeunea stephanii Schifn. ex Benedix, Feddes Repert. Spec. Nov. Regni Veg.
Beih. 134: 40, 1953 (Benedix 1953).
** Cololejeunea subloccosa Mizut., J. Hattori Bot. Lab. 57: 168, 1984 (Mizutani 1984b).
** Cololejeunea subocelloides Mizut., J. Hattori Bot. Lab. 57: 163, 1984 (Mizutani 1984b).
287 Cololejeunea polisiana may be conspeciic with Cololejeunea perainis.
340
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*
Cololejeunea tamdaoensis Tixier, Rev. Bryol. Lichénol. 36 (3/4): 567, 1969 [1970]
(Tixier 1969).
** Cololejeunea verdoornii (S.Hatt.) S.Hatt., J. Hattori Bot. Lab. 17: 75, 1956 [1957]
(Hattori 1956b). Bas.: Taeniolejeunea verdoornii S.Hatt., J. Jap. Bot. 17: 459,
1941 (Hattori 1941).
** Cololejeunea yipii R.L.Zhu, Beih. Nova Hedwigia 121: 346, 2001 (Zhu and So 2001).
** Cololejeunea zantenorum Pócs, Acta Bryolichenol. Asiat. 4: 127, 2011 (Pócs and
Piippo 2011).
Incertae sedis
** Cololejeunea conchifolia (Gottsche) Gradst., J. Hattori Bot. Lab. 45: 109, 1979
(Gradstein and Hekking 1979). Bas.: Lejeunea conchifolia Gottsche, Ann. Sci. Nat.
Bot. (sér. 5) 1: 163, 1864 (Gottsche 1864). 288
** Cololejeunea dankiaensis Tixier, Phytotaxa 220 (2): 199, 2015 (Söderström et al.
2015d). Based on: Cololejeunea dankiaensis Tixier, Rev. Bryol. Lichénol. 36 (3/4):
581, 1969 [1970] (Tixier 1969), nom. inval.
** Cololejeunea ensifera Tixier, Phytotaxa 220 (2): 199, 2015 (Söderström et al.
2015d). Based on: Cololejeunea ensifera Tixier, Rev. Bryol. Lichénol. 36 (3/4):
562, 1969 [1970] (Tixier 1969), nom. inval.
** Cololejeunea hamata Steph., Hedwigia 34 (5): 248, 1895 (Stephani 1895b).
** Cololejeunea herzogii K.I.Goebel, Biblioth. Bot. 87 (2): 269, 1916 (Stephani 1916a).
** Cololejeunea jamesii (Austin) M.E.Reiner et Pócs, Phytotaxa 208 (1): 98, 2015
(Pócs et al. 2015a). Bas.: Lejeunea jamesii Austin, Bull. Torrey Bot. Club 6 (30):
158, 1877 (Austin 1877).
** Cololejeunea sublatistyla Jian Wang bis et R.L.Zhu, Phytotaxa 161 (2): 165, 2014
(Wang et al. 2014a).
** Cololejeunea tixieri Onr., Bull. Jard. Bot. Natl. Belg. 59 (3/4): 436, 1989 (Onraedt
1989).
* Cololejeunea variifolia (Mitt.) Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 309, 1896 (Stephani
1896a). Bas.: Lejeunea variifolia Mitt., Fl. vit.: 415, 1871 [1873] (Mitten 1871).
*** Colura (Dumort.) Dumort., Recueil Observ. Jungerm.: 12, 1835 (Dumortier
1835). Bas.: Lejeunea sect. Colura Dumort., Syll. Jungerm. Europ.: 32, 1831 (Dumortier 1831).
** subg. Colura
** sect. Colura
*** Colura berghenii Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 245, 1953 [1954]
(Jovet-Ast 1953).
288 Cololejeunea conchifolia is probably a Myriocoleopsis species (Yu et al. 2014).
World checklist of hornworts and liverworts
341
*** Colura calyptrifolia (Hook.) Dumort., Recueil Observ. Jungerm.: 12, 1835 (Dumortier 1835). Bas.: Jungermannia calyptrifolia Hook., Brit. Jungermann.: tab. 43,
1813 (Hooker 1813).
*** Colura hedbergiana Pócs, J. Bryol. 14 (3): 499, 1987 (Jones and Pócs 1987).
*** Colura humbertii Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 251, 1953 [1954] (Jovet-Ast 1953).
*** Colura irrorata (Spruce) Heinrichs, Y.Yu, Schäf.-Verw. et Pócs, Phytotaxa 66: 58,
2012 (Heinrichs et al. 2012c). Bas.: Myriocolea irrorata Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 305, 1884 (Spruce 1884).
** Colura junghuhniana (Prantl) Steph., Sp. Hepat. (Stephani) 5: 938, 1916 (Stephani
1916b). Bas.: Lejeunea junghuhniana Prantl, Hedwigia 29: xvi, 1890 (Prantl 1890).
** Colura medusa J.Eggers et Pócs, Chenia 11: 22, 2013 (Pócs 2013).
** Colura mizutanii Pócs, Chenia 11: 22, 2013 (Pócs 2013).
*** Colura rhynchophora Jovet-Ast, Rev. Bryol. Lichénol. 17 (1/4): 27, 1948 [1949]
(Jovet-Ast 1948).
*** Colura tenuicornis (A.Evans) Steph., Sp. Hepat. (Stephani) 5: 942, 1916 (Stephani
1916b), nom. conserv. Bas.: Colurolejeunea tenuicornis A.Evans, Trans. Connecticut Acad. Arts 10 (8): 455, 1900 (Evans 1900a).
** sect. Gamolepis Jovet-Ast, Cryptog. Bryol. Lichénol. 4 (3): 207, 1983 (Jovet-Ast
1983).
*** Colura cristata Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 291, 1953 [1954] (Jovet-Ast 1953).
*** Colura greig-smithii Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 293, 1953 [1954]
(Jovet-Ast 1953).
*** Colura inlata K.I.Goebel, Ann. Jard. Bot. Buitenzorg 39: 11, 1928 (Goebel 1928).
*** Colura jovet-astiae Grolle, J. Hattori Bot. Lab. 28: 44, 1965 (Grolle 1965b). Nom.
nov. pro Colura undulata Jovet-Ast, Rev. Bryol. Lichénol. 30 (1/2): 7, 1961 (Jovet-Ast 1961), nom. illeg.
*** Colura meijeri Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 290, 1953 [1954] (Jovet-Ast
1953).
*** Colura obvoluta Jovet-Ast, Cryptog. Bryol. Lichénol. 4 (3): 207, 1983 (Jovet-Ast
1983).
*** Colura ornata K.I.Goebel, Ann. Jard. Bot. Buitenzorg 9 (1): 26, 1890 [1891]
(Goebel 1890).
*** Colura palawanensis Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 305, 1953 [1954]
(Jovet-Ast 1953).
*** Colura valida Jovet-Ast, Rev. Bryol. Lichénol. 30 (1/2): 6, 1961 (Jovet-Ast 1961).
*** Colura verdoornii Herzog et Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 288, 1953
[1954] (Jovet-Ast 1953).
** sect. Harmophyllum Grolle, J. Hattori Bot. Lab. 28: 44, 1965 (Grolle 1965b).
*** Colura andoi Gradst. et Jovet-Ast, Hikobia 9: 355, 1986 (Gradstein 1986).
342
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Colura ari (Steph.) Steph., Sp. Hepat. (Stephani) 5: 936, 1916 (Stephani 1916b).
Bas.: Colurolejeunea ari Steph., Hedwigia 35 (3): 73, 1896 (Stephani 1896b).
*** Colura australiensis Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 260, 1953 [1954]
(Jovet-Ast 1953).
*** Colura brevistyla Herzog, Beih. Bot. Centralbl. 38 (2): 331, 1921 (Herzog 1921).
*** Colura calderae Pócs, J. Bryol. 29 (2): 84, 2007 (Müller and Pócs 2007).
*** Colura clementis Grolle, J. Hattori Bot. Lab. 28: 45, 1965 (Grolle 1965b).
*** Colura conica (Sande Lac.) K.I.Goebel, Ann. Jard. Bot. Buitenzorg 39: 3, 1928
(Goebel 1928). Bas.: Lejeunea conica Sande Lac., Ann. Mus. Bot. Lugduno-Batavi
1: 311, 1864 (Sande Lacoste 1864).
*** Colura corynophora (Nees, Lindenb. et Gottsche) Trevis., Mem. Reale Ist. Lombardo
Sci. (Ser. 3), C. Sci. Mat. 4 (13): 402, 1877 (Trevisan 1877). Bas.: Lejeunea corynophora Nees, Lindenb. et Gottsche, Observ. bot.: 474, 1843 (Gottsche et al. 1843).
*** Colura crenulata Grolle, J. Hattori Bot. Lab. 28: 46, 1965 (Grolle 1965b).
*** Colura crispiloba Jovet-Ast, Cryptog. Bryol. Lichénol. 4 (3): 205, 1983 (Jovet-Ast
1983).
** Colura cylindrica Herzog, Svensk Bot. Tidskr. 46 (1): 106, 1952 (Herzog 1952e).
*** Colura cymbalifera Herzog et Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 268, 1953
[1954] (Jovet-Ast 1953).
*** Colura digitalis (Mitt.) Steph., Sp. Hepat. (Stephani) 5: 931, 1916 (Stephani
1916b). Bas.: Lejeunea digitalis Mitt., J. Linn. Soc., Bot. 22 (146): 325, 1886
(Mitten 1886b).
** Colura digitalis var. mucronata Pócs, Fragm. Florist. Geobot. 40 (1): 263, 1995
(Pócs 1995).
*** Colura dusenii Steph., Sp. Hepat. (Stephani) 5: 931, 1916 (Stephani 1916b). Based
on: Colurolejeunea dusenii Steph., Hedwigia 31 (4): 168, 1892 (Stephani 1892g).
*** Colura fastigiata Jovet-Ast, Rev. Bryol. Lichénol. 27 (1/2): 28, 1958 (Jovet-Ast 1958).
*** Colura istulosa Jovet-Ast, Cryptog. Bryol. Lichénol. 4 (3): 211, 1983 (Jovet-Ast 1983).
*** Colura hattoriana Pócs, J. Hattori Bot. Lab. 74: 47, 1993 (Pócs 1993).
*** Colura heimii Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 275, 1953 [1954] (Jovet-Ast 1953).
*** Colura hemisphaerica Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 267, 1953 [1954]
(Jovet-Ast 1953).
*** Colura herzogii Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 261, 1953 [1954] (JovetAst 1953).
*** Colura inuii Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 1: 68, 1931
(Horikawa 1931a).
** Colura koponenii Pócs, Chenia 11: 29, 2013 (Pócs 2013).
** Colura mauritiana Pócs, Cryptog. Bryol. Lichénol. 18 (3): 196, 1997 (Pócs 1997a).
*** Colura maxima Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 284, 1953 [1954]
(Jovet-Ast 1953).
*** Colura mosenii Steph., Sp. Hepat. (Stephani) 5: 940, 1916 (Stephani 1916b).
** Colura norrisii Pócs, Chenia 11: 29, 2013 (Pócs 2013).
World checklist of hornworts and liverworts
343
*** Colura obesa Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 273, 1953 [1954] (JovetAst 1953).
*** Colura pallida Steph., Sp. Hepat. (Stephani) 5: 941, 1916 (Stephani 1916b).
*** Colura pluridentata Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 265, 1953 [1954]
(Jovet-Ast 1953).
*** Colura speciosa Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 307, 1953 [1954]
(Jovet-Ast 1953).
** Colura streimannii Pócs, Polish Bot. J. 60 (1): 7, 2015 (Pócs 2015b).
*** Colura superba (Mont.) Steph., Sp. Hepat. (Stephani) 5: 941, 1916 (Stephani
1916b). Bas.: Lejeunea superba Mont., Ann. Sci. Nat. Bot. (sér. 3) 10: 115, 1848
(Montagne 1848).
*** Colura thomeensis Pócs, Bryologist 114 (2): 363, 2011 (Pócs 2011b).
*** Colura tortifolia (Nees et Mont.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 402, 1877 (Trevisan 1877). Bas.: Lejeunea tortifolia Nees et
Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 265, 1843 (Montagne 1843).
*** Colura tutuilana (Pearson) H.A.Mill., Phytologia 47 (4): 322, 1981 (Miller 1981).
Bas.: Colurolejeunea tutuilana Pearson, Amer. Samoa: 151, 1924 (Pearson 1924a).
*** Colura ulei Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 270, 1953 [1954] (Jovet-Ast
1953).
*** Colura vitiensis Pócs et J.Eggers, Polish Bot. J. 52 (2): 88, 2007 (Pócs and Eggers 2007).
** sect. Oidocorys Jovet-Ast ex Grolle, J. Hattori Bot. Lab. 32: 140, 1969 (Grolle 1969c).
*** Colura itatyana Steph., Sp. Hepat. (Stephani) 5: 932, 1916 (Stephani 1916b).
*** Colura naumannii (Schifn.) Steph., Sp. Hepat. (Stephani) 5: 935, 1916 (Stephani 1916b). Bas.: Colurolejeunea naumannii Schifn., Hepat. (Engl.-Prantl): 121,
1893 (Schifner 1893b).
*** Colura ornithocephala Herzog, Svensk Bot. Tidskr. 46 (1): 107, 1952 (Herzog 1952e).
*** Colura pulcherrima Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 235, 1953 [1954]
(Jovet-Ast 1953).
** Colura pulcherrima var. bartlettii Jovet-Ast, Cryptog. Bryol. Lichénol. 1 (3): 278,
1980 (Jovet-Ast 1980).
*** Colura saccophylla E.A.Hodgs. et Herzog, Trans. & Proc. Roy. Soc. New Zealand
77 (2): 253, 1949 (Herzog 1949a).
*** Colura schusteri Grolle, J. Hattori Bot. Lab. 32: 140, 1969 (Grolle 1969c).
** subg. Glotta Grolle et R.L.Zhu, J. Hattori Bot. Lab. 92: 187, 2002 (Grolle and
Zhu 2002).
** sect. Glotta Grolle et R.L.Zhu, J. Hattori Bot. Lab. 92: 187, 2002 (Grolle and
Zhu 2002).
*** Colura bicornis Jovet-Ast, Rev. Bryol. Lichénol. 25 (3/4): 272, 1956 [1957]
(Jovet-Ast 1956).
*** Colura bisvoluta Herzog et Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 228, 1953
[1954] (Jovet-Ast 1953).
344
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Colura inornata Jovet-Ast, Rev. Bryol. Lichénol. 25 (3/4): 275, 1956 [1957]
(Jovet-Ast 1956).
*** Colura karstenii K.I.Goebel, Planzenbiol. Schilderungen 2 (1): 153, 1891 (Goebel
1891).
*** Colura queenslandica B.M.hiers, Brittonia 39 (2): 175, 1987 (hiers 1987b).
*** Colura saroltae Pócs, J. Bryol. 14 (3): 497, 1987 (Jones and Pócs 1987).
*** Colura strophiolata Jovet-Ast, Rev. Bryol. Lichénol. 42 (4): 917, 1976 (Jovet-Ast 1976).
*** Colura usambarica E.W.Jones, J. Bryol. 14 (3): 495, 1987 (Jones and Pócs 1987).
** sect. Heterophyllum Jovet-Ast, Cryptog. Bryol. Lichénol. 4 (3): 213, 1983 (JovetAst 1983).
*** Colura acroloba (Prantl) Jovet-Ast, Rev. Bryol. Lichénol. 22 (2/3): 297, 1953
[1954] (Jovet-Ast 1953). Bas.: Lejeunea acroloba Prantl, Hedwigia 29: xiv, 1890
(Prantl 1890).
*** Colura corniantha Grolle, J. Hattori Bot. Lab. 28: 44, 1965 (Grolle 1965b). Nom.
nov. pro Colura cornuta Jovet-Ast, Rev. Bryol. Lichénol. 30 (1/2): 11, 1961 (Jovet-Ast
1961), nom. illeg.
** Colura denticulata Jovet-Ast, Rev. Bryol. Lichénol. 23 (1/2): 2, 1954 (Jovet-Ast 1954).
*** Colura galeata Jovet-Ast, Rev. Bryol. Lichénol. 35 (1/4): 146, 1967 [1968] (JovetAst 1967a).
** Colura imperfecta Steph., Sp. Hepat. (Stephani) 5: 938, 1916 (Stephani 1916b).
*** Colura siamensis Jovet-Ast, Rev. Bryol. Lichénol. 35 (1/4): 139, 1967 [1968]
(Jovet-Ast 1967b).
*** Colura tixieri Jovet-Ast, Rev. Bryol. Lichénol. 30 (1/2): 9, 1961 (Jovet-Ast 1961).
*** Colura vietnamensis Jovet-Ast et Tixier, Rev. Bryol. Lichénol. 27 (3/4): 205, 1958
[1959] (Jovet-Ast and Tixier 1958).
Incertae sedis
** Colura amboinensis Steph., Sp. Hepat. (Stephani) 5: 935, 1916 (Stephani 1916b).
*** Colura hirta Steph., Sp. Hepat. (Stephani) 5: 932, 1916 (Stephani 1916b).
*** Diplasiolejeunea (Spruce) Schifn., Hepat. (Engl.-Prantl): 121, 1893 (Schifner
1893b). Bas.: Lejeunea subg. Diplasiolejeunea Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 301, 1884 (Spruce 1884). 289
*** Diplasiolejeunea utriculata Steph., Sp. Hepat. (Stephani) 5: 920, 1916 (Stephani 1916b).
** subg. Austrolejeuneopsis R.M.Schust., Bull. Torrey Bot. Club 97 (6): 349, 1970
(Schuster 1970b).
*** Diplasiolejeunea alata Jovet-Ast, Rev. Bryol. Lichénol. 17 (1/4): 31, 1948 [1949]
(Jovet-Ast 1948).
289 Diplasiolejeunea is here only provisionally accepted with subgenera following Schuster (1970b), but
they are not supported by the molecular phylogeny by Dong et al. (2012) where the type species of
subg. Austrolejeuneopsis and Physolejeunea are nested in subg. Diplasiolejeunea.
World checklist of hornworts and liverworts
345
*** Diplasiolejeunea eggersii Pócs, Bryologist 109 (3): 408, 2006 (Pócs 2006b).
*** Diplasiolejeunea erostrata Schäf.-Verw., Cryptog. Bryol. 25 (1): 3, 2004 (SchäferVerwimp 2004).
* Diplasiolejeunea guadalupensis Steph., Sp. Hepat. (Stephani) 5: 924, 1916 (Stephani
1916b). 290
* Diplasiolejeunea heimii Jovet-Ast, Rev. Bryol. Lichénol. 29 (1/2): 39, 1960 (Jovet-Ast 1960). 291
** Diplasiolejeunea involuta S.Winkl., Rev. Bryol. Lichénol. 35 (1/4): 320, 1967
[1968] (Winkler 1967).
** Diplasiolejeunea involuta subsp. andicola Pócs, Cryptog. Bryol. Lichénol. 19 (1):
12, 1998 (León et al. 1998).
*** Diplasiolejeunea johnsonii A.Evans, Bull. Torrey Bot. Club 39 (12): 603, 1912
[1913] (Evans 1912d).
** Diplasiolejeunea johnsonii var. mexicana Jovet-Ast, Rev. Bryol. Lichénol. 29 (1/2):
39, 1960 (Jovet-Ast 1960).
* Diplasiolejeunea montecristensis S.Winkl., Rev. Bryol. Lichénol. 35 (1/4): 321,
1967 [1968] (Winkler 1967). 292
*** Diplasiolejeunea papilionacea R.M.Schust., Phytologia 39 (6): 431, 1978 (Schuster
1978b).
*** Diplasiolejeunea pauckertii (Nees) Steph., Sp. Hepat. (Stephani) 5: 924, 1916 (Stephani 1916b). Bas.: Lejeunea pauckertii Nees, Syn. Hepat. 3: 392, 1845 (Gottsche
et al. 1845b).
*** Diplasiolejeunea pluridentata Schäf.-Verw., Haussknechtia 8: 71, 2001 (SchäferVerwimp 2001b).
*** Diplasiolejeunea replicata (Spruce) Steph., Sp. Hepat. (Stephani) 5: 926, 1916
(Stephani 1916b). Bas.: Lejeunea replicata Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 302, 1884 (Spruce 1884).
** Diplasiolejeunea rudolphiana Steph., Hedwigia 35 (3): 79, 1896 (Stephani 1896b). 293
*** Diplasiolejeunea runssorensis Steph., Bot. Jahrb. Syst. 20 (3): 318, 1895 (Stephani
1895a).
*** Diplasiolejeunea unidentata (Lehm. et Lindenb.) Schifn., Bot. Jahrb. Syst. 23 (5):
583, 1897 (Schifner 1897). Bas.: Jungermannia unidentata Lehm. et Lindenb.,
Nov. Stirp. Pug. 6: 48, 1834 (Lehmann 1834).
** subg. Diplasiolejeunea
*** Diplasiolejeunea armatiloba Steph., Hedwigia 35 (3): 80, 1896 (Stephani 1896b).
290 Diplasiolejeunea guadalupensis is a poorly known taxon with some similarities to the Diplasiolejeunea
replicata species complex (Schäfer-Verwimp and Reiner-Drehwald 2009).
291 Diplasiolejeunea heimii is possibly conspeciic with Diplasiolejeunea replicata (Schäfer-Verwimp and
Pócs 2009).
292 Diplasiolejeunea montecristensis is possibly conspeciic with Diplasiolejeunea replicata.
293 Diplasiolejeunea rudolphiana and Diplasiolejeunea unidentata are genetically very close and may be
conspeciic (Dong et al. 2012).
346
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Diplasiolejeunea borhidiana Reyes Montoya, Acta Bot. Acad. Sci. Hung. 28 (1/2):
177, 1982 [1983] (Reyes 1982).
*** Diplasiolejeunea brunnea Steph., Sp. Hepat. (Stephani) 5: 922, 1916 (Stephani
1916b).
** Diplasiolejeunea caribea Tixier, Bryophyt. Biblioth. 27: 377, 1985 (Tixier 1985a).
*** Diplasiolejeunea cavifolia Steph., Sp. Hepat. (Stephani) 5: 918, 1916 (Stephani
1916b). Nom. nov. pro Lejeunea cavifolia Steph., Bot. Jahrb. Syst. 8 (2): 89, 1886
(Stephani 1886d), nom. illeg.
*** Diplasiolejeunea cobrensis Steph., Sp. Hepat. (Stephani) 5: 923, 1916 (Stephani 1916b).
** Diplasiolejeunea cobrensis subsp. antsirananae Pócs, Candollea 56 (1): 72, 2001
(Pócs 2001).
** Diplasiolejeunea cogoensis M.Infante, Heras et Pócs, Trop. Bryol. 17: 9, 1999 (Infante et al. 1999).
** Diplasiolejeunea cubensis Tixier, Bryophyt. Biblioth. 27: 389, 1985 (Tixier 1985a).
* Diplasiolejeunea glaziovii Tixier, Bryophyt. Biblioth. 27: 397, 1985 (Tixier 1985a). 294
*** Diplasiolejeunea ingekarolae Schäf.-Verw., Herzogia 19: 239, 2006 (Schäfer-Verwimp 2006).
** Diplasiolejeunea jonesii Tixier, Acta Bot. Hung. 30 (1/2): 21, 1984 (Tixier 1984).
*** Diplasiolejeunea leiocarpa Jovet-Ast, Rev. Bryol. Lichénol. 16 (1/2): 34, 1947
[1948] (Jovet-Ast 1947b).
*** Diplasiolejeunea malleiformis (A.Evans) Tixier, Bryophyt. Biblioth. 27: 351, 1985
(Tixier 1985a). Bas.: Diplasiolejeunea pellucida var. malleiformis A.Evans, Bull.
Torrey Bot. Club 39 (5): 215, 1912 (Evans 1912c).
** Diplasiolejeunea palustrium Tixier, Acta Bot. Hung. 30 (1/2): 19, 1984 (Tixier 1984).
*** Diplasiolejeunea pellucida (C.F.W.Meissn. ex Spreng.) Schifn., Hepat. (Engl.Prantl): 121, 1893 (Schifner 1893b). Bas.: Jungermannia pellucida C.F.W.Meissn.
ex Spreng. Syst. Veg. (ed. 16) [Sprengel] 4 (2): 325, 1827 (Sprengel 1827b).
** Diplasiolejeunea phyllarthronii Tixier, Acta Bot. Hung. 30 (1/2): 19, 1984 (Tixier 1984).
*** Diplasiolejeunea pocsii Reyes Montoya, Acta Bot. Acad. Sci. Hung. 28 (1/2): 173,
1982 [1983] (Reyes 1982).
** subg. Physolejeunea R.M.Schust., Bull. Torrey Bot. Club 97 (6): 348, 1970
(Schuster 1970b).
*** Diplasiolejeunea andringitrae Schäf.-Verw., Cryptog. Bryol. 27 (4): 447, 2006
(Pócs and Schäfer-Verwimp 2006).
*** Diplasiolejeunea cornuta Steph., Sp. Hepat. (Stephani) 5: 918, 1916 (Stephani 1916b).
*** Diplasiolejeunea jovet-astiae Grolle, Feddes Repert. 73 (2): 84, 1966 (Grolle 1966b).
*** Diplasiolejeunea kraussiana (Lindenb.) Steph., Sp. Hepat. (Stephani) 5: 919, 1916
(Stephani 1916b). Bas.: Lejeunea kraussiana Lindenb., Syn. Hepat. 3: 393, 1845
(Gottsche et al. 1845b).
*** Diplasiolejeunea ornata Pócs et Schäf.-Verw., Cryptog. Bryol. 27 (4): 440, 2006
(Pócs and Schäfer-Verwimp 2006).
294 Diplasiolejeunea glaziovii may be conspeciic with Diplasiolejeunea cavifolia (Dong et al. 2012).
World checklist of hornworts and liverworts
347
*** Diplasiolejeunea patelligera Herzog, Svensk Bot. Tidskr. 42 (3): 240, 1948 (Herzog
1948).
*** Diplasiolejeunea plicatiloba (Hook.f. et Taylor) Grolle, J. Hattori Bot. Lab. 45:
175, 1979 (Grolle 1979c). Bas.: Jungermannia plicatiloba Hook.f. et Taylor, London J. Bot. 4: 92, 1845 (Hooker and Taylor 1845).
*** Diplasiolejeunea pusilla Grolle, Feddes Repert. 86 (1/2): 79, 1975 (Grolle 1975a).
*** Diplasiolejeunea ranomafanae Pócs, Cryptog. Bryol. 27 (4): 444, 2006 (Pócs and
Schäfer-Verwimp 2006).
*** Diplasiolejeunea villaumei Steph., Sp. Hepat. (Stephani) 5: 921, 1916 (Stephani 1916b).
*** Diplasiolejeunea zakiae Tixier, Lindbergia 4 (1/2): 123, 1977 (Tixier 1977b).
***
***
**
***
**
*
**
***
**
*
**
***
***
**
**
*
***
Incertae sedis
Diplasiolejeunea albifolia (Taylor) E.W.Jones, Trans. Brit. Bryol. Soc. 2 (3): 393,
1954 (Jones 1954a). Bas.: Lejeunea albifolia Taylor, London J. Bot. 5: 399, 1846
(Taylor 1846b).
Diplasiolejeunea aulae E.W.Jones, J. Bryol. 7 (4): 552, 1973 [1974] (Jones 1973).
Diplasiolejeunea auriculata Tixier, Rev. Bryol. Lichénol. 45 (2): 210, 1979 (Tixier
1979c).
Diplasiolejeunea buckii Grolle, Beitr. Phytotax. 15: 105, 1992 (Grolle 1992a).
Diplasiolejeunea columbica Tixier, Cryptog. Bryol. Lichénol. 4 (3): 233, 1983
(Tixier 1983a).
Diplasiolejeunea comorensis Tixier, Rev. Bryol. Lichénol. 45 (2): 213, 1979 (Tixier
1979c).
Diplasiolejeunea cyanguguensis Tixier, Trop. Bryol. 11: 67, 1995 (Tixier 1995b).
Diplasiolejeunea deslooveri Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 47 (1/2):
217, 1977 (Vanden Berghen 1977).
Diplasiolejeunea ensifera Tixier, Rev. Bryol. Lichénol. 45 (2): 215, 1979 (Tixier 1979c).
Diplasiolejeunea evansii Tixier, Bryophyt. Biblioth. 27: 360, 1985 (Tixier 1985a). 295
Diplasiolejeunea gradsteinii Tixier, Trop. Bryol. 11: 67, 1995 (Tixier 1995b).
Diplasiolejeunea grandirostrata Schäf.-Verw., Cryptog. Bryol. 25 (1): 7, 2004
(Schäfer-Verwimp 2004).
Diplasiolejeunea grolleana Reyes Montoya, Acta Bot. Acad. Sci. Hung. 28 (1/2):
175, 1982 [1983] (Reyes 1982).
Diplasiolejeunea hamata Tixier, Rev. Bryol. Lichénol. 45 (2): 217, 1979 (Tixier
1979c).
Diplasiolejeunea insignis Tixier, Lindbergia 4 (1/2): 120, 1977 (Tixier 1977b).
Diplasiolejeunea integerrima Tixier, Rev. Bryol. Lichénol. 45 (2): 219, 1979 (Tixier
1979c). 296
Diplasiolejeunea lanceolata Grolle, Beitr. Phytotax. 15: 107, 1992 (Grolle 1992a).
295 Diplasiolejeunea evansii is a doubtful taxon and the type specimen could not be found in PC (SchäferVerwimp 2004).
296 Diplasiolejeunea integerrima is a juvenile Acrolejeunea species, but the type is too poor to allow identiication (Grolle 1995).
348
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Diplasiolejeunea latipuensis Tixier, Cryptog. Bryol. Lichénol. 16 (3): 230, 1995
(Tixier 1995a). Based on: Diplasiolejeunea latipuensis Tixier, Candollea 46 (2):
294, 1991 (Tixier 1991), nom. inval.
** Diplasiolejeunea lemuriana Tixier, Lindbergia 4 (1/2): 120, 1977 (Tixier 1977b).
** Diplasiolejeunea longilobula Herzog, Trans. Brit. Bryol. Soc. 1 (4): 325, 1950
(Herzog 1950a).
** Diplasiolejeunea magnistipula Tixier, Ann. Fac. Sci. Yaoundé 20: 7, 1975 (Tixier 1975b).
*** Diplasiolejeunea mayaykuensis Schäf.-Verw. et Heinrichs, Polish Bot. J. 58 (1):
143, 2013 (Schäfer-Verwimp et al. 2013).
*** Diplasiolejeunea onraedtii Grolle, Feddes Repert. 89 (5/6): 301, 1978 (Grolle 1978a).
** Diplasiolejeunea ramicola Tixier, Rev. Bryol. Lichénol. 45 (2): 222, 1979 (Tixier 1979c).
** Diplasiolejeunea riclefgrollei Schäf.-Verw., Cryptog. Bryol. 26 (1): 37, 2005
(Schäfer-Verwimp 2005).
** Diplasiolejeunea subcornuta Tixier, Lindbergia 4 (1/2): 122, 1977 (Tixier 1977b).
*** Diplasiolejeunea symoensii Vanden Berghen, Bull. Soc. Roy. Bot. Belgique 92: 126,
1960 (Vanden Berghen 1960b).
** Haplolejeunea Grolle, J. Hattori Bot. Lab. 39: 205, 1975 (Grolle 1975b).
*** Haplolejeunea cucullata (Steph.) Grolle, J. Hattori Bot. Lab. 45: 176, 1979 (Grolle
1979c). Bas.: Cheilolejeunea cucullata Steph., Sp. Hepat. (Stephani) 5: 644, 1914
(Stephani 1914b).
** Haplolejeunea sticta Grolle, J. Hattori Bot. Lab. 39: 205, 1975 (Grolle 1975b).
** Macrocolura R.M.Schust., J. Hattori Bot. Lab. 75: 233, 1994 (Schuster 1994).
*** Macrocolura sagittistipula (Spruce) R.M.Schust., J. Hattori Bot. Lab. 75: 233,
1994 (Schuster 1994). Bas.: Lejeunea sagittistipula Spruce, Trans. & Proc. Bot.
Soc. Edinburgh 15: 304, 1884 (Spruce 1884).
*** Myriocoleopsis Schifn., Hedwigia 81 (5/6): 234, 1944 (Schifner 1944).
*** Myriocoleopsis luviatilis (Steph.) M.E.Reiner et Gradst., J. Bryol. 19 (3): 639,
1997 (Reiner-Drehwald and Gradstein 1997). Bas.: Cololejeunea luviatilis Steph.,
Hedwigia 34 (5): 248, 1895 (Stephani 1895b).
*** Myriocoleopsis gymnocolea (Spruce) M.E.Reiner et Gradst., J. Bryol. 19 (3): 640,
1997 (Reiner-Drehwald and Gradstein 1997). Bas.: Lejeunea gymnocolea Spruce,
Trans. & Proc. Bot. Soc. Edinburgh 15: 296, 1884 (Spruce 1884).
*** Myriocoleopsis minutissima (Sm.) R.L.Zhu, Y.Yu et Pócs, Phytotaxa 183 (4): 293,
2014 (Yu et al. 2014). Bas.: Jungermannia minutissima Sm., Engl. Bot. 23: tab.
1633, 1806 (Smith and Sowerby 1806).
*** Myriocoleopsis minutissima subsp. myriocarpa (Nees et Mont.) R.L.Zhu, Y.Yu et
Pócs, Phytotaxa 183 (4): 294, 2014 (Yu et al. 2014). Bas.: Lejeunea myriocarpa
Nees et Mont., Hist. Phys. Cuba, Bot., Pl. Cell.: 473, 1842 (Montagne 1842a).
World checklist of hornworts and liverworts
349
** Myriocoleopsis vuquangensis (Pócs et Ninh) Pócs, Trop. Bryol. 31: 124, 2010 (Pócs
2010a). Bas.: Cololejeunea vuquangensis Pócs et Ninh, Acta Bot. Hung. 47 (1/2):
156, 2005 (Pócs and Ninh 2005).
** Nephelolejeunea Grolle, J. Hattori Bot. Lab. 37: 252, 1973 (Grolle 1973b).
** Nephelolejeunea bidentata B.M.hiers ex L.Söderstr. et A.Hagborg, Phytotaxa 202
(1): 65, 2015 (Pócs et al. 2015c). Based on: Austrolejeunea bidentata B.M.hiers,
Bryologist 88 (4): 350, 1985 [1986] (hiers 1985), nom. inval.
** Nephelolejeunea carcharias M.A.M.Renner, Syst. Bot. 34 (4): 621, 2009 (Renner et
al. 2009).
*** Nephelolejeunea conchophylla Grolle, J. Hattori Bot. Lab. 48: 164, 1980 (Grolle 1980a).
** Nephelolejeunea fragilis (R.M.Schust.) L.Söderstr. et A.Hagborg, Phytotaxa 202 (1):
65, 2015 (Pócs et al. 2015c). Bas.: Cololejeunea fragilis R.M.Schust., Phytologia 56
(7): 458, 1985 (Schuster 1985c).
*** Nephelolejeunea hamata Grolle, J. Hattori Bot. Lab. 48: 167, 1980 (Grolle 1980a).
*** Nephelolejeunea hispida R.M.Schust. ex L.Söderstr. et A.Hagborg, Phytotaxa 202
(1): 66, 2015 (Pócs et al. 2015c). Based on: Austrolejeunea hispida R.M.Schust.,
Phytologia 47 (4): 305, 1981 (Schuster 1981a), nom. inval.
*** Nephelolejeunea jarmaniana Grolle ex L.Söderstr. et A.Hagborg, Phytotaxa 202
(1): 66, 2015 (Pócs et al. 2015c). Based on: Austrolejeunea jarmaniana Grolle,
Nova Hedwigia 55 (1/2): 112, 1992 (Grolle 1992b), nom. inval.
*** Nephelolejeunea nudipes (Hook.f. et Taylor) L.Söderstr. et A.Hagborg, Phytotaxa
202 (1): 66, 2015 (Pócs et al. 2015c). Bas.: Jungermannia nudipes Hook.f. et Taylor, London J. Bot. 3: 568, 1844 (Hooker and Taylor 1844d).
** Nephelolejeunea occidentalis Pócs ex L.Söderstr. et A.Hagborg, Phytotaxa 202 (1):
66, 2015 (Pócs et al. 2015c). Based on: Austrolejeunea occidentalis Pócs, J. Hattori
Bot. Lab. 99: 187, 2006 (Pócs 2006d), nom. inval.
*** Nephelolejeunea papillosa Glenny, New Zealand J. Bot. 34 (2): 195, 1996 (Glenny
1996).
*** Nephelolejeunea radulifolia (C.Massal.) L.Söderstr. et A.Hagborg, Phytotaxa 202
(1): 66, 2015 (Pócs et al. 2015c). Bas.: Lejeunea radulifolia C.Massal., Nuovo
Giorn. Bot. Ital. 17 (3): 248, 1885 (Massalongo 1885).
** Nephelolejeunea secunda M.A.M.Renner ex L.Söderstr. et A.Hagborg, Phytotaxa 202 (1): 66, 2015 (Pócs et al. 2015c). Based on: Austrolejeunea secunda
M.A.M.Renner, Bryologist 113 (4): 782, 2010 (Renner 2010), nom. inval.
*** Nephelolejeunea talinayi (S.W.Arnell) Grolle, J. Hattori Bot. Lab. 37: 253, 1973
(Grolle 1973b). Bas.: Harpalejeunea talinayi S.W.Arnell, Svensk Bot. Tidskr. 50
(2): 309, 1956 (Arnell 1956d).
** Schusterolejeunea Grolle, J. Bryol. 11 (1): 105, 1980 (Grolle 1980b). Nom. nov.
pro Cladocolea R.M.Schust., Beih. Nova Hedwigia 9: 155, 1963 (Schuster 1963a).
350
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Schusterolejeunea inundata (Spruce) Grolle, J. Bryol. 11 (1): 105, 1980 (Grolle
1980b). Bas.: Lejeunea inundata Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:
278, 1884 (Spruce 1884).
** Siphonolejeunea Herzog, Nat. Hist. Juan Fernandez (Botany) 2 (5): 744, 1942
(Herzog 1942a).
** Siphonolejeunea elegantissima (Steph.) Grolle, J. Hattori Bot. Lab. 41: 405, 1976
(Grolle 1976b). Bas.: Trachylejeunea elegantissima Steph., Hedwigia 28 (4): 262,
1889 (Stephani 1889c).
** Siphonolejeunea neesii (Mont.) Bischl., Nova Hedwigia 17: 338, 1969 (Bischler
1969). Bas.: Lejeunea neesii Mont., Ann. Sci. Nat. Bot. (sér. 2) 5: 62, 1836 (Nees
and Montagne 1836).
*** Siphonolejeunea schifneri (Schifn.) Herzog, Svensk Bot. Tidskr. 42 (3): 230, 1948
(Herzog 1948). Bas.: Pycnolejeunea schifneri Schifn., Consp. Hepat. Arch. Ind.:
260, 1898 (Schifner 1898b).
** Tuyamaella S.Hatt., J. Hattori Bot. Lab. 5: 60, 1951 (Hattori 1951d).
** Tuyamaella angulistipa (Steph.) R.M.Schust. et Kachroo, J. Linn. Soc., Bot. 56
(368): 508, 1961 (Kachroo and Schuster 1961). Bas.: Pycnolejeunea angulistipa
Steph., Hedwigia 35 (3): 123, 1896 (Stephani 1896b).
** Tuyamaella borneensis Tixier, Rev. Bryol. Lichénol. 39 (2): 241, 1973 (Tixier
1973a).
** Tuyamaella hattorii Tixier, Rev. Bryol. Lichénol. 31 (3/4): 188, 1962 [1963] (Tixier
1962).
** Tuyamaella jackii (Steph.) Tixier, Rev. Bryol. Lichénol. 39 (2): 235, 1973 (Tixier
1973a). Bas.: Pycnolejeunea jackii Steph., Sp. Hepat. (Stephani) 5: 611, 1914 (Stephani 1914b).
*** Tuyamaella molischii (Schifn.) S.Hatt., J. Hattori Bot. Lab. 5: 62, 1951 (Hattori
1951d). Bas.: Pycnolejeunea molischii Schifn., Ann. Bryol. 2: 97, 1929 (Schifner
1929).
** Tuyamaella molischii var. brevistipa P.C.Wu et P.J.Lin, Acta Phytotax. Sin. 16 (2):
65, 1978 (Wu and Lin 1978).
** Tuyamaella molischii var. taiwanensis R.L.Zhu et M.L.So, Nova Hedwigia 70
(1/2): 190, 2000 (Zhu and So 2000).
** Tuyamaella serratistipa S.Hatt., Bot. Mag. (Tokyo) 64 (755/756): 118, 1951 (Hattori 1951c).
**
subtrib. Cyclolejeuneinae Gradst.
*** Bromeliophila R.M.Schust., J. Hattori Bot. Lab. 75: 226, 1994 (Schuster 1994).
*** Bromeliophila helenae Gradst., Cryptog. Bryol. Lichénol. 18 (3): 218, 1997 (Gradstein 1997).
World checklist of hornworts and liverworts
351
*** Bromeliophila natans (Steph.) R.M.Schust., J. Hattori Bot. Lab. 75: 226, 1994
(Schuster 1994). Bas.: Peltolejeunea natans Steph., Hedwigia 44 (4): 228, 1905
(Stephani 1905a).
*** Cyclolejeunea A.Evans, Bull. Torrey Bot. Club 31 (4): 193, 1904 (Evans 1904a).
*** Cyclolejeunea accedens (Gottsche) A.Evans, Bull. Torrey Bot. Club 31 (4): 201,
1904 (Evans 1904a). Bas.: Lejeunea accedens Gottsche, Syn. Hepat. 3: 339, 1845
(Gottsche et al. 1845b).
*** Cyclolejeunea chitonia (Taylor) A.Evans, Bull. Torrey Bot. Club 31 (4): 194,
1904 (Evans 1904a). Bas.: Lejeunea chitonia Taylor, Nov. Stirp. Pug. 8: 27, 1844
(Lehmann 1844).
*** Cyclolejeunea convexistipa (Lehm. et Lindenb.) A.Evans, Bull. Torrey Bot. Club 31
(4): 198, 1904 (Evans 1904a). Bas.: Jungermannia convexistipa Lehm. et Lindenb.,
Nov. Stirp. Pug. 6: 43, 1834 (Lehmann 1834).
* Cyclolejeunea ecuadorensis Steph., Sp. Hepat. (Stephani) 5: 194, 1913 (Stephani 1913a).
*** Cyclolejeunea foliorum (Nees) Grolle, J. Hattori Bot. Lab. 65: 403, 1988 (Grolle 1988c).
Bas.: Lejeunea foliorum Nees, Syn. Hepat. 3: 326, 1845 (Gottsche et al. 1845b).
* Cyclolejeunea integerrima (Steph.) Steph., Sp. Hepat. (Stephani) 5: 195, 1913
(Stephani 1913a). Bas.: Odontolejeunea integerrima Steph., Hedwigia 44 (4): 227,
1905 (Stephani 1905a).
*** Cyclolejeunea luteola (Spruce) Grolle, Wiss. Z. Friedrich-Schiller-Univ. Jena,
Math.-Naturwiss. Reihe 33: 761, 1984 (Grolle 1984b). Bas.: Lejeunea luteola
Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 205, 1884 (Spruce 1884).
*** Cyclolejeunea peruviana (Lehm. et Lindenb.) A.Evans, Bull. Torrey Bot. Club 31
(4): 196, 1904 (Evans 1904a). Bas.: Jungermannia peruviana Lehm. et Lindenb.,
Nov. Stirp. Pug. 5: 18, 1833 (Lehmann 1833).
* Cyclolejeunea spectabilis Steph., Sp. Hepat. (Stephani) 5: 193, 1913 (Stephani 1913a).
*** Prionolejeunea (Spruce) Schifn., Hepat. (Engl.-Prantl): 127, 1893 (Schifner
1893b). Bas.: Lejeunea subg. Prionolejeunea Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 152, 1884 (Spruce 1884).
*** Prionolejeunea aemula (Gottsche) A.Evans, Bull. Torrey Bot. Club 31 (4): 219,
1904 (Evans 1904a). Bas.: Lejeunea aemula Gottsche, Syn. Hepat. 3: 338, 1845
(Gottsche et al. 1845b).
*** Prionolejeunea ampliretis Herzog, Feddes Repert. Spec. Nov. Regni Veg. 57 (1/2):
178, 1955 (Herzog 1955).
*** Prionolejeunea arguta (Nees) Steph., Sp. Hepat. (Stephani) 5: 204, 1913 (Stephani
1913a). Bas.: Lejeunea arguta Nees, Syn. Hepat. 3: 338, 1845 (Gottsche et al. 1845b).
*** Prionolejeunea decora (Taylor) Steph., Sp. Hepat. (Stephani) 5: 207, 1913 (Stephani
1913a). Bas.: Lejeunea decora Taylor, London J. Bot. 5: 393, 1846 (Taylor 1846b).
*** Prionolejeunea denticulata (F.Weber) Schifn., Hepat. (Engl.-Prantl): 127, 1893
(Schifner 1893b). Bas.: Jungermannia denticulata F.Weber, Hist. Musc. Hepat.
Prodr.: 30, 1815 (Weber 1815).
352
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Prionolejeunea diversitexta (Hampe et Gottsche) Steph., Sp. Hepat. (Stephani) 5:
209, 1913 (Stephani 1913a). Bas.: Lejeunea diversitexta Hampe et Gottsche, Linnaea 25 (3): 357, 1852 [1853] (Hampe and Gottsche 1852).
*** Prionolejeunea exauriculata A.Evans, Bull. Torrey Bot. Club 31 (4): 223, 1904
(Evans 1904a).
*** Prionolejeunea galliotii Steph., Sp. Hepat. (Stephani) 6: 388, 1923 (Stephani
1923).
*** Prionolejeunea grata (Gottsche) Schifn., Hepat. (Engl.-Prantl): 127, 1893
(Schifner 1893b). Bas.: Lejeunea grata Gottsche, Abh. Naturwiss. Vereins Bremen
7: 359, 1882 (Gottsche 1882).
*** Prionolejeunea grollei Ilk.-Borg. et Schäf.-Verw., Cryptog. Bryol. 26 (1): 29, 2005
(Ilkiu-Borges and Schäfer-Verwimp 2005).
*** Prionolejeunea guadalupensis (Lindenb.) Steph., Sp. Hepat. (Stephani) 5: 211,
1913 (Stephani 1913a). Bas.: Lejeunea guadalupensis Lindenb., Syn. Hepat. 3:
340, 1845 (Gottsche et al. 1845b).
*** Prionolejeunea limpida Herzog, Hedwigia 67 (6): 252, 1927 (Herzog 1927).
*** Prionolejeunea magnistipula Herzog, Feddes Repert. Spec. Nov. Regni Veg. 57
(1/2): 176, 1955 (Herzog 1955).
*** Prionolejeunea meissneri (Gottsche) Steph., Sp. Hepat. (Stephani) 5: 212, 1913
(Stephani 1913a). Bas.: Lejeunea meissneri Gottsche, Syn. Hepat. 3: 340, 1845
(Gottsche et al. 1845b).
*** Prionolejeunea mucronata (Sande Lac.) Steph., Hedwigia 35 (3): 119, 1896 (Stephani 1896b). Bas.: Lejeunea mucronata Sande Lac., Syn. hepat. jav.: 106, 1856
[1857] (Sande Lacoste 1856b).
*** Prionolejeunea muricatoserrulata (Spruce) Steph., Sp. Hepat. (Stephani) 5: 223,
1913 (Stephani 1913a). Bas.: Lejeunea muricatoserrulata Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 155, 1884 (Spruce 1884).
*** Prionolejeunea principensis Vanden Berghen, Rev. Bryol. Lichénol. 29 (1/2): 65,
1960 (Vanden Berghen 1960a).
*** Prionolejeunea recurvula (Spruce) Steph., Sp. Hepat. (Stephani) 5: 224, 1913
(Stephani 1913a). Bas.: Lejeunea recurvula Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 155, 1884 (Spruce 1884).
*** Prionolejeunea scaberula (Spruce) Steph., Sp. Hepat. (Stephani) 5: 214, 1913
(Stephani 1913a). Bas.: Lejeunea scaberula Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 159, 1884 (Spruce 1884).
*** Prionolejeunea schlimiana (Gottsche) Steph., Sp. Hepat. (Stephani) 5: 214, 1913
(Stephani 1913a). Bas.: Lejeunea schlimiana Gottsche, Ann. Sci. Nat. Bot. (sér. 5)
1: 154, 1864 (Gottsche 1864).
*** Prionolejeunea trachyodes (Spruce) Steph., Sp. Hepat. (Stephani) 5: 215, 1913
(Stephani 1913a). Bas.: Lejeunea trachyodes Spruce, J. Linn. Soc., Bot. 30 (210):
338, 1895 (Gepp 1895b).
World checklist of hornworts and liverworts
*
*
**
353
Excluded from the genus
Prionolejeunea corbisieri Pearson, Natuurw. Tijdschr. 4 (5/6): 128, 1922 (Pearson
1922a). 297
Prionolejeunea maculata Herzog, Feddes Repert. Spec. Nov. Regni Veg. 57 (1/2):
180, 1955 (Herzog 1955). 298
subtrib. Drepanolejeuneinae Gradst.
** Capillolejeunea S.W.Arnell, Svensk Bot. Tidskr. 59 (1): 69, 1965 (Arnell 1965).
** Capillolejeunea geisslerae (Pócs) R.L.Zhu, Qiong He, Y.M.Wei et Pócs, Phytotaxa
175 (3): 167, 2014 (He et al. 2014a). Bas.: Drepanolejeunea geisslerae Pócs, Candollea 56 (1): 70, 2001 (Pócs 2001).
** Capillolejeunea mascarena S.W.Arnell, Svensk Bot. Tidskr. 59 (1): 69, 1965 (Arnell
1965).
*** Drepanolejeunea (Spruce) Steph., Hedwigia 30 (5): 209, 1891 (Stephani 1891a).
Bas.: Lejeunea subg. Drepanolejeunea Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 186, 1884 (Spruce 1884).
** subg. Acantholejeunea R.M.Schust., Beih. Nova Hedwigia 9: 115, 1963 (Schuster
1963a).
** Drepanolejeunea dentistipula Steph., Sp. Hepat. (Stephani) 5: 343, 1913 (Stephani
1913a).
** Drepanolejeunea spinistipula Herzog, Svensk Bot. Tidskr. 42 (3): 238, 1948 (Herzog 1948).
*** subg. Drepanolejeunea
*** Drepanolejeunea aculeata Bischl., Rev. Bryol. Lichénol. 33 (1/2): 68, 1964 (Bischler
1964).
** Drepanolejeunea anderssonii (Ångstr.) A.Evans, Trans. Connecticut Acad. Arts 10
(8): 429, 1900 (Evans 1900a). Bas.: Lejeunea anderssonii Ångstr., Öfvers. Kongl.
Vetensk.-Akad. Förh. 29 (4): 24, 1872 (Ångström 1872).
*** Drepanolejeunea andina Herzog, Svensk Bot. Tidskr. 51 (1): 196, 1957 (Herzog 1957a).
*** Drepanolejeunea angustifolia (Mitt.) Grolle, J. Jap. Bot. 40 (7): 206, 1965 (Grolle
1965d). Bas.: Lejeunea angustifolia Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 116,
1860 [1861] (Mitten 1860c).
*** Drepanolejeunea ankasica E.W.Jones, Bull. Brit. Mus. (Nat. Hist.), Bot. 11 (3):
245, 1983 (Jones and Harrington 1983).
297 Prionolejeunea corbisieri is a Cheilolejeunea species (Ilkiu-Borges 2006).
298 Prionolejeunea maculata is a Cyclolejeunea species (Ilkiu-Borges 2006).
354
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Drepanolejeunea anoplantha (Spruce) Steph., Sp. Hepat. (Stephani) 5: 325, 1913
(Stephani 1913a). Bas.: Lejeunea anoplantha Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 189, 1884 (Spruce 1884).
*** Drepanolejeunea appalachiana R.M.Schust., J. Elisha Mitchell Sci. Soc. 83 (4):
219, 1967 (Schuster 1967a).
*** Drepanolejeunea araucariae Steph., Hedwigia 35 (3): 80, 1896 (Stephani 1896b).
** Drepanolejeunea araucariae var. chilensis Herzog, Acta Horti Gotob. 15: 161, 1943
(Herzog 1943a).
** Drepanolejeunea aucklandica Steph., Sp. Hepat. (Stephani) 5: 358, 1913 (Stephani
1913a).
*** Drepanolejeunea aurita Bischl., Rev. Bryol. Lichénol. 33 (1/2): 146, 1964 (Bischler
1964).
** Drepanolejeunea bakeri Herzog, Ann. Bryol. 3: 148, 1930 (Herzog 1930b).
*** Drepanolejeunea bidens (Prantl) A.Evans, Bull. Torrey Bot. Club 30 (1): 29, 1903
(Evans 1903b). Bas.: Lejeunea bidens Prantl, Hedwigia 29: xiv, 1890 (Prantl 1890).
*** Drepanolejeunea biocellata A.Evans, Bull. Torrey Bot. Club 30 (1): 22, 1903 (Evans
1903b).
** Drepanolejeunea blumei Steph., Hedwigia 35 (3): 81, 1896 (Stephani 1896b).
** Drepanolejeunea brunnea Mizut., J. Hattori Bot. Lab. 33: 231, 1970 (Mizutani 1970).
** Drepanolejeunea caledonica Steph., Sp. Hepat. (Stephani) 5: 342, 1913 (Stephani
1913a).
*** Drepanolejeunea campanulata (Spruce) Steph., Sp. Hepat. (Stephani) 5: 328, 1913
(Stephani 1913a). Bas.: Lejeunea campanulata Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 192, 1884 (Spruce 1884).
** Drepanolejeunea canceroides H.A.Mill. et Bonner, Beih. Nova Hedwigia 11: 53,
1963 (Miller et al. 1963).
** Drepanolejeunea capulata (Taylor) J.B.Jack et Steph., Hedwigia 31 (1): 13, 1892
(Jack and Stephani 1892). Bas.: Lejeunea capulata Taylor, London J. Bot. 5: 394,
1846 (Taylor 1846b).
** Drepanolejeunea capulata var. lagellifera S.W.Arnell, Österr. Akad. Wiss., Math.Naturwiss. Kl., Denkschr. 111: 100, 1964 (Schifner and Arnell 1964).
** Drepanolejeunea ciliata Mizut., J. Hattori Bot. Lab. 33: 232, 1970 (Mizutani 1970).
*** Drepanolejeunea crassiretis A.Evans, Bull. Torrey Bot. Club 30 (1): 25, 1903 (Evans
1903b).
*** Drepanolejeunea crucianella (Taylor) A.Evans, Bull. Torrey Bot. Club 30 (1): 33,
1903 (Evans 1903b). Bas.: Lejeunea crucianella Taylor, London J. Bot. 5: 393,
1846 (Taylor 1846b).
*** Drepanolejeunea cultrella (Mitt.) Steph., Sp. Hepat. (Stephani) 5: 324, 1913 (Stephani 1913a). Bas.: Lejeunea cultrella Mitt., J. Proc. Linn. Soc., Bot. 7 (27): 168,
1863 (Mitten 1863).
** Drepanolejeunea cutervoensis (Loitl.) Grolle, J. Hattori Bot. Lab. 69: 186, 1991
(Grolle 1991). Bas.: Lejeunea cutervoensis Loitl., Diagn. pl. nov.: 21, 1894 (Loitlesberger 1894).
World checklist of hornworts and liverworts
355
** Drepanolejeunea decurviloba Steph., Sp. Hepat. (Stephani) 6: 397, 1923 (Stephani
1923).
* Drepanolejeunea deslooveri Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 47 (1/2):
210, 1977 (Vanden Berghen 1977). 299
* Drepanolejeunea dissitifolia A.Evans, Bull. Torrey Bot. Club 30 (1): 28, 1903 (Evans
1903b).
** Drepanolejeunea elegans Herzog, Ann. Bryol. 9: 128, 1936 [1937] (Herzog 1936a).
** Drepanolejeunea erecta (Steph.) Mizut., J. Hattori Bot. Lab. 40: 442, 1976 (Mizutani 1976b). Bas.: Leptolejeunea erecta Steph., Bull. Soc. Roy. Bot. Belgique 38 (1):
44, 1899 (Stephani 1899h).
** Drepanolejeunea evansii Bischl. ex L.Söderstr., A.Hagborg et von Konrat, Phytotaxa 65: 47, 2012 (Söderström et al. 2012c). Based on: Drepanolejeunea evansii
Bischl., Rev. Bryol. Lichénol. 33 (1/2): 75, 1964 (Bischler 1964), nom. inval.
*** Drepanolejeunea fragilis Bischl. ex L.Söderstr., A.Hagborg et von Konrat, Phytotaxa 65: 47, 2012 (Söderström et al. 2012c). Based on: Drepanolejeunea fragilis
Bischl., Rev. Bryol. Lichénol. 33 (1/2): 123, 1964 (Bischler 1964), nom. inval.
** Drepanolejeunea fulfordiae L.Söderstr., Phytotaxa 208 (1): 98, 2015 (Pócs et al.
2015a). Nom. nov. pro Drepanolejeunea papillosa Fulford, Mem. New York Bot.
Gard. 23: 843, 1972 (Fulford 1972), nom. illeg.
*** Drepanolejeunea granatensis (Prantl) Bischl., Rev. Bryol. Lichénol. 33 (1/2): 150,
1964 (Bischler 1964). Bas.: Lejeunea granatensis Prantl, Hedwigia 31: xvi, 1892
(Prantl 1892).
** Drepanolejeunea grandis Herzog, Ann. Bryol. 12: 113, 1939 (Herzog 1939a).
*** Drepanolejeunea grollei M.E.Reiner et Schäf.-Verw., Candollea 51 (2): 475, 1996
(Reiner-Drehwald and Schäfer-Verwimp 1996).
** Drepanolejeunea grossidens Steph., Sp. Hepat. (Stephani) 5: 358, 1913 (Stephani
1913a). Based on: Drepanolejeunea grossidens Steph., Hedwigia 28 (3): 168, 1889
(Stephani 1889d), nom. inval.
*** Drepanolejeunea hamatifolia (Hook.) Schifn., Hepat. (Engl.-Prantl): 126, 1893
(Schifner 1893b). Bas.: Jungermannia hamatifolia Hook., Brit. Jungermann.: tab.
51, 1813 (Hooker 1813).
** Drepanolejeunea hamulata Steph., Sp. Hepat. (Stephani) 5: 331, 1913 (Stephani
1913a).
*** Drepanolejeunea helenae Pócs, Cryptog. Bryol. Lichénol. 18 (3): 198, 1997 (Pócs
1997a).
** Drepanolejeunea herzogii R.L.Zhu et M.L.So, Beih. Nova Hedwigia 121: 181,
2001 (Zhu and So 2001). Nom. nov. pro Strepsilejeunea ocellata Herzog, Memoranda Soc. Fauna Fl. Fennica 26: 57, 1950 [1951] (Herzog 1950b).
299 Drepanolejeunea deslooveri was treated as conspeciic with Drepanolejeunea hamatifolia by Tixier
(1995b) and accepted uncritically as such by Wigginton and Grolle (1996) and Wigginton (2002).
However, no published supporting evidence for the synonymy has been found, and it was therefore
reinstated pending further studies (Wigginton 2009).
356
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Drepanolejeunea inchoata (C.F.W.Meissn.) Steph., Primit. l. costar.: 115, 1892
[1893] (Stephani 1892e). Bas.: Jungermannia inchoata C.F.W.Meissn., Nov.
Stirp. Pug. 5: 19, 1833 (Lehmann 1833).
** Drepanolejeunea inchoata var. palmicola Pócs, Acta Bot. Hung. 51 (3/4): 381,
2009 (Schäfer-Verwimp and Pócs 2009).
** Drepanolejeunea inchoata var. roraimae (Steph. ex Zwickel) Bischl., Rev. Bryol. Lichénol. 33 (1/2): 48, 1964 (Bischler 1964). Bas.: Drepanolejeunea roraimae Steph.
ex Zwickel, Ann. Bryol. 6: 121, 1933 (Zwickel 1933).
*** Drepanolejeunea infundibulata (Spruce) A.Evans, Bull. Torrey Bot. Club 30 (1):
35, 1903 (Evans 1903b). Bas.: Lejeunea infundibulata Spruce, Trans. & Proc. Bot.
Soc. Edinburgh 15: 191, 1884 (Spruce 1884).
*** Drepanolejeunea integribracteata Bischl., Rev. Bryol. Lichénol. 33 (1/2): 142, 1964
(Bischler 1964).
** Drepanolejeunea laevis (Mitt.) Steph., Sp. Hepat. (Stephani) 5: 357, 1913
(Stephani 1913a). Bas.: Lejeunea laevis Mitt., Fl. vit.: 416, 1871 [1873] (Mitten
1871).
* Drepanolejeunea lancifolia (Gottsche) J.B.Jack et Steph., Hedwigia 31 (1): 13,
1892 (Jack and Stephani 1892). Bas.: Lejeunea lancifolia Gottsche, Ann. Sci. Nat.
Bot. (sér. 5) 1: 155, 1864 (Gottsche 1864). 300
*** Drepanolejeunea lichenicola (Spruce) Steph., Sp. Hepat. (Stephani) 5: 335, 1913
(Stephani 1913a). Bas.: Lejeunea lichenicola Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 191, 1884 (Spruce 1884).
** Drepanolejeunea longii Grolle et R.L.Zhu, Ann. Bot. Fenn. 36 (2): 115, 1999
(Grolle and Zhu 1999).
** Drepanolejeunea macrodonta (Mitt.) Steph., Sp. Hepat. (Stephani) 5: 357, 1913
(Stephani 1913a). Bas.: Lejeunea macrodonta Mitt., J. Proc. Linn. Soc., Bot. 5
(18): 116, 1860 [1861] (Mitten 1860c).
** Drepanolejeunea mawtmiana Ajit P.Singh et V.Nath, Hepat. Khasi Jaintia Hills: E.
Himal.: 251, 2007 (Singh and Nath 2007b).
** Drepanolejeunea microcarpa Pearson, J. Linn. Soc., Bot. 46 (305): 36, 1922 (Pearson
1922b).
** Drepanolejeunea moluccensis Herzog, Ann. Bryol. 7: 88, 1934 (Herzog 1934b).
*** Drepanolejeunea mosenii (Steph.) Bischl., Rev. Bryol. Lichénol. 35 (1/4): 118,
1967 [1968] (Bischler 1967). Bas.: Leptolejeunea mosenii Steph., Sp. Hepat. (Stephani) 5: 372, 1913 (Stephani 1913a).
* Drepanolejeunea obliqua Steph., Hedwigia 35 (3): 82, 1896 (Stephani 1896b). 301
** Drepanolejeunea obtriangulata T.Kodama, J. Hattori Bot. Lab. 41: 381, 1976 (Kodama 1976).
300 Drepanolejeunea lancifolia resembles Drepanolejeunea bidens and Drepanolejeunea araucariae, but the
type material has not been found (Bischler 1964).
301 Drepanolejeunea obliqua is possibly conspeciic with Drepanolejeunea ternatensis (Söderström et al.
2010a).
World checklist of hornworts and liverworts
357
** Drepanolejeunea obtusifolia T.Yamag., J. Jap. Bot. 59 (11): 332, 1984 (Yamaguchi
1984).
*** Drepanolejeunea orthophylla (Nees et Mont.) Bischl., Rev. Bryol. Lichénol. 35
(1/4): 102, 1967 [1968] (Bischler 1967). Bas.: Lejeunea orthophylla Nees et Mont.,
Ann. Sci. Nat. Bot. (sér. 2) 19: 265, 1843 (Montagne 1843).
*** Drepanolejeunea palmifolia (Nees) Schifn., Hepat. (Engl.-Prantl): 126, 1893
(Schifner 1893b). Bas.: Jungermannia palmifolia Nees, Fl. Bras. (Martius) 1 (1):
366, 1833 (Nees 1833a).
*** Drepanolejeunea pentadactyla (Mont.) Steph., Sp. Hepat. (Stephani) 5: 357, 1913
(Stephani 1913a). Bas.: Lejeunea pentadactyla Mont., Ann. Sci. Nat. Bot. (sér. 3)
10: 113, 1848 (Montagne 1848).
** Drepanolejeunea perissodonta (Spruce) Bischl., Rev. Bryol. Lichénol. 33 (1/2): 73,
1964 (Bischler 1964). Bas.: Lejeunea inchoata var. perissodonta Spruce, J. Linn.
Soc., Bot. 30 (210): 340, 1895 (Gepp 1895b).
*** Drepanolejeunea physifolia (Gottsche) Pearson, Forh. Vidensk.-Selsk. Kristiania
1892 (8): 8, 1892 (Pearson 1892). Bas.: Lejeunea physifolia Gottsche, Abh. Naturwiss. Vereins Bremen 7: 357, 1882 (Gottsche 1882).
** Drepanolejeunea pinnatiloba Schifn., Bot. Jahrb. Syst. 23 (5): 591, 1897 (Schifner
1897).
** Drepanolejeunea pleiodictya Herzog, Ann. Bryol. 7: 89, 1934 (Herzog 1934b).
** Drepanolejeunea propagulifera Herzog, Mitt. Inst. Allg. Bot. Hamburg 7 (3): 204,
1931 (Herzog 1931a).
** Drepanolejeunea pseudoneura (A.Evans) Grolle, J. Hattori Bot. Lab. 65: 405, 1988
(Grolle 1988c). Bas.: Harpalejeunea pseudoneura A.Evans, Trans. Connecticut
Acad. Arts 10 (8): 427, 1900 (Evans 1900a).
** Drepanolejeunea pterocalyx (Herzog) Bischl., Rev. Bryol. Lichénol. 35 (1/4): 114,
1967 [1968] (Bischler 1967). Bas.: Leptolejeunea pterocalyx Herzog, Feddes Repert. Spec. Nov. Regni Veg. 57 (1/2): 187, 1955 (Herzog 1955).
** Drepanolejeunea pungens Bischl., Rev. Bryol. Lichénol. 33 (1/2): 104, 1964
(Bischler 1964).
** Drepanolejeunea ramentilora Steph., Sp. Hepat. (Stephani) 5: 338, 1913 (Stephani
1913a).
* Drepanolejeunea ruandensis Vanden Berghen, Bull. Soc. Roy. Bot. Belgique 93: 63,
1961 (Vanden Berghen 1961). 302
*** Drepanolejeunea senticosa Bischl., Rev. Bryol. Lichénol. 33 (1/2): 96, 1964 (Bischler
1964).
** Drepanolejeunea sikkimensis (Udar et U.S.Awasthi) Grolle, J. Hattori Bot. Lab. 55:
503, 1984 (Grolle 1984a). Bas.: Leptolejeunea sikkimensis Udar et U.S.Awasthi,
Misc. Bryol. Lichenol. 8 (6): 115, 1979 (Udar and Awasthi 1979).
302 Drepanolejeunea ruandensis is conspeciic with Drepanolejeunea cultrella in Vanden Berghen (1972a),
but it was accepted by Tixier (1995b) and Wigginton and Grolle (1996).
358
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Drepanolejeunea spinosa Herzog, Feddes Repert. Spec. Nov. Regni Veg. 57 (1/2):
185, 1955 (Herzog 1955).
** Drepanolejeunea spinosocornuta Steph., Sp. Hepat. (Stephani) 5: 351, 1913
(Stephani 1913a).
* Drepanolejeunea subdissitifolia Herzog, Memoranda Soc. Fauna Fl. Fennica 25: 65,
1950 (Herzog 1950c).
** Drepanolejeunea submuricata R.M.Schust., Phytotaxa 208 (1): 98, 2015 (Pócs et
al. 2015a). Based on: Drepanolejeunea submuricata R.M.Schust., Nova Hedwigia
62 (1/2): 34, 1996 (Schuster 1996c), nom. inval.
** Drepanolejeunea subquadrata (Mitt.) Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 311,
1896 (Stephani 1896a). Bas.: Lejeunea subquadrata Mitt., Fl. vit.: 416, 1871
[1873] (Mitten 1871).
** Drepanolejeunea subvittata (Herzog) Grolle, J. Hattori Bot. Lab. 69: 186, 1991
(Grolle 1991). Bas.: Harpalejeunea subvittata Herzog, Svensk Bot. Tidskr. 51 (1):
195, 1957 (Herzog 1957a).
* Drepanolejeunea tenax K.I.Goebel, Ann. Jard. Bot. Buitenzorg 39: 15, 1928 (Goebel
1928).
** Drepanolejeunea tenera K.I.Goebel, Ann. Jard. Bot. Buitenzorg 39: 20, 1928 (Goebel
1928). 303
* Drepanolejeunea tenera var. litoceras Herzog, Ann. Bryol. 7: 85, 1934 (Herzog
1934b).
*** Drepanolejeunea ternatensis (Gottsche) Schifn., Hepat. (Engl.-Prantl): 126, 1893
(Schifner 1893b). Bas.: Lejeunea ternatensis Gottsche, Syn. Hepat. 3: 346, 1845
(Gottsche et al. 1845b).
** Drepanolejeunea teysmannii (Gottsche) Steph., Hedwigia 35 (3): 84, 1896 (Stephani
1896b). Bas.: Lejeunea teysmannii Gottsche, Abh. Naturwiss. Vereins Bremen 7:
360, 1882 (Gottsche 1882).
** Drepanolejeunea tridactyla (Gottsche) Steph., Sp. Hepat. (Stephani) 5: 354, 1913
(Stephani 1913a). Bas.: Lejeunea tridactyla Gottsche, Syn. Hepat. 3: 347, 1845
(Gottsche et al. 1845b).
*** Drepanolejeunea trigonophylla Steph., Hedwigia 35 (3): 85, 1896 (Stephani 1896b).
** Drepanolejeunea tristaniana S.W.Arnell, Results Norweg. Sci. Exped. Tristan da
Cunha 42: 7, 1958 (Arnell 1958b).
** Drepanolejeunea tuyamae S.Hatt., Bot. Mag. (Tokyo) 64 (755/756): 116, 1951
(Hattori 1951c).
** Drepanolejeunea ualanensis Inoue et H.A.Mill., Bull. Natl. Sci. Mus. Tokyo (n.ser.)
8 (2): 148, 1965 (Inoue and Miller 1965).
** Drepanolejeunea ungulata (Steph.) Grolle, J. Hattori Bot. Lab. 69: 187, 1991
(Grolle 1991). Bas.: Harpalejeunea ungulata Steph., Sp. Hepat. (Stephani) 5: 264,
1913 (Stephani 1913a).
303 Drepanolejeunea tenera is possibly conspeciic with Drepanolejeunea pentadactyla (Söderström et al.
2010a).
World checklist of hornworts and liverworts
359
** Drepanolejeunea urceolata R.M.Schust., Phytologia 39 (6): 427, 1978 (Schuster 1978b).
** Drepanolejeunea valiae Jovet-Ast, Rev. Bryol. Lichénol. 18 (1/2): 38, 1949 (JovetAst 1949a).
** Drepanolejeunea vandenberghenii Buchb. et Eb.Fisch., J. Bryol. 26 (4): 273, 2004
(Buchbender and Fischer 2004).
*** Drepanolejeunea vesiculosa (Mitt.) Steph., Sp. Hepat. (Stephani) 5: 356, 1913
(Stephani 1913a). Bas.: Lejeunea vesiculosa Mitt., J. Proc. Linn. Soc., Bot. 5 (18):
116, 1860 [1861] (Mitten 1860c).
** Drepanolejeunea yulensis Steph., Sp. Hepat. (Stephani) 5: 356, 1913 (Stephani
1913a).
***
***
***
***
***
**
**
***
subg. Kolpolejeunea Grolle, J. Hattori Bot. Lab. 40: 193, 1976 (Grolle 1976c).
Drepanolejeunea intermedia Zwickel, Ann. Bryol. 6: 119, 1933 (Zwickel 1933).
Drepanolejeunea lyrata Grolle, J. Hattori Bot. Lab. 40: 199, 1976 (Grolle 1976c).
Drepanolejeunea madagascariensis (Steph.) Grolle, Lindbergia 2 (3/4): 232, 1974
(Grolle and Onraedt 1974). Bas.: Leptolejeunea madagascariensis Steph., Sp. Hepat.
(Stephani) 5: 363, 1913 (Stephani 1913a).
Drepanolejeunea pocsii Grolle, J. Hattori Bot. Lab. 40: 209, 1976 (Grolle 1976c).
Drepanolejeunea symoensii Vanden Berghen et Grolle, J. Hattori Bot. Lab. 49: 86,
1981 (Grolle 1981). Based on: Leptolejeunea symoensii Vanden Berghen, Bull. Soc.
Roy. Bot. Belgique 93: 58, 1961 (Vanden Berghen 1961), nom. inval.
Drepanolejeunea symoensii var. minor Tixier, Trop. Bryol. 11: 25, 1995 (Tixier
1995b).
Drepanolejeunea trematodes (Nees) Bischl., Rev. Bryol. Lichénol. 35 (1/4): 125,
1967 [1968] (Bischler 1967). Bas.: Lejeunea trematodes Nees, Ann. Sci. Nat. Bot.
(sér. 2) 5: 63, 1836 (Nees and Montagne 1836).
** subg. Pristolejeunea Grolle, J. Hattori Bot. Lab. 40: 193, 1976 (Grolle 1976c).
** Drepanolejeunea actinogyna Inuthai, R.L.Zhu et Chantanaorr., Bryologist 117 (2):
165, 2014 (Inuthai et al. 2014).
*** Drepanolejeunea issicornua Steph., Sp. Hepat. (Stephani) 5: 344, 1913 (Stephani
1913a).
* Drepanolejeunea hampeana Steph., Sp. Hepat. (Stephani) 5: 345, 1913 (Stephani
1913a). Nom. nov. pro Drepanolejeunea hampeana Steph., Hedwigia 29 (2): 70,
1890 (Stephani 1890b), nom. inval.
** Drepanolejeunea laciniata Qiong He et R.L.Zhu, Cryptog. Bryol. 33 (3): 292,
2012 (He et al. 2012a).
*** Drepanolejeunea levicornua Steph., Sp. Hepat. (Stephani) 5: 347, 1913 (Stephani
1913a).
* Drepanolejeunea longicornua (Herzog) Mizut., J. Hattori Bot. Lab. 68: 368, 1990
(Mizutani 1990). Bas.: Drepanolejeunea levicornua var. longicornua Herzog, Ann.
Bryol. 3: 142, 1930 (Herzog 1930b).
* Drepanolejeunea nymanii Steph., Sp. Hepat. (Stephani) 5: 348, 1913 (Stephani 1913a).
360
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Drepanolejeunea pulla (Mitt.) Grolle, J. Hattori Bot. Lab. 46: 349, 1979 (Grolle
1979d). Bas.: Lejeunea pulla Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 116, 1860
[1861] (Mitten 1860c).
* Drepanolejeunea serricalyx Herzog, Ann. Bryol. 9: 126, 1936 [1937] (Herzog 1936a).
*** Drepanolejeunea thwaitesiana (Mitt.) Steph., Sp. Hepat. (Stephani) 5: 350, 1913
(Stephani 1913a). Bas.: Lejeunea thwaitesiana Mitt., J. Proc. Linn. Soc., Bot. 5
(18): 117, 1860 [1861] (Mitten 1860c). 304
** Drepanolejeunea thwaitesiana var. zhengii R.L.Zhu, Beih. Nova Hedwigia 121:
197, 2001 (Zhu and So 2001).
*** Drepanolejeunea tricornua Herzog, Ann. Bryol. 9: 124, 1936 [1937] (Herzog 1936a).
*** subg. Rhaphidolejeunea (Herzog) Grolle et R.L.Zhu, Nova Hedwigia 70 (3/4):
376, 2000 (Grolle and Zhu 2000). Bas.: Rhaphidolejeunea Herzog, Mitth. hüring.
Bot. Vereins 50: 104, 1943 (Herzog 1943c).
** Drepanolejeunea bidoupensis Pócs, Cryptog. Bryol. 34 (3): 293, 2013 (Pócs et al. 2013).
*** Drepanolejeunea bischlerae (Grolle) Grolle et R.L.Zhu, Nova Hedwigia 70 (3/4):
391, 2000 (Grolle and Zhu 2000). Bas.: Rhaphidolejeunea bischlerae Grolle, J.
Hattori Bot. Lab. 38: 653, 1974 (Grolle 1974b).
*** Drepanolejeunea commutata Grolle et R.L.Zhu, Nova Hedwigia 70 (3/4): 377,
2000 (Grolle and Zhu 2000).
*** Drepanolejeunea cyclops (Sande Lac.) Grolle et R.L.Zhu, Nova Hedwigia 70 (3/4):
391, 2000 (Grolle and Zhu 2000). Bas.: Lejeunea cyclops Sande Lac., Ann. Mus.
Bot. Lugduno-Batavi 1: 310, 1864 (Sande Lacoste 1864).
*** Drepanolejeunea leischeri (Steph.) Grolle et R.L.Zhu, Nova Hedwigia 70 (3/4):
379, 2000 (Grolle and Zhu 2000). Bas.: Leptolejeunea leischeri Steph., Sp. Hepat.
(Stephani) 5: 382, 1913 (Stephani 1913a).
** Drepanolejeunea foliicola Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 1:
85, 1932 (Horikawa 1932a).
** Drepanolejeunea longicruris (Steph.) Grolle et R.L.Zhu, Nova Hedwigia 70 (3/4):
392, 2000 (Grolle and Zhu 2000). Bas.: Leptolejeunea longicruris Steph., Hedwigia
35 (3): 106, 1896 (Stephani 1896b).
*** Drepanolejeunea polyrhiza (Nees) Grolle et R.L.Zhu, Nova Hedwigia 70 (3/4):
392, 2000 (Grolle and Zhu 2000). Bas.: Lejeunea polyrhiza Nees, Syn. Hepat. 3:
403, 1845 (Gottsche et al. 1845b).
*** Drepanolejeunea siamensis (Bischl.) Grolle et R.L.Zhu, Nova Hedwigia 70 (3/4):
393, 2000 (Grolle and Zhu 2000). Bas.: Rhaphidolejeunea siamensis Bischl., Rev.
Bryol. Lichénol. 36 (1/2): 86, 1968 [1969] (Bischler 1968).
304 Drepanolejeunea thwaitesiana is a species complex also including Drepanolejeunea hampeana,
Drepanolejeunea longicornua, Drepanolejeunea nymannii and Drepanolejeunea serricalyx.
World checklist of hornworts and liverworts
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*** Drepanolejeunea spicata (Steph.) Grolle et R.L.Zhu, Nova Hedwigia 70 (3/4): 384,
2000 (Grolle and Zhu 2000). Bas.: Leptolejeunea spicata Steph., Hedwigia 35 (3):
108, 1896 (Stephani 1896b).
*** Drepanolejeunea tibetana (P.C.Wu et J.S.Lou) Grolle et R.L.Zhu, Nova Hedwigia 70 (3/4): 386, 2000 (Grolle and Zhu 2000). Bas.: Rhaphidolejeunea tibetana
P.C.Wu et J.S.Lou, Acta Phytotax. Sin. 16 (4): 102, 1978 (Wu and Lou 1978).
*** Drepanolejeunea yunnanensis (P.C.Chen) Grolle et R.L.Zhu, Nova Hedwigia
70 (3/4): 388, 2000 (Grolle and Zhu 2000). Bas.: Rhaphidolejeunea yunnanensis
P.C.Chen, Feddes Repert. Spec. Nov. Regni Veg. 58: 44, 1955 (Chen 1955).
Incertae sedis
*** Drepanolejeunea dactylophora (Nees, Lindenb. et Gottsche) J.B.Jack et Steph.,
Hedwigia 31 (1): 12, 1892 (Jack and Stephani 1892). Bas.: Lejeunea dactylophora
Nees, Lindenb. et Gottsche, Observ. bot.: 473, 1843 (Gottsche et al. 1843).
*** Drepanolejeunea dactylophora var. submuricata Herzog, Ann. Bryol. 4: 92, 1931
(Herzog 1931b).
* Drepanolejeunea devendrae Sushil K.Singh et M.Dey, Nelumbo 54: 20, 2012
(Singh and Dey 2012).
* Drepanolejeunea integerrima Herzog, Rev. Bryol. Lichénol. 20 (1/2): 146, 1951
[1952] (Herzog 1951a). 305
** Vitalianthus R.M.Schust. et Giancotti, Nova Hedwigia 57 (3/4): 447, 1993
(Schuster and Giancotti 1993).
*** Vitalianthus bischlerianus (K.C.Pôrto et Grolle) R.M.Schust. et Giancotti, Nova
Hedwigia 57 (3/4): 448, 1993 (Schuster and Giancotti 1993). Bas.: Drepanolejeunea
bischleriana K.C.Pôrto et Grolle, Cryptog. Bryol. Lichénol. 8 (4): 301, 1987 (Pôrto
and Grolle 1987).
** Vitalianthus guangxianus R.L.Zhu, Qiong He et Y.M.Wei, J. Bryol. 34 (1): 32,
2012 (He et al. 2012b).
**
subtrib. Echinolejeuneinae Gradst.
*** Anoplolejeunea (Spruce) Schifn., Hepat. (Engl.-Prantl): 131, 1893 (Schifner
1893b). Bas.: Lejeunea subg. Anoplolejeunea Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 129, 1884 (Spruce 1884).
*** Anoplolejeunea conferta (C.F.W.Meissn. ex Spreng.) A.Evans, Bull. Torrey Bot.
Club 35 (4): 175, 1908 (Evans 1908a). Bas.: Jungermannia conferta C.F.W.Meissn.
ex Spreng. Syst. Veg. (ed. 16) [Sprengel] 4 (2): 325, 1827 (Sprengel 1827b).
305 Drepanolejeunea integerrima is a Leptolejeunea species (Bischler 1964), but it is probably a synonym
since she did not transfer it.
362
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Echinolejeunea R.M.Schust., Beih. Nova Hedwigia 9: 187, 1963 (Schuster 1963a).
*** Echinolejeunea papillata (Mitt.) R.M.Schust. ex Hamlin, Rec. Domin. Mus. 7:
260, 1972 (Hamlin 1972). Bas.: Lejeunea papillata Mitt., Bot. antarct. voy. II (Fl.
Nov.-Zel. 2): 158, 1854 (Mitten 1854).
** Kymatolejeunea Grolle, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Reihe 32 (6): 1005, 1984 (Grolle 1984c).
*** Kymatolejeunea bartlettii Grolle, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.Naturwiss. Reihe 32 (6): 1005, 1984 (Grolle 1984c).
**
subtrib. Leiolejeuneinae Schäf.-Verw. et Heinrichs
** Leiolejeunea A.Evans, Bull. Torrey Bot. Club 35 (8): 377, 1908 (Evans 1908b).
*** Leiolejeunea grandilora A.Evans, Bull. Torrey Bot. Club 35 (8): 378, 1908 (Evans
1908b).
*** subtrib. Lejeuneinae Gradst.
*** Harpalejeunea (Spruce) Schifn., Hepat. (Engl.-Prantl): 126, 1893 (Schifner
1893b). Bas.: Lejeunea subg. Harpalejeunea Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 164, 1884 (Spruce 1884).
** subg. Cleeiolejeunea Grolle et M.E.Reiner, J. Bryol. 21 (1): 33, 1999 (Grolle
and Reiner-Drehwald 1999).
*** Harpalejeunea grandis Grolle et M.E.Reiner, J. Bryol. 21 (1): 32, 1999 (Grolle and
Reiner-Drehwald 1999).
** subg. Harpalejeunea
* Harpalejeunea acuta S.Winkl., Rev. Bryol. Lichénol. 42 (3): 812, 1976 (Winkler
1976). 306
*** Harpalejeunea ancistrodes (Spruce) Schifn., Hepat. (Engl.-Prantl): 127, 1893
(Schifner 1893b). Bas.: Lejeunea ancistrodes Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 169, 1884 (Spruce 1884).
** Harpalejeunea buenaventurae Herzog, Feddes Repert. Spec. Nov. Regni Veg. 57
(1/2): 183, 1955 (Herzog 1955).
*** Harpalejeunea cinchonae (Nees) Schifn., Hepat. (Engl.-Prantl): 127, 1893
(Schifner 1893b). Bas.: Lejeunea cinchonae Nees, Syn. Hepat. 3: 342, 1845
(Gottsche et al. 1845b).
306 Harpalejeunea acuta is possibly conspeciic with Harpalejeunea cinchonae (Grolle and ReinerDrehwald 1999).
World checklist of hornworts and liverworts
363
** Harpalejeunea cinchonae var. strigulosa Herzog, Svensk Bot. Tidskr. 51 (1): 192,
1957 (Herzog 1957a).
** Harpalejeunea decurvicuspis (Besch. et C.Massal.) P.Syd., Just’s Bot. Jahresber. 19:
246, 1894 (Sydow 1894). Bas.: Lejeunea decurvicuspis Besch. et C.Massal., Bull.
Mens. Soc. Linn. Paris 1 (79): 639, 1886 (Bescherelle and Massalongo 1886).
** Harpalejeunea emarginata Jovet-Ast, Rev. Bryol. Lichénol. 16 (1/2): 38, 1947
[1948] (Jovet-Ast 1947b).
** Harpalejeunea exocellata Herzog, Rev. Bryol. Lichénol. 20 (1/2): 144, 1951 [1952]
(Herzog 1951a).
** Harpalejeunea grandistipula R.M.Schust., J. Hattori Bot. Lab. 87: 290, 1999
(Schuster 1999c).
*** Harpalejeunea harpaphylla (Herzog) Bischl., Rev. Bryol. Lichénol. 33 (1/2): 164,
1964 (Bischler 1964). Bas.: Drepanolejeunea harpaphylla Herzog, Svensk Bot.
Tidskr. 46 (1): 93, 1952 (Herzog 1952e).
** Harpalejeunea herzogii Jovet-Ast, Feddes Repert. Spec. Nov. Regni Veg. 58: 19,
1955 (Jovet-Ast 1955).
** Harpalejeunea longibracteata (Spruce) Steph., Sp. Hepat. (Stephani) 5: 254, 1913
(Stephani 1913a). Bas.: Lejeunea longibracteata Spruce, Bull. Soc. Bot. France
(Congr. Bot.) 36: cciii, 1889 [1890] (Spruce 1889).
** Harpalejeunea marginalis (Hook.f. et Taylor) Steph., Sp. Hepat. (Stephani) 5:
271, 1913 (Stephani 1913a). Bas.: Jungermannia marginalis Hook.f. et Taylor,
London J. Bot. 4: 91, 1845 (Hooker and Taylor 1845).
*** Harpalejeunea molleri (Steph.) Grolle, Taxon 38 (1): 89, 1989 (Grolle 1989c).
Bas.: Lejeunea molleri Steph., Hedwigia 26 (1): 3, 1887 (Stephani 1887).
** Harpalejeunea molleri subsp. integra (R.M.Schust.) Damsh., Ill. Fl. Nord. Liverw.
Hornw.: 615, 2002 (Damsholt 2002). Bas.: Harpalejeunea ovata subsp. integra
R.M.Schust., J. Elisha Mitchell Sci. Soc. 83 (4): 199, 1967 (Schuster 1967a).
*** Harpalejeunea oxyphylla (Nees et Mont.) Steph., Sp. Hepat. (Stephani) 5: 255,
1913 (Stephani 1913a). Bas.: Lejeunea oxyphylla Nees et Mont., Ann. Sci. Nat.
Bot. (sér. 2) 19: 264, 1843 (Montagne 1843).
*** Harpalejeunea parasitica (Hook.f. et Taylor) Steph., Sp. Hepat. (Stephani) 5: 268,
1913 (Stephani 1913a). Bas.: Jungermannia parasitica Hook.f. et Taylor, London
J. Bot. 3: 477, 1844 (Hooker and Taylor 1844b).
** Harpalejeunea pinaundensis Grolle, J. Hattori Bot. Lab. 46: 44, 1979 (Grolle 1979a).
** Harpalejeunea relexula A.Evans, Bull. Torrey Bot. Club 35 (8): 375, 1908 (Evans
1908b).
** Harpalejeunea scabra Gradst. et Schäf.-Verw., Cryptog. Bryol. 32 (2): 102, 2011
(Gradstein and Schäfer-Verwimp 2011).
*** Harpalejeunea schifneri S.W.Arnell, Österr. Akad. Wiss., Math.-Naturwiss. Kl.,
Denkschr. 111: 102, 1964 (Schifner and Arnell 1964).
* Harpalejeunea solitaria (Gottsche) Steph., Sp. Hepat. (Stephani) 5: 246, 1913
(Stephani 1913a). Bas.: Lejeunea solitaria Gottsche, Abh. Naturwiss. Vereins Bremen 7: 356, 1882 (Gottsche 1882).
364
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*** Harpalejeunea stricta (Lindenb. et Gottsche) Steph., Sp. Hepat. (Stephani) 5: 259,
1913 (Stephani 1913a). Bas.: Lejeunea stricta Lindenb. et Gottsche, Syn. Hepat. 5:
756, 1847 (Gottsche et al. 1847).
** Harpalejeunea subacuta A.Evans, Bull. Torrey Bot. Club 30 (10): 547, 1903 (Evans 1903c).
* Harpalejeunea tenuicuspis (Spruce) Schifn., Hepat. (Engl.-Prantl): 127, 1893
(Schifner 1893b). Bas.: Lejeunea tenuicuspis Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 170, 1884 (Spruce 1884). 307
*** Harpalejeunea tridens (Besch. et Spruce) Steph., Sp. Hepat. (Stephani) 5: 263,
1913 (Stephani 1913a). Bas.: Lejeunea tridens Besch. et Spruce, Bull. Soc. Bot.
France (Congr. Bot.) 36: clxxxi, 1889 [1890] (Bescherelle and Spruce 1889).
** Harpalejeunea uncinata Steph., Hedwigia 35 (3): 97, 1896 (Stephani 1896b).
* Harpalejeunea uncinata var. setulosa Herzog, Feddes Repert. Spec. Nov. Regni
Veg. 57 (1/2): 183, 1955 (Herzog 1955).
*
*
*
*
Incertae sedis
Harpalejeunea grossearmata Steph., Biblioth. Bot. 87 (2): 256, 1916 (Stephani
1916a). 308
Harpalejeunea renneri Herzog, Memoranda Soc. Fauna Fl. Fennica 26: 54, 1950
[1951] (Herzog 1950b). 309
Harpalejeunea spruceana Steph., Biblioth. Bot. 87 (2): 257, 1916 (Stephani 1916a). 310
Harpalejeunea vitrea Herzog, Memoranda Soc. Fauna Fl. Fennica 27: 94, 1952
(Herzog 1952c). 311
** Hattoriolejeunea Mizut., J. Hattori Bot. Lab. 61: 303, 1986 [1987] (Mizutani 1986b).
** Hattoriolejeunea akiyamae Mizut., J. Hattori Bot. Lab. 61: 303, 1986 [1987]
(Mizutani 1986b).
*** Lejeunea Lib., Ann. Gen. Sci. Phys. 6: 373, 1820 (Libert 1820) nom. conserv. 312
307 Harpalejeunea tenuicuspis is possibly conspeciic with Harpalejeunea oxyphylla (Gradstein and Costa
2003, Grolle and Reiner-Drehwald 1999).
308 Harpalejeunea grossearmata is a Lejeunea species (Grolle and Reiner-Drehwald 1999). It was treated
as a doubtful taxon by Gradstein et al. (2003).
309 Harpalejeunea renneri is a Lejeunea species and most likely conspeciic with something (Grolle and
Reiner-Drehwald 1999).
310 Harpalejeunea spruceana is a Lejeunea species (non Lejeunea spruceana C.Massal. 1885) (Grolle and
Reiner-Drehwald 1999). It was treated as a doubtful taxon by Gradstein et al. (2003).
311 Harpalejeunea vitrea is a Lejeunea species (non Lejeunea vitrea Nees, Lindenb. et Gottsche 1843)
(Grolle and Reiner-Drehwald 1999).
312 Lejeunea is a large genus and its infrageneric classiication, which has not been studied in a rigorous
manner, needs investigation although Heinrichs et al. (2013) showed two main clades on molecular
grounds, provisionally named Lejeunea subg. Lejeunea and Lejeunea subg. Crossotolejeunea. However,
their morphology is not yet understood. he subgenera listed here are recently accepted ones. Crossotolejeunea, Eulejeunea, Hygrolejeunea and Taxilejeunea also belong here, but several taxa have neither
World checklist of hornworts and liverworts
365
** Lejeunea tunquiniensis M.E.Reiner et Drehwald, Nova Hedwigia 100 (3/4): 584,
2015 (Reiner-Drehwald 2015).
** subg. Lejeunea
*** Lejeunea abyssinica (Gola) Cufod., Phyton (Horn) 4: 75, 1952 (Cufodontis 1952).
Bas.: Eulejeunea abyssinica Gola, Ann. Bot. (Rome) 13 (1): 70, 1914 (Gola 1914a).
*** Lejeunea acanthogona Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 177, 1884
(Spruce 1884).
** Lejeunea acuminata (Lehm. et Lindenb.) Lehm., Nov. Stirp. Pug. 7: 22, 1838
(Lehmann 1838). Bas.: Jungermannia acuminata Lehm. et Lindenb., Nov. Stirp.
Pug. 6: 49, 1834 (Lehmann 1834).
** Lejeunea acuta Mitt., J. Proc. Linn. Soc., Bot. 7 (27): 167, 1863 (Mitten 1863).
** Lejeunea acutata (Steph.) Solari, J. Hattori Bot. Lab. 54: 541, 1983 (Solari 1983a).
Bas.: Strepsilejeunea acutata Steph., Hedwigia 35 (3): 127, 1896 (Stephani 1896b).
*** Lejeunea adpressa Nees, Repert. Pharm. 76: 45, 1842 (von Flotow et al. 1842). 313
*** Lejeunea aethiopica E.W.Jones, J. Bryol. 13 (3): 387, 1985 (Jones 1985).
** Lejeunea alaskana (R.M.Schust. et Steere) Inoue et Steere, J. Hattori Bot. Lab. 44:
330, 1978 (Steere and Inoue 1978). Bas.: Hygrolejeunea alaskana R.M.Schust. et
Steere, Bull. Torrey Bot. Club 85 (3): 190, 1958 (Schuster and Steere 1958).
*** Lejeunea alata Gottsche, Syn. Hepat. 3: 406, 1845 (Gottsche et al. 1845b).
** Lejeunea alata var. patriciae Pócs, Candollea 56 (1): 72, 2001 (Pócs 2001).
*** Lejeunea albescens (Steph.) Mizut., J. Hattori Bot. Lab. 33: 245, 1970 (Mizutani 1970).
Bas.: Taxilejeunea albescens Steph., Hedwigia 35 (3): 132, 1896 (Stephani 1896b).
* Lejeunea albilora Colenso, Trans. & Proc. New Zealand Inst. 21: 72, 1889 (Colenso
1889).
** Lejeunea aloba Sande Lac., Plagiochila Sandei: 10, 1856 (Sande Lacoste 1856c).
** Lejeunea alobifolia H.A.Mill., Phytologia 47 (4): 323, 1981 (Miller 1981). Nom.
nov. pro Lejeunea aloba Steph., Sp. Hepat. (Stephani) 5: 767, 1915 (Stephani
1915b), nom. illeg.
** Lejeunea amaniensis E.W.Jones, J. Bryol. 13 (3): 392, 1985 (Jones 1985).
** Lejeunea ambigua Lindenb. et Gottsche, Syn. Hepat. 5: 764, 1847 (Gottsche et al.
1847).
* Lejeunea amentulifera Steph., Sp. Hepat. (Stephani) 5: 707, 1915 (Stephani 1915b).
** Lejeunea androgyna R.M.Schust., Phytologia 45 (5): 432, 1980 (Schuster 1980b).
* Lejeunea angulifolia Mitt., Philos. Trans. 168: 400, 1879 (Mitten 1879). 314
*** Lejeunea angusta (Lehm. et Lindenb.) Mont., Hist. Phys. Cuba, Bot., Pl. Cell.:
469, 1842 (Montagne 1842a). Bas.: Jungermannia angusta Lehm. et Lindenb.,
Nov. Stirp. Pug. 4: 52, 1832 (Lehmann 1832).
been transferred nor synonymized. hey are listed in the “Names in genera not currently accepted”
section below.
313 Lejeunea adpressa from South America is very similar to the palaeotropical Lejeunea anisophylla. he
two taxa probably represent a single pantropical species (possibly with two subspecies).
314 Lejeunea angulifolia may be conspeciic with Lejeunea cocoes (Pócs 2011a).
366
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Lejeunea anisophylla Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 263, 1843 (Montagne
1843).
*** Lejeunea anomala Lindenb. et Gottsche, Linnaea 24 (6): 636, 1851 [1852] (Lindenberg and Gottsche 1851a).
* Lejeunea antillana Steph., Hedwigia 27 (11/12): 281, 1888 (Stephani 1888c).
** Lejeunea aphanes Spruce, J. Bot. 19: 36, 1881 (Spruce 1881b).
*** Lejeunea apiculata Sande Lac., Ned. Kruidk. Arch. 3: 421, 1854 [1855] (Sande
Lacoste 1854).
** Lejeunea aquatica Horik., Sci. Rep. Tôhoku Imp. Univ., Ser. 4, Biol. 5 (4): 643,
1929 [1930] (Horikawa 1929c).
** Lejeunea aquatica var. apiculata S.Hatt., Bull. Tokyo Sci. Mus. 11: 108, 1944
(Hattori 1944d).
** Lejeunea armitii (Steph.) Steph., Sp. Hepat. (Stephani) 5: 768, 1915 (Stephani 1915b).
Bas.: Eulejeunea armitii Steph., Hedwigia 28 (3): 169, 1889 (Stephani 1889d).
* Lejeunea asperifolia Steph., Sp. Hepat. (Stephani) 5: 708, 1915 (Stephani 1915b).
*** Lejeunea asperrima Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 160, 1884
(Spruce 1884).
** Lejeunea asperula (Steph.) Mizut., J. Hattori Bot. Lab. 33: 236, 1970 (Mizutani
1970). Bas.: Taxilejeunea asperula Steph., Sp. Hepat. (Stephani) 5: 499, 1914
(Stephani 1914b).
** Lejeunea asprella Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 175, 1884
(Spruce 1884).
* Lejeunea atheatostipa Spruce, J. Bot. 33: 83, 1895 (Gepp 1895a).
** Lejeunea barbata (Herzog) R.L.Zhu et M.J.Lai, Ann. Bot. Fenn. 48 (5): 376, 2011
(Wang et al. 2011). Bas.: Rectolejeunea barbata Herzog, J. Hattori Bot. Lab. 14:
49, 1955 (Herzog and Noguchi 1955).
*** Lejeunea bermudiana (A.Evans) R.M.Schust., Hepat. Anthocerotae N. Amer. 4:
1105, 1980 (Schuster 1980c). Bas.: Crossotolejeunea bermudiana A.Evans, Bull.
Torrey Bot. Club 33 (3): 132, 1906 (Evans 1906c).
** Lejeunea bidentula Herzog, Symb. Sin. 5: 51, 1930 (Nicholson et al. 1930).
** Lejeunea biformis Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 162 (68), 1864 (Gottsche
1864).
** Lejeunea blepharogona Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 178, 1884
(Spruce 1884).
** Lejeunea blomquistii R.M.Schust., J. Elisha Mitchell Sci. Soc. 78 (1): 64, 1962
(Schuster 1962b).
*** Lejeunea boliviensis (Steph.) R.L.Zhu et M.E.Reiner, Bryologist 107 (2): 237,
2004 (Zhu and Reiner-Drehwald 2004). Bas.: Strepsilejeunea boliviensis Steph.,
Biblioth. Bot. 87 (2): 257, 1916 (Stephani 1916a).
** Lejeunea boryana Mont., Ann. Sci. Nat. Bot. (sér. 2) 9: 47, 1838 (Montagne 1838).
** Lejeunea brenanii E.W.Jones, J. Bryol. 10 (4): 391, 1979 (Jones 1979).
*** Lejeunea calcicola R.M.Schust., J. Elisha Mitchell Sci. Soc. 73 (2): 404, 1957
(Schuster 1957d).
World checklist of hornworts and liverworts
367
** Lejeunea calcicola var. mexicana R.M.Schust., J. Elisha Mitchell Sci. Soc. 73 (2):
408, 1957 (Schuster 1957d).
** Lejeunea canariensis (Steph.) Steph., Sp. Hepat. (Stephani) 5: 802, 1915 (Stephani
1915b). Bas.: Eulejeunea canariensis Steph., Mém. Soc. Bot. France 7: 42, 1907
(Pitard and Corbière 1907).
*** Lejeunea cancellata Nees et Mont., Hist. Phys. Cuba, Bot., Pl. Cell.: 472, 1842
(Montagne 1842a).
** Lejeunea cantabrigiensis E.W.Jones, J. Bryol. 15 (4): 669, 1989 (Jones 1989).
*** Lejeunea capensis Gottsche, Syn. Hepat. 3: 374, 1845 (Gottsche et al. 1845b).
* Lejeunea caroliniana Austin, Bot. Bull. (Hanover) 1 (8): 36, 1876 (Austin 1876a). 315
*** Lejeunea catinulifera Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 233, 1884
(Spruce 1884).
*** Lejeunea caulicalyx (Steph.) M.E.Reiner et Goda, J. Hattori Bot. Lab. 89: 13, 2000
(Reiner-Drehwald and Goda 2000). Bas.: Crossotolejeunea caulicalyx Steph., Sp.
Hepat. (Stephani) 5: 237, 1913 (Stephani 1913a).
*** Lejeunea cavifolia (Ehrh.) Lindb., Revis. crit. icon.: 43, 1871 (Lindberg 1871). Bas.:
Jungermannia cavifolia Ehrh., Beitr. Naturk. (Ehrhart) 4: 45, 1789 (Ehrhart 1789).
** Lejeunea caviloba (Steph.) Besch., J. Bot. (Morot) 12: 140, 1898 (Bescherelle 1898).
Bas.: Eulejeunea caviloba Steph., Hedwigia 35 (3): 86, 1896 (Stephani 1896b).
*** Lejeunea cerina (Lehm. et Lindenb.) Lehm. et Lindenb., Syn. Hepat. 3: 391, 1845
(Gottsche et al. 1845b). Bas.: Jungermannia cerina Lehm. et Lindenb., Nov. Stirp.
Pug. 5: 16, 1833 (Lehmann 1833).
** Lejeunea chaishanensis S.H.Lin, Yushania 9: 7, 1992 (Lin and Yang 1992).
** Lejeunea cladogyna A.Evans, Amer. J. Bot. 5 (3): 134, 1918 (Evans 1918).
** Lejeunea clavata Lindenb., Syn. Hepat. 3: 379, 1845 (Gottsche et al. 1845b).
** Lejeunea claviformis Lindenb. ex Steph., Sp. Hepat. (Stephani) 5: 727, 1915
(Stephani 1915b).
** Lejeunea cochleata Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 273, 1884
(Spruce 1884).
** Lejeunea cocoes Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 114, 1860 [1861] (Mitten 1860c).
*** Lejeunea colensoana (Steph.) M.A.M.Renner, Austral. Syst. Bot. 23 (6): 455, 2010
(Renner et al. 2010b). Bas.: Taxilejeunea colensoana Steph., Hedwigia 35 (3): 132,
1896 (Stephani 1896b).
** Lejeunea compacta (Steph.) Steph., Sp. Hepat. (Stephani) 5: 771, 1915 (Stephani
1915b). Bas.: Eulejeunea compacta Steph., Bull. Herb. Boissier 5 (2): 93, 1897
(Stephani 1897b).
** Lejeunea concinnula Spruce et Steph., J. Bot. 25: 39, 1887 (Spruce 1887a).
** Lejeunea connatistipula (Steph.) Steph., Sp. Hepat. (Stephani) 5: 772, 1915
(Stephani 1915b). Bas.: Eulejeunea connatistipula Steph., Hedwigia 35 (3): 87,
1896 (Stephani 1896b).
315 Lejeunea caroliniana was poorly described and the type specimen so poor that Evans (1902a) could
not identify it.
368
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Lejeunea contracta Mizut., J. Hattori Bot. Lab. 33: 248, 1970 (Mizutani 1970).
** Lejeunea controversa Gottsche, Hepat. Eur., Leberm. 56-57: no 556, 1873
(Gottsche and Rabenhorst 1873b).
** Lejeunea convexiloba M.L.So et R.L.Zhu, Bryologist 101 (1): 137, 1998 (So and
Zhu 1998).
** Lejeunea corcovadae (Steph.) Bischl., Nova Hedwigia 5 (1/2): 406, 1963 (Bischler
et al. 1963). Bas.: Microlejeunea corcovadae Steph., Sp. Hepat. (Stephani) 5: 820,
1915 (Stephani 1915b).
** Lejeunea cordilora Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 283, 1884
(Spruce 1884).
** Lejeunea corralensis A.Evans, Ann. Bryol. 3: 86, 1930 (Evans 1930b).
*** Lejeunea corynantha Spruce, J. Linn. Soc., Bot. 30 (210): 344, 1895 (Gepp 1895b).
** Lejeunea crassiretis Mitt., Fl. vit.: 414, 1871 [1873] (Mitten 1871).
*** Lejeunea cristulata (Steph.) M.E.Reiner et Goda, J. Hattori Bot. Lab. 89: 21, 2000
(Reiner-Drehwald and Goda 2000). Bas.: Crossotolejeunea cristulata Steph., Hedwigia 35 (3): 75, 1896 (Stephani 1896b).
*** Lejeunea cristulilora (Steph.) M.E.Reiner et Goda, J. Hattori Bot. Lab. 89: 19,
2000 (Reiner-Drehwald and Goda 2000). Bas.: Crossotolejeunea cristulilora Steph.,
Sp. Hepat. (Stephani) 5: 231, 1913 (Stephani 1913a).
** Lejeunea cuspidistipula (Steph.) Steph. ex Watts, Proc. Linn. Soc. New South
Wales (ser. 2) 27 (108): 493, 1903 (Watts 1903). Bas.: Eulejeunea cuspidistipula
Steph., Hedwigia 35 (3): 88, 1896 (Stephani 1896b).
** Lejeunea cyanomontana R.M.Schust., Phytologia 45 (5): 432, 1980 (Schuster 1980b).
** Lejeunea cyanophora R.M.Schust., J. Hattori Bot. Lab. 26: 246, 1963 (Schuster
1963b).
** Lejeunea cyathearum E.W.Jones, J. Bryol. 8 (1): 86, 1974 (Jones 1974).
*** Lejeunea cyathophora Mitt., Hooker’s J. Bot. Kew Gard. Misc. 3: 359, 1851 (Mitten
1851).
** Lejeunea denticalyx (Steph.) Steph., Sp. Hepat. (Stephani) 5: 794, 1915 (Stephani
1915b). Bas.: Eulejeunea denticalyx Steph., Hedwigia 28 (3): 169, 1889 (Stephani
1889d).
** Lejeunea denticuspis (Steph.) Mizut., J. Hattori Bot. Lab. 36: 160, 1972 [1973]
(Mizutani 1972a). Bas.: Strepsilejeunea denticuspis Steph., Hedwigia 35 (3): 129,
1896 (Stephani 1896b).
** Lejeunea denudata (Pearson) J.J.Engel, Bryologist 78 (3): 361, 1975 (Engel 1975).
Bas.: Eulejeunea denudata Pearson, J. Linn. Soc., Bot. 46 (305): 39, 1922 (Pearson
1922b).
*** Lejeunea deplanata Nees, Syn. Hepat. 3: 368, 1845 (Gottsche et al. 1845b).
** Lejeunea deplanata var. cuspidata (Steph.) M.E.Reiner, Nova Hedwigia 91 (3/4):
529, 2010 (Reiner-Drehwald 2010). Bas.: Pycnolejeunea cuspidata Steph., Sp. Hepat. (Stephani) 5: 605, 1914 (Stephani 1914b).
** Lejeunea diaphana Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 168, 1884
(Spruce 1884).
World checklist of hornworts and liverworts
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369
Lejeunea dimorpha T.Kodama, J. Hattori Bot. Lab. 41: 384, 1976 (Kodama 1976).
Lejeunea dipterocarpa E.W.Jones, J. Bryol. 7 (1): 44, 1972 (Jones 1972).
Lejeunea discreta Lindenb., Syn. Hepat. 3: 361, 1845 (Gottsche et al. 1845b).
Lejeunea disjecta Spruce, J. Linn. Soc., Bot. 30 (210): 347, 1895 (Gepp 1895b).
Lejeunea diversicuspis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 176, 1884
(Spruce 1884).
Lejeunea drummondii Taylor, London J. Bot. 5: 400, 1846 (Taylor 1846b).
Lejeunea ecarinata (Steph.) J.M.Coult., Barnes et Arthur, Bot. Gaz. 15 (12): 349,
1890 (Coulter et al. 1890). Bas.: Eulejeunea ecarinata Steph., Bot. Gaz. 15 (11):
283, 1890 (Stephani 1890c).
Lejeunea eckloniana Lindenb., Syn. Hepat. 3: 381, 1845 (Gottsche et al. 1845b).
Lejeunea eifrigii Mizut., J. Hattori Bot. Lab. 33: 244, 1970 (Mizutani 1970). Nom.
nov. pro Taxilejeunea acutiloba Eifrig, Monogr. Stud. Indomal. Art. Taxilejeunea:
94, 1937 (Eifrig 1937).
Lejeunea elongella Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 161, 1864 (Gottsche 1864).
Lejeunea erostrata M.E.Reiner et Goda, J. Hattori Bot. Lab. 89: 25, 2000 (Reiner-Drehwald and Goda 2000). Nom. nov. pro Crossotolejeunea parva Steph., Sp.
Hepat. (Stephani) 5: 241, 1913 (Stephani 1913a).
Lejeunea exilis (Reinw., Blume et Nees) Grolle, J. Hattori Bot. Lab. 46: 353, 1979
(Grolle 1979d). Bas.: Jungermannia exilis Reinw., Blume et Nees, Nova Acta
Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 227, 1824 [1825] (Reinwardt et al. 1824a).
Lejeunea exilis var. abnormis (Herzog) G.E.Lee, Polish Bot. J. 58 (1): 61, 2013 (Lee
and Gradstein 2013). Bas.: Byssolejeunea abnormis Herzog, Hedwigia 80 (1/2): 84,
1941 (Herzog 1941b).
Lejeunea fernandeziana S.W.Arnell, Ark. Bot. (n.ser.) 4 (1): 16, 1957 (Arnell 1957b).
Lejeunea irma Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 112, 1860 [1861] (Mitten
1860c).
Lejeunea issistipula (Steph.) Steph., Sp. Hepat. (Stephani) 5: 775, 1915 (Stephani
1915b). Bas.: Eulejeunea issistipula Steph., Hedwigia 35 (3): 88, 1896 (Stephani
1896b).
Lejeunea lagellaris Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 273, 1884
(Spruce 1884).
Lejeunea lava (Sw.) Nees, Naturgesch. Eur. Leberm. 3: 277, 1838 (Nees 1838b).
Bas.: Jungermannia lava Sw., Prodr. (Swartz): 144, 1788 (Swartz 1788). 316
Lejeunea lava subsp. moorei (Lindb.) R.M.Schust., J. Elisha Mitchell Sci. Soc.
73 (1): 161, 1957 (Schuster 1957a). Bas.: Lejeunea moorei Lindb., Acta Soc. Sci.
Fenn. 10: 487, 1875 (Lindberg 1875).
Lejeunea lava subsp. orientalis R.M.Schust., J. Elisha Mitchell Sci. Soc. 73 (1):
161, 1957 (Schuster 1957a).
316 Lejeunea lava is a species complex also including Lejeunea grossecristata.
370
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Lejeunea lava var. pellucida Lindenb. et Gottsche, Linnaea 24 (6): 634, 1851
[1852] (Lindenberg and Gottsche 1851a).
** Lejeunea lava subsp. tabularis (Spreng.) S.W.Arnell, Hepat. South Africa: 199,
1963 (Arnell 1963b). Bas.: Jungermannia tabularis Spreng. Syst. Veg. (ed. 16)
[Sprengel] 4 (2): 325, 1827 (Sprengel 1827b).
*** Lejeunea lavovirens Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 30 (5): 144,
1873 (Ångström 1873).
** Lejeunea leischeri (Steph.) Mizut., J. Hattori Bot. Lab. 33: 238, 1970 (Mizutani
1970). Bas.: Hygrolejeunea leischeri Steph., Sp. Hepat. (Stephani) 5: 560, 1914
(Stephani 1914b).
** Lejeunea loridana A.Evans, Bull. Torrey Bot. Club 32 (4): 185, 1905 (Evans 1905b).
*** Lejeunea fulfordiae (Jovet-Ast) R.L.Zhu, Syst. Bot. 33 (4): 617, 2008 (Zhu and
Cheng 2008). Bas.: Amblyolejeunea fulfordiae Jovet-Ast, Rev. Bryol. Lichénol. 17
(1/4): 25, 1948 [1949] (Jovet-Ast 1948).
** Lejeunea fusagasugana Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 158, 1864
(Gottsche 1864).
** Lejeunea galeata Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 172, 1884
(Spruce 1884).
** Lejeunea gayana Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 157, 1864 (Gottsche 1864).
** Lejeunea gibbiloba (Steph.) H.A.Mill., Phytologia 47 (4): 323, 1981 (Miller 1981).
Bas.: Eulejeunea gibbiloba Steph., Sp. Hepat. (Stephani) 6: 418, 1923 (Stephani 1923).
*** Lejeunea glaucescens Gottsche, Syn. Hepat. 3: 378, 1845 (Gottsche et al. 1845b).
** Lejeunea glaucescens var. acrogyna R.M.Schust., J. Elisha Mitchell Sci. Soc. 73 (2):
400, 1957 (Schuster 1957d).
* Lejeunea glaucescens var. obsoleta R.M.Schust., J. Elisha Mitchell Sci. Soc. 73 (2):
395, 1957 (Schuster 1957d).
** Lejeunea globosilora (Steph.) Steph., Sp. Hepat. (Stephani) 5: 795, 1915 (Stephani
1915b). Bas.: Eulejeunea globosilora Steph., Bih. Kongl. Svenska Vetensk.-Akad.
Handl. 26 (III, 6): 65, 1900 (Stephani 1900b).
** Lejeunea gomphocalyx Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 174, 1884
(Spruce 1884).
** Lejeunea gracilicaulis Spruce, Mem. Torrey Bot. Club 1 (3): 126, 1890 (Spruce 1890).
** Lejeunea gracilipes (Taylor) Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 213,
1884 (Spruce 1884). Bas.: Omphalanthus gracilipes Taylor, London J. Bot. 5: 385,
1846 (Taylor 1846b).
** Lejeunea gracilis Steph., Sp. Hepat. (Stephani) 5: 777, 1915 (Stephani 1915b).
** Lejeunea gradsteiniana Pócs, Acta Biol. Pl. Agr. 1: 55, 2010 [2011] (Pócs 2010d).
Nom. nov. pro Ceratolejeunea aberrans Steph., Sp. Hepat. (Stephani) 6: 399, 1923
(Stephani 1923).
** Lejeunea gradsteinii G.E.Lee, Damanhuri et Latif, Acta Biol. Pl. Agr. 1: 29, 2010
[2011] (Lee et al. 2010).
* Lejeunea grossecristata (Steph.) E.W.Jones, Trans. Brit. Bryol. Soc. 5 (3): 556,
1968 (Jones 1968). Bas.: Hygrolejeunea grossecristata Steph., Hedwigia 35 (3): 102,
1896 (Stephani 1896b).
World checklist of hornworts and liverworts
371
*** Lejeunea grossiretis (Steph.) M.E.Reiner et Goda, J. Hattori Bot. Lab. 89: 27, 2000
(Reiner-Drehwald and Goda 2000). Bas.: Crossotolejeunea grossiretis Steph., Hedwigia 35 (3): 75, 1896 (Stephani 1896b).
** Lejeunea grossistipula Steph., Sp. Hepat. (Stephani) 5: 739, 1915 (Stephani 1915b).
*** Lejeunea grossitexta (Steph.) M.E.Reiner et Goda, J. Hattori Bot. Lab. 89: 29,
2000 (Reiner-Drehwald and Goda 2000). Bas.: Crossotolejeunea grossitexta Steph.,
Sp. Hepat. (Stephani) 5: 240, 1913 (Stephani 1913a).
* Lejeunea grossiuscula Gottsche ex Steph., Sp. Hepat. (Stephani) 5: 739, 1915
(Stephani 1915b).
** Lejeunea hahnii Solari, J. Hattori Bot. Lab. 54: 543, 1983 (Solari 1983a). Nom. nov.
pro Microlejeunea grandistipula Steph., Hedwigia 35 (3): 114, 1896 (Stephani 1896b).
* Lejeunea haitica Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 263, 1843 (Montagne 1843).
** Lejeunea hawaikiana M.A.M.Renner et de Lange, New Zealand J. Bot. 49 (3):
431, 2011 (Renner and de Lange 2011). Nom. nov. pro Stenolejeunea acuminata
R.M.Schust., J. Hattori Bot. Lab. 89: 156, 2000 (Schuster 2000b).
** Lejeunea helenae Pearson, Forh. Vidensk.-Selsk. Kristiania 1886 (3): 6, 1886
(Pearson 1886).
** Lejeunea helmsiana (Steph.) Steph., Sp. Hepat. (Stephani) 5: 796, 1915 (Stephani
1915b). Bas.: Eulejeunea helmsiana Steph., Hedwigia 35 (3): 89, 1896 (Stephani 1896b).
*** Lejeunea hepaticola (Steph.) Steph., Sp. Hepat. (Stephani) 5: 714, 1915 (Stephani
1915b). Bas.: Eulejeunea hepaticola Steph., Hedwigia 27 (2): 60, 1888 (Stephani 1888a).
** Lejeunea hibernica Bischl., H.A.Mill. et Bonner ex Grolle, Lindbergia 3 (1/2): 48,
1975 [1976] (Grolle 1975c). Based on: Lejeunea hibernica Bischl., H.A.Mill. et
Bonner, Nova Hedwigia 3 (4): 455, 1961 [1962] (Bischler et al. 1961), nom. inval.
** Lejeunea hodgsoniana Grolle ex R.J.Lewington, Bever. et M.A.M.Renner, PhytoKeys
29: 2, 2013 (Lewington et al. 2013).
** Lejeunea howeana Grolle, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Reihe 31 (2): 219, 1982 (Grolle 1982). Nom. nov. pro Cheilolejeunea wattsiana Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2): 102, 1914 (Stephani
and Watts 1914).
** Lejeunea hui R.L.Zhu, Beih. Nova Hedwigia 121: 134, 2001 (Zhu and So 2001).
** Lejeunea humefacta Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 275, 1884
(Spruce 1884).
** Lejeunea hyalina (Steph.) L.Söderstr. et A.Hagborg, Phytotaxa 220 (2): 188, 2015
(Söderström et al. 2015a). Bas.: Pycnolejeunea hyalina Steph., Sp. Hepat. (Stephani) 5: 614, 1914 (Stephani 1914b).
** Lejeunea ibadana A.J.Harr. et E.W.Jones, J. Bryol. 12 (1): 40, 1982 (Jones 1982).
** Lejeunea immersa Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 186, 1884
(Spruce 1884).
** Lejeunea increscens Spruce, Mem. Torrey Bot. Club 1 (3): 124, 1890 (Spruce 1890).
* Lejeunea inlatiloba (Steph.) H.A.Mill., Bonner et Bischl., Nova Hedwigia 14 (1):
67, 1967 (Miller et al. 1967). Bas.: Microlejeunea inlatiloba Steph., Sp. Hepat.
(Stephani) 6: 422, 1923 (Stephani 1923).
372
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Lejeunea inlexiloba Prantl, Hedwigia 31: xvi, 1892 (Prantl 1892). Based on: Crossotolejeunea inlexiloba J.B.Jack et Steph., Hedwigia 31 (1): 16, 1892 (Jack and
Stephani 1892), nom. inval.
*** Lejeunea intricata Prantl, Hedwigia 31: xvi, 1892 (Prantl 1892). Based on: Crossotolejeunea intricata J.B.Jack et Steph., Hedwigia 31 (1): 17, 1892 (Jack and
Stephani 1892), nom. inval.
*** Lejeunea isophylla E.W.Jones, Trans. Brit. Bryol. Soc. 5 (3): 559, 1968 (Jones 1968).
** Lejeunea japonica Mitt., Trans. Linn. Soc. London, Bot. 3 (3): 203, 1891 (Mitten
1891).
** Lejeunea jardinii Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: ccv, 1889 [1890]
(Spruce 1889).
** Lejeunea julacea Steph., Sp. Hepat. (Stephani) 5: 715, 1915 (Stephani 1915b).
*** Lejeunea jungneri (Steph.) Steph., Cat. Afr. Pl. (Hiern) 2 (2): 318, 1901 (Stephani
1901d). Bas.: Eulejeunea jungneri Steph., Hedwigia 35 (3): 90, 1896 (Stephani 1896b).
** Lejeunea kashyapii M.Dey, D.K.Singh et D.Singh, J. Bryol. 30 (2): 126, 2008
(Dey et al. 2008).
* Lejeunea kilimandjarensis Steph., Sp. Hepat. (Stephani) 5: 716, 1915 (Stephani 1915b).
** Lejeunea kinabalensis Mizut., J. Hattori Bot. Lab. 33: 246, 1970 (Mizutani 1970).
** Lejeunea kodamae Ikegami et Inoue, J. Jap. Bot. 36 (1): 7, 1961 (Inoue 1961a).
** Lejeunea konosensis Mizut., J. Hattori Bot. Lab. 71: 127, 1992 (Mizutani 1992).
** Lejeunea kuerschneriana Pócs, Beih. Nova Hedwigia 138: 100, 2010 (Pócs 2010b).
*** Lejeunea laeta (Lehm. et Lindenb.) Lehm. et Lindenb., Syn. Hepat. 3: 380, 1845
(Gottsche et al. 1845b). Bas.: Jungermannia laeta Lehm. et Lindenb., Nov. Stirp.
Pug. 6: 45, 1834 (Lehmann 1834).
** Lejeunea laii R.L.Zhu, J. Bryol. 30 (2): 173, 2008 (Wang and Zhu 2008). Nom.
nov. pro Microlejeunea ramulosa Herzog, J. Hattori Bot. Lab. 14: 51, 1955 (Herzog
and Noguchi 1955).
*** Lejeunea lamacerina (Steph.) Schifn., Hedwigia 41 (5): 278, 1902 (Schifner 1902).
Bas.: Eulejeunea lamacerina Steph., Hedwigia 35 (3): 91, 1896 (Stephani 1896b). 317
** Lejeunea lamacerina subsp. gemminata R.M.Schust., J. Elisha Mitchell Sci. Soc. 73
(1): 168, 1957 (Schuster 1957a).
** Lejeunea latilobula (Herzog) R.L.Zhu et M.L.So, J. Bryol. 24 (2): 168, 2002 (Zhu
and So 2002). Bas.: Taxilejeunea latilobula Herzog, Symb. Sin. 5: 50, 1930 (Nicholson et al. 1930).
** Lejeunea leptalea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 272, 1884
(Spruce 1884).
* Lejeunea leratii (Steph.) Mizut., J. Hattori Bot. Lab. 33: 243, 1970 (Mizutani
1970). Bas.: Hygrolejeunea leratii Steph., Sp. Hepat. (Stephani) 5: 562, 1914 (Stephani 1914b). 318
317 Lejeunea lamacerina forms a North American and a Macaronesian/European clade and the separation
of two subspecies are justiied if not treating them at species level (Heinrichs et al. 2013).
318 Lejeunea leratii is probably distinct from Lejeunea mimula (Lee 2013).
World checklist of hornworts and liverworts
373
** Lejeunea leucosis Besch. et Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: clxxxviii, 1889 [1890] (Bescherelle and Spruce 1889).
* Lejeunea litoralis Steph., Sp. Hepat. (Stephani) 5: 778, 1915 (Stephani 1915b).
** Lejeunea lomana E.W.Jones, Bull. Brit. Mus. (Nat. Hist.), Bot. 11 (3): 257, 1983
(Jones and Harrington 1983).
** Lejeunea longicollis S.W.Arnell, Results Norweg. Sci. Exped. Tristan da Cunha 42:
12, 1958 (Arnell 1958b).
** Lejeunea longilobula Pócs, Beih. Nova Hedwigia 138: 112, 2010 (Pócs 2010b).
Nom. nov. pro Lejeunea halei subsp. africana Pócs, J. Bryol. 29 (2): 89, 2007 (Müller and Pócs 2007).
** Lejeunea lowriana Steph., Sp. Hepat. (Stephani) 5: 779, 1915 (Stephani 1915b).
*** Lejeunea lumbricoides (Nees) Nees, Syn. Hepat. 3: 342, 1845 (Gottsche et al. 1845b).
Bas.: Jungermannia lumbricoides Nees, Enum. Pl. Crypt. Javae: 40, 1830 (Nees 1830).
** Lejeunea lunatigastria Tixier, Ann. Hist.-Nat. Mus. Natl. Hung. 66: 98, 1974
(Tixier 1974).
** Lejeunea lyratilora Prantl, Hedwigia 31: xvi, 1892 (Prantl 1892). Based on: Hygrolejeunea lyratilora Steph., Hedwigia 31 (4): 169, 1892 (Stephani 1892g), nom. inval.
** Lejeunea magohukui Mizut., Misc. Bryol. Lichenol. 7 (7): 133, 1977 (Mizutani 1977).
** Lejeunea mandonii (Steph.) Müll.Frib., Leberm. Eur. 2 (9): 1281, 1958 (Müller 1958).
Bas.: Microlejeunea mandonii Steph., Hedwigia 35 (3): 115, 1896 (Stephani 1896b).
*** Lejeunea masoalae Pócs, Beih. Nova Hedwigia 138: 103, 2010 (Pócs 2010b).
** Lejeunea massalongoana (Schifn. ex P.Syd.) Solari, J. Hattori Bot. Lab. 54: 542,
1983 (Solari 1983a).Bas.: Harpalejeunea massalongoana Schifn. ex P.Syd., Just’s
Bot. Jahresber. 19: 246, 1894 (Sydow 1894).
** Lejeunea megalantha Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 172, 1884
(Spruce 1884).
** Lejeunea mehrana M.Dey, D.K.Singh et D.Singh, J. Bryol. 30 (2): 128, 2008 (Dey
et al. 2008).
*** Lejeunea meridensis Ilk.-Borg., Nova Hedwigia 80 (1/2): 59, 2005 (Ilkiu-Borges 2005).
** Lejeunea micholitzii Mizut., J. Hattori Bot. Lab. 33: 236, 1970 (Mizutani 1970).
Nom. nov. pro Hygrolejeunea parvisaccata Steph., Sp. Hepat. (Stephani) 5: 567,
1914 (Stephani 1914b).
** Lejeunea microloba Taylor, London J. Bot. 5: 399, 1846 (Taylor 1846b).
*** Lejeunea mimula Hürl., Bauhinia 11 (1): 12, 1993 (Hürlimann 1993). Nom. nov. pro
Hygrolejeunea luteola Steph., Sp. Hepat. (Stephani) 5: 553, 1914 (Stephani 1914b).
*** Lejeunea minutiloba A.Evans, Bull. Torrey Bot. Club 44 (11): 525, 1917 (Evans
1917d).
** Lejeunea minutiloba var. heterogyna R.M.Schust., J. Elisha Mitchell Sci. Soc. 73
(2): 425, 1957 (Schuster 1957d).
** Lejeunea mizutanii Grolle, J. Hattori Bot. Lab. 45: 178, 1979 (Grolle 1979c). Nom.
nov. pro Cheilolejeunea zollingeri Steph., Hedwigia 34 (5): 245, 1895 (Stephani 1895b).
** Lejeunea molkenboeriana Sande Lac., Ned. Kruidk. Arch. 3: 421, 1854 [1855]
(Sande Lacoste 1854).
374
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Lejeunea monimiae (Steph.) Steph., Sp. Hepat. (Stephani) 5: 747, 1915 (Stephani
1915b). Bas.: Eulejeunea monimiae Steph., Hedwigia 35 (3): 91, 1896 (Stephani 1896b).
*** Lejeunea multidentata M.E.Reiner et Mustelier, J. Bryol. 26 (2): 103, 2004 (Reiner-Drehwald and Mustelier 2004).
* Lejeunea musae (Spreng.) Gottsche, Lindenb. et Nees, Syn. Hepat. 3: 407, 1845
(Gottsche et al. 1845b). Bas.: Jungermannia musae Spreng. Ann. Wetterauischen
Ges. Gesammte Naturk. 1: 25, 1809 (Sprengel 1809).
** Lejeunea muscicola Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 281, 1884
(Spruce 1884).
* Lejeunea muscicola var. palmicola Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:
282, 1884 (Spruce 1884).
** Lejeunea neelgherriana Gottsche, Syn. Hepat. 3: 354, 1845 (Gottsche et al. 1845b).
** Lejeunea nemoralis Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 159 (65), 1864
(Gottsche 1864).
** Lejeunea nepalensis Steph., Sp. Hepat. (Stephani) 5: 780, 1915 (Stephani 1915b).
* Lejeunea nesiotica Besch. et Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: clxxxviii, 1889 [1890] (Bescherelle and Spruce 1889).
** Lejeunea neumanniana Nees, Repert. Pharm. 76: 44, 1842 (von Flotow et al. 1842).
** Lejeunea nietneri (Steph.) Steph. ex Watts, Proc. Linn. Soc. New South Wales (ser.
2) 26 (104): 633, 1902 (Watts 1902). Bas.: Eulejeunea nietneri Steph., Hedwigia
35 (3): 91, 1896 (Stephani 1896b).
** Lejeunea notata Gottsche, Abh. Naturwiss. Vereins Bremen 7: 361, 1882 (Gottsche
1882).
** Lejeunea nymannii Steph., Sp. Hepat. (Stephani) 5: 781, 1915 (Stephani 1915b).
** Lejeunea obfusca Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 114, 1860 [1861] (Mitten
1860c).
** Lejeunea obidensis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 277, 1884
(Spruce 1884).
* Lejeunea obscura Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 112, 1860 [1861] (Mitten
1860c). 319
** Lejeunea obtusata Gottsche, Abh. Naturwiss. Vereins Bremen 7: 354, 1882
(Gottsche 1882).
** Lejeunea okomuensis E.W.Jones, Trans. Brit. Bryol. Soc. 5 (4): 787, 1969 (Jones
1969).
*** Lejeunea oligoclada Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: cxcix, 1889
[1890] (Spruce 1889).
*** Lejeunea oracola M.A.M.Renner, Austral. Syst. Bot. 23 (6): 448, 2010 (Renner et
al. 2010b).
** Lejeunea osculatiana De Not., Mem. Reale Accad. Sci. Torino (ser. 2) 16: 233,
1857 (De Notaris 1857).
** Lejeunea otiana S.Hatt., Bot. Mag. (Tokyo) 65 (763/764): 15, 1952 (Hattori 1952a).
319 Lejeunea obscura is possibly conspeciic with Lejeunea aloba (Söderström et al. 2010a).
World checklist of hornworts and liverworts
**
**
**
**
*
***
*
**
**
***
***
***
***
**
***
*
***
*
***
**
**
375
Lejeunea ovalifolia Steph., Sp. Hepat. (Stephani) 5: 751, 1915 (Stephani 1915b).
Lejeunea paciica Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 262, 1843 (Montagne 1843).
Lejeunea pallescens Mitt., Hooker’s J. Bot. Kew Gard. Misc. 3: 360, 1851 (Mitten 1851).
Lejeunea pallida Lindenb. et Gottsche, Syn. Hepat. 5: 762, 1847 (Gottsche et al. 1847).
Lejeunea pallidissima Gola, Nuovo Giorn. Bot. Ital. (n.ser.) 27 (2/4): 247, 1920
(Gola 1920). 320
Lejeunea papilionacea Prantl, Hedwigia 31: xvii, 1892 (Prantl 1892). Based on:
Hygrolejeunea papilionacea Steph., Hedwigia 31 (4): 169, 1892 (Stephani 1892g),
nom. inval.
Lejeunea paratropa Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 176, 1884
(Spruce 1884). 321
Lejeunea patagonica (Steph.) Steph., Sp. Hepat. (Stephani) 5: 797, 1915 (Stephani
1915b). Bas.: Eulejeunea patagonica Steph., Bih. Kongl. Svenska Vetensk.-Akad.
Handl. 26 (III, 6): 66, 1900 (Stephani 1900b).
Lejeunea patens Lindb., Acta Soc. Sci. Fenn. 10: 482, 1875 (Lindberg 1875).
Lejeunea patersonii (Steph.) Steph., Sp. Hepat. (Stephani) 5: 784, 1915 (Stephani
1915b). Bas.: Eulejeunea patersonii Steph., Hedwigia 35 (3): 92, 1896 (Stephani
1896b).
Lejeunea patriciae Schäf.-Verw., Candollea 56 (1): 64, 2001 (Schäfer-Verwimp
2001a). Nom. nov. pro Lejeunea pilifera Tixier, Gard. Bull. Singapore 25 (3): 351,
1971 (Tixier 1971), nom. illeg.
Lejeunea paucidentata (Steph.) Grolle, J. Hattori Bot. Lab. 69: 191, 1991 (Grolle
1991). Bas.: Crossotolejeunea paucidentata Steph., Hedwigia 35 (3): 76, 1896
(Stephani 1896b).
Lejeunea pectinella Mizut., J. Hattori Bot. Lab. 33: 239, 1970 (Mizutani 1970).
Lejeunea perigonialis Gottsche, Mexik. Leverm.: 223, 1863 (Gottsche 1863).
Lejeunea perpapillosa M.E.Reiner et K.C.Pôrto, Nova Hedwigia 85 (3/4): 542,
2007 (Reiner-Drehwald and Pôrto 2007).
Lejeunea pertusa (Corda ex Nees et Mont.) Gottsche, Lindenb. et Nees, Syn. Hepat.
3: 407, 1845 (Gottsche et al. 1845b). Bas.: Jungermannia pertusa Corda ex Nees et
Mont., Ann. Sci. Nat. Bot. (sér. 2) 5: 63, 1836 (Nees and Montagne 1836).
Lejeunea phyllobola Nees et Mont., Hist. Phys. Cuba, Bot., Pl. Cell.: 471, 1842
(Montagne 1842a).
Lejeunea phyllobola var. turgidula Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:
259, 1884 (Spruce 1884).
Lejeunea planiloba A.Evans, Proc. Wash. Acad. Sci. 8: 147, 1906 (Evans 1906b).
Lejeunea polilloensis Steph., Sp. Hepat. (Stephani) 5: 786, 1915 (Stephani 1915b).
Lejeunea praetervisa Steph., Sp. Hepat. (Stephani) 5: 752, 1915 (Stephani 1915b).
320 Lejeunea pallidissima is possibly conspeciic with Lejeunea lavovirens (Vanden Berghen 1972b).
321 Lejeunea paratropa is closely related to or perhaps conspeciic with Lejeunea raddiana Lindenb.
(Grolle and Reiner-Drehwald 1999).
376
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Lejeunea primordialis (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat.
3: 375, 1845 (Gottsche et al. 1845b). Bas.: Jungermannia primordialis Hook.f. et
Taylor, London J. Bot. 4: 92, 1845 (Hooker and Taylor 1845).
** Lejeunea princeps (Steph.) Mizut., J. Hattori Bot. Lab. 34: 454, 1971 (Mizutani
1971a). Bas.: Hygrolejeunea princeps Steph., Sp. Hepat. (Stephani) 5: 568, 1914
(Stephani 1914b).
** Lejeunea procumbens Mitt., Fl. vit.: 413, 1871 [1873] (Mitten 1871).
** Lejeunea propagulifera Gradst., Phytotaxa 9: 54, 2010 (Söderström et al. 2010a).
Nom. nov. pro Trachylejeunea schifneri Herzog, Svensk Bot. Tidskr. 42 (3): 239,
1948 (Herzog 1948).
** Lejeunea pteridis Besch. et Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: clxxxvii, 1889 [1890] (Bescherelle and Spruce 1889).
*** Lejeunea ptosimophylla C.Massal., Nuovo Giorn. Bot. Ital. 13 (2): 123, 1881
(Massalongo 1881).
*** Lejeunea puiggariana Steph., Sp. Hepat. (Stephani) 5: 754, 1915 (Stephani 1915b).
*** Lejeunea pulverulenta (Steph.) M.E.Reiner, Cryptog. Bryol. 26 (1): 60, 2005
(Reiner-Drehwald 2005b). Bas.: Taxilejeunea pulverulenta Steph., Sp. Hepat.
(Stephani) 5: 477, 1914 (Stephani 1914b).
*** Lejeunea pulvinata Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 5: 61, 1836 (Nees
and Montagne 1836). Nom. nov. pro Jungermannia pulvinata Lehm. et Lindenb.,
Nov. Stirp. Pug. 5: 15, 1833 (Lehmann 1833), nom. illeg.
*** Lejeunea raddiana Lindenb., Syn. Hepat. 3: 342, 1845 (Gottsche et al. 1845b).
** Lejeunea radicans Lindenb. et Gottsche, Syn. Hepat. 5: 766, 1847 (Gottsche et al. 1847).
** Lejeunea ramosissima Steph., Bot. Jahrb. Syst. 8 (2): 88, 1886 (Stephani 1886d).
** Lejeunea rara Steph., Sp. Hepat. (Stephani) 5: 798, 1915 (Stephani 1915b). 322
* Lejeunea ravenelii Austin, Bot. Bull. (Hanover) 1 (8): 35, 1876 (Austin 1876a). 323
** Lejeunea recurva M.E.Reiner, Polish Bot. J. 58 (2): 423, 2013 (Reiner-Drehwald
et al. 2013). Nom. nov. pro Hygrolejeunea herzogii Steph., Biblioth. Bot. 87 (2):
265, 1916 (Stephani 1916a), nom. illeg.
*** Lejeunea relexistipula (Lehm. et Lindenb.) Lehm. et Lindenb., Syn. Hepat. 3: 335,
1845 (Gottsche et al. 1845b). Bas.: Jungermannia relexistipula Lehm. et Lindenb.,
Nov. Stirp. Pug. 5: 10, 1833 (Lehmann 1833).
** Lejeunea relexistipula var. costaricensis (Steph.) M.E.Reiner, Nova Hedwigia
81 (3/4): 408, 2005 (Reiner-Drehwald 2005a). Bas.: Hygrolejeunea costaricensis
Steph., Hedwigia 35 (3): 100, 1896 (Stephani 1896b).
*** Lejeunea reinerae Ilk.-Borg., Nova Hedwigia 80 (1/2): 61, 2005 (Ilkiu-Borges
2005). Nom. nov. pro Echinocolea herzogii Mizut. et Grolle, Bot. Mag. (Tokyo) 77
(915): 333, 1964 (Grolle 1964d).
322 Lejeunea rara is the valid name for Lejeunea sinclairii Spruce (1884) hom. illeg. (non Mitten 1862
= hysananthus fruticosus), but Scott and Bradshaw (1985) confused the taxa and considered them
conspeciic. Grolle (1982) accepted Spruce’s taxon without commenting on any homonym problem.
323 Lejeunea ravenelii was poorly described and the type specimen so poor that Evans (1902a) could not
identify it.
World checklist of hornworts and liverworts
377
** Lejeunea resupinata (Steph.) Steph., Sp. Hepat. (Stephani) 5: 757, 1915 (Stephani
1915b). Bas.: Eulejeunea resupinata Steph., Bih. Kongl. Svenska Vetensk.-Akad.
Handl. 23 (III, 2): 22, 1897 (Stephani 1897a).
** Lejeunea reticulata Herzog, Nat. Hist. Juan Fernandez (Botany) 2 (5): 747, 1942
(Herzog 1942a).
*** Lejeunea rhigophila M.A.M.Renner, Austral. Syst. Bot. 23 (6): 453, 2010 (Renner
et al. 2010b).
** Lejeunea rhodesiae (Sim) R.M.Schust., J. Hattori Bot. Lab. 25: 71, 1962 (Schuster
1962a). Bas.: Stylolejeunea rhodesiae Sim, Trans. Roy. Soc. South Africa 15 (1): 68,
1926 (Sim 1926).
*** Lejeunea rionegrensis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 579, 1885
(Spruce 1885). Nom. nov. pro Lejeunea implexa Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 240, 1884 (Spruce 1884), nom. inval.
** Lejeunea riparia Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 113, 1860 [1861] (Mitten
1860c).
* Lejeunea rothii (Schwägr.) Gottsche, Lindenb. et Nees, Syn. Hepat. 3: 407, 1845
(Gottsche et al. 1845b). Bas.: Jungermannia rothii Schwägr., Hist. Musc. Hepat.
Prodr.: 17, 1814 (Schwägrichen 1814).
*** Lejeunea rotundifolia Mitt., Hooker’s J. Bot. Kew Gard. Misc. 3: 359, 1851 (Mitten
1851).
** Lejeunea sanctae-helenae M.Wigginton, J. Bryol. 29 (1): 12, 2007 (Wigginton
2007).
** Lejeunea schusteri Grolle, Haussknechtia 8: 60, 2001 (Grolle 2001). Nom. nov. pro Rectolejeunea denudata R.M.Schust., J. Hattori Bot. Lab. 89: 143, 2000 (Schuster 2000c).
** Lejeunea semiscabrida Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 154 (60), 1864
(Gottsche 1864).
** Lejeunea semperi Steph., Sp. Hepat. (Stephani) 5: 788, 1915 (Stephani 1915b).
** Lejeunea seriata Lindenb. et Gottsche, Syn. Hepat. 5: 762, 1847 (Gottsche et al. 1847).
** Lejeunea sessililora (Steph.) Grolle, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.Naturwiss. Reihe 37: 171, 1988 (Grolle 1988b). Bas.: Macrolejeunea sessililora
Steph., Sp. Hepat. (Stephani) 5: 512, 1914 (Stephani 1914b).
** Lejeunea setacea (Steph.) Steph., Sp. Hepat. (Stephani) 5: 719, 1914 (Stephani
1914b). Bas.: Eulejeunea setacea Steph., Bull. Mus. Natl. Hist. Nat. 18 (2): 120,
1912 (Corbière 1912).
** Lejeunea setiloba Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 281, 1884
(Spruce 1884).
** Lejeunea sharpii (R.M.Schust.) R.M.Schust., Hepat. Anthocerotae N. Amer. 4:
991, 1980 (Schuster 1980c). Bas.: Taxilejeunea sharpii R.M.Schust., J. Elisha
Mitchell Sci. Soc. 81 (1): 41, 1965 (Schuster 1965a).
** Lejeunea siccata Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 284, 1884
(Spruce 1884).
** Lejeunea silvatica Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 159 (65), 1864 (Gottsche
1864).
378
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Lejeunea smaragdina Besch. et Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36:
clxxxii, 1889 [1890] (Bescherelle and Spruce 1889).
*** Lejeunea soae R.L.Zhu, Y.M.Wei, L.Söderstr., A.Hagborg et von Konrat, Phytotaxa 81 (1): 1, 2013 (Zhu et al. 2013). Nom. nov. pro Trachylejeunea chinensis
Herzog, Symb. Sin. 5: 49, 1930 (Nicholson et al. 1930).
** Lejeunea solanicola Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 280, 1884
(Spruce 1884).
*** Lejeunea sordida (Nees) Nees, Naturgesch. Eur. Leberm. 3: 278, 1838 (Nees 1838b).
Bas.: Jungermannia sordida Nees, Enum. Pl. Crypt. Javae: 41, 1830 (Nees 1830).
*** Lejeunea spiniloba Lindenb. et Gottsche, Syn. Hepat. 5: 770, 1847 (Gottsche et al.
1847).
** Lejeunea spinulilora Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 177, 1884
(Spruce 1884).
** Lejeunea sporadica Besch. et Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: clxxx,
1889 [1890] (Bescherelle and Spruce 1889).
* Lejeunea squarrosa (Steph.) Steph., Sp. Hepat. (Stephani) 5: 719, 1915 (Stephani
1915b). Bas.: Eulejeunea squarrosa Steph., Bot. Jahrb. Syst. 20 (3): 317, 1895
(Stephani 1895a).
** Lejeunea squarrosula (Herzog) Solari, J. Hattori Bot. Lab. 54: 543, 1983 (Solari
1983a). Bas.: Strepsilejeunea squarrosula Herzog, Nat. Hist. Juan Fernandez (Botany) 2 (5): 741, 1942 (Herzog 1942a).
* Lejeunea stephaniana Mizut., J. Hattori Bot. Lab. 27: 143, 1964 (Mizutani 1964a).
Nom. nov. pro Strepsilejeunea heterophylla Steph., Sp. Hepat. (Stephani) 6: 395,
1923 (Stephani 1923). 324
** Lejeunea stevensiana (Steph.) Mizut., J. Hattori Bot. Lab. 34: 452, 1971 (Mizutani 1971a). Bas.: Taxilejeunea stevensiana Steph., Hedwigia 35 (3): 136, 1896
(Stephani 1896b).
** Lejeunea subacuta Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 113, 1860 [1861] (Mitten 1860c).
* Lejeunea subaquatica Schifn., Arch. Hydrobiol., suppl. 21 (3/4): 397, 1955
(Schifner 1955).
* Lejeunea subbiida Steph. ex Duss, Enum. musc. Antilles franç., Hép.: 10, 1903
(Duss 1903). Nom. nov. pro Odontolejeunea subbiida Steph., Symb. Antill. (Urban) 3 (2): 277, 1902 (Stephani 1902e), nom. illeg.
** Lejeunea subigiensis (Steph.) Steph., Hedwigia 35 (3): 94, 1896 (Stephani 1896b).
Bas.: Eulejeunea subigiensis Steph., Hedwigia 35 (3): 94, 1896 (Stephani 1896b).
** Lejeunea subrufula Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 289, 1884
(Spruce 1884).
** Lejeunea subsessilis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 282, 1884
(Spruce 1884).
324 Lejeunea stephaniana is doubtfully distinct from Lejeunea discreta (Söderström et al. 2010a).
World checklist of hornworts and liverworts
379
*** Lejeunea subspathulata Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 173, 1884
(Spruce 1884).
** Lejeunea succulenta Herzog, Rev. Bryol. Lichénol. 20 (1/2): 171, 1951 [1952]
(Herzog 1951a).
** Lejeunea sufruticola Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 279, 1884
(Spruce 1884).
** Lejeunea syoshii Inoue, Bull. Natl. Sci. Mus. Tokyo, B 3 (4): 143, 1977 (Inoue 1977b).
*** Lejeunea talamancensis M.E.Reiner et Schäf.-Verw., Nova Hedwigia 87 (3/4):
414, 2008 (Reiner-Drehwald and Schäfer-Verwimp 2008).
*** Lejeunea tamasii M.E.Reiner, N.Salazar et C.Chung, Polish Bot. J. 58 (2): 420,
2013 (Reiner-Drehwald et al. 2013).
** Lejeunea tamaspocsii G.E.Lee, Polish Bot. J. 58 (1): 66, 2013 (Lee and Gradstein 2013).
*** Lejeunea tapajosensis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 223, 1884
(Spruce 1884).
** Lejeunea tarapotensis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 282, 1884
(Spruce 1884).
** Lejeunea tenella Taylor, London J. Bot. 5: 398, 1846 (Taylor 1846b).
** Lejeunea thallophora (Eifrig) Gradst., Phytotaxa 9: 54, 2010 (Söderström et al.
2010a). Bas.: Taxilejeunea thallophora Eifrig, Monogr. Stud. Indomal. Art. Taxilejeunea: 99, 1937 (Eifrig 1937).
*** Lejeunea tonduzana (Steph.) M.E.Reiner, Polish Bot. J. 58 (2): 421, 2013 (ReinerDrehwald et al. 2013). Bas.: Hygrolejeunea tonduzana Steph., Hedwigia 35 (3):
105, 1896 (Stephani 1896b).
** Lejeunea trachygona Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 175, 1884
(Spruce 1884).
*** Lejeunea trinitensis Lindenb., Syn. Hepat. 3: 381, 1845 (Gottsche et al. 1845b).
** Lejeunea trukensis H.A.Mill. et Bonner, Beih. Nova Hedwigia 11: 56, 1963 (Miller
et al. 1963).
** Lejeunea tuberculosa Steph., Sp. Hepat. (Stephani) 5: 790, 1915 (Stephani 1915b).
** Lejeunea tumida Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 157, 1854 (Mitten 1854).
** Lejeunea uleana (Steph.) Steph., Sp. Hepat. (Stephani) 5: 765, 1915 (Stephani
1915b). Bas.: Eulejeunea uleana Steph., Hedwigia 35 (3): 94, 1896 (Stephani 1896b).
*** Lejeunea umbilicata (Nees) Nees, Observ. bot.: 472, 1843 (Gottsche et al. 1843).
Bas.: Jungermannia umbilicata Nees, Enum. Pl. Crypt. Javae: 42, 1830 (Nees 1830).
** Lejeunea utriculata (Steph.) Mizut., J. Hattori Bot. Lab. 33: 242, 1970 (Mizutani 1970).
Bas.: Pycnolejeunea utriculata Steph., Hedwigia 35 (3): 126, 1896 (Stephani 1896b).
** Lejeunea vesicata Mitt., Fl. vit.: 415, 1871 [1873] (Mitten 1871).
*** Lejeunea villaumei (Steph.) Grolle, J. Bryol. 9 (4): 536, 1977 [1978] (Grolle
1977a). Bas.: Otigoniolejeunea villaumei Steph., Sp. Hepat. (Stephani) 5: 516,
1914 (Stephani 1914b).
** Lejeunea vojtkoi Pócs, Beih. Nova Hedwigia 138: 107, 2010 (Pócs 2010b).
** Lejeunea vulgariformis Gottsche, Abh. Naturwiss. Vereins Bremen 7: 355, 1882
(Gottsche 1882).
380
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Lejeunea wattsiana (Steph.) H.A.Mill., Bonner et Bischl., Nova Hedwigia 14 (1):
66, 1967 (Miller et al. 1967). Bas.: Microlejeunea wattsiana Steph., Sp. Hepat.
(Stephani) 5: 834, 1916 (Stephani 1916b).
* Lejeunea wichurae (Steph.) Steph., Sp. Hepat. (Stephani) 5: 792, 1915 (Stephani
1915b). Bas.: Eulejeunea wichurae Steph., Hedwigia 35 (3): 94, 1896 (Stephani 1896b).
** Lejeunea wightii Lindenb., Syn. Hepat. 3: 379, 1845 (Gottsche et al. 1845b).
*** Lejeunea xiphophylla (Herzog) M.E.Reiner, Nova Hedwigia 87 (3/4): 409, 2008
(Reiner-Drehwald and Schäfer-Verwimp 2008). Bas.: Taxilejeunea xiphophylla
Herzog, Svensk Bot. Tidskr. 46 (1): 99, 1952 (Herzog 1952e).
** subg. Nanolejeunea R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 1092, 1980
(Schuster 1980c).
** Lejeunea curviloba Steph., Sp. Hepat. (Stephani) 5: 774, 1915 (Stephani
1915b).
*** Lejeunea laetevirens Nees et Mont., Hist. Phys. Cuba, Bot., Pl. Cell.: 469, 1842
(Montagne 1842a).
** Lejeunea pallidevirens S.Hatt., J. Hattori Bot. Lab. 12: 80, 1954 (Hattori 1954).
Nom. nov. pro Microlejeunea rotundistipula var. pallida S.Hatt., J. Hattori Bot.
Lab. 5: 53, 1951 (Hattori 1951d).
** Lejeunea parva (S.Hatt.) Mizut., Misc. Bryol. Lichenol. 5 (10/12): 178, 1971
(Mizutani 1971b). Bas.: Microlejeunea rotundistipula f. parva S.Hatt., Bull. Tokyo
Sci. Mus. 11: 123, 1944 (Hattori 1944d).
*** Lejeunea prionoides Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 238, 1884
(Spruce 1884).
*** Lejeunea ramulosa Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 274, 1884
(Spruce 1884).
*** subg. Neopotamolejeunea (M.E.Reiner) Gradst. et M.E.Reiner, Syst. Bot. 32
(3): 487, 2007 (Gradstein and Reiner-Drehwald 2007). Bas.: Neopotamolejeunea
M.E.Reiner, Nova Hedwigia 71 (3/4): 449, 2000 (Reiner-Drehwald 2000).
** Lejeunea juruana Gradst. et M.E.Reiner, Syst. Bot. 32 (3): 488, 2007 (Gradstein
and Reiner-Drehwald 2007). Nom. nov. pro Potamolejeunea uleana Steph., Sp.
Hepat. (Stephani) 5: 641, 1914 (Stephani 1914b).
*** Lejeunea polyantha Mont., Ann. Sci. Nat. Bot. (sér. 4) 5: 350, 1856 (Montagne 1856c).
** Lejeunea tenera (Sw.) Gottsche, Lindenb. et Nees, Syn. Hepat. 3: 406, 1845
(Gottsche et al. 1845b). Bas.: Jungermannia tenera Sw., Prodr. (Swartz): 143, 1788
(Swartz 1788).
*** Lejeunea topoensis Gradst. et M.E.Reiner, Syst. Bot. 32 (3): 488, 2007 (Gradstein
and Reiner-Drehwald 2007).
** subg. Papillolejeunea (Pócs) R.M.Schust., J. Hattori Bot. Lab. 85: 84, 1998
(Schuster 1998a). Bas.: Papillolejeunea Pócs, Trop. Bryol. 13: 2, 1997 (Pócs 1997b).
** Lejeunea balazsii (Pócs) R.M.Schust., J. Hattori Bot. Lab. 85: 84, 1998 (Schuster
1998a). Bas.: Papillolejeunea balazsii Pócs, Trop. Bryol. 13: 3, 1997 (Pócs 1997b).
World checklist of hornworts and liverworts
381
** Lejeunea candida (Pócs) R.M.Schust., J. Hattori Bot. Lab. 85: 84, 1998 (Schuster
1998a). Bas.: Papillolejeunea candida Pócs, Trop. Bryol. 13: 8, 1997 (Pócs 1997b).
** Lejeunea falcata (Pócs et J.Eggers) Pócs, Phytotaxa 208 (1): 99, 2015 (Pócs et al.
2015a). Bas.: Papillolejeunea falcata Pócs et J.Eggers, Bryobrothera 5: 163, 1999
(Pócs and Eggers 1999).
** Lejeunea koponenii (Pócs et J.Eggers) Pócs, Phytotaxa 208 (1): 99, 2015 (Pócs et
al. 2015a). Bas.: Papillolejeunea koponenii Pócs et J.Eggers, Bryobrothera 5: 159,
1999 (Pócs and Eggers 1999).
** Lejeunea papuana (Pócs) R.M.Schust., J. Hattori Bot. Lab. 85: 84, 1998 (Schuster
1998a). Bas.: Papillolejeunea papuana Pócs, Trop. Bryol. 13: 14, 1997 (Pócs 1997b).
** Lejeunea touwii (Pócs) R.M.Schust., J. Hattori Bot. Lab. 85: 84, 1998 (Schuster
1998a). Bas.: Papillolejeunea touwii Pócs, Trop. Bryol. 13: 11, 1997 (Pócs 1997b).
*
*
**
*
**
**
**
**
**
**
***
**
***
**
**
***
Incertae sedis
Lejeunea apiahyna (Steph.) Sushil K.Singh, Phytotaxa 96 (1): 63, 2013 (Singh
2013). Bas.: Otigoniolejeunea apiahyna Steph., Sp. Hepat. (Stephani) 5: 514, 1914
(Stephani 1914b).
Lejeunea aptycta Gottsche, Syn. Hepat. 3: 369, 1845 (Gottsche et al. 1845b).
Lejeunea asthenica Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 222, 1884
(Spruce 1884).
Lejeunea bethanica Gottsche, Syn. Hepat. 3: 381, 1845 (Gottsche et al. 1945b)
Lejeunea bombonasensis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 222,
1884 (Spruce 1884).
Lejeunea bornmuelleri (Steph.) M.E.Reiner, Nova Hedwigia 95 (3/4): 471, 2012
(Reiner-Drehwald and Grolle 2012). Bas.: Rectolejeunea bornmuelleri Steph., Sp.
Hepat. (Stephani) 5: 682, 1914 (Stephani 1914b).
Lejeunea caracensis Lindenb., Syn. Hepat. 3: 355, 1845 (Gottsche et al. 1845b).
Lejeunea caripensis Lindenb. et Gottsche, Syn. Hepat. 5: 758, 1847 (Gottsche et al.
1847).
Lejeunea chamissonis Lindenb., Syn. Hepat. 3: 378, 1845 (Gottsche et al. 1845b).
Lejeunea chimborazensis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 215,
1884 (Spruce 1884).
Lejeunea combuensis O.S.Moura, Ilk.-Borg. et M.E.Reiner, Nova Hedwigia 95
(1/2): 198, 2012 (Moura et al. 2012).
Lejeunea concava Lindenb. et Gottsche, Syn. Hepat. 5: 759, 1847 (Gottsche et al. 1847).
Lejeunea conformis Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 260, 1843
(Montagne 1843).
Lejeunea cordistipula Lindenb. et Gottsche, Syn. Hepat. 5: 758, 1847 (Gottsche et
al. 1847).
Lejeunea cyrtotis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 229, 1884
(Spruce 1884).
Lejeunea debilis (Lehm. et Lindenb.) Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2)
5: 60, 1836 (Nees and Montagne 1836). Bas.: Jungermannia debilis Lehm. et Lindenb., Nov. Stirp. Pug. 4: 51, 1832 (Lehmann 1832).
382
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Lejeunea devendrae (Sushil K.Singh) P.K.Verma et K.K.Rawat, J. Bryol. 36 (2):
161, 2014 (Verma and Rawat 2014). Bas.: Rectolejeunea devendrae Sushil K.Singh,
Indian J. Forest. 34 (3): 341, 2011 (Singh 2011).
** Lejeunea dictyocalyx Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 218, 1884
(Spruce 1884).
** Lejeunea dipterota (Eifrig) G.E.Lee, Polish Bot. J. 58 (1): 61, 2013 (Lee and Gradstein 2013). Bas.: Taxilejeunea dipterota Eifrig, Monogr. Stud. Indomal. Art. Taxilejeunea: 96, 1937 (Eifrig 1937).
*** Lejeunea drehwaldii Heinrichs et Schäf.-Verw., Phytotaxa 69: 14, 2012 (Heinrichs
et al. 2012b). Nom. nov. pro Sphaerolejeunea umbilicata Herzog, Ann. Bryol. 11:
88, 1938 (Herzog 1938b).
** Lejeunea duncaniae (Sim) M.E.Reiner, Phytotaxa 208 (1): 98, 2015 (Pócs et al.
2015a). Bas.: Stylolejeunea duncaniae Sim, Trans. Roy. Soc. South Africa 15 (1):
68, 1926 (Sim 1926).
** Lejeunea edentata L.Söderstr., Phytotaxa 208 (1): 98, 2015 (Pócs et al. 2015a).
Nom. nov. pro Cyclolejeunea marginata R.M.Schust., Phytologia 39 (6): 430, 1978
(Schuster 1978b).
* Lejeunea emarginulilora Gottsche ex Steph., Sp. Hepat. (Stephani) 5: 734, 1915
(Stephani 1915b). 325
* Lejeunea epibrya Taylor, London J. Bot. 7: 199, 1848 (Taylor 1848a).
** Lejeunea estrellamontana M.A.M.Renner et Pócs, Phytotaxa 81 (1): 9, 2013 (Renner et al. 2013d). Nom. nov. pro Stenolejeunea issistipula R.M.Schust., J. Hattori
Bot. Lab. 89: 167, 2000 (Schuster 2000b).
* Lejeunea fawcettiae D.J.Carr, Proc. Roy. Soc. Victoria 117 (2): 325, 2005 (Carr
2005). Based on: Lejeunea fawcettiae D.J.Carr, Proc. Roy. Soc. Victoria 116 (2):
229, 2004 (Carr 2004), nom. inval.
** Lejeunea laccida Lindenb. et Gottsche, Syn. Hepat. 5: 758, 1847 (Gottsche et al. 1847).
** Lejeunea lavida Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 113, 1860 [1861] (Mitten
1860c).
** Lejeunea lorida Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 221, 1884
(Spruce 1884).
** Lejeunea fulva Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 237, 1884 (Spruce
1884).
** Lejeunea gottscheana Lindenb., Syn. Hepat. 3: 382, 1845 (Gottsche et al. 1845b).
** Lejeunea graminicolor Spruce, J. Linn. Soc., Bot. 30 (210): 343, 1895 (Gepp 1895b).
*** Lejeunea grolleana (Bernecker) R.L.Zhu et W.Ye, J. Syst. Evol. 51 (4): 472, 2013
(Ye et al. 2013b). Bas.: Oryzolejeunea grolleana Bernecker, Haussknechtia, Beih. 9:
37, 1999 (Bernecker-Lücking 1999).
*** Lejeunea herminieri (Steph.) R.L.Zhu, Phytotaxa 208 (1): 99, 2015 (Pócs et al.
2015a). Bas.: Archilejeunea herminieri Steph., Sp. Hepat. (Stephani) 4: 714, 1911
(Stephani 1911e).
325 Lejeunea emarginulilora is a Lejeunea (subg. Heterolejeunea) species (Reiner-Drehwald and Grolle 2012).
World checklist of hornworts and liverworts
383
** Lejeunea heterocheila Taylor, London J. Bot. 5: 394, 1846 (Taylor 1846b).
* Lejeunea hygrophila Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 157, 1864 (Gottsche
1864).
** Lejeunea laevicalyx Gottsche, Mexik. Leverm.: 221, 1863 (Gottsche 1863).
* Lejeunea laxa (Nees) Lindenb., Syn. Hepat. 3: 378, 1845 (Gottsche et al. 1845b). Bas.:
Jungermannia thymifolia δ laxa Nees, Enum. Pl. Crypt. Javae: 43, 1830 (Nees 1830).
** Lejeunea leiantha Spruce, J. Linn. Soc., Bot. 30 (210): 345, 1895 (Gepp 1895b).
** Lejeunea leptoscypha Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: ccv, 1889
[1890] (Spruce 1889).
*** Lejeunea lusoria (Lindenb. et Gottsche) Steph., Hedwigia 29 (3): 141, 1890 (Stephani 1890d). Bas.: Omphalanthus lusorius Lindenb. et Gottsche, Syn. Hepat. 5:
747, 1847 (Gottsche et al. 1847).
** Lejeunea luzonensis (Steph.) R.L.Zhu et M.J.Lai, Ann. Bot. Fenn. 48 (5): 376,
2011 (Wang et al. 2011). Bas.: Taxilejeunea luzonensis Steph., Hedwigia 35 (3):
134, 1896 (Stephani 1896b).
** Lejeunea macrorhyncha Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 220, 1884
(Spruce 1884).
** Lejeunea malangensis (Herzog) R.L.Zhu et Y.M.Wei, J. Bryol. 34 (4): 319, 2012
(Zhu and Wei 2012). Bas.: Trachylejeunea malangensis Herzog, Mitt. Inst. Allg.
Bot. Hamburg 7 (3): 203, 1931 (Herzog 1931a).
** Lejeunea marasmodes Spruce, Mem. Torrey Bot. Club 1 (3): 125, 1890 (Spruce 1890).
** Lejeunea morobensis (Grolle) M.A.M.Renner et Pócs, Phytotaxa 81 (1): 9, 2013
(Renner et al. 2013d). Bas.: Stenolejeunea morobensis Grolle, J. Hattori Bot. Lab.
29: 76, 1966 (Grolle 1966e).
** Lejeunea novoguineensis Schifn., Nova Acta Acad. Caes. Leop.-Carol. German.
Nat. Cur. 60 (2): 238, 1893 (Schifner 1893a).
*** Lejeunea obtusangula Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 221, 1884
(Spruce 1884).
* Lejeunea oerstediana Lindenb. et Hampe, Linnaea 24 (6): 641, 1851 [1852] (Hampe
1851a). 326
** Lejeunea paraensis Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 224, 1884
(Spruce 1884).
** Lejeunea parviloba Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 33 (7): 87, 1876
[1877] (Ångström 1876).
* Lejeunea pleidereri Sushil K.Singh, Phytotaxa 96 (1): 63, 2013 (Singh 2013).
Nom. nov. pro Otigoniolejeunea indica Steph., Sp. Hepat. (Stephani) 6: 408, 1923
(Stephani 1923).
* Lejeunea prominula Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 157, 1864 (Gottsche 1864).
326 Lejeunea oerstediana is a poorly known taxon often placed in Euosmolejeunea (=Cheilolejeunea). We
can not tell where it belongs until the type specimen is re-studied (Söderström et al. 2011a).
384
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Lejeunea pterigonia (Lehm. et Lindenb.) Mont., Ann. Sci. Nat. Bot. (sér. 2) 14:
337, 1840 (Montagne 1840a). Bas.: Jungermannia pterigonia Lehm. et Lindenb.,
Nov. Stirp. Pug. 6: 44, 1834 (Lehmann 1834).
** Lejeunea quinqueumbonata Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 230,
1884 (Spruce 1884).
** Lejeunea quinqueumbonata var. rotundata (Herzog) Sushil K.Singh, Phytotaxa 96
(1): 64, 2013 (Singh 2013). Bas.: Otigoniolejeunea quinqueumbonata var. rotundata Herzog, Rev. Bryol. Lichénol. 20 (1/2): 154, 1951 [1952] (Herzog 1951a).
** Lejeunea remotifolia Hampe ex Steph., Sp. Hepat. (Stephani) 5: 756, 1915
(Stephani 1915b).
*** Lejeunea ruthii (A.Evans) R.M.Schust., J. Hattori Bot. Lab. 25: 23, 1962 (Schuster 1962a). Bas.: Microlejeunea ruthii A.Evans, Mem. Torrey Bot. Club 8 (2): 161,
1902 (Evans 1902a).
** Lejeunea ruthii var. alata R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 1063,
1980 (Schuster 1980c).
** Lejeunea sikorae (Steph.) Steph., Bull. Soc. Roy. Bot. Belgique, Mém. 32: 120,
1893 [1894] (Renauld and Cardot 1893). Bas.: Taxilejeunea sikorae Steph., Bull.
Soc. Roy. Bot. Belgique, Compt. Rend. 32 (2): 34, 1893 [1894] (Stephani 1893e).
** Lejeunea srivastavae P.K.Verma et K.K.Rawat, Taiwania 58 (1): 8, 2013 (Verma
and Rawat 2013).
** Lejeunea stenodentata M.A.M.Renner et Pócs, Phytotaxa 81 (1): 8, 2013 (Renner
et al. 2013d). Nom. nov. pro Drepanolejeunea dentata Steph., Hedwigia 35 (3): 82,
1896 (Stephani 1896b).
*** Lejeunea subelobata Carrington et Pearson, Proc. Linn. Soc. New South Wales
(ser. 2) 2 (4): 1039, 1888 (Carrington and Pearson 1888a).
** Lejeunea subolivacea Mizut., J. Hattori Bot. Lab. 28: 121, 1965 (Mizutani 1965). Nom.
nov. pro Eulejeunea olivacea Steph., Hedwigia 29 (2): 85, 1890 (Stephani 1890b).
** Lejeunea subplana (Steph.) C.J.Bastos, J. Bryol. 36 (3): 249, 2014 (Bastos 2014). Bas.:
Trachylejeunea subplana Steph., Sp. Hepat. (Stephani) 5: 310, 1913 (Stephani 1913a).
*** Lejeunea sulphurea (Lehm. et Lindenb.) Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 217, 1884 (Spruce 1884). Bas.: Jungermannia sulphurea Lehm. et Lindenb., Nov. Stirp. Pug. 5: 14, 1833 (Lehmann 1833).
** Lejeunea terricola Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: cxci, 1889
[1890] (Spruce 1889).
** Lejeunea urbanii (Steph.) Steph., Sp. Hepat. (Stephani) 5: 766, 1915 (Stephani 1915b).
Bas.: Eulejeunea urbanii Steph., Hedwigia 27 (11/12): 301, 1888 (Stephani 1888b).
** Lejeunea venezuelana (R.M.Schust.) R.L.Zhu et W.Ye, J. Syst. Evol. 51 (4): 473,
2013 (Ye et al. 2013b). Bas.: Cyrtolejeunea venezuelana R.M.Schust., Phytologia
39 (6): 426, 1978 (Schuster 1978b).
** Lejeunea viridis R.M.Schust. ex L.Söderstr. et A.Hagborg, Phytotaxa 208 (1): 99,
2015 (Pócs et al. 2015a). Based on: Prionocolea viridissima R.M.Schust., J. Hattori
Bot. Lab. 75: 215, 1994 (Schuster 1994), nom. inval.
World checklist of hornworts and liverworts
*
*
*
*
*
*
385
Lejeunea zacuapana (Steph.) Prantl, Hedwigia 29: xviii, 1890 (Prantl 1890). Bas.:
Eulejeunea zacuapana Steph., Hedwigia 29 (2): 87, 1890 (Stephani 1890b).
Excluded from the genus
Lejeunea elegans Gottsche, Syn. Hepat. 3: 364, 1845 (Gottsche et al. 1845b). 327
Lejeunea hieronymii Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: cciii, 1889
[1890] (Spruce 1889). 328
Lejeunea proboscidea Gottsche, Mexik. Leverm.: 225, 1863 (Gottsche 1863). 329
Lejeunea scabrilora Loitl., Diagn. pl. nov.: 22, 1894 (Szyszylowicz 1894). 330
Lejeunea trochantha Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: cxcii, 1889
[1890] (Spruce 1889). 331
** Microlejeunea (Spruce) Steph., Hedwigia 27 (2): 61, 1888 (Stephani 1888a).
Bas.: Lejeunea subg. Microlejeunea Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 286, 1884 (Spruce 1884).
*** Microlejeunea acutifolia Steph., Hedwigia 35 (3): 113, 1896 (Stephani 1896b).
** Microlejeunea africana Steph., Hedwigia 27 (2): 61, 1888 (Stephani 1888a).
** Microlejeunea aligera (Mitt.) Steph., Sp. Hepat. (Stephani) 5: 827, 1915 (Stephani
1915b). Bas.: Lejeunea aligera Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 113, 1860
[1861] (Mitten 1860c).
** Microlejeunea ankasica E.W.Jones, J. Bryol. 10 (4): 394, 1979 (Jones 1979).
** Microlejeunea aphanella (Spruce) Steph., Sp. Hepat. (Stephani) 5: 816, 1915
(Stephani 1915b). Bas.: Lejeunea aphanella Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 290, 1884 (Spruce 1884).
** Microlejeunea atsuana Steph., Hedwigia 35 (3): 113, 1896 (Stephani 1896b).
** Microlejeunea bischlerae (B.M.hiers) B.M.hiers, Phytotaxa 65: 59, 2012 (hiers
et al. 2012). Bas.: Lejeunea bischlerae B.M.hiers, Cryptog. Bryol. Lichénol. 18
(3): 223, 1997 (hiers 1997b).
*** Microlejeunea bullata (Taylor) Steph., Hedwigia 29 (2): 90, 1890 (Stephani 1890b).
Bas.: Lejeunea bullata Taylor, London J. Bot. 5: 398, 1846 (Taylor 1846b).
* Microlejeunea byssoides (Gottsche) Pearson, Forh. Vidensk.-Selsk. Kristiania 1892
(8): 9, 1892 (Pearson 1892). Bas.: Lejeunea byssoides Gottsche, Abh. Naturwiss.
Vereins Bremen 7: 358, 1882 (Gottsche 1882).
** Microlejeunea capillaris (Gottsche) Steph., Sp. Hepat. (Stephani) 5: 819, 1915
(Stephani 1915b). Bas.: Lejeunea capillaris Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1:
163, 1864 (Gottsche 1864).
327
328
329
330
Lejeunea elegans is a Cheilolejeunea species (Reiner-Drehwald 2006).
Lejeunea hieronymii is a Cheilolejeunea species of uncertain status.
Lejeunea proboscidea is a Drepanolejeunea species, but Bischler (1964) did not place it to any species.
Lejeunea scabrilora was rejected from Dicranolejeunea by Kruijt (1988), but he did not refer it to any
other genus.
331 Lejeunea trochantha is a Cheilolejeunea species (Reiner-Drehwald 2006).
386
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Microlejeunea cochlearifolia Steph., Hedwigia 27 (3/4): 113, 1888 (Stephani 1888d). 332
** Microlejeunea colombiana Bischl., Nova Hedwigia 5 (1/2): 373, 1963 (Bischler et
al. 1963).
** Microlejeunea constricta (Grolle) Grolle, J. Bryol. 21 (1): 41, 1999 (Grolle and
Reiner-Drehwald 1999). Bas.: Harpalejeunea constricta Grolle, Nova Hedwigia 16:
150, 1968 (Grolle 1968d).
** Microlejeunea crenulifolia (Gottsche) Steph., Hedwigia 35 (3): 114, 1896 (Stephani
1896b). Bas.: Lejeunea crenulifolia Gottsche, Mexik. Leverm.: 227, 1863 (Gottsche
1863).
*** Microlejeunea cystifera Herzog, Memoranda Soc. Fauna Fl. Fennica 25: 68, 1950
(Herzog 1950c).
** Microlejeunea dispar Jovet-Ast, Rev. Bryol. Lichénol. 27 (3/4): 191, 1959 (Jovet-Ast
1959).
*** Microlejeunea epiphylla Bischl., Nova Hedwigia 5 (1/2): 378, 1963 (Bischler et al. 1963).
*** Microlejeunea ilicuspis (Steph.) Heinrichs, Schäf.-Verw., Pócs et S.Dong, Phytotaxa 85 (2): 52, 2013 (Dong et al. 2013). Bas.: Drepanolejeunea ilicuspis Steph.,
Sp. Hepat. (Stephani) 5: 344, 1913 (Stephani 1913a).
*** Microlejeunea ischeri (Tixier) Heinrichs, Schäf.-Verw., Pócs et S.Dong, Phytotaxa
85 (2): 52, 2013 (Dong et al. 2013). Bas.: Harpalejeunea ischeri Tixier, Trop.
Bryol. 11: 29, 1995 (Tixier 1995b).
** Microlejeunea issistipula Steph., Sp. Hepat. (Stephani) 5: 810, 1915 (Stephani
1915b).
*** Microlejeunea globosa (Spruce) Steph., Sp. Hepat. (Stephani) 5: 821, 1915 (Stephani 1915b). Bas.: Lejeunea globosa Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36:
cxciii, 1889 [1890] (Spruce 1889).
** Microlejeunea herzogiana Steph., Biblioth. Bot. 87 (2): 266, 1916 (Stephani 1916a).
** Microlejeunea indica (Udar et U.S.Awasthi) Y.M.Wei et R.L.Zhu, Phytotaxa 97
(2): 63, 2013 (Wei and Zhu 2013b). Bas.: Lejeunea indica Udar et U.S.Awasthi,
Cryptog. Bryol. Lichénol. 2 (3): 345, 1981 (Udar and Awasthi 1981).
** Microlejeunea inlata Steph., Sp. Hepat. (Stephani) 5: 811, 1915 (Stephani 1915b).
** Microlejeunea kinabaluensis (Mizut.) Grolle, J. Bryol. 21 (1): 42, 1999 (Grolle and
Reiner-Drehwald 1999). Bas.: Harpalejeunea kinabaluensis Mizut., J. Hattori Bot.
Lab. 37: 200, 1973 (Mizutani 1973).
*** Microlejeunea latitans (Hook.f. et Taylor) Heinrichs, Schäf.-Verw., Pócs et
S.Dong, Phytotaxa 85 (2): 52, 2013 (Dong et al. 2013). Bas.: Jungermannia latitans Hook.f. et Taylor, London J. Bot. 3: 399, 1844 (Hooker and Taylor 1844a).
** Microlejeunea lunulatiloba Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 1:
27, 1931 (Horikawa 1931b). 333
* Microlejeunea magnilobula Gola, Ann. Bot. (Rome) 6 (2): 274, 1907 (Gola 1907).
332 Microlejeunea cochlearifolia is conspeciic with Microlejeunea kamerunensis in Jones (1969), but he
used the younger name Microlejeunea kamerunensis Steph. 1915.
333 Microlejeunea lunulatiloba is conspeciic with Microlejeunea ulicina in Schuster (1962a), but that is
not followed by many recent authors.
World checklist of hornworts and liverworts
387
** Microlejeunea mammillosa (Mizut.) Grolle, J. Bryol. 21 (1): 42, 1999 (Grolle and
Reiner-Drehwald 1999). Bas.: Harpalejeunea mammillosa Mizut., J. Hattori Bot.
Lab. 37: 192, 1973 (Mizutani 1973).
** Microlejeunea minutissima (Mizut.) Grolle, J. Bryol. 21 (1): 42, 1999 (Grolle and
Reiner-Drehwald 1999). Bas.: Harpalejeunea minutissima Mizut., J. Hattori Bot.
Lab. 37: 202, 1973 (Mizutani 1973).
* Microlejeunea minutistipula Steph., Wiss. Ergebn. Deut. Zentr.-Afr. Exped. (1907–08),
Bot. 2: 131, 1911 (Stephani 1911a). 334
*** Microlejeunea moniliata (Mizut.) R.L.Zhu et Y.M.Wei, Cryptog. Bryol. 34 (3):
308, 2013 (Wei and Zhu 2013a). Bas.: Lejeunea moniliata Mizut., J. Hattori Bot.
Lab. 46: 357, 1979 (Mizutani 1979b).
* Microlejeunea nepalensis Steph., Sp. Hepat. (Stephani) 5: 832, 1915 (Stephani 1915b).
** Microlejeunea nyandaruensis Pócs, Polish Bot. J. 47 (1): 14, 2002 (Pócs 2002c).
** Microlejeunea oblongistipula (Gottsche) Pearson, Forh. Vidensk.-Selsk. Kristiania
1892 (8): 9, 1892 (Pearson 1892). Bas.: Lejeunea oblongistipula Gottsche, Abh.
Naturwiss. Vereins Bremen 7: 357, 1882 (Gottsche 1882).
** Microlejeunea ocellata (Herzog) Grolle, Haussknechtia 8: 60, 2001 (Grolle 2001).
Bas.: Rectolejeunea ocellata Herzog, Trans. & Proc. Roy. Soc. New Zealand 77 (2):
255, 1949 (Herzog 1949a).
* Microlejeunea ovistipula Steph., Wiss. Ergebn. Deut. Zentr.-Afr. Exped. (190708), Bot. 2: 131, 1911 (Stephani 1911a). 335
** Microlejeunea papulosa (Gottsche) Pearson, Forh. Vidensk.-Selsk. Kristiania 1892
(8): 9, 1892 (Pearson 1892). Bas.: Lejeunea papulosa Gottsche, Abh. Naturwiss.
Vereins Bremen 7: 358, 1882 (Gottsche 1882).
** Microlejeunea perpusilla (Spruce) Steph., Hedwigia 35 (3): 113, 1896 (Stephani
1896b). Bas.: Lejeunea perpusilla Spruce, J. Bot. 19: 36, 1881 (Spruce 1881b).
** Microlejeunea punctiformis (Taylor) Steph., Hedwigia 29 (2): 90, 1890 (Stephani
1890b). Bas.: Lejeunea punctiformis Taylor, London J. Bot. 5: 398, 1846 (Taylor
1846b). 336
** Microlejeunea pyriformis (Lindenb. et Gottsche) Steph., Sp. Hepat. (Stephani) 5:
824, 1915 (Stephani 1915b). Bas.: Lejeunea pyriformis Lindenb. et Gottsche, Syn.
Hepat. 5: 767, 1847 (Gottsche et al. 1847).
** Microlejeunea spinosa (Mizut.) Grolle, J. Bryol. 21 (1): 43, 1999 (Grolle and Reiner-Drehwald 1999). Bas.: Harpalejeunea spinosa Mizut., J. Hattori Bot. Lab. 37:
196, 1973 (Mizutani 1973).
*** Microlejeunea squarrosa (Steph.) Heinrichs, Schäf.-Verw., Pócs et S.Dong, Phytotaxa 85 (2): 51, 2013 (Dong et al. 2013). Bas.: Strepsilejeunea squarrosa Steph.,
Hedwigia 35 (3): 130, 1896 (Stephani 1896b).
334 Microlejeunea minutistipula is probably either Microlejeunea cochlearifolia or Microlejeunea africana
(Jones 1969).
335 Microlejeunea ovistipula is possibly conspeciic with Lejeunea hepaticola (Jones (1985, Wigginton and
Grolle 1996).
336 Microlejeunea punctiformis was treated as conspeciic with Microlejeunea ulicina by Long and Grolle
(1990), Fang et al. (1998) and Piippo (1990), but Zhu and So (2001) showed that it merits distinction.
388
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Microlejeunea strasbergii Bardat et Ah-Peng, Bryologist 114 (4): 669, 2011 (AhPeng and Bardat 2011).
*** Microlejeunea subulistipa Steph., Hedwigia 35 (3): 115, 1896 (Stephani 1896b).
* Microlejeunea szechuanensis P.C.Chen, Feddes Repert. Spec. Nov. Regni Veg. 58:
46, 1955 (Chen 1955). 337
** Microlejeunea udarii P.K.Verma et S.C.Srivast., J. Bombay Nat. Hist. Soc. 108
(2): 122, 2011 [2012] (Verma and Srivastava 2011).
*** Microlejeunea ulicina (Taylor) Steph., Hedwigia 29 (2): 88, 1890 (Stephani
1890b). Bas.: Jungermannia ulicina Taylor, Trans. Bot. Soc. Edinburgh 1 (1/4):
115, 1844 (Taylor 1844a).
* Microlejeunea usambarensis Steph., Akad. Wiss. Wien, Math.-Naturwiss. Kl.,
Denkschr. 88: 730, 1913 (Stephani 1913b).
** Microlejeunea valenciana Steph., Sp. Hepat. (Stephani) 5: 815, 1915 (Stephani 1915b).
* Microlejeunea victoriensis D.J.Carr, Proc. Roy. Soc. Victoria 117 (2): 322, 2005
(Carr 2005).
*** Microlejeunea wallichiana (Lehm.) R.L.Zhu et Y.M.Wei, Cryptog. Bryol. 34 (3):
308, 2013 (Wei and Zhu 2013a). Bas.: Jungermannia wallichiana Lehm., Nov.
Stirp. Pug. 3: 5, 1831 (Lehmann 1831).
*
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Taxilejeunea (Spruce) Steph., Hedwigia 28 (4): 262, 1889 (Stephani 1889c)
nom. rejic. Bas.: Lejeunea subg. Taxilejeunea Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 212, 1884 (Spruce 1884).
Taxilejeunea acutifolia (Steph.) Steph., Sp. Hepat. (Stephani) 5: 481, 1914
(Stephani 1914b). Bas.: Dicranolejeunea acutifolia Steph., Hedwigia 35 (3): 76,
1896 (Stephani 1896b).
Taxilejeunea antillana Steph., Sp. Hepat. (Stephani) 5: 482, 1914 (Stephani 1914b).
Taxilejeunea apiculata (Gottsche) J.B.Jack et Steph., Hedwigia 31 (1): 13, 1892
(Jack and Stephani 1892). Bas.: Omphalanthus apiculatus Gottsche, Ann. Sci. Nat.
Bot. (sér. 5) 1: 149, 1864 (Gottsche 1864).
Taxilejeunea argentina Steph., Sp. Hepat. (Stephani) 5: 482, 1914 (Stephani 1914b).
Taxilejeunea arsenii Steph., Sp. Hepat. (Stephani) 6: 400, 1923 (Stephani 1923).
Taxilejeunea auriculata Steph., Sp. Hepat. (Stephani) 5: 459, 1914 (Stephani 1914b).
Taxilejeunea berteroana Steph., Sp. Hepat. (Stephani) 5: 483, 1914 (Stephani
1914b).
Taxilejeunea beyrichiana Steph., Sp. Hepat. (Stephani) 5: 460, 1914 (Stephani 1914b).
Taxilejeunea biapiculata Steph., Sp. Hepat. (Stephani) 5: 460, 1914 (Stephani 1914b).
Taxilejeunea boliviana Steph., Biblioth. Bot. 87 (2): 260, 1916 (Stephani 1916a).
Taxilejeunea brasiliensis Steph., Hedwigia 35 (3): 132, 1896 (Stephani 1896b).
Taxilejeunea coilantha Herzog, Rev. Bryol. Lichénol. 20 (1/2): 159, 1951 [1952]
(Herzog 1951a).
337 Microlejeunea szechuanensis may be conspeciic with Microlejeunea punctiformis.
World checklist of hornworts and liverworts
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389
Taxilejeunea compressiuscula Steph., Sp. Hepat. (Stephani) 5: 501, 1914 (Stephani
1914b).
Taxilejeunea convoluta Herzog, Biblioth. Bot. 88: 31, 1920 [1921] (Herzog 1920).
Taxilejeunea cuervi (Gottsche) Steph., Sp. Hepat. (Stephani) 5: 486, 1914 (Stephani
1914b). Bas.: Omphalanthus cuervi Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 147,
1864 (Gottsche 1864).
Taxilejeunea cuneistipula Steph., Sp. Hepat. (Stephani) 6: 401, 1923 (Stephani 1923).
Taxilejeunea cuspidata Steph., Biblioth. Bot. 87 (2): 260, 1916 (Stephani 1916a).
Taxilejeunea decurrens Steph., Sp. Hepat. (Stephani) 5: 487, 1914 (Stephani 1914b).
Taxilejeunea delexa Steph., Sp. Hepat. (Stephani) 5: 502, 1914 (Stephani 1914b).
Taxilejeunea densilora A.Evans, Bull. Torrey Bot. Club 48 (4): 121, 1921 (Evans 1921b).
Taxilejeunea diaphana (Lehm.) Steph., Sp. Hepat. (Stephani) 5: 463, 1914
(Stephani 1914b). Bas.: Omphalanthus diaphanus Lehm., Nov. Stirp. Pug. 10: 12,
1857 (Lehmann 1857).
Taxilejeunea dissitifolia Steph., Symb. Antill. 2: 472, 1901 (Stephani 1901f).
Taxilejeunea eggersiana Schifn., Bot. Jahrb. Syst. 23 (5): 579, 1897 (Schifner
1897). Based on: Taxilejeunea eggersiana Steph., Hedwigia 27 (11/12): 285, 1888
(Stephani 1888c), nom. inval.
Taxilejeunea elobulata Sim, Trans. Roy. Soc. South Africa 15 (1): 66, 1926 (Sim 1926).
Taxilejeunea issistipula Steph., Sp. Hepat. (Stephani) 5: 488, 1914 (Stephani
1914b).
Taxilejeunea foliicola Steph., Sp. Hepat. (Stephani) 5: 466, 1914 (Stephani 1914b).
Taxilejeunea furcicornuta Grolle, J. Bryol. 8 (1): 93, 1974 (Grolle 1974a).
Taxilejeunea fuscorufa Steph., Hedwigia 35 (3): 133, 1896 (Stephani 1896b).
Taxilejeunea galapagensis Onr., Misc. Bryol. Lichenol. 9 (6): 117, 1982 (Onraedt 1982).
Taxilejeunea ghatensis P.K.Verma et S.C.Srivast., Proc. Natl. Acad. Sci. India, B 77
(2): 211, 2007 (Verma and Srivastava 2007).
Taxilejeunea giulianettii Steph., Sp. Hepat. (Stephani) 5: 502, 1914 (Stephani 1914b).
Taxilejeunea gomphocalyx Herzog, Beih. Bot. Centralbl. 61B (3): 580, 1942 (Herzog
1942d).
Taxilejeunea grandifolia Steph., Biblioth. Bot. 87 (2): 261, 1916 (Stephani 1916a).
Taxilejeunea grandistipula Steph., Sp. Hepat. (Stephani) 5: 504, 1914 (Stephani
1914b).
Taxilejeunea hamatifolia Steph., Biblioth. Bot. 87 (2): 261, 1916 (Stephani 1916a).
Taxilejeunea himalayensis Herzog, Ann. Bryol. 12: 86, 1939 (Herzog 1939b).
Taxilejeunea immersa Eifrig, Monogr. Stud. Indomal. Art. Taxilejeunea: 96, 1937
(Eifrig 1937).
Taxilejeunea irregularis Steph., Sp. Hepat. (Stephani) 5: 490, 1914 (Stephani 1914b).
Taxilejeunea jamaicensis A.Evans, Bull. Torrey Bot. Club 48 (4): 117, 1921 (Evans
1921b).
Taxilejeunea jeringii Steph., Hedwigia 35 (3): 134, 1896 (Stephani 1896b).
Taxilejeunea killipii Herzog, Feddes Repert. Spec. Nov. Regni Veg. 57 (1/2): 190,
1955 (Herzog 1955).
390
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Taxilejeunea laevis (Gottsche) Steph., Hedwigia 44 (4): 229, 1905 (Stephani
1905a). Bas.: Omphalanthus laevis Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 148,
1864 (Gottsche 1864).
Taxilejeunea langiana Pearson, Ann. Bryol. 4: 102, 1931 (Pearson 1931b).
Taxilejeunea lindbergiana Steph., Sp. Hepat. (Stephani) 5: 491, 1914 (Stephani 1914b).
Taxilejeunea linguifolia Steph., Sp. Hepat. (Stephani) 5: 471, 1914 (Stephani 1914b).
Taxilejeunea maxima Steph., Sp. Hepat. (Stephani) 5: 473, 1914 (Stephani 1914b).
Taxilejeunea mexicana Steph., Sp. Hepat. (Stephani) 6: 402, 1923 (Stephani 1923).
Taxilejeunea microstipula Steph., Sp. Hepat. (Stephani) 5: 493, 1914 (Stephani 1914b).
Taxilejeunea mucronata Steph., Sp. Hepat. (Stephani) 5: 473, 1914 (Stephani 1914b).
Taxilejeunea multilora Steph., Hedwigia 35 (3): 135, 1896 (Stephani 1896b).
Taxilejeunea muscicola Steph., Biblioth. Bot. 87 (2): 263, 1916 (Stephani 1916a).
Taxilejeunea nilgiriensis P.K.Verma et S.C.Srivast., Proc. Natl. Acad. Sci. India, B
77 (2): 207, 2007 (Verma and Srivastava 2007).
Taxilejeunea nymannii Steph., Sp. Hepat. (Stephani) 5: 507, 1914 (Stephani 1914b).
Taxilejeunea obtusifolia Steph., Sp. Hepat. (Stephani) 5: 474, 1914 (Stephani 1914b).
Taxilejeunea papulilora Steph., Sp. Hepat. (Stephani) 5: 494, 1914 (Stephani 1914b).
Taxilejeunea parvibracteata Steph., Hedwigia 35 (3): 136, 1896 (Stephani 1896b).
Taxilejeunea parvistipula Steph., Sp. Hepat. (Stephani) 6: 403, 1923 (Stephani 1923).
Taxilejeunea paucidens Steph., Biblioth. Bot. 87 (2): 263, 1916 (Stephani 1916a).
Taxilejeunea pendula Steph., Biblioth. Bot. 87 (2): 263, 1916 (Stephani 1916a).
Taxilejeunea peruviana Steph., Sp. Hepat. (Stephani) 5: 475, 1914 (Stephani 1914b).
Taxilejeunea planilobula Herzog, Rev. Bryol. Lichénol. 20 (1/2): 160, 1951 [1952]
(Herzog 1951a).
Taxilejeunea pulchrilora Pearson, Ark. Bot. 19 (5): 15, 1924 (Pearson 1924b).
Taxilejeunea pusilla Steph., Biblioth. Bot. 87 (2): 264, 1916 (Stephani 1916a).
Taxilejeunea renistipula (Lindenb.) Steph., Hedwigia 29 (3): 142, 1890 (Stephani
1890d). Bas.: Omphalanthus renistipulus Lindenb., Syn. Hepat. 3: 308, 1845 (Gottsche
et al. 1845b).
Taxilejeunea rufescens Steph., Biblioth. Bot. 87 (2): 264, 1916 (Stephani 1916a).
Taxilejeunea serpillifolioides (Raddi) D.P.Costa, J. Bryol. 31 (4): 230, 2009 (Costa
2009). Bas.: Jungermannia serpillifolioides Raddi, Critt. Brasil.: 17, 1822 (Raddi 1822).
Taxilejeunea setchellii Pearson, Amer. Samoa: 146, 1924 (Pearson 1924a).
Taxilejeunea speciosa Herzog, Feddes Repert. Spec. Nov. Regni Veg. 57 (1/2): 188,
1955 (Herzog 1955).
Taxilejeunea splendida Eifrig, Monogr. Stud. Indomal. Art. Taxilejeunea: 88, 1937
(Eifrig 1937).
Taxilejeunea stephanii Eifrig, Monogr. Stud. Indomal. Art. Taxilejeunea: 90, 1937
(Eifrig 1937). Nom. nov. pro Hygrolejeunea nymanii Steph., Sp. Hepat. (Stephani)
5: 564, 1914 (Stephani 1914b).
Taxilejeunea steyermarkii H.Rob., Phytologia 34 (1): 67, 1976 (Robinson 1976a).
Taxilejeunea surinamensis (Lindenb. et Gottsche) Steph., Hedwigia 29 (3): 142,
1890 (Stephani 1890d). Bas.: Omphalanthus surinamensis Lindenb. et Gottsche,
Linnaea 24 (6): 628, 1851 [1852] (Lindenberg and Gottsche 1851a).
World checklist of hornworts and liverworts
391
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*
Taxilejeunea suringarii Steph., Sp. Hepat. (Stephani) 5: 479, 1914 (Stephani 1914b).
Taxilejeunea tenerrima Steph., Sp. Hepat. (Stephani) 6: 406, 1923 (Stephani 1923).
Taxilejeunea tenuiplica Steph., Sp. Hepat. (Stephani) 5: 480, 1914 (Stephani 1914b).
Taxilejeunea tjibodensis (Steph.) Eifrig, Monogr. Stud. Indomal. Art. Taxilejeunea:
91, 1937 (Eifrig 1937). Bas.: Hygrolejeunea tjibodensis Steph., Sp. Hepat. (Stephani)
5: 571, 1914 (Stephani 1914b).
** Taxilejeunea tonduzana Steph., Sp. Hepat. (Stephani) 5: 498, 1914 (Stephani 1914b).
* Taxilejeunea uleana Steph., Hedwigia 35 (3): 136, 1896 (Stephani 1896b).
* Taxilejeunea umbonata Steph., Sp. Hepat. (Stephani) 5: 481, 1914 (Stephani 1914b).
* Taxilejeunea urbanii Steph., Symb. Antill. (Urban) 3 (2): 278, 1902 (Stephani 1902e).
** Taxilejeunea vallis-gratiae Steph., Hedwigia 35 (3): 137, 1896 (Stephani 1896b).
**
subtrib. Lepidolejeuneinae Gradst.
*** Lepidolejeunea R.M.Schust., Beih. Nova Hedwigia 9: 139, 1963 (Schuster
1963a). 338
** subg. Kingiolejeunea (H.Rob.) R.M.Schust., Phytologia 45 (5): 425, 1980
(Schuster 1980b). Bas.: Kingiolejeunea H.Rob., Bryologist 70 (1): 53, 1967
(Robinson 1967).
** Lepidolejeunea auriculata Schäf.-Verw. et Heinrichs, Taxon 64 (2): 224, 2015
(Heinrichs et al. 2015).
*** Lepidolejeunea cordiissa (Taylor) M.E.Reiner, Nova Hedwigia 83 (3/4): 478, 2006
(Reiner-Drehwald 2006). Bas.: Lejeunea cordiissa Taylor, London J. Bot. 5: 395,
1846 (Taylor 1846b).
*** Lepidolejeunea grossepapulosa (Steph.) Piippo, Acta Bot. Fenn. 132: 49, 1986 (Piippo 1986a). Bas.: Prionolejeunea grossepapulosa Steph., Sp. Hepat. (Stephani) 5:
220, 1913 (Stephani 1913a).
*** Lepidolejeunea involuta (Gottsche) Grolle, J. Hattori Bot. Lab. 55: 504, 1984
(Grolle 1984a). Bas.: Lejeunea involuta Gottsche, Syn. Hepat. 3: 350, 1845
(Gottsche et al. 1845b).
** subg. Lepidolejeunea
*** Lepidolejeunea bidentula (Steph.) R.M.Schust., Phytologia 45 (5): 425, 1980
(Schuster 1980b). Bas.: Pycnolejeunea bidentula Steph., Bot. Centralbl. 60 (4):
107, 1894 (Jack and Stephani 1894).
*** Lepidolejeunea bidentula var. novae-caledoniae Piippo, Acta Bot. Fenn. 132: 26,
1986 (Piippo 1986a).
*** Lepidolejeunea falcata (Herzog) R.M.Schust., Beih. Nova Hedwigia 9: 139, 1963
(Schuster 1963a). Bas.: Hygrolejeunea falcata Herzog, Ark. Bot. (n.ser.) 3 (3): 57,
1953 (Herzog 1953a).
338 Lepidolejeunea includes Kingiolejeunea, but one taxon has neither been transferred nor synonymized.
It is listed in the “Names in genera not currently accepted” section below.
392
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Lepidolejeunea graefei (J.B.Jack et Steph.) R.M.Schust., Phytologia 45 (5): 425,
1980 (Schuster 1980b). Bas.: Archilejeunea graefei J.B.Jack et Steph., Bot. Centralbl. 60 (4): 104, 1894 (Jack and Stephani 1894).
*** Lepidolejeunea integristipula (J.B.Jack et Steph.) R.M.Schust., Phytologia 45 (5):
425, 1980 (Schuster 1980b). Bas.: Pycnolejeunea integristipula J.B.Jack et Steph.,
Bot. Centralbl. 60 (4): 107, 1894 (Jack and Stephani 1894).
** subg. Perilejeunea (Kachroo et R.M.Schust.) R.M.Schust., Phytologia 45 (5):
424, 1980 (Schuster 1980b). Bas.: Pycnolejeunea subg. Perilejeunea Kachroo et
R.M.Schust., J. Linn. Soc., Bot. 56 (368): 493, 1961 (Kachroo and Schuster 1961).
** Lepidolejeunea cuspidata (Gottsche) Heinrichs et Schäf.-Verw., Taxon 64 (2): 224,
2015 (Heinrichs et al. 2015). Bas.: Lejeunea cuspidata Gottsche, Syn. Hepat. 3:
351, 1845 (Gottsche et al. 1845b).
*** Lepidolejeunea delessertii (Nees et Mont.) Grolle, J. Hattori Bot. Lab. 55: 505,
1984 (Grolle 1984a). Bas.: Lejeunea delessertii Nees et Mont., Ann. Sci. Nat. Bot.
(sér. 2) 19: 260, 1843 (Montagne 1843).
*** Lepidolejeunea eluta (Nees) R.M.Schust., Beih. Nova Hedwigia 9: 139, 1963
(Schuster 1963a). Bas.: Jungermannia eluta Nees, Fl. Bras. (Martius) 1 (1): 362,
1833 (Nees 1833a).
*** Lepidolejeunea serrulata (Steph.) Grolle, J. Hattori Bot. Lab. 55: 505, 1984 (Grolle
1984a). Bas.: Trachylejeunea serrulata Steph., Sp. Hepat. (Stephani) 5: 300, 1913
(Stephani 1913a).
*** Lepidolejeunea sullivantii (Gottsche) M.E.Reiner, Nova Hedwigia 83 (3/4): 479,
2006 (Reiner-Drehwald 2006). Bas.: Lejeunea sullivantii Gottsche, Mexik. Leverm.: 196, 1863 (Gottsche 1863).
Incertae sedis
*** Lepidolejeunea borneensis (Steph.) R.M.Schust., Phytologia 45 (5): 425, 1980
(Schuster 1980b). Bas.: Hygrolejeunea borneensis Steph., Sp. Hepat. (Stephani) 5:
557, 1914 (Stephani 1914b).
*** Lepidolejeunea longilobula (Amakawa) Xiao L.He, Acta Bot. Fenn. 163: 59, 1999
(He 1999). Bas.: Pycnolejeunea longilobula Amakawa, J. Jap. Bot. 40 (10): 307,
1965 (Amakawa 1965).
*** Otolejeunea Grolle et Tixier, Nova Hedwigia 32: 609, 1980 (Tixier 1980b).
*** subg. Allorgella (Tixier) Grolle, Haussknechtia 2: 53, 1985 (Grolle 1985a). Bas.:
Allorgella Tixier, Nova Hedwigia 32: 612, 1980 (Tixier 1980b).
*** Otolejeunea australiensis B.M.hiers, Brittonia 44 (2): 162, 1992 (hiers 1992a).
*** Otolejeunea hoana (Tixier) Grolle, Haussknechtia 2: 54, 1985 [1986] (Grolle
1985a). Bas.: Allorgella hoana Tixier, Nova Hedwigia 32: 615, 1980 (Tixier 1980b).
*** Otolejeunea rabenorii Tixier, Nova Hedwigia 46 (3/4): 376, 1988 (Tixier 1988).
*** Otolejeunea schmidii (Tixier) Grolle, Haussknechtia 2: 54, 1985 [1986] (Grolle
1985a). Bas.: Allorgella schmidii Tixier, Nova Hedwigia 32: 613, 1980 (Tixier 1980b).
World checklist of hornworts and liverworts
393
*** Otolejeunea schnellii (Tixier) R.L.Zhu et M.L.So, Ann. Bot. Fenn. 34 (4): 287,
1997 (Zhu and So 1997a). Bas.: Allorgella schnellii Tixier, Cryptog. Bryol. Lichenol. 16 (3): 230, 1995 (Tixier 1995a).
*** Otolejeunea semperiana (Steph.) Grolle, Haussknechtia 2: 53, 1985 [1986] (Grolle 1985a). Bas.: Prionolejeunea semperiana Steph., Sp. Hepat. (Stephani) 5: 227,
1913 (Stephani 1913a).
** Otolejeunea subana Pocs, Acta Acad. Ped. Agr., Sect. Biol. 25: 50, 2004 (Pocs 2004).
*** Otolejeunea zantenii Grolle, Haussknechtia 2: 54, 1985 [1986] (Grolle 1985a).
*** subg. Otolejeunea
*** Otolejeunea moniliata Grolle, Nova Hedwigia 32: 609, 1980 (Tixier 1980b). World checklist of hornworts and liverworts 353
*** subg. Phoxolejeunea Grolle, Haussknechtia 2: 49, 1985 (Grolle 1985a).
** Otolejeunea philippinensis R.L.Zhu et M.L.So, Syst. Bot. 23 (2): 231, 1998 (Zhu
and So 1998).
*** Otolejeunea streimannii Grolle, Haussknechtia 2: 49, 1985 [1986] (Grolle 1985a).
*** Rectolejeunea A.Evans, Bull. Torrey Bot. Club 33 (1): 8, 1906 (Evans 1906a).
* Rectolejeunea colombiana R.M.Schust., J. Hattori Bot. Lab. 89: 146, 2000 (Schuster 2000c).
*** Rectolejeunea emarginulilora (Schifn.) A.Evans, Bull. Torrey Bot. Club 33 (1): 14,
1906 (Evans 1906a). Bas.: Cheilolejeunea emarginulilora Schifn., Bot. Jahrb. Syst.
23 (5): 585, 1897 (Schifner 1897).
*** Rectolejeunea lagelliformis A.Evans, Bull. Torrey Bot. Club 33 (1): 9, 1906 (Evans
1906a).
* Rectolejeunea lagelliformis subsp. hamata R.M.Schust., J. Hattori Bot. Lab. 89:
137, 2000 (Schuster 2000c).
*** Rectolejeunea queenslandica (B.M.hiers) Xiao L.He, Ann. Bot. Fenn. 34 (2): 129,
1997 (He 1997). Bas.: Lepidolejeunea queenslandica B.M.hiers, Mem. New York
Bot. Gard. 45: 556, 1987 (hiers 1987a).
** Rectolejeunea truncatilobula C.J.Bastos, J. Bryol. 34 (2): 144, 2012 (Bastos 2012a).
*** Rectolejeunea versifolia (Schifn.) L.Söderstr. et A.Hagborg, Phytotaxa 220 (2):
188, 2015 (Söderström et al. 2015a). Bas.: Cheilolejeunea versifolia Schifn., Bot.
Jahrb. Syst. 23 (5): 597, 1897 (Schifner 1897).
*
Excluded from the genus
Rectolejeunea lindenbergii Steph., Sp. Hepat. (Stephani) 5: 689, 1914 (Stephani
1914b). 339
339 Rectolejeunea lindenbergii is a Lejeunea species (Reiner-Drehwald and Grolle 2012; non Lejeunea
lindenbergii Gottsche). However, it may be conspeciic with something else.
394
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Rectolejeunea lindigiana Steph., Sp. Hepat. (Stephani) 5: 690, 1914 (Stephani
1914b). 340
Rectolejeunea pachyderma R.M.Schust., J. Hattori Bot. Lab. 89: 148, 2000 (Schuster
2000c). 341
subtrib. Leptolejeuneinae Heinrichs et Schäf.-Verw.
*** Leptolejeunea (Spruce) Steph., Hedwigia 30 (6): 270, 1891 (Stephani 1891c).
Bas.: Lejeunea subg. Leptolejeunea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:
193, 1884 (Spruce 1884).
*** Leptolejeunea amphiophthalma Zwickel, Ann. Bryol. 6: 117, 1933 (Zwickel 1933).
** Leptolejeunea apiculata (Horik.) S.Hatt., J. Hattori Bot. Lab. 5: 46, 1951 (Hattori
1951d). Bas.: Drepanolejeunea apiculata Horik., J. Sci. Hiroshima Univ., Ser. B,
Div. 2, Bot. 2: 266, 1934 (Horikawa 1934).
** Leptolejeunea arunachalensis Sudipa Das et D.K.Singh, J. Jap. Bot. 83 (6): 343,
2008 (Das and Singh 2008).
*** Leptolejeunea astroidea (Mitt.) Steph., Sp. Hepat. (Stephani) 5: 363, 1913 (Stephani 1913a). Bas.: Lejeunea astroidea Mitt., Trans. Linn. Soc. London 23 (1): 58,
1860 (Mitten 1860a).
** Leptolejeunea australis Steph., Sp. Hepat. (Stephani) 5: 389, 1913 (Stephani
1913a). Based on: Leptolejeunea australis Steph., Hedwigia 28 (3): 173, 1889
(Stephani 1889d), nom. inval.
** Leptolejeunea balansae Steph., Hedwigia 35 (3): 105, 1896 (Stephani 1896b).
* Leptolejeunea borneensis Herzog, Flora 135: 394, 1942 (Herzog 1942c). 342
** Leptolejeunea brasiliensis Bischl., Nova Hedwigia 17: 301, 1969 (Bischler 1969).
* Leptolejeunea convexistipa Bischl., Nova Hedwigia 17: 325, 1969 (Bischler 1969).
* Leptolejeunea curvatifolia Steph., Sp. Hepat. (Stephani) 6: 398, 1923 (Stephani
1923). 343
** Leptolejeunea denticulata Steph., Sp. Hepat. (Stephani) 5: 389, 1913 (Stephani
1913a). Based on: Leptolejeunea denticulata Steph., Hedwigia 28 (3): 174, 1889
(Stephani 1889d), nom. inval.
** Leptolejeunea dentistipula Steph., Sp. Hepat. (Stephani) 5: 379, 1913 (Stephani 1913a).
** Leptolejeunea diversilobulata Bischl., Nova Hedwigia 17: 313, 1969 (Bischler 1969).
** Leptolejeunea dolabriformis Pearson, J. Linn. Soc., Bot. 46 (305): 37, 1922 (Pearson
1922b).
340
341
342
343
Rectolejeunea lindigiana is a Lejeunea species of doubtful status (Reiner-Drehwald and Grolle 2012).
Rectolejeunea pachyderma is a Lejeunea species of doubtful status (Reiner-Drehwald and Grolle 2012).
Leptolejeunea borneensis may be conspeciic with Leptolejeunea vitrea.
Leptolejeunea curvatifolia is possibly conspeciic with Drepanolejeunea thwaitesiana, but the type
specimen was burned in B and the identity can not be ascertained (Grolle and Piippo 1984).
World checklist of hornworts and liverworts
395
*** Leptolejeunea elliptica (Lehm. et Lindenb.) Besch., Rev. Bryol. 19 (1): 14, 1892
(Bescherelle 1892). Bas.: Jungermannia elliptica Lehm. et Lindenb., Nov. Stirp.
Pug. 5: 13, 1833 (Lehmann 1833).
* Leptolejeunea emarginata (Horik.) S.Hatt., J. Hattori Bot. Lab. 5: 46, 1951 (Hattori 1951d). Bas.: Drepanolejeunea emarginata Horik., J. Sci. Hiroshima Univ.,
Ser. B, Div. 2, Bot. 2: 267, 1934 (Horikawa 1934). 344
*** Leptolejeunea epiphylla (Mitt.) Steph., Sp. Hepat. (Stephani) 5: 380, 1913 (Stephani 1913a). Bas.: Lejeunea epiphylla Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 118,
1860 [1861] (Mitten 1860c).
*** Leptolejeunea exocellata (Spruce) A.Evans, Bull. Torrey Bot. Club 29 (8): 498,
1902 (Evans 1902c). Bas.: Lejeunea exocellata Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 195, 1884 (Spruce 1884).
** Leptolejeunea foliicola Steph., Hedwigia 35 (3): 106, 1896 (Stephani 1896b).
** Leptolejeunea integristipula Steph., Sp. Hepat. (Stephani) 6: 398, 1923 (Stephani 1923).
* Leptolejeunea jamaicensis R.M.Schust., J. Elisha Mitchell Sci. Soc. 83 (4): 229,
1967 (Schuster 1967a).
** Leptolejeunea lancifolia Steph., Sp. Hepat. (Stephani) 5: 382, 1913 (Stephani
1913a). Nom. nov. pro Lejeunea lancifolia Mitt., Fl. vit.: 415, 1871 [1873] (Mitten
1871), nom. illeg.
** Leptolejeunea latifolia Herzog, Memoranda Soc. Fauna Fl. Fennica 26: 58, 1950
[1951] (Herzog 1950b).
** Leptolejeunea lepinii Steph., Sp. Hepat. (Stephani) 5: 383, 1913 (Stephani 1913a).
** Leptolejeunea ligulata Herzog, Flora 135: 429, 1942 (Herzog 1942c).
*** Leptolejeunea maculata (Mitt.) Schifn., Consp. Hepat. Arch. Ind.: 275, 1898
(Schifner 1898b). Bas.: Lejeunea maculata Mitt., J. Proc. Linn. Soc., Bot. 5 (18):
118, 1860 [1861] (Mitten 1860c).
* Leptolejeunea massartiana Schifn. ex Herzog, Flora 135: 421, 1942 (Herzog 1942c). 345
** Leptolejeunea micronesica Inoue et H.A.Mill., Bull. Natl. Sci. Mus. Tokyo (n.ser.)
8 (2): 149, 1965 (Inoue and Miller 1965).
** Leptolejeunea minima Herzog, Memoranda Soc. Fauna Fl. Fennica 26: 60, 1950
[1951] (Herzog 1950b).
** Leptolejeunea mirikana M.Dey et D.K.Singh, Taiwania 55 (4): 355, 2010 (Dey
and Singh 2010).
*** Leptolejeunea moniliata Steph., Sp. Hepat. (Stephani) 5: 371, 1913 (Stephani 1913a).
*** Leptolejeunea obfuscata (Spruce) Steph., Sp. Hepat. (Stephani) 5: 373, 1913
(Stephani 1913a). Bas.: Lejeunea obfuscata Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 579, 1885 (Spruce 1885).
* Leptolejeunea punctata Herzog, Flora 135: 432, 1942 (Herzog 1942c).
344 Leptolejeunea emarginata is possibly conspeciic with Leptolejeunea apiculata (Zhu and So 2001).
345 Leptolejeunea massartiana is very possibly conspeciic with Leptolejeunea elliptica judging from the
original description and illustration (Söderström et al. 2010a).
396
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Leptolejeunea radicosa (Nees ex Mont.) Grolle, J. Hattori Bot. Lab. 45: 178, 1979
(Grolle 1979c). Bas.: Lejeunea radicosa Nees ex Mont., Hist. Phys. Cuba, Bot., Pl.
Cell.: 475, 1842 (Montagne 1842a).
* Leptolejeunea renneri Herzog, Flora 135: 422, 1942 (Herzog 1942c).
* Leptolejeunea revoluta P.C.Chen, Feddes Repert. Spec. Nov. Regni Veg. 58: 44,
1955 (Chen 1955). 346
** Leptolejeunea rosulans Steph., Sp. Hepat. (Stephani) 5: 390, 1913 (Stephani
1913a). Based on: Leptolejeunea rosulans Steph., Hedwigia 28 (3): 174, 1889
(Stephani 1889d), nom. inval.
** Leptolejeunea serratifolia Schifn., Bot. Jahrb. Syst. 23 (5): 594, 1897 (Schifner 1897).
** Leptolejeunea serrulata Herzog, Flora 135: 426, 1942 (Herzog 1942c).
** Leptolejeunea spinistipula (Mizut.) Xiao L.He, Ann. Bot. Fenn. 34 (2): 127, 1997
(He 1997). Bas.: Pycnolejeunea spinistipula Mizut., J. Hattori Bot. Lab. 33: 255,
1970 (Mizutani 1970).
** Leptolejeunea subdentata Schifn. ex Herzog, Flora 135: 403, 1942 (Herzog 1942c).
*** Leptolejeunea subrotundifolia Herzog, Flora 135: 424, 1942 (Herzog 1942c).
*** Leptolejeunea tridentata Bischl., Nova Hedwigia 17: 335, 1969 (Bischler 1969).
* Leptolejeunea trigonostipa (Spruce) Steph., Sp. Hepat. (Stephani) 5: 376, 1913
(Stephani 1913a). Bas.: Lejeunea trigonostipa Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 197, 1884 (Spruce 1884).
** Leptolejeunea tripuncta (Mitt.) Steph., Sp. Hepat. (Stephani) 5: 388, 1913 (Stephani 1913a). Bas.: Lejeunea tripuncta Mitt., Fl. vit.: 415, 1871 [1873] (Mitten 1871).
** Leptolejeunea truncatifolia Steph., Sp. Hepat. (Stephani) 5: 388, 1913 (Stephani
1913a).
* Leptolejeunea udarii M.Dey et D.K.Singh, Taiwania 55 (4): 359, 2010 (Dey and
Singh 2010). 347
*** Leptolejeunea vitrea (Nees) Schifn., Hepat. (Engl.-Prantl): 126, 1893 (Schifner 1893b).
Bas.: Jungermannia vitrea Nees, Enum. Pl. Crypt. Javae: 56, 1830 (Nees 1830).
**
subtrib. Pycnolejeuneinae Heinrichs et Schäf.Verw.
** Pycnolejeunea (Spruce) Schifn., Hepat. (Engl.-Prantl): 124, 1893 (Schifner
1893b). Bas.: Lejeunea subg. Pycnolejeunea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 246, 1884 (Spruce 1884).
* Pycnolejeunea anotomensis Tixier, Bull. Soc. Hist. Nat. Afrique N. 63: 10, 1972
(Tixier 1972a). 348
** Pycnolejeunea borneensis Steph., Sp. Hepat. (Stephani) 5: 632, 1914 (Stephani
1914b).
346 Leptolejeunea revoluta may be conspeciic with Leptolejeunea elliptica (Zhu and So 2001).
347 Leptolejeunea udarii is possibly conspeciic with Leptolejeunea latifolia (He et al. 2013).
348 Pycnolejeunea anotomensis is a Cheilolejeunea species.
World checklist of hornworts and liverworts
*
*
***
*
***
***
**
**
*
***
***
**
*
***
**
**
*
397
Pycnolejeunea cavistipula (Steph.) Mizut., J. Hattori Bot. Lab. 36: 161, 1972
[1973] (Mizutani 1972a). Bas.: Strepsilejeunea cavistipula Steph., Hedwigia 35 (3):
128, 1896 (Stephani 1896b). 349
Pycnolejeunea connivens Schifn. ex P.Syd., Just’s Bot. Jahresber. 19: 246, 1894
(Sydow 1894). Based on: Pycnolejeunea connivens Gottsche ex Schifn., Leberm.,
Forschungsr. Gazelle 4 (4): 32, 1890 (Schifner 1890), nom. inval.
Pycnolejeunea contigua (Nees) Grolle, J. Hattori Bot. Lab. 45: 179, 1979 (Grolle
1979c). Bas.: Jungermannia contigua Nees, Fl. Bras. (Martius) 1 (1): 360, 1833
(Nees 1833a).
Pycnolejeunea convexifolia (Mitt.) Steph., Sp. Hepat. (Stephani) 5: 636, 1914
(Stephani 1914b). Bas.: Lejeunea convexifolia Mitt., Fl. vit.: 416, 1871 [1873]
(Mitten 1871). 350
Pycnolejeunea decurviloba Steph., Hedwigia 35 (3): 125, 1896 (Stephani 1896b).
Pycnolejeunea densistipula (Lehm. et Lindenb.) Steph., Sp. Hepat. (Stephani) 5:
602, 1914 (Stephani 1914b). Bas.: Lejeunea densistipula Lehm. et Lindenb., Nov.
Stirp. Pug. 7: 20, 1838 (Lehmann 1838).
Pycnolejeunea gradsteinii Ilk.-Borg., Bol. Inst. Bot. (São Paulo) 21 (1): 1, 2011
(Ilkiu-Borges 2011).
Pycnolejeunea grandiocellata Steph., Bot. Tidsskr. 24 (3): 279, 1902 (Stephani
1902b).
Pycnolejeunea grossiloba Steph., Sp. Hepat. (Stephani) 5: 629, 1914 (Stephani
1914b). 351
Pycnolejeunea macroloba (Nees et Mont.) Schifn., Hepat. (Engl.-Prantl): 124,
1893 (Schifner 1893b). Bas.: Lejeunea macroloba Nees et Mont., Ann. Sci. Nat.
Bot. (sér. 2) 19: 260, 1843 (Montagne 1843).
Pycnolejeunea monophthalma (R.M.Schust.) Xiao L.He, Acta Bot. Fenn. 163: 52,
1999 (He 1999). Bas.: Trachylejeunea monophthalma R.M.Schust., Bull. Torrey
Bot. Club 97 (6): 345, 1970 [1971] (Schuster 1970b).
Pycnolejeunea novae-caledoniae (Steph.) Horik., Acta Phytotax. Geobot. 13: 214,
1943 (Horikawa 1943). Bas.: Archilejeunea novae-caledoniae Steph., Sp. Hepat.
(Stephani) 4: 729, 1911 (Stephani 1911e).
Pycnolejeunea palmicola Steph., Sp. Hepat. (Stephani) 6: 413, 1923 (Stephani 1923). 352
Pycnolejeunea papillosa Xiao L.He, Acta Bot. Fenn. 163: 55, 1999 (He 1999).
Pycnolejeunea porrectilobula C.J.Bastos et O.Yano, Nova Hedwigia 74 (3/4): 440,
2002 (Bastos and Yano 2002).
Pycnolejeunea retusa R.M.Schust., J. Hattori Bot. Lab. 100: 402, 2006 (Schuster 2006).
Pycnolejeunea schlimiana Steph., Sp. Hepat. (Stephani) 5: 615, 1914 (Stephani
1914b). 353
349 Pycnolejeunea cavistipula is possibly a Cheilolejeunea species.
350 Pycnolejeunea convexifolia is possibly conspeciic with Cheilolejeunea imbricata.
351 Pycnolejeunea grossiloba is a Cheilolejeunea species, but its taxonomic status is unclear (Grolle and
Piippo 1984).
352 Pycnolejeunea palmicola is not a Pycnolejeunea species.
353 Pycnolejeunea schlimiana is not a Pycnolejeunea species.
398
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Pycnolejeunea schwaneckei (Steph.) Schifn. ex P.Syd., Just’s Bot. Jahresber. 19:
246, 1894 (Sydow 1894). Bas.: Lejeunea schwaneckei Steph., Hedwigia 27 (11/12):
290, 1888 (Stephani 1888c).
** Pycnolejeunea sphaeroides (Sande Lac.) J.B.Jack et Steph., Bot. Centralbl. 60 (4):
107, 1894 (Jack and Stephani 1894). Bas.: Lejeunea sphaeroides Sande Lac., Ann.
Mus. Bot. Lugduno-Batavi 1: 309, 1864 (Sande Lacoste 1864).
**
subtrib. Xylolejeuneinae Heinrichs et Schäf.Verw.
*** Xylolejeunea Xiao L.He et Grolle, Ann. Bot. Fenn. 38 (1): 27, 2001 (He and
Grolle 2001).
*** Xylolejeunea aquarius (Spruce) Xiao L.He et Grolle, Ann. Bot. Fenn. 38 (1): 29,
2001 (He and Grolle 2001). Bas.: Lejeunea aquarius Spruce, Trans. & Proc. Bot.
Soc. Edinburgh 15: 185, 1884 (Spruce 1884).
*** Xylolejeunea crenata (Nees et Mont.) Xiao L.He et Grolle, Ann. Bot. Fenn. 38 (1):
36, 2001 (He and Grolle 2001). Bas.: Lejeunea crenata Nees et Mont., Ann. Sci.
Nat. Bot. (sér. 2) 9: 48, 1838 (Montagne 1838).
*** Xylolejeunea grolleana (Pócs) Xiao L.He et Grolle, Ann. Bot. Fenn. 38 (1): 32,
2001 (He and Grolle 2001). Bas.: Trachylejeunea grolleana Pócs, Haussknechtia,
Beih. 9: 285, 1999 (Pócs 1999).
*** Xylolejeunea muricella Xiao L.He et Grolle, Ann. Bot. Fenn. 38 (1): 34, 2001 (He
and Grolle 2001).
**
trib. Symbiezidieae Gradst.
*** Symbiezidium Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci. Mat. 4
(13): 402, 1877 (Trevisan 1877).
*** Symbiezidium barbilorum (Lindenb. et Gottsche) A.Evans, Bull. Torrey Bot. Club
34 (11): 540, 1907 [1908] (Evans 1907a). Bas.: Lejeunea barbilora Lindenb. et
Gottsche, Linnaea 24 (6): 630, 1851 [1852] (Lindenberg and Gottsche 1851a).
*** Symbiezidium dentatum Herzog, Feddes Repert. Spec. Nov. Regni Veg. 57 (1/2):
175, 1955 (Herzog 1955).
*** Symbiezidium madagascariense Steph., Sp. Hepat. (Stephani) 5: 99, 1912 (Stephani
1912c).
*** Symbiezidium transversale (Sw.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 403, 1877 (Trevisan 1877). Bas.: Jungermannia transversalis
Sw., Prodr. (Swartz): 144, 1788 (Swartz 1788).
*** Symbiezidium transversale var. hookerianum (Gottsche, Lindenb. et Nees) Gradst.
et J.Beek, Beih. Nova Hedwigia 80: 237, 1985 (Gradstein and van Beek 1985).
Bas.: Lejeunea transversalis β hookeriana Gottsche, Lindenb. et Nees, Syn. Hepat.
3: 311, 1845 (Gottsche et al. 1845b).
World checklist of hornworts and liverworts
***
399
Ptychanthoideae Mizut.
*** Acrolejeunea (Spruce) Schifn., Hepat. (Engl.-Prantl): 128, 1893 (Schifner
1893b) nom. conserv. Bas.: Lejeunea subg. Acrolejeunea Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 115, 1884 (Spruce 1884).
*** Acrolejeunea sandvicensis (Gottsche) Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 312,
1896 (Stephani 1896a). Bas.: Phragmicoma sandvicensis Gottsche, Ann. Sci. Nat.
Bot. (sér. 4) 8: 344, 1857 (Gottsche 1857).
** sect. Acrolejeunea
*** Acrolejeunea emergens (Mitt.) Steph., Planzenw. Ost-Afrikas C: 65, 1895 (Stephani
1895d). Bas.: Phragmicoma emergens Mitt., Philos. Trans. 168: 397, 1879 (Mitten
1879).
*** Acrolejeunea emergens var. confertissima (Steph.) Gradst., Bryophyt. Biblioth. 4: 76,
1975 (Gradstein 1975). Bas.: Acrolejeunea confertissima Steph., Hedwigia 31 (4):
165, 1892 (Stephani 1892g).
*** Acrolejeunea heterophylla (A.Evans) Grolle et Gradst., J. Hattori Bot. Lab. 38: 332,
1974 (Gradstein 1974a). Bas.: Ptychocoleus heterophyllus A.Evans, Amer. J. Bot. 5
(3): 144, 1918 (Evans 1918).
*** Acrolejeunea torulosa (Lehm. et Lindenb.) Schifn., Hepat. (Engl.-Prantl): 128,
1893 (Schifner 1893b). Bas.: Jungermannia torulosa Lehm. et Lindenb., Nov.
Stirp. Pug. 6: 41, 1834 (Lehmann 1834).
** sect. Minores (Verd.) L.Söderstr. et A.Hagborg, Bryophyte Diversity Evol. 36
(1): 41, 2014 (Wang et al. 2014b). Bas.: Ptychocoleus sect. Minores Verd., Ann.
Bryol., Suppl. 4: 132, 1934 (Verdoorn 1934a).
*** Acrolejeunea arcuata (Nees) Grolle et Gradst., J. Hattori Bot. Lab. 38: 332, 1974
(Gradstein 1974a). Bas.: Jungermannia arcuata Nees, Enum. Pl. Crypt. Javae: 38,
1830 (Nees 1830).
** Acrolejeunea arcuata subsp. gradsteinii M.A.M.Renner, Phytotaxa 83 (1): 42, 2013
(Renner 2013).
*** Acrolejeunea fertilis (Reinw., Blume et Nees) Schifn., Hepat. (Engl.-Prantl): 128,
1893 (Schifner 1893b). Bas.: Jungermannia fertilis Reinw., Blume et Nees, Nova
Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 211, 1824 [1825]
(Reinwardt et al. 1824a).
*** Acrolejeunea parvula (Mizut.) Gradst., Bryophyt. Biblioth. 4: 115, 1975 (Gradstein 1975). Bas.: Ptychocoleus parvulus Mizut., Dansk Bot. Ark. 27 (1): 97, 1969
(Hattori and Mizutani 1969).
*** Acrolejeunea pycnoclada (Taylor) Schifn., Hepat. (Engl.-Prantl): 128, 1893
(Schifner 1893b). Bas.: Ptychanthus pycnocladus Taylor, London J. Bot. 5: 385,
1846 (Taylor 1846b).
*** Acrolejeunea pycnoclada subsp. latistipula Gradst., Bryophyt. Biblioth. 4: 113,
1975 (Gradstein 1975).
400
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Acrolejeunea tjibodensis (Verd.) Grolle et Gradst., J. Hattori Bot. Lab. 38: 333,
1974 (Gradstein 1974a). Bas.: Ptychocoleus tjibodensis Verd., Recueil Trav. Bot.
Neerl. 30: 227, 1933 (Verdoorn 1933b).
** sect. Pusillae Gradst., Bryophyt. Biblioth. 4: 59, 1975 (Gradstein 1975).
*** Acrolejeunea pusilla (Steph.) Grolle et Gradst., J. Hattori Bot. Lab. 38: 332, 1974
(Gradstein 1974a). Bas.: Archilejeunea pusilla Steph., Sp. Hepat. (Stephani) 4:
731, 1911 (Stephani 1911e).
*** Acrolejeunea sikkimensis (Mizut.) Gradst., Bryophyt. Biblioth. 4: 83, 1975 (Gradstein 1975). Bas.: Ptychocoleus sikkimensis Mizut., Fl. E. Himalaya: 532, 1966
(Hattori 1966c).
** sect. Recurvatae Jian Wang bis et Gradst., Bryophyte Diversity Evol. 36 (1): 39,
2014 (Wang et al. 2014b).
*** Acrolejeunea recurvata Gradst., Bryophyt. Biblioth. 4: 79, 1975 (Gradstein 1975).
** sect. Regulares (Verd.) Gradst., Bryophyt. Biblioth. 4: 63, 1975 (Gradstein
1975). Bas.: Ptychocoleus sect. Regulares Verd., Ann. Bryol., Suppl. 4: 143, 1934
(Verdoorn 1934a).
*** Acrolejeunea allisonii Gradst., Bryophyt. Biblioth. 4: 103, 1975 (Gradstein 1975).
*** Acrolejeunea aulacophora (Mont.) Steph., Bot. Jahrb. Syst. 20 (3): 317, 1895
(Stephani 1895a). Bas.: Phragmicoma aulacophora Mont., Ann. Sci. Nat. Bot. (sér.
2) 19: 259, 1843 (Montagne 1843).
*** Acrolejeunea mollis (Hook.f. et Taylor) Schifn., Hedwigia 33 (4): 178, 1894
(Schifner 1894). Bas.: Ptychanthus mollis Hook.f. et Taylor, London J. Bot. 5:
384, 1846 (Taylor 1846b).
*** Acrolejeunea securifolia (Nees) Steph., Hedwigia 34 (2): 59, 1895 (Stephani
1895c). Bas.: Jungermannia securifolia Nees, Prodr. Fl. Norfolk.: 5, 1833 (Endlicher 1833).
*** Acrolejeunea securifolia subsp. caledonica (Steph.) Gradst., Bryophyt. Biblioth.
4: 100, 1975 (Gradstein 1975). Bas.: Ptychocoleus caledonicus Steph., Sp. Hepat.
(Stephani) 5: 39, 1912 (Stephani 1912c).
*** Acrolejeunea securifolia subsp. hartmannii (Steph.) Gradst., Bryophyt. Biblioth.
4: 99, 1975 (Gradstein 1975). Bas.: Ptychocoleus hartmannii Steph., Sp. Hepat.
(Stephani) 5: 44, 1912 (Stephani 1912c).
*** Acrolejeunea securifolia subsp. pallida (Ångstr.) Gradst., Bryophyt. Biblioth. 4:
101, 1975 (Gradstein 1975). Bas.: Phragmicoma pallida Ångstr., Öfvers. Kongl.
Vetensk.-Akad. Förh. 30 (5): 132, 1873 (Ångström 1873).
** sect. Trocholejeunea (Schifn.) Jian Wang bis et Gradst., Bryophyte Diversity
Evol. 36 (1): 38, 2014 (Wang et al. 2014b). Bas.: Trocholejeunea Schifn., Ann.
Bryol. 5: 160, 1932 (Dixon et al. 1932).
World checklist of hornworts and liverworts
401
*** Acrolejeunea crassicaulis (Steph.) Jian Wang bis et Gradst., Bryophyte Diversity
Evol. 36 (1): 38, 2014 (Wang et al. 2014b). Bas.: Hygrolejeunea crassicaulis Steph.,
Sp. Hepat. (Stephani) 5: 550, 1914 (Stephani 1914b).
*** Acrolejeunea infuscata (Mitt.) Jian Wang bis et Gradst., Bryophyte Diversity Evol.
36 (1): 38, 2014 (Wang et al. 2014b). Bas.: Lejeunea infuscata Mitt., J. Proc. Linn.
Soc., Bot. 5 (18): 111, 1860 [1861] (Mitten 1860c).
** Acrolejeunea meghalayensis (Ajit P.Singh et V.Nath) Jian Wang bis et Gradst., Bryophyte
Diversity Evol. 36 (1): 39, 2014 (Wang et al. 2014b). Bas.: Trocholejeunea meghalayensis Ajit P.Singh et V.Nath, J. Jap. Bot. 83 (1): 2, 2008 (Singh and Nath 2008).
** Acrolejeunea sinensis (Jian Wang bis, R.L.Zhu et Gradst.) Jian Wang bis et Gradst.,
Bryophyte Diversity Evol. 36 (1): 39, 2014 (Wang et al. 2014b). Bas.: Trocholejeunea sinensis Jian Wang bis, R.L.Zhu et Gradst., Phytotaxa 174 (5): 296, 2014
(Wang et al. 2014c).
*
*
*
Excluded from the genus
Acrolejeunea abnormis (Gottsche) Pearson, Forh. Vidensk.-Selsk. Kristiania 1892
(8): 4, 1892 (Pearson 1892). Bas.: Phragmicoma abnormis Gottsche, Abh. Naturwiss. Vereins Bremen 7: 352, 1882 (Gottsche 1882). 354
Acrolejeunea comptonii Pearson, J. Linn. Soc., Bot. 46 (305): 33, 1922 (Pearson
1922b). 355
Acrolejeunea inlexa (Gottsche) Pearson, Forh. Vidensk.-Selsk. Kristiania 1892 (8):
4, 1892 (Pearson 1892). Bas.: Phragmicoma inlexa Gottsche, Abh. Naturwiss.
Vereins Bremen 7: 351, 1882 (Gottsche 1882). 356
*** Archilejeunea (Spruce) Steph., Hedwigia 27 (3/4): 113, 1888 (Stephani 1888d).
Bas.: Lejeunea subg. Archilejeunea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:
88, 1884 (Spruce 1884). 357
*** Archilejeunea abbreviata (Mitt.) Vanden Berghen, Rev. Bryol. Lichénol. 20 (1/2):
117, 1951 (Vanden Berghen 1951c). Bas.: Lejeunea abbreviata Mitt., J. Proc.
Linn. Soc., Bot. 7 (27): 167, 1863 (Mitten 1863).
** Archilejeunea africana Steph., Sp. Hepat. (Stephani) 4: 705, 1911 (Stephani 1911e).
** Archilejeunea alata Steph., Bull. Soc. Roy. Bot. Belgique, Compt. Rend. 32 (2):
33, 1893 [1894] (Stephani 1893e).
354 Acrolejeunea abnormis was excluded from Acrolejeunea by Gradstein (1975), but the type specimen
was not localized.
355 Acrolejeunea comptonii is a Mastigolejeunea species (Gradstein 1975). he type specimen has not been
localized.
356 Acrolejeunea inlexa was excluded from Acrolejeunea by Gradstein (1975), but the type specimen was
not localized.
357 Archilejeunea is polyphyletic and several nomenclatural and taxonomic will be proposed (Shi et al. 2015a).
402
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Archilejeunea amakawana Inoue, J. Jap. Bot. 41 (1): 16, 1966 (Inoue 1966d).
Nom. nov. pro Archilejeunea falcata Amakawa, J. Jap. Bot. 39 (5): 137, 1964 (Amakawa 1964b), nom. illeg.
*** Archilejeunea auberiana (Mont.) Steph., Hedwigia 29 (3): 134, 1890 (Stephani
1890d). Bas.: Lejeunea auberiana Mont., Hist. Phys. Cuba, Bot., Pl. Cell.: 483,
1842 (Montagne 1842a).
*** Archilejeunea autoica Vanden Berghen, Rev. Bryol. Lichénol. 20 (1/2): 119, 1951
(Vanden Bergen 1951b).
*** Archilejeunea badia (Spruce) Steph., Sp. Hepat. (Stephani) 4: 711, 1911 (Stephani
1911e). Bas.: Lejeunea badia Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 92,
1884 (Spruce 1884).
* Archilejeunea bilabiata (Mitt.) Steph., Sp. Hepat. (Stephani) 4: 723, 1911 (Stephani
1911e). Bas.: Phragmicoma bilabiata Mitt., Fl. vit.: 412, 1871 [1873] (Mitten 1871). 358
*** Archilejeunea bischleriana Gradst., Fl. Neotrop. Monogr. 62: 62, 1994 (Gradstein
1994).
** Archilejeunea bongardii Steph., Hedwigia 29 (1): 20, 1890 (Stephani 1890a).
** Archilejeunea brachyantha J.B.Jack et Steph., Bot. Centralbl. 60 (4): 104, 1894
(Jack and Stephani 1894).
** Archilejeunea brevilobula Steph., Sp. Hepat. (Stephani) 4: 706, 1911 (Stephani 1911e).
*** Archilejeunea crispistipula (Spruce) Steph., Sp. Hepat. (Stephani) 4: 712, 1911
(Stephani 1911e). Bas.: Lejeunea crispistipula Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 93, 1884 (Spruce 1884).
* Archilejeunea eberhardtii Steph., Sp. Hepat. (Stephani) 4: 725, 1911 (Stephani 1911e).
** Archilejeunea elobulata Steph., Sp. Hepat. (Stephani) 4: 707, 1911 (Stephani 1911e).
** Archilejeunea gradsteinii X.Q.Shi et R.L.Zhu, Nova Hedwigia 100 (3/4): 592,
2015 (Shi and Zhu 2015).
** Archilejeunea incrassata Steph., Rev. Bryol. 35 (2): 30, 1908 (Stephani 1908l).
** Archilejeunea jonesii Vanden Berghen, Rev. Bryol. Lichénol. 20 (1/2): 116, 1951
(Vanden Berghen 1951c).
*** Archilejeunea juliformis (Nees) Gradst., Bryophyt. Biblioth. 4: 127, 1975 (Gradstein 1975). Bas.: Jungermannia juliformis Nees, Fl. Bras. (Martius) 1 (1): 351,
1833 (Nees 1833a).
*** Archilejeunea kiushiana (Horik.) Verd., Ann. Bryol., Suppl. 4: 46, 1934 (Verdoorn
1934a). Bas.: Lopholejeunea kiushiana Horik., J. Sci. Hiroshima Univ., Ser. B, Div.
2, Bot. 1: 129, 1932 (Horikawa 1932c).
*** Archilejeunea ludoviciana (De Not.) P.Geissler et Gradst., J. Hattori Bot. Lab. 75:
202, 1994 (Geissler and Gradstein 1994). Bas.: Phragmicoma ludoviciana De Not.,
Nov. Stirp. Pug. 10: 11, 1857 (Lehmann 1857).
*** Archilejeunea ludoviciana subsp. porelloides (Spruce) Gradst., Fl. Neotrop. Monogr. 62: 58, 1994 (Gradstein 1994). Bas.: Lejeunea porelloides Spruce, Trans. &
Proc. Bot. Soc. Edinburgh 15: 90, 1884 (Spruce 1884).
358 Archilejeunea bilabiata is a doubtful taxon, the type specimen could not be found (Geissler and Gradstein 1994).
World checklist of hornworts and liverworts
403
** Archilejeunea nebeliana Gradst. et Schäf.-Verw., Cryptog. Bryol. 33 (2): 108, 2012
(Gradstein and Schäfer-Verwimp 2012).
*** Archilejeunea olivacea (Hook.f. et Taylor) Steph., Hedwigia 29 (3): 134, 1890
(Stephani 1890d). Bas.: Jungermannia olivacea Hook.f. et Taylor, London J. Bot.
3: 568, 1844 (Hooker and Taylor 1844d).
*** Archilejeunea parvilora (Nees) Steph., Hedwigia 29 (3): 134, 1890 (Stephani
1890d). Bas.: Jungermannia parvilora Nees, Fl. Bras. (Martius) 1 (1): 353, 1833
(Nees 1833a).
** Archilejeunea planifolia (Horik.) Mizut., J. Hattori Bot. Lab. 73: 175, 1993
(Mizutani 1993). Bas.: Leucolejeunea planifolia Horik., J. Sci. Hiroshima Univ.,
Ser. B, Div. 2, Bot. 1: 199, 1933 (Horikawa 1933). 359
*** Archilejeunea planiuscula (Mitt.) Steph., Sp. Hepat. (Stephani) 4: 731, 1911
(Stephani 1911e). Bas.: Lejeunea planiuscula Mitt., J. Proc. Linn. Soc., Bot. 5 (18):
111, 1860 [1861] (Mitten 1860c).
*
*
Excluded from the genus
Archilejeunea negrensis Steph., Sp. Hepat. (Stephani) 4: 716, 1911 (Stephani 1911e). 360
Archilejeunea ovata Herzog, Rev. Bryol. Lichénol. 20 (1/2): 130, 1951 [1952]
(Herzog 1951a). 361
*** Bryopteris (Nees) Lindenb., Syn. Hepat. 2: 284, 1845 (Gottsche et al. 1845a).
Bas.: Frullania subg. Bryopteris Nees, Naturgesch. Eur. Leberm. 3: 211, 1838
(Nees 1838b).
*** Bryopteris difusa (Sw.) Nees, Syn. Hepat. 2: 286, 1845 (Gottsche et al. 1845a).
Bas.: Jungermannia difusa Sw., Prodr. (Swartz): 144, 1788 (Swartz 1788).
*** Bryopteris ilicina (Sw.) Nees, Syn. Hepat. 2: 284, 1845 (Gottsche et al. 1845a).
Bas.: Jungermannia ilicina Sw., Prodr. (Swartz): 145, 1788 (Swartz 1788).
* Bryopteris issiloba Steph., Sp. Hepat. (Stephani) 6: 568, 1924 (Stephani 1924). 362
*** Bryopteris gaudichaudii Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 8: 340, 1857
(Gottsche 1857).
*** Caudalejeunea Schifn., Hepat. (Engl.-Prantl): 129, 1893 (Schifner 1893b).
** subg. Acaudalejeunea R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 779, 1980
(Schuster 1980c).
*** Caudalejeunea grolleana Gradst., Acta Bot. Neerl. 23 (3): 334, 1974 (Gradstein 1974b).
359 Archilejeunea planifolia is conspeciic with Archilejeunea kiushiana in Hattori (1952c) and Gradstein
and Geissler (1997), but it is a separate species in Mizutani (1993).
360 Archilejeunea negrensis is probably not an Archilejeunea species, but the type specimen is too poor to
permit identiication (Gradstein and Buskes 1985).
361 Archilejeunea ovata is conspeciic with Lopholejeunea nigricans or Lopholejeunea subfusca (Gradstein 1994).
362 Bryopteris issiloba is probably conspeciic with Bryopteris difusa (Gradstein and Costa 2003). It was
not accepted by Hartmann et al. (2006).
404
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** subg. Caudalejeunea
** Caudalejeunea africana (Steph.) Schifn., Hepat. (Engl.-Prantl): 129, 1893
(Schifner 1893b). Bas.: hysananthus africanus Steph., Bot. Jahrb. Syst. 8 (2): 93,
1886 (Stephani 1886d).
*** Caudalejeunea hanningtonii (Mitt.) Schifn., Hepat. (Engl.-Prantl): 129, 1893
(Schifner 1893b). Bas.: Lejeunea hanningtonii Mitt., J. Linn. Soc., Bot. 22 (146):
324, 1886 (Mitten 1886b).
** Caudalejeunea katangensis Vanden Berghen, Explor. Hydrobiol. Lac Bangweolo
Luapula: 94, 1972 (Vanden Berghen 1972b).
*** Caudalejeunea lehmanniana (Gottsche) A.Evans, Bull. Torrey Bot. Club 34 (11):
554, 1907 [1908] (Evans 1907a). Bas.: Lejeunea lehmanniana Gottsche, Syn. Hepat. 3: 325, 1845 (Gottsche et al. 1845b).
** Caudalejeunea lewallei Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 42 (4): 434,
1972 (Vanden Berghen 1972a).
** subg. Vermilejeunea R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 778, 1980
(Schuster 1980c).
*** Caudalejeunea cristiloba (Steph.) Gradst., Acta Bot. Neerl. 23 (3): 340, 1974
(Gradstein 1974b). Bas.: Acrolejeunea cristiloba Steph., Hedwigia 34 (2): 56, 1895
(Stephani 1895c).
*** Caudalejeunea cristiloba subsp. samoana (Steph.) Gradst., Acta Bot. Neerl. 23 (3):
343, 1974 (Gradstein 1974b). Bas.: Caudalejeunea samoana Steph., Akad. Wiss.
Wien, Math.-Naturwiss. Kl., Denkschr. 81: 296, 1907 (Stephani 1907a).
** Caudalejeunea dusenii Steph., Sp. Hepat. (Stephani) 5: 11, 1912 (Stephani 1912c).
*** Caudalejeunea reniloba (Gottsche) Steph., Sp. Hepat. (Stephani) 5: 16, 1912 (Stephani 1912c). Bas.: Phragmicoma reniloba Gottsche, Syn. Hepat. 2: 301, 1845
(Gottsche et al. 1845a).
** Caudalejeunea yangambiensis (Vanden Berghen) E.W.Jones, Trans. Brit. Bryol.
Soc. 3 (2): 192, 1957 (Jones 1957b). Bas.: Ptychocoleus yangambiensis Vanden
Berghen, Bull. Soc. Roy. Bot. Belgique 84: 61, 1951 (Vanden Berghen 1951a).
Incertae sedis
** Caudalejeunea acutifolia Gerola, Lav. Bot. Ist. Bot. Univ. Padova 12: 478, 1947
(Gerola 1947).
* Caudalejeunea mauritiana Horik., Acta Phytotax. Geobot. 13: 214, 1943 (Horikawa 1943). Nom. nov. pro Dicranolejeunea africana Steph., Sp. Hepat. (Stephani)
5: 158, 1912 (Stephani 1912c).
** Caudalejeunea streimannii Gyarmati, Trop. Bryol. 22: 129, 2002 (Sass-Gyarmati
2002).
** Caudalejeunea tridentata R.L.Zhu, Y.M.Wei et Qiong He, Bryologist 114 (3):
469, 2011 (Zhu et al. 2011).
World checklist of hornworts and liverworts
405
** Cephalantholejeunea R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 798, 1980
(Schuster 1980c).
*** Cephalantholejeunea temnanthoides (R.M.Schust.) R.M.Schust., Hepat. Anthocerotae N. Amer. 4: 807, 1980 (Schuster 1980c). Bas.: Potamolejeunea temnanthoides
R.M.Schust., Beih. Nova Hedwigia 9: 123, 1963 (Schuster 1963a).
** Cephalolejeunea Mizut., J. Hattori Bot. Lab. 46: 359, 1979 (Mizutani 1979b).
*** Cephalolejeunea parvilobula Mizut., J. Hattori Bot. Lab. 46: 359, 1979 (Mizutani
1979b).
*** Frullanoides Raddi, Critt. Brasil.: 13, 1822 (Raddi 1822).
*** Frullanoides bahamensis (A.Evans) van Slageren, Meded. Bot. Mus. Herb. Rijks
Univ. Utrecht 544: 81, 1985 (van Slageren 1985). Bas.: Brachiolejeunea bahamensis A.Evans, Bull. Torrey Bot. Club 35 (8): 383, 1908 (Evans 1908b).
*** Frullanoides corticalis (Lehm. et Lindenb.) van Slageren, Meded. Bot. Mus. Herb.
Rijks Univ. Utrecht 544: 84, 1985 (van Slageren 1985). Bas.: Jungermannia corticalis Lehm. et Lindenb., Nov. Stirp. Pug. 4: 50, 1832 (Lehmann 1832).
*** Frullanoides densifolia Raddi, Critt. Brasil.: 14, 1822 (Raddi 1822).
*** Frullanoides densifolia subsp. grandidentata (L.Clark) van Slageren, Meded. Bot.
Mus. Herb. Rijks Univ. Utrecht 544: 95, 1985 (van Slageren 1985). Bas.: Brachiolejeunea grandidentata L.Clark, Proc. Calif. Acad. Sci. (ser. 4) 27 (18): 595, 1953
(Clark 1953).
*** Frullanoides laciniatilora (Loitl.) van Slageren, Meded. Bot. Mus. Herb. Rijks
Univ. Utrecht 544: 100, 1985 (van Slageren 1985). Bas.: Lejeunea laciniatilora
Loitl., Diagn. pl. nov.: 19, 1894 (Loitlesberger 1894).
*** Frullanoides liebmaniana (Lindenb. et Gottsche) van Slageren, Meded. Bot. Mus.
Herb. Rijks Univ. Utrecht 544: 102, 1985 (van Slageren 1985). Bas.: Phragmicoma
liebmaniana Lindenb. et Gottsche, Syn. Hepat. 5: 744, 1847 (Gottsche et al. 1847).
*** Frullanoides mexicana van Slageren, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht
544: 106, 1985 (van Slageren 1985).
*** Frullanoides tristis (Steph.) van Slageren, Meded. Bot. Mus. Herb. Rijks Univ.
Utrecht 544: 110, 1985 (van Slageren 1985). Bas.: Brachiolejeunea tristis Steph.,
Sp. Hepat. (Stephani) 5: 112, 1912 (Stephani 1912c).
*** Fulfordianthus Gradst., Bryologist 95 (1): 44, 1992 (Gradstein 1992a).
*** Fulfordianthus evansii (Fulford) Gradst., Bryologist 95 (1): 46, 1992 (Gradstein
1992a). Bas.: hysananthus evansii Fulford, Bull. Torrey Bot. Club 68 (1): 34,
1941 (Fulford 1941).
*** Fulfordianthus pterobryoides (Spruce) Gradst., Bryologist 95 (1): 44, 1992 (Gradstein 1992a). Bas.: Lejeunea pterobryoides Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 109, 1884 (Spruce 1884).
406
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Lopholejeunea (Spruce) Steph., Bot. Gaz. 15 (11): 285, 1890 (Stephani 1890c)
nom. rejic. Bas.: Lejeunea subg. Lopholejeunea Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 119, 1884 (Spruce 1884).
** subg. Lopholejeunea
** Lopholejeunea borbonica Steph., Hedwigia 35 (3): 109, 1896 (Stephani 1896b).
** Lopholejeunea jonesii Vanden Berghen, Bull. Jard. Bot. État Bruxelles 20 (2): 178,
1950 (Vanden Berghen 1950).
** Lopholejeunea laciniata E.W.Jones, Trans. Brit. Bryol. Soc. 3 (2): 194, 1957 (Jones
1957b).
** Lopholejeunea minima Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 54 (3/4): 437,
1984 (Vanden Berghen 1984a).
* Lopholejeunea obtusilacera Herzog, Bull. Jard. Bot. État Bruxelles 20 (2): 172,
1950 (Vanden Berghen 1950).
* Lopholejeunea paramultilacera Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 54
(3/4): 435, 1984 (Vanden Berghen 1984a).
* Lopholejeunea quinquecarinata Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 54
(3/4): 408, 1984 (Vanden Berghen 1984a).
** Lopholejeunea renistipula (Mitt.) Steph., Sp. Hepat. (Stephani) 5: 94, 1912
(Stephani 1912c). Bas.: Phragmicoma renistipula Mitt., Fl. vit.: 413, 1871 [1873]
(Mitten 1871).
** Lopholejeunea revoluta E.W.Jones, Trans. Brit. Bryol. Soc. 3 (2): 207, 1957 (Jones
1957b).
** sect. Eulophae Verd., Ann. Bryol., Suppl. 4: 87, 1934 (Verdoorn 1934a).
*** Lopholejeunea applanata (Reinw., Blume et Nees) Schifn., Hepat. (Engl.-Prantl):
129, 1893 (Schifner 1893b). Bas.: Jungermannia applanata Reinw., Blume et
Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 210,
1824 [1825] (Reinwardt et al. 1824a).
*** Lopholejeunea borneensis (Steph.) Verd., Ann. Bryol., Suppl. 4: 83, 1934 (Verdoorn 1934a). Bas.: Mastigolejeunea borneensis Steph., Sp. Hepat. (Stephani) 4:
777, 1912 (Stephani 1912b).
*** Lopholejeunea erugata B.M.hiers, Brittonia 36 (2): 174, 1984 (hiers 1984). Nom.
nov. pro Ptychocoleus inermis Steph., Sp. Hepat. (Stephani) 5: 27, 1912 (Stephani
1912c).
*** Lopholejeunea eulopha (Taylor) Schifn., Hepat. (Engl.-Prantl): 129, 1893
(Schifner 1893b). Bas.: Lejeunea eulopha Taylor, London J. Bot. 5: 391, 1846
(Taylor 1846b).
*** Lopholejeunea evansiana Verd., Nova Guinea 18: 4, 1934 (Verdoorn 1934d).
*** Lopholejeunea grollei R.L.Zhu et Gradst., Monogr. Syst. Bot. Missouri Bot. Gard.
74: 36, 2005 (Zhu and Gradstein 2005).
*** Lopholejeunea herzogiana Verd., Recueil Trav. Bot. Neerl. 30: 217, 1933 (Verdoorn 1933b).
World checklist of hornworts and liverworts
407
*** Lopholejeunea hispidissima Steph., Sp. Hepat. (Stephani) 5: 80, 1912 (Stephani
1912c).
*** Lopholejeunea loheri Steph., Sp. Hepat. (Stephani) 5: 77, 1912 (Stephani 1912c).
*** Lopholejeunea minuta R.L.Zhu et Gradst., Nova Hedwigia 78 (3/4): 436, 2004
(Zhu and Gradstein 2004).
*** Lopholejeunea nigricans (Lindenb.) Schifn., Consp. Hepat. Arch. Ind.: 293, 1898
(Schifner 1898b). Bas.: Lejeunea nigricans Lindenb., Syn. Hepat. 3: 316, 1845
(Gottsche et al. 1845b).
*** Lopholejeunea plicatiscypha (Hook.f. et Taylor) Steph., Sp. Hepat. (Stephani) 5:
96, 1912 (Stephani 1912c). Bas.: Phragmicoma plicatiscypha Hook.f. et Taylor,
London J. Bot. 5: 386, 1846 (Taylor 1846b).
* Lopholejeunea proxima Steph., Sp. Hepat. (Stephani) 5: 89, 1912 (Stephani 1912c). 363
*** Lopholejeunea streimannii B.M.hiers et Gradst., Mem. New York Bot. Gard. 52:
37, 1989 (hiers and Gradstein 1989).
** Lopholejeunea vojtkoana Gyarmati, Nova Hedwigia 87 (3/4): 480, 2008 (SassGyarmati 2008).
**
***
***
***
***
***
***
***
**
***
*
***
sect. Lopholejeunea
Lopholejeunea ceylanica Steph., Sp. Hepat. (Stephani) 5: 86, 1912 (Stephani 1912c).
Lopholejeunea horticola Schifn., Ann. Bryol. 6: 133, 1933 (Herzog et al. 1933).
Lopholejeunea latilobula Verd., Nova Guinea 18: 4, 1934 (Verdoorn 1934d).
Lopholejeunea magna Mizut., J. Hattori Bot. Lab. 32: 131, 1969 (Mizutani 1969).
Lopholejeunea recurvata Mizut., J. Hattori Bot. Lab. 46: 369, 1979 (Mizutani 1979b).
Lopholejeunea soae R.L.Zhu et Gradst., Monogr. Syst. Bot. Missouri Bot. Gard.
74: 69, 2005 (Zhu and Gradstein 2005).
Lopholejeunea subfusca (Nees) Schifn., Bot. Jahrb. Syst. 23 (5): 593, 1897
(Schifner 1897). Bas.: Jungermannia subfusca Nees, Enum. Pl. Crypt. Javae: 36,
1830 (Nees 1830).
Lopholejeunea subfusca var. elongata Vanden Berghen, Bull. Jard. Bot. Natl. Belg.
54 (3/4): 445, 1984 (Vanden Berghen 1984a).
Lopholejeunea wiltensii Steph., Hedwigia 35 (3): 112, 1896 (Stephani 1896b).
Lopholejeunea yapensis Steph., Sp. Hepat. (Stephani) 5: 81, 1912 (Stephani 1912c). 364
Lopholejeunea zollingeri (Steph.) Schifn., Consp. Hepat. Arch. Ind.: 296, 1898
(Schifner 1898b). Bas.: Lejeunea zollingeri Steph., Hedwigia 29 (1): 14, 1890
(Stephani 1890a).
** subg. Pholianthus B.M.hiers et Gradst., Mem. New York Bot. Gard. 52: 25,
1989 (hiers and Gradstein 1989).
*** Lopholejeunea colensoi Steph., Sp. Hepat. (Stephani) 5: 97, 1912 (Stephani 1912c).
*** Lopholejeunea pocsii Gyarmati, Cryptog. Bryol. 26 (4): 404, 2005 (Sass-Gyarmati
2005).
363 Lopholejeunea proxima is possibly conspeciic with Lopholejeunea nigricans (Zhu and Gradstein 2005).
364 Lopholejeunea yapensis is possibly conspeciic with Lopholejeunea subfusca (Mizutani 1985).
408
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** subg. Pteryganthus B.M.hiers, Brittonia 35 (1): 85, 1983 (hiers 1983).
** Lopholejeunea grandicrista Steph., Bull. Soc. Roy. Bot. Belgique, Compt. Rend. 32
(2): 34, 1893 [1894] (Stephani 1893e).
*** Lopholejeunea leioptera Gyarmati, Candollea 56 (1): 80, 2001 (Sass-Gyarmati 2001).
** Lopholejeunea onraedtii Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 54 (3/4):
452, 1984 (Vanden Berghen 1984a).
** Lopholejeunea sphaerophora (Lehm. et Lindenb.) Steph., Sp. Hepat. (Stephani) 5:
68, 1912 (Stephani 1912c). Bas.: Jungermannia sphaerophora Lehm. et Lindenb.,
Nov. Stirp. Pug. 5: 9, 1833 (Lehmann 1833).
* Lopholejeunea tixieriana Vanden Berghen, Bull. Jard. Bot. Natl. Belg. 54 (3/4):
454, 1984 (Vanden Berghen 1984a).
** Lopholejeunea utriculata Steph., Sp. Hepat. (Stephani) 5: 69, 1912 (Stephani 1912c).
Incertae sedis
** Lopholejeunea lepidoscypha Kiaer et Pearson, Forh. Vidensk.-Selsk. Kristiania 1892
(8): 5, 1892 (Pearson 1892).
* Lopholejeunea multilacera Steph., Bot. Gaz. 15 (11): 285, 1890 (Stephani 1890c).
** Lopholejeunea udarii M.Dey et D.K.Singh, Nelumbo 53: 197, 2011 (Dey and
Singh 2011).
*
*
Excluded from the genus
Lopholejeunea aberrantia Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2:
256, 1934 (Horikawa 1934). 365
Lopholejeunea vietnamica Tixier, Ann. Hist.-Nat. Mus. Natl. Hung. 66: 99, 1974
(Tixier 1974). 366
*** Marchesinia Gray, Nat. Arr. Brit. Pl. 1: 689, 1821 (Gray 1821) nom. conserv.
*** subg. Marchesinia
*** Marchesinia mackaii (Hook.) Gray, Nat. Arr. Brit. Pl. 1: 689, 1821 (Gray 1821).
Bas.: Jungermannia mackaii Hook., Brit. Jungermann.: tab. 53, 1813 (Hooker
1813).
*** subg. Marchesiniopsis R.M.Schust., J. Hattori Bot. Lab. 72: 358, 1992 (Schuster
1992a).
*** Marchesinia bongardiana (Lehm. et Lindenb.) Trevis., Mem. Reale Ist. Lombardo
Sci. (Ser. 3), C. Sci. Mat. 4 (13): 405, 1877 (Trevisan 1877). Bas.: Lejeunea bongardiana Lehm. et Lindenb., Nov. Stirp. Pug. 7: 18, 1838 (Lehmann 1838).
365 Lopholejeunea aberrantia is probably an Archilejeunea species, but the type specimen in HIRO is
destroyed (Zhu and Gradstein 2005).
366 Lopholejeunea vietnamica is probably an Archilejeunea species (Zhu and Gradstein 2005). he type
specimen was not found in PC.
World checklist of hornworts and liverworts
409
*** Marchesinia brachiata (Sw.) Schifn., Hepat. (Engl.-Prantl): 128, 1893 (Schifner
1893b). Bas.: Jungermannia brachiata Sw., Prodr. (Swartz): 144, 1788 (Swartz
1788).
*** Marchesinia deslooveri Vanden Berghen, Rev. Bryol. Lichénol. 42 (4): 926, 1976
(Vanden Berghen 1976a).
*** Marchesinia excavata (Mitt.) Schifn., Hepat. (Engl.-Prantl): 128, 1893 (Schifner
1893b). Bas.: Lejeunea excavata Mitt., Trans. Linn. Soc. London 23 (1): 58, 1860
(Mitten 1860a).
*** Marchesinia languida (Nees et Mont.) Steph., Sp. Hepat. (Stephani) 5: 149, 1912
(Stephani 1912c). Bas.: Lejeunea languida Nees et Mont., Ann. Sci. Nat. Bot. (sér.
2) 5: 59, 1836 (Nees and Montagne 1836).
*** Marchesinia nobilis (Gottsche) X.Q.Shi, R.L.Zhu et Gradst., Phytotaxa 195 (3):
249, 2015 (Shi et al. 2015b). Bas.: Lejeunea nobilis Gottsche, Abh. Naturwiss.
Vereins Bremen 7: 353, 1882 (Gottsche 1882).
*** Marchesinia robusta (Mitt.) Schifn., Hepat. (Engl.-Prantl): 128, 1893 (Schifner
1893b). Bas.: Lejeunea robusta Mitt., Hooker’s J. Bot. Kew Gard. Misc. 3: 359,
1851 (Mitten 1851).
** Mastigolejeunea (Spruce) Steph., Hedwigia 30 (5): 206, 1891 (Stephani 1891a).
Bas.: Lejeunea subg. Mastigolejeunea Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 100, 1884 (Spruce 1884).
*** Mastigolejeunea auriculata (Wilson et Hook.) Steph., Bot. Gaz. 17 (6): 171, 1892
(Stephani 1892f). Bas.: Jungermannia auriculata Wilson et Hook., Musci Amer.,
S. States: no. 170, 1841 (Wilson 1841; non vidi).
** Mastigolejeunea auriculata var. rhodesica (Vanden Berghen) Sukkharak et Gradst.,
Nova Hedwigia 99 (3/4): 297, 2014 (Sukkharak and Gradstein 2014). Bas.:
Brachiolejeunea rhodesica Vanden Berghen, Bull. Jard. Bot. État Bruxelles 21 (1/2):
94, 1951 (Vanden Berghen 1951d).
*** Mastigolejeunea calcarata (Steph.) Verd., Blumea 1 (1): 230, 1934 (Verdoorn
1934b). Bas.: Archilejeunea calcarata Steph., Sp. Hepat. (Stephani) 4: 724, 1911
(Stephani 1911e).
*** Mastigolejeunea lorea (Mitt.) Paris, Rev. Bryol. 33 (3): 42, 1906 (Paris 1906b).
Bas.: Phragmicoma lorea Mitt., J. Linn. Soc., Bot. 22 (146): 323, 1886 (Mitten
1886b).
*** Mastigolejeunea frauenfeldii (Reichardt) Verd., Blumea 1 (1): 230, 1934 (Verdoorn 1934b). Bas.: hysananthus frauenfeldii Reichardt, Verh. K.K. Zool.-Bot.
Ges. Wien 16: 958, 1866 (Reichardt 1866).
** Mastigolejeunea gradsteinii Sukkharak, J. Bryol. 36 (1): 56, 2014 (Sukkharak 2014).
410
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Mastigolejeunea humilis (Gottsche) Schifn., Hepat. (Engl.-Prantl): 129, 1893
(Schifner 1893b). Bas.: Phragmicoma humilis Gottsche, Syn. Hepat. 2: 299, 1845
(Gottsche et al. 1845a). 367
*** Mastigolejeunea indica Steph., Sp. Hepat. (Stephani) 4: 776, 1912 (Stephani 1912b).
*** Mastigolejeunea innovans (Spruce) Steph., Sp. Hepat. (Stephani) 4: 765, 1912
(Stephani 1912b). Bas.: Lejeunea innovans Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 103, 1884 (Spruce 1884).
*** Mastigolejeunea ligulata (Lehm. et Lindenb.) Schifn., Hepat. (Engl.-Prantl): 129,
1893 (Schifner 1893b). Bas.: Jungermannia ligulata Lehm. et Lindenb., Nov.
Stirp. Pug. 6: 39, 1834 (Lehmann 1834).
*** Mastigolejeunea nigra Steph., Hedwigia 30 (5): 206, 1891 (Stephani 1891a).
*** Mastigolejeunea plicatilora (Spruce) Steph., Sp. Hepat. (Stephani) 4: 766, 1912
(Stephani 1912b). Bas.: Lejeunea plicatilora Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 104, 1884 (Spruce 1884).
*** Mastigolejeunea recondita (Steph.) Mizut., J. Hattori Bot. Lab. 32: 134, 1969 (Mizutani 1969). Bas.: Ptycholejeunea recondita Steph., Hedwigia 35 (3): 122, 1896
(Stephani 1896b).
*** Mastigolejeunea repleta (Taylor) A.Evans, Mem. Torrey Bot. Club 8 (2): 131, 1902
(Evans 1902a). Bas.: Lejeunea repleta Taylor, London J. Bot. 5: 392, 1846 (Taylor
1846b).
*** Mastigolejeunea truncata Mizut., J. Hattori Bot. Lab. 61: 292, 1986 [1987] (Mizutani 1986c).
** Mastigolejeunea turgida Steph., Hedwigia 31 (4): 170, 1892 (Stephani 1892g).
*** Mastigolejeunea virens (Ångstr.) Steph., Sp. Hepat. (Stephani) 4: 776, 1912
(Stephani 1912b), nom. conserv. Bas.: hysananthus virens Ångstr., Öfvers. Kongl.
Vetensk.-Akad. Förh. 30 (5): 131, 1873 (Ångström 1873).
** Phaeolejeunea Mizut., J. Hattori Bot. Lab. 31: 130, 1968 (Mizutani 1968).
*** Phaeolejeunea amicorum (Hürl.) Pócs, Fieldiana, Bot. (n.ser.) 47: 140, 2008 (Pócs
2008b). Bas.: Phaeolejeunea etesseana subsp. amicorum Hürl., Bauhinia 9 (4): 263,
1991 (Hürlimann 1991).
** Phaeolejeunea etesseana (Steph.) Mizut., J. Hattori Bot. Lab. 31: 133, 1968 (Mizutani 1968). Bas.: Brachiolejeunea etesseana Steph., Sp. Hepat. (Stephani) 5: 133,
1912 (Stephani 1912c).
** Phaeolejeunea inermis (Steph.) Mizut., J. Hattori Bot. Lab. 31: 134, 1968 (Mizutani 1968). Bas.: Lopholejeunea inermis Steph., Sp. Hepat. (Stephani) 5: 92, 1912
(Stephani 1912c).
*** Phaeolejeunea latistipula (Schifn. ex P.Syd.) Mizut., J. Hattori Bot. Lab. 31: 131,
1968 (Mizutani 1968). Bas.: Hygrolejeunea latistipula Schifn. ex P.Syd., Just’s
Bot. Jahresber. 19: 246, 1894 (Sydow 1894).
367 Mastigolejeunea humilis has been treated as conspeciic with Mastigolejeunea auriculata by many recent authors, but it merits recognition based on molecular and morphological evidence (Sukkharak
et al. 2011).
World checklist of hornworts and liverworts
411
*** Ptychanthus Nees, Naturgesch. Eur. Leberm. 3: 211, 1838 (Nees 1838b).
** Ptychanthus africanus Steph., Wiss. Ergebn. Deut. Zentr.-Afr. Exped. (1907-08),
Bot. 2: 131, 1911 (Stephani 1911a).
* Ptychanthus stephensonianus (Mitt.) Steph., Sp. Hepat. (Stephani) 4: 754, 1912
(Stephani 1912b). Bas.: Lejeunea stephensoniana Mitt., Bot. antarct. voy. II (Fl.
Nov.-Zel. 2): 155, 1854 (Mitten 1854). 368
*** Ptychanthus striatus (Lehm. et Lindenb.) Nees, Naturgesch. Eur. Leberm. 3: 212,
1838 (Nees 1838b). Bas.: Jungermannia striata Lehm. et Lindenb., Nov. Stirp.
Pug. 4: 16, 1832 (Lehmann 1832).
** Ptychanthus striatus var. intermedius (Gottsche) Verd., Ann. Bryol., Suppl. 4: 122,
1934 (Verdoorn 1934a). Bas.: Ptychanthus intermedius Gottsche, Natuurk. Tijdschr. Ned.-Indië 4: 576, 1853 (Gottsche 1853).
*** Schifneriolejeunea Verd., Ann. Bryol. 6: 89, 1933 (Verdoorn 1933a).
** sect. Pappeanae R.M.Schust. ex Gradst. et Vanden Berghen, Beih. Nova Hedwigia 80: 174, 1985 (Gradstein and Vanden Berghen 1985). Based on: Phragmilejeunea R.M.Schust., J. Hattori Bot. Lab. 11: 27, 1954 (Schuster and Hattori
1954).
*** Schifneriolejeunea fragilis Gradst. et E.W.Jones, J. Bryol. 12 (1): 45, 1982 (Jones 1982).
*** Schifneriolejeunea madagascariensis (Steph.) Gradst., J. Hattori Bot. Lab. 38: 333,
1974 (Gradstein 1974a). Bas.: Ptychocoleus madagascariensis Steph., Sp. Hepat.
(Stephani) 5: 27, 1912 (Stephani 1912c).
*** Schifneriolejeunea pappeana var. bidentata Gradst. et Vanden Berghen, Beih. Nova
Hedwigia 80: 182, 1985 (Gradstein and Vanden Berghen 1985).
*** Schifneriolejeunea pappeana var. integra Gradst. et Vanden Berghen, Beih. Nova
Hedwigia 80: 182, 1985 (Gradstein and Vanden Berghen 1985).
*** Schifneriolejeunea parviloba (Steph.) Gradst., J. Hattori Bot. Lab. 38: 335, 1974
(Gradstein 1974a). Bas.: Acrolejeunea parviloba Steph., Bot. Gaz. 15 (11): 286,
1890 (Stephani 1890c).
*** Schifneriolejeunea pappeana (Nees) Gradst., J. Hattori Bot. Lab. 38: 335, 1974
(Gradstein 1974a). Bas.: Phragmicoma pappeana Nees, Syn. Hepat. 2: 296, 1845
(Gottsche et al. 1845a).
** sect. Schifneriolejeunea, Occas. Pap. Farlow Herb. Cryptog. Bot. 16: 72, 1981
(Gradstein and Terken 1981).
*** Schifneriolejeunea altimontana Vanden Berghen, Rev. Bryol. Lichénol. 42 (4):
923, 1976 (Vanden Berghen 1976a).
*** Schifneriolejeunea amazonica Gradst., Beih. Nova Hedwigia 80: 25, 1985 (Gradstein
1985a).
368 Ptychanthus stephensonianus was tentatively treated as a separate species by hiers and Gradstein
(1989). Further work on the infraspeciic variation of Ptychanthus striatus is needed to determine the
correct status.
412
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*** Schifneriolejeunea cumingiana (Mont.) Gradst., J. Hattori Bot. Lab. 38: 333, 1974
(Gradstein 1974a). Bas.: Phragmicoma cumingiana Mont., London J. Bot. 4: 7,
1845 (Montagne 1845a).
*** Schifneriolejeunea ferruginea (Steph.) Gradst., J. Hattori Bot. Lab. 38: 333, 1974
(Gradstein 1974a). Bas.: Acrolejeunea ferruginea Steph., Hedwigia 34 (2): 57, 1895
(Stephani 1895c).
*** Schifneriolejeunea occulta (Steph.) Gradst., J. Hattori Bot. Lab. 38: 333, 1974
(Gradstein 1974a). Bas.: Ptychocoleus occultus Steph., Sp. Hepat. (Stephani) 5: 25,
1912 (Stephani 1912c).
*** Schifneriolejeunea omphalanthoides Verd., Ann. Bryol. 6: 91, 1933 (Verdoorn 1933c).
*** Schifneriolejeunea tumida (Nees) Gradst., J. Hattori Bot. Lab. 38: 335, 1974
(Gradstein 1974a). Bas.: Ptychanthus tumidus Nees, Naturgesch. Eur. Leberm. 3:
213, 1838 (Nees 1838b).
*** Schifneriolejeunea tumida var. haskarliana (Gottsche) Gradst. et Terken, Occas.
Pap. Farlow Herb. Cryptog. Bot. 16: 77, 1981 (Gradstein and Terken 1981).
Bas.: Phragmicoma haskarliana Gottsche, Syn. Hepat. 2: 299, 1845 (Gottsche et
al. 1845a).
*** Schifneriolejeunea nymannii (Steph.) Gradst. et Terken, Occas. Pap. Farlow Herb.
Cryptog. Bot. 16: 79, 1981 (Gradstein and Terken 1981). Bas.: Archilejeunea nymannii Steph., Sp. Hepat. (Stephani) 4: 730, 1911 (Stephani 1911e).
*** Schifneriolejeunea polycarpa (Nees) Gradst., J. Hattori Bot. Lab. 38: 335, 1974
(Gradstein 1974a). Bas.: Jungermannia polycarpa Nees, Fl. Bras. (Martius) 1 (1):
350, 1833 (Nees 1833a).
*** Schifneriolejeunea pulopenangensis (Gottsche) Gradst., J. Hattori Bot. Lab. 38:
335, 1974 (Gradstein 1974a). Bas.: Phragmicoma pulopenangensis Gottsche, Syn.
Hepat. 2: 299, 1845 (Gottsche et al. 1845a).
** Spruceanthus Verd., Ann. Bryol., Suppl. 4: 151, 1934 (Verdoorn 1934a).
*** Spruceanthus macrostipulus (Steph.) Gradst., Trop. Bryol. 4: 13, 1991 (Gradstein
1991). Bas.: Mastigolejeunea macrostipula Steph., Sp. Hepat. (Stephani) 4: 767,
1912 (Stephani 1912b).
** Spruceanthus mamillilobulus (Herzog) Verd., Hepat. Select. Crit. 9: no. 447, 1936
(Verdoorn 1936; non vidi). Bas.: Ptychanthus mamillilobulus Herzog, Symb. Sin.
5: 44, 1930 (Nicholson et al. 1930).
*** Spruceanthus pluriplicatus (Steph.) Gradst., Schriftenreihe Mensch, Kultur Umwelt
z. Bergl. W Neug. 7: 14, 1981 (Hiepko and Schultze-Motel 1981). Bas.: Brachiolejeunea pluriplicata Steph., Sp. Hepat. (Stephani) 5: 135, 1912 (Stephani 1912c).
*** Spruceanthus polymorphus (Sande Lac.) Verd., Ann. Bryol., Suppl. 4: 155, 1934
(Verdoorn 1934a). Bas.: Phragmicoma polymorpha Sande Lac., Ned. Kruidk. Arch.
3: 420, 1854 [1855] (Sande Lacoste 1854).
*** Spruceanthus semirepandus (Nees) Verd., Ann. Bryol., Suppl. 4: 153, 1934 (Verdoorn 1934a). Bas.: Jungermannia semirepanda Nees, Enum. Pl. Crypt. Javae: 39,
1830 (Nees 1830).
World checklist of hornworts and liverworts
413
*** Spruceanthus sulcatus (Nees) Gradst., Beih. Nova Hedwigia 80: 26, 1985 (Gradstein 1985a). Bas.: Jungermannia sulcata Nees, Enum. Pl. Crypt. Javae: 36, 1830
(Nees 1830).
*** Spruceanthus theobromae (Spruce) Gradst., Beih. Nova Hedwigia 80: 26, 1985
(Gradstein 1985a). Bas.: Lejeunea theobromae Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 99, 1884 (Spruce 1884).
*** Spruceanthus thozetianus (Gottsche et F.Muell.) B.M.hiers et Gradst., Mem. New
York Bot. Gard. 52: 62, 1989 (hiers and Gradstein 1989). Bas.: Phragmicoma
thozetiana Gottsche et F.Muell., Fragm. (Mueller): 63, 1880 (Gottsche 1880).
*** hysananthus Lindenb., Nov. Stirp. Pug. 8: 24, 1844 (Lehmann 1844).
** sect. hysananthus
** subsect. Anguiformes Sukkharak, Phytotaxa 193 (1): 37, 2015 (Sukkharak 2015).
*** hysananthus anguiformis (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn.
Hepat. 2: 289, 1845 (Gottsche et al. 1845a). Bas.: Jungermannia anguiformis
Hook.f. et Taylor, London J. Bot. 3: 567, 1844 (Hooker and Taylor 1844d).
*** hysananthus pancheri (Steph.) Hürl., Bauhinia 9 (2): 167, 1989 (Hürlimann
1989). Bas.: Mastigolejeunea pancheri Steph., Sp. Hepat. (Stephani) 4: 771, 1912
(Stephani 1912b).
** subsect. hysananthus
*** hysananthus aculeatus Herzog, Ann. Bryol. 4: 89, 1931 (Herzog 1931b).
*** hysananthus amazonicus (Spruce) Schifn., Hepat. (Engl.-Prantl): 130, 1893
(Schifner 1893b). Bas.: Lejeunea amazonica Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 106, 1884 (Spruce 1884).
*** hysananthus appendiculatus Steph., Sp. Hepat. (Stephani) 4: 794, 1912 (Stephani 1912b).
** hysananthus ciliaris (Sande Lac.) Sukkharak, Nova Hedwigia 99 (3/4): 339, 2014
(Sukkharak and Gradstein 2014). Bas.: Phragmicoma ciliaris Sande Lac., Ann.
Mus. Bot. Lugduno-Batavi 1: 307, 1864 (Sande Lacoste 1864).
*** hysananthus comosus Lindenb., Nov. Stirp. Pug. 8: 25, 1844 (Lehmann 1844).
*** hysananthus convolutus Lindenb., Syn. Hepat. 2: 288, 1845 (Gottsche et al. 1845a).
** hysananthus convolutus var. laceratus (Steph.) Sukkharak, Phytotaxa 193 (1): 30,
2015 (Sukkharak 2015). Bas.: hysananthus laceratus Steph., Sp. Hepat. (Stephani)
4: 796, 1912 (Stephani 1912b).
*** hysananthus discretus Sukkharak et Gradst., Cryptog. Bryol. 31 (2): 113, 2010
(Sukkharak and Gradstein 2010).
*** hysananthus gottschei (J.B.Jack et Steph.) Steph., Sp. Hepat. (Stephani) 4: 787,
1912 (Stephani 1912b). Bas.: hysanolejeunea gottschei J.B.Jack et Steph., Hedwigia 31 (1): 20, 1892 (Jack and Stephani 1892).
** hysananthus gottschei var. continuus Sukkharak, Phytotaxa 193 (1): 33, 2015
(Sukkharak 2015).
414
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** hysananthus spathulistipus (Reinw., Blume et Nees) Lindenb., Syn. Hepat. 2: 287,
1845 (Gottsche et al. 1845a). Bas.: Jungermannia spathulistipa Reinw., Blume et
Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 212,
1824 [1825] (Reinwardt et al. 1824a).
** sect. Vittatae Verd., Ann. Bryol., Suppl. 4: 182, 1934 (Verdoorn 1934a).
** subsect. Sandeanthus (B.M.hiers et Gradst.) Sukkharak, Phytotaxa 193 (1):
43, 2015 (Sukkharak 2015). Bas.: hysananthus subg. Sandeanthus B.M.hiers et
Gradst., Mem. New York Bot. Gard. 52: 66, 1989 (hiers and Gradstein 1989).
*** hysananthus mollis Steph., Sp. Hepat. (Stephani) 4: 798, 1912 (Stephani 1912b).
*** hysananthus montanus Gradst., Xiao L.He et Piippo, Acta Bot. Fenn. 174: 77,
2002 (Gradstein et al. 2002).
*** hysananthus retusus (Reinw., Blume et Nees) B.M.hiers et Gradst., Mem. New
York Bot. Gard. 52: 67, 1989 (hiers and Gradstein 1989). Bas.: Jungermannia
retusa Reinw., Blume et Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol.
Nat. Cur. 12 (1): 214, 1824 [1825] (Reinwardt et al. 1824a).
** hysananthus retusus subsp. sellingii (Herzog) Sukkharak, Phytotaxa 193 (1): 47,
2015 (Sukkharak 2015). Bas.: Mastigolejeunea sellingii Herzog, Ark. Bot. (n.ser.) 3
(3): 60, 1953 (Herzog 1953a).
** subsect. Vittatae (Verd.) Sukkharak, Phytotaxa 193 (1): 40, 2015 (Sukkharak
2015). Bas.: hysananthus sect. Vittatae Verd., Ann. Bryol., Suppl. 4: 182, 1934
(Verdoorn 1934a).
*** hysananthus fruticosus (Lindenb. et Gottsche) Schifn., Hepat. (Engl.-Prantl):
130, 1893 (Schifner 1893b). Bas.: Bryopteris fruticosa Lindenb. et Gottsche, Syn.
Hepat. 5: 737, 1847 (Gottsche et al. 1847).
** Tuzibeanthus S.Hatt., Biosphaera 1 (1): 7, 1947 (Hattori 1947a).
*** Tuzibeanthus chinensis (Steph.) Mizut., J. Hattori Bot. Lab. 24: 151, 1961 (Mizutani 1961). Bas.: Ptychanthus chinensis Steph., Sp. Hepat. (Stephani) 4: 744, 1911
(Stephani 1911e).
** Verdoornianthus Gradst., Bryologist 80 (4): 607, 1977 [1978] (Gradstein 1977).
*** Verdoornianthus griinii Gradst., Bryologist 80 (4): 609, 1977 [1978] (Gradstein 1977).
*** Verdoornianthus marsupiifolius (Spruce) Gradst., Bryologist 80 (4): 609, 1977
[1978] (Gradstein 1977). Bas.: Lejeunea marsupiifolia Spruce, Trans. & Proc. Bot.
Soc. Edinburgh 15: 118, 1884 (Spruce 1884).
World checklist of hornworts and liverworts
415
Porellineae R.M.Schust.
***
Goebeliellaceae Verd.
by M. von Konrat and M.A.M. Renner
*** Goebeliella Steph., Hedwigia 51 (1): 61, 1911 (Stephani 1911c).
*** Goebeliella cornigera (Mitt.) Steph., Hedwigia 51 (1): 62, 1911 (Stephani 1911c). Bas.:
Frullania cornigera Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 163, 1855 (Mitten 1855).
***
Lepidolaenaceae Nakai
by M. von Konrat
he generic composition of Lepidolaenaceae follows Crandall-Stotler et al. (2009), except that the genus Jubulopsis was reduced to a synonym of Lepidolaena by von Konrat
et al. (2012a).
*** Gackstroemia Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci. Mat. 4
(13): 397, 1877 (Trevisan 1877).
** subg. Gackstroemia
** Gackstroemia ljungneri (Herzog) Grolle, J. Hattori Bot. Lab. 30: 17, 1967
(Grolle 1967a). Bas.: Lepidolaena ljungneri Herzog, Ark. Bot. 29A (21): 6, 1940
(Herzog 1940).
*** Gackstroemia magellanica (Lam.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 397, 1877 (Trevisan 1877). Bas.: Jungermannia magellanica
Lam., Encycl. (Lamarck) 3 (1): 284, 1789 (Lamarck 1789).
*** Gackstroemia novae-zelandiae R.M.Schust. et J.J.Engel, Phytotaxa 118 (1): 10,
2013 (Engel 2013a).
*** Gackstroemia weindorferi (Herzog) Grolle, J. Hattori Bot. Lab. 30: 20, 1967
(Grolle 1967a). Bas.: Lepidolaena weindorferi Herzog, Ann. Bryol. 6: 103, 1933
(Verdoorn 1933a).
** subg. Hariotiella (Besch. et C.Massal. ex Schifn.) Grolle, J. Hattori Bot. Lab.
30: 12, 1967 (Grolle 1967a). Bas.: Lepidolaena subg. Hariotiella Besch. et C.Massal. ex Schifn., Hepat. (Engl.-Prantl): 110, 1893 (Schifner 1893b).
*** Gackstroemia hariotiana (Besch. et C.Massal.) Grolle, J. Hattori Bot. Lab. 30: 12,
1967 (Grolle 1967a). Bas.: Polyotus hariotianus Besch. et C.Massal., Bull. Mens.
Soc. Linn. Paris 1 (79): 639, 1886 (Bescherelle and Massalongo 1886).
416
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Gackstroemia patagonica (Steph.) Grolle, J. Hattori Bot. Lab. 30: 14, 1967 (Grolle
1967a). Bas.: Lepidolaena patagonica Steph., Kungl. Svenska Vetensk.-Akad. Handl.
(n.ser.) 46 (9): 76, 1911 (Stephani 1911b).
** Gackstroemia schwabei (Herzog) Grolle, J. Hattori Bot. Lab. 30: 16, 1967 (Grolle
1967a). Bas.: Lepidolaena schwabei Herzog, Rev. Bryol. Lichénol. 29 (3/4): 191,
1960 [1961] (Herzog 1960).
Incertae sedis
*** Gackstroemia alpina R.M.Schust., J. Hattori Bot. Lab. 36: 349, 1972 [1973]
(Schuster 1972).
*** Lepidogyna R.M.Schust., Phytologia 45 (5): 419, 1980 (Schuster 1980b).
*** Lepidogyna hodgsoniae (Grolle) R.M.Schust., Phytologia 45 (5): 419, 1980 (Schuster
1980b). Bas.: Lepidolaena hodgsoniae Grolle, J. Hattori Bot. Lab. 30: 29, 1967 (Grolle
1967a).
*** Lepidogyna menziesii (Hook.) R.M.Schust., Phytologia 45 (5): 419, 1980 (Schuster 1980b). Bas.: Jungermannia menziesii Hook., Musci Exot. 2: tab. 118, 1820
(Hooker 1820).
*** Lepidolaena Dumort., Recueil Observ. Jungerm.: 13, 1835 (Dumortier 1835). 369
*** Lepidolaena berggrenii E.A.Hodgs., Trans. Roy. Soc. New Zealand 87 (3/4): 205,
1959 (Hodgson 1959).
** Lepidolaena brachyclada (Lehm.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser.
3), C. Sci. Mat. 4 (13): 393, 1877 (Trevisan 1877). Bas.: Frullania brachyclada
Lehm., Nov. Stirp. Pug. 8: 21, 1844 (Lehmann 1844).
*** Lepidolaena clavigera (Hook.) Dumort. ex Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 393, 1877 (Trevisan 1877). Bas.: Jungermannia clavigera Hook., Musci Exot. 1: tab. 70, 1818 (Hooker 1818).
*** Lepidolaena novae-zelandiae (E.A.Hodgs. et S.W.Arnell) von Konrat, L.Söderstr.
et A.Hagborg, Phytotaxa 65: 51, 2012 (von Konrat et al. 2012a). Bas.: Jubula
novae-zelandiae E.A.Hodgs. et S.W.Arnell, Trans. Roy. Soc. New Zealand, Bot. 3
(4): 90, 1965 (Hodgson 1965).
*** Lepidolaena palpebrifolia (Hook.) Dumort. ex Trevis., Mem. Reale Ist. Lombardo
Sci. (Ser. 3), C. Sci. Mat. 4 (13): 393, 1877 (Trevisan 1877). Bas.: Jungermannia
palpebrifolia Hook., Musci Exot. 1: tab. 71, 1818 (Hooker 1818).
*** Lepidolaena reticulata (Hook.f. et Taylor) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 393, 1877 (Trevisan 1877). Bas.: Jungermannia reticulata
Hook.f. et Taylor, London J. Bot. 3: 395, 1844 (Hooker and Taylor 1844a).
369 Lepidolaena includes Polyotus, but a few taxa have neither been transferred nor synonymized. hey are
listed in the “Names in genera not currently accepted” section below.
World checklist of hornworts and liverworts
417
*** Lepidolaena taylorii (Gottsche) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 393, 1877 (Trevisan 1877). Bas.: Polyotus taylorii Gottsche, Syn.
Hepat. 2: 246, 1845 (Gottsche et al. 1845a).
***
Porellaceae Cavers nom. conserv.
*** Ascidiota C.Massal., Nuovo Giorn. Bot. Ital. (n.ser.) 5 (2): 256, 1898 (Massalongo 1898).
*** Ascidiota blepharophylla C.Massal., Nuovo Giorn. Bot. Ital. (n.ser.) 5 (2): 257,
1898 (Massalongo 1898).
** Ascidiota blepharophylla subsp. alaskana Steere et R.M.Schust., Bull. Torrey Bot.
Club 87 (3): 213, 1960 (Steere and Schuster 1960).
*** Porella L., Sp. Pl. 1: 1106, 1753 (Linnaeus 1753).
** Porella abyssinica (Nees) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 407, 1877 (Trevisan 1877). Bas.: Madotheca abyssinica Nees, Syn.
Hepat. 2: 281, 1845 (Gottsche et al. 1845a).
** Porella abyssinica var. hoehnelii (Steph.) Pócs, Fragm. Florist. Geobot. 39 (1): 229,
1994 (Pócs 1994a). Bas.: Porella hoehnelii Steph., Hedwigia 30 (6): 266, 1891
(Stephani 1891c).
** Porella acutifolia (Lehm. et Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser.
3), C. Sci. Mat. 4 (13): 408, 1877 (Trevisan 1877). Bas.: Madotheca acutifolia
Lehm. et Lindenb., Nov. Stirp. Pug. 7: 8, 1838 (Lehmann 1838).
** Porella acutifolia var. hattoriana (Pócs) S.Hatt., Misc. Bryol. Lichenol. 8 (4): 79,
1979 (Hattori 1979c). Bas.: Porella plumosa var. hattoriana Pócs, J. Hattori Bot.
Lab. 31: 82, 1968 (Pócs 1968).
* Porella acutifolia var. linguifolia (Steph.) M.L.So, Syst. Bot. 27 (1): 5, 2002 (So
2002a). Bas.: Madotheca linguifolia Steph., Sp. Hepat. (Stephani) 4: 291, 1910
(Stephani 1910b). 370
** Porella acutifolia subsp. tosana (Steph.) S.Hatt., J. Hattori Bot. Lab. 44: 100, 1978
(Hattori 1978a). Bas.: Madotheca tosana Steph., Bull. Herb. Boissier 5 (2): 97,
1897 (Stephani 1897b).
* Porella andica (Gottsche) Hässel, Beih. Nova Hedwigia 134: 452, 2009 (Hässel
and Rubies 2009). Bas.: Madotheca andica Gottsche, Ann. Sci. Nat. Bot. (sér. 4) 8:
339, 1857 (Gottsche 1857). 371
370 Porella acutifolia var. linguifolia was accepted by So (2002a), but it is conspeciic with Plagiochila
viridissima in Hattori (1976f, 1986d). his synonymy is doubtful and further study is necessary to
clarify the status.
371 Madotheca andica was transferred to Porella by Hässel (in Hässel and Rubies 2009), but she did not
study the type specimen. It may be conspeciic with Plagiochila subsquarrosa.
418
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Porella arboris-vitae (With.) Grolle, Trans. Brit. Bryol. Soc. 5 (4): 770, 1969
(Grolle 1969b). Bas.: Jungermannia arboris-vitae With., Bot. arr. veg. Gr. Brit. 2:
697, 1776 (Withering 1776).
** Porella arboris-vitae subsp. nitidula (C.Massal.) S.Hatt., J. Hattori Bot. Lab. 40:
123, 1976 (Hattori 1976f). Bas.: Madotheca nitidula C.Massal., Bull. Soc. Bot.
Ital. 1906: 141, 1906 (Massalongo 1906b).
** Porella baueri (Schifn.) C.E.O.Jensen, Danmarks mosser: 240, 1915 (Jensen
1915). Bas.: Madotheca baueri Schifn., Sitzungsber. deutsch. naturwiss.-med. Vereins Böhmen “Lotos” Prag 48: 346, 1900 (Schifner 1900d).
** Porella bolanderi (Austin) Pearson, List. Canad. Hepat.: 7, 1890 (Pearson 1890).
Bas.: Madotheca bolanderi Austin, Bull. Torrey Bot. Club 3 (3): 14, 1872 (Austin
1872).
** Porella borellii (Gola) Parihar, Univ. Allahabad Stud., Bot. 1961-2: 15, 1962
(Parihar 1962). Bas.: Madotheca borellii Gola, Atti Reale Accad. Sci. Torino, Cl.
Sci. Fis. Mat. Nat. 49: 760, 1914 (Gola 1914b).
*** Porella brachiata (Taylor) Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 334,
1885 (Spruce 1885). Bas.: Madotheca brachiata Taylor, London J. Bot. 6: 341,
1847 (Taylor 1847b).
*** Porella brasiliensis (Raddi) Schifn., Nova Acta Acad. Caes. Leop.-Carol. German.
Nat. Cur. 60 (2): 246, 1893 (Schifner 1893a). Bas.: Schulthesia brasiliensis Raddi,
Critt. Brasil.: 10, 1822 (Raddi 1822).
* Porella brasiliensis var. ciliata (Gottsche, Lindenb. et Nees) Schifn., Nova Acta
Acad. Caes. Leop.-Carol. German. Nat. Cur. 60 (2): 246, 1893 (Schifner 1893a).
Bas.: Madotheca brasiliensis β ciliata Gottsche, Lindenb. et Nees, Syn. Hepat. 2:
271, 1845 (Gottsche et al. 1845a).
* Porella brasiliensis var. laevior (Gottsche, Lindenb. et Nees) Schifn., Nova Acta
Acad. Caes. Leop.-Carol. German. Nat. Cur. 60 (2): 246, 1893 (Schifner 1893a).
Bas.: Madotheca brasiliensis α laevior Gottsche, Lindenb. et Nees, Syn. Hepat. 2:
271, 1845 (Gottsche et al. 1845a).
** Porella caespitans (Steph.) S.Hatt., J. Hattori Bot. Lab. 33: 50, 1970 (Hattori
1970). Bas.: Madotheca caespitans Steph., Mém. Soc. Nat. Sci. Nat. Math. Cherbourg 29: 218, 1894 (Stephani 1894b). 372
** Porella caespitans var. cordifolia (Steph.) S.Hatt. ex T.Katag. et T.Yamag., Bryol.
Res. 10 (5): 133, 2011 (Katagiri and Yamaguchi 2011). Bas.: Madotheca cordifolia
Steph., Sp. Hepat. (Stephani) 4: 315, 1910 (Stephani 1910b).
** Porella caespitans subsp. latior (S.Hatt.) S.Hatt., J. Hattori Bot. Lab. 40: 127, 1976
(Hattori 1976f). Bas.: Porella acutifolia subsp. latior S.Hatt., J. Hattori Bot. Lab.
32: 325, 1969 (Hattori 1969).
** Porella caespitans var. nipponica S.Hatt., J. Hattori Bot. Lab. 33: 57, 1970 (Hattori
1970).
372 Porella caespitans was resolved as polyphyletic in the phylogeny of Hentschel et al. (2007b), suggesting
that the species is a complex deserving further study.
World checklist of hornworts and liverworts
419
** Porella caespitans var. relexigastria (Pócs) S.Hatt., J. Hattori Bot. Lab. 40: 127,
1976 (Hattori 1976f). Bas.: Porella relexigastria Pócs, J. Hattori Bot. Lab. 31: 71,
1968 (Pócs 1968).
** Porella campylophylla (Lehm. et Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 408, 1877 (Trevisan 1877). Bas.: Jungermannia campylophylla Lehm. et Lindenb., Nov. Stirp. Pug. 6: 40, 1834 (Lehmann 1834).
** Porella campylophylla subsp. lancistipula (Steph.) S.Hatt., J. Hattori Bot. Lab. 44:
102, 1978 (Hattori 1978a). Bas.: Madotheca lancistipula Steph., Sp. Hepat. (Stephani) 6: 524, 1924 (Stephani 1924).
** Porella campylophylla var. ligulifera (Taylor) S.Hatt., J. Hattori Bot. Lab. 32: 333,
1969 (Hattori 1969). Bas.: Madotheca ligulifera Taylor, Nov. Stirp. Pug. 8: 10,
1844 (Lehmann 1844).
** Porella campylophylla var. tixieri (Pócs) S.Hatt., J. Hattori Bot. Lab. 40: 128, 1976
(Hattori 1976f). Bas.: Porella plumosa var. tixieri Pócs, J. Hattori Bot. Lab. 31: 82,
1968 (Pócs 1968).
** Porella canariensis (F.Weber) Underw., Rep. (Annual) Missouri Bot. Gard. 8: 186,
1897 (Trelease 1897). Bas.: Jungermannia platyphylla var. canariensis F.Weber,
Hist. Musc. Hepat. Prodr.: 16, 1815 (Weber 1815).
* Porella capehorniensis Swails, Nova Hedwigia 19: 244, 1970 (Swails 1970). 373
** Porella capensis (Gottsche) Mitt., J. Linn. Soc., Bot. 22 (146): 323, 1886 (Mitten
1886b). Bas.: Madotheca capensis Gottsche, Syn. Hepat. 2: 270, 1845 (Gottsche et
al. 1845a).
* Porella caucasica Steph., Bot. Centralbl. 50 (3): 71, 1892 (Stephani 1892a).
** Porella chenii S.Hatt., J. Hattori Bot. Lab. 30: 129, 1967 (Hattori 1967).
*** Porella chilensis (Lehm. et Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser.
3), C. Sci. Mat. 4 (13): 407, 1877 (Trevisan 1877). Bas.: Jungermannia chilensis
Lehm. et Lindenb., Nov. Stirp. Pug. 6: 36, 1834 (Lehmann 1834).
* Porella chilensis var. antucensis (Gottsche) Hässel, Beih. Nova Hedwigia 134: 452,
2009 (Hässel and Rubies 2009). Bas.: Madotheca chilensis β antucensis Gottsche,
Ann. Sci. Nat. Bot. (sér. 4) 8: 340, 1857 (Gottsche 1857).
** Porella chilensis var. fernandeziensis (Herzog) Swails, Nova Hedwigia 19: 236,
1970 (Swails 1970). Bas.: Madotheca chilensis f. fernandeziensis Herzog, Nat. Hist.
Juan Fernandez (Botany) 2 (5): 736, 1942 (Herzog 1942a).
** Porella chilensis var. microloba (Herzog) Swails, Nova Hedwigia 19: 236, 1970
(Swails 1970). Bas.: Madotheca chilensis f. microloba Herzog, Nat. Hist. Juan Fernandez (Botany) 2 (5): 736, 1942 (Herzog 1942a).
** Porella chinensis (Steph.) S.Hatt., J. Hattori Bot. Lab. 30: 131, 1967 (Hattori
1967). Bas.: Madotheca chinensis Steph., Mém. Soc. Nat. Sci. Nat. Math. Cherbourg 29: 218, 1894 (Stephani 1894b).
373 Porella capehorniensis may be conspeciic with Plagiochila setigera and the type specimen is probably
not from South America (Hässel and Rubies 2009).
420
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Porella chinensis var. crispata Udar et Shaheen, Misc. Bryol. Lichenol. 9 (4): 74,
1982 (Udar and Shaheen 1982).
** Porella chinensis var. decurrens (Steph.) S.Hatt., J. Hattori Bot. Lab. 44: 102, 1978
(Hattori 1978a). Bas.: Madotheca decurrens Steph., Sp. Hepat. (Stephani) 4: 289,
1910 (Stephani 1910b).
** Porella chinensis var. hattorii Udar et Shaheen, Misc. Bryol. Lichenol. 9 (7): 146,
1983 (Udar and Shaheen 1983a).
** Porella chinensis var. irregularis (Steph.) S.Hatt., J. Hattori Bot. Lab. 39: 270, 1975
(Hattori 1975c). Bas.: Madotheca irregularis Steph., Sp. Hepat. (Stephani) 4: 304,
1910 (Stephani 1910b).
** Porella circinnata Lindb., Not. Sällsk. Fauna Fl. Fenn. Förh. 13: 355, 1874 (Lindberg 1874a).
*** Porella cordaeana (Huebener) Moore, Proc. Roy. Irish Acad. (ser. 2) 2: 618, 1877
(Moore 1877). Bas.: Jungermannia cordaeana Huebener, Hepaticol. germ.: 291,
1834 (Hübener 1834).
** Porella cranfordii Steph., Hedwigia 28 (4): 270, 1889 (Stephani 1889c).
*** Porella crispata (Hook.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 407, 1877 (Trevisan 1877). Bas.: Jungermannia crispata Hook., Pl.
crypt. (Hooker): tab. 4b, 1816 (Hooker 1816b).
** Porella cucullistipula Steph., Bull. Soc. Roy. Bot. Belgique, Compt. Rend. 32 (2):
38, 1893 [1894] (Stephani 1893e).
** Porella densifolia (Steph.) S.Hatt., J. Jap. Bot. 20: 109, 1944 (Hattori 1944c). Bas.:
Madotheca densifolia Steph., Mém. Soc. Nat. Sci. Nat. Math. Cherbourg 29: 219,
1894 (Stephani 1894b). 374
** Porella densifolia subsp. andamana S.Hatt., J. Hattori Bot. Lab. 32: 346, 1969
(Hattori 1969).
** Porella densifolia subsp. appendiculata (Steph.) S.Hatt., J. Hattori Bot. Lab. 32:
343, 1969 (Hattori 1969). Bas.: Madotheca appendiculata Steph., Sp. Hepat. (Stephani) 4: 301, 1910 (Stephani 1910b).
** Porella densifolia var. oviloba (Steph.) N.Kitag., Acta Phytotax. Geobot. 19 (2/3):
64, 1962 (Kitagawa 1962b). Bas.: Madotheca oviloba Steph., Sp. Hepat. (Stephani) 4: 312, 1910 (Stephani 1910b).
** Porella densifolia var. paraphyllina (P.C.Chen) Pócs, J. Hattori Bot. Lab. 31: 84,
1968 (Pócs 1968). Bas.: Madotheca paraphyllina P.C.Chen, Feddes Repert. Spec.
Nov. Regni Veg. 58: 42, 1955 (Chen 1955).
** Porella densifolia var. pilosa S.Hatt. et K.C.Chang, Bull. Bot. Res., Harbin 8 (2):
44, 1988 (Chang 1988).
** Porella densifolia var. robusta (Steph.) S.Hatt., J. Hattori Bot. Lab. 32: 343, 1969
(Hattori 1969). Bas.: Madotheca robusta Steph., Sp. Hepat. (Stephani) 4: 313,
1910 (Stephani 1910b).
374 Porella densifolia was resolved as paraphyletic in the phylogeny of Hentschel et al. (2007b), suggesting
that the species is a complex deserving further study.
World checklist of hornworts and liverworts
421
*** Porella elegantula (Mont.) E.A.Hodgs., Svensk Bot. Tidskr. 42 (3): 277, 1948
(Hodgson and Sainsbury 1948). Bas.: Madotheca elegantula Mont., Ann. Sci. Nat.
Bot. (sér. 2) 19: 255, 1843 (Montagne 1843).
** Porella faurieri (Steph.) S.Hatt., J. Jap. Bot. 20: 109, 1944 (Hattori 1944c). Bas.:
Madotheca faurieri Steph., Sp. Hepat. (Stephani) 4: 315, 1910 (Stephani 1910b).
** Porella fengii P.C.Chen et S.Hatt., J. Hattori Bot. Lab. 30: 133, 1967 (Hattori 1967).
** Porella geheebii (Steph.) S.Hatt., Bot. Mag. (Tokyo) 64 (755/756): 114, 1951
(Hattori 1951c). Bas.: Madotheca geheebii Steph., Sp. Hepat. (Stephani) 4: 290,
1910 (Stephani 1910b).
** Porella gracillima Mitt., Trans. Linn. Soc. London, Bot. 3 (3): 202, 1891 (Mitten 1891).
** Porella grandifolia (Steph.) S.Hatt., J. Hattori Bot. Lab. 30: 136, 1967 (Hattori
1967). Bas.: Madotheca grandifolia Steph., Sp. Hepat. (Stephani) 4: 289, 1910
(Stephani 1910b).
** Porella grandiloba Lindb., Acta Soc. Sci. Fenn. 10: 234, 1872 [1873] (Lindberg 1872b).
** Porella grollei S.Hatt., J. Hattori Bot. Lab. 34: 411, 1971 (Hattori 1971b).
** Porella handelii S.Hatt., J. Hattori Bot. Lab. 33: 65, 1970 (Hattori 1970).
** Porella hattorii Udar et Shaheen, Lindbergia 9 (1): 70, 1983 (Udar and Shaheen 1983b).
** Porella hoeana S.Hatt., Misc. Bryol. Lichenol. 7 (5): 86, 1976 (Hattori 1976c).
* Porella imbricata Lour., Fl. Cochinch. 2: 683, 1790 (Loureiro 1790).
** Porella inaequalis (Gottsche) Perss., Arch. Soc. Zool. Bot. Fenn. “Vanamo”, suppl.
9: 225, 1955 (Persson 1955). Bas.: Madotheca inaequalis Gottsche, Sp. Hepat.
(Stephani) 4: 251, 1910 (Stephani 1910b).
** Porella japonica (Sande Lac.) Mitt., Trans. Linn. Soc. London, Bot. 3 (3): 202,
1891 (Mitten 1891). Bas.: Madotheca japonica Sande Lac., Syn. hepat. jav.: 105,
1856 [1857] (Sande Lacoste 1856b).
** Porella japonica subsp. appalachiana R.M.Schust., Hepat. Anthocerotae N. Amer.
4: 682, 1980 (Schuster 1980c).
** Porella japonica var. densespinosa S.Hatt. et M.X.Zhang, J. Jap. Bot. 60 (11): 324,
1985 (Hattori and Zhang 1985).
** Porella javanica (Gottsche) Inoue, J. Hattori Bot. Lab. 30: 60, 1967 (Inoue 1967a).
Bas.: Madotheca javanica Gottsche, Sp. Hepat. (Stephani) 4: 290, 1910 (Stephani
1910b).
** Porella latifolia J.S.Lou et Q.Li, Acta Phytotax. Sin. 25 (6): 482, 1987 (Lou 1987).
*** Porella leiboldii (Lehm.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 407, 1877 (Trevisan 1877). Bas.: Madotheca leiboldii Lehm., Nov.
Stirp. Pug. 8: 11, 1844 (Lehmann 1844).
** Porella longifolia (Steph.) S.Hatt., J. Hattori Bot. Lab. 32: 351, 1969 (Hattori
1969). Bas.: Madotheca longifolia Steph., Sp. Hepat. (Stephani) 4: 305, 1910 (Stephani 1910b).
** Porella madagascariensis (Nees et Mont.) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 407, 1877 (Trevisan 1877). Bas.: Lejeunea madagascariensis Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 5: 57, 1836 (Nees and Montagne
1836).
422
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**
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**
**
**
**
**
**
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**
***
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Porella maxima (Steph.) M.L.So, Syst. Bot. 27 (1): 11, 2002 (So 2002a). Bas.:
Madotheca maxima Steph., Sp. Hepat. (Stephani) 4: 291, 1910 (Stephani 1910b). 375
Porella mexicana (Hampe ex Gottsche, Lindenb. et Nees) Trevis., Mem. Reale
Ist. Lombardo Sci. (Ser. 3), C. Sci. Mat. 4 (13): 407, 1877 (Trevisan 1877). Bas.:
Madotheca mexicana Hampe ex Gottsche, Lindenb. et Nees, Syn. Hepat. 2: 273,
1845 (Gottsche et al. 1845a).
Porella montantii (Steph.) E.W.Jones, Trans. Brit. Bryol. Soc. 4 (3): 460, 1963
(Jones 1963). Bas.: Madotheca montantii Steph., Sp. Hepat. (Stephani) 4: 259,
1910 (Stephani 1910b).
Porella navicularis (Lehm. et Lindenb.) Pfeif., Fl. Niederhessen 2: 234, 1855
(Pfeifer 1855). Bas.: Jungermannia navicularis Lehm. et Lindenb., Nov. Stirp.
Pug. 6: 38, 1834 (Lehmann 1834).
Porella nitens (Steph.) S.Hatt., Fl. E. Himalaya: 525, 1966 (Hattori 1966c). Bas.:
Madotheca nitens Steph., Mém. Soc. Nat. Sci. Nat. Math. Cherbourg 29: 220,
1894 (Stephani 1894b).
Porella oblongifolia S.Hatt., J. Jap. Bot. 19 (7): 200, 1943 (Hattori 1943c). 376
Porella obtusata (Taylor) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 407, 1877 (Trevisan 1877). Bas.: Madotheca obtusata Taylor, London
J. Bot. 5: 380, 1846 (Taylor 1846b).
Porella obtusata var. macroloba (Steph.) S.Hatt. et M.X.Zhang, J. Jap. Bot. 60
(11): 325, 1985 (Hattori and Zhang 1985). Bas.: Madotheca macroloba Steph., Sp.
Hepat. (Stephani) 4: 292, 1910 (Stephani 1910b).
Porella obtusiloba S.Hatt., J. Hattori Bot. Lab. 33: 69, 1970 (Hattori 1970).
Porella perrottetiana (Mont.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 408, 1877 (Trevisan 1877). Bas.: Madotheca perrottetiana Mont.,
Ann. Sci. Nat. Bot. (sér. 2) 18: 15, 1842 (Montagne 1842b).
Porella perrottetiana var. angustifolia Pócs, J. Hattori Bot. Lab. 31: 75, 1968 (Pócs 1968).
Porella perrottetiana var. ciliatodentata (P.C.Chen et P.C.Wu) S.Hatt., J. Hattori
Bot. Lab. 30: 144, 1967 (Hattori 1967). Bas.: Porella ciliatodentata P.C.Chen et
P.C.Wu, Obs. l. Hwangs.: 8, 1965 (Chen and Wu 1965).
Porella perrottetiana var. triciliata (Steph.) Pócs, J. Hattori Bot. Lab. 31: 75, 1968
(Pócs 1968). Bas.: Madotheca triciliata Steph., Sp. Hepat. (Stephani) 4: 308, 1910
(Stephani 1910b).
Porella pinnata L., Sp. Pl. 1: 1106, 1753 (Linnaeus 1753).
Porella planifolia J.S.Lou, Coll. Pap. Quing-Zang Huang-Den 1: 277, 1983 (Lou
and Wang 1983).
Porella platyphylla (L.) Pfeif., Fl. Niederhessen 2: 234, 1855 (Pfeifer 1855). Bas.:
Jungermannia platyphylla L., Sp. Pl. 1: 1134, 1753 (Linnaeus 1753).
375 Porella maxima was accepted by So (2002a), but it was treated as conspeciic with Porella viridissima
in Hattori (1976f, 1986d). his synonymy is doubtful and further study is necessary to clarify the
status.
376 Porella oblongifolia is closely related to Porella densifolia (Hentschel et al. 2007b).
World checklist of hornworts and liverworts
423
*** Porella platyphylloidea (Schwein.) Lindb., Morgonbladet (Helsinki) 1876 (287, 10
Dec.): 1, 1876 (Lindberg 1876c). Bas.: Jungermannia platyphylloidea Schwein.,
Spec. Fl. Amer. Crypt.: 9, 1821 (Schweinitz 1821).
** Porella plicata J.S.Lou, Acta Phytotax. Sin. 18 (1): 119, 1980 (Lou and Wu 1980).
** Porella plumosa (Mitt.) Parihar, Univ. Allahabad Stud., Bot. 1961-2: 17, 1962
(Parihar 1962). Bas.: Madotheca plumosa Mitt., J. Proc. Linn. Soc., Bot. 5 (18):
108, 1860 [1861] (Mitten 1860c).
** Porella prolixa (Gottsche) E.W.Jones, Trans. Brit. Bryol. Soc. 4 (3): 460, 1963 (Jones 1963). Bas.: Madotheca prolixa Gottsche, Sp. Hepat. (Stephani) 4: 260, 1910
(Stephani 1910b).
** Porella pulcherrima Herzog et S.Hatt., Bull. Natl. Sci. Mus. Tokyo, B 12 (1): 34,
1986 (Hattori 1986a).
*** Porella relexa (Lehm. et Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser.
3), C. Sci. Mat. 4 (13): 408, 1877 (Trevisan 1877). Bas.: Jungermannia relexa
Lehm. et Lindenb., Nov. Stirp. Pug. 5: 5, 1833 (Lehmann 1833).
** Porella revoluta (Lehm. et Lindenb.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser.
3), C. Sci. Mat. 4 (13): 407, 1877 (Trevisan 1877). Bas.: Jungermannia revoluta
Lehm. et Lindenb., Nov. Stirp. Pug. 4: 18, 1832 (Lehmann 1832).
** Porella revoluta var. propinqua (C.Massal.) S.Hatt., J. Hattori Bot. Lab. 30: 148,
1967 (Hattori 1967). Bas.: Madotheca propinqua C.Massal., Hepat. Shen-si: 27,
1897 (Massalongo 1897).
** Porella roellii Steph., Bot. Centralbl. 45: 203, 1891 (Röll 1891).
*** Porella saccata M.L.So, New Zealand J. Bot. 43 (1): 302, 2005 (So 2005b).
** Porella sichuanensis S.Hatt. et K.C.Chang, Bull. Bot. Res., Harbin 8 (2): 43, 1988
(Chang 1988).
** Porella spinulosa (Steph.) S.Hatt., J. Hattori Bot. Lab. 33: 74, 1970 (Hattori 1970).
Bas.: Madotheca spinulosa Steph., Sp. Hepat. (Stephani) 6: 529, 1924 (Stephani 1924).
*** Porella squamulifera (Taylor) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 407, 1877 (Trevisan 1877). Bas.: Madotheca squamulifera Taylor,
London J. Bot. 5: 378, 1846 (Taylor 1846b).
** Porella stephaniana (C.Massal.) S.Hatt., J. Hattori Bot. Lab. 5: 81, 1951 (Hattori
1951b). Bas.: Madotheca stephaniana C.Massal., Hepat. Shen-si: 23, 1897 (Massalongo 1897). 377
** Porella subdentata (Mitt.) Steph., Hedwigia 30 (5): 203, 1891 (Stephani 1891a). Bas.:
Madotheca subdentata Mitt., J. Proc. Linn. Soc., Bot. 7 (27): 167, 1863 (Mitten 1863).
** Porella subdentata var. camerunensis E.W.Jones, Trans. Brit. Bryol. Soc. 4 (3): 456,
1963 (Jones 1963).
** Porella subobtusa (Steph.) S.Hatt., J. Jap. Bot. 20: 111, 1944 (Hattori 1944c). Bas.:
Madotheca subobtusa Steph., Sp. Hepat. (Stephani) 4: 311, 1910 (Stephani 1910b).
** Porella subparaphyllina J.S.Lou, Acta Phytotax. Sin. 25 (6): 483, 1987 (Lou 1987).
377 Porella stephaniana is closely related to Porella densifolia (Hentschel et al. 2007b).
424
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Porella subsquarrosa (Nees et Mont.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 407, 1877 (Trevisan 1877). Bas.: Lejeunea subsquarrosa Nees et
Mont., Ann. Sci. Nat. Bot. (sér. 2) 5: 57, 1836 (Nees and Montagne 1836).
*** Porella swailsii Grolle, J. Bryol. 10 (3): 270, 1979 (Grolle 1979b). Nom. nov. pro
Madotheca apiculata Herzog, Feddes Repert. Spec. Nov. Regni Veg. 57 (1/2): 197,
1955 (Herzog 1955).
*** Porella swartziana (F.Weber) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3),
C. Sci. Mat. 4 (13): 407, 1877 (Trevisan 1877). Bas.: Jungermannia swartziana
F.Weber, Hist. Musc. Hepat. Prodr.: 18, 1815 (Weber 1815).
** Porella triquetra (Steph.) E.W.Jones, Trans. Brit. Bryol. Soc. 4 (3): 454, 1963
(Jones 1963). Bas.: Madotheca triquetra Steph., Bot. Jahrb. Syst. 20 (3): 321, 1895
(Stephani 1895a).
** Porella truncata J.S.Lou, Acta Phytotax. Sin. 18 (1): 119, 1980 (Lou and Wu 1980).
** Porella ulophylla (Steph.) S.Hatt., Bull. Tokyo Sci. Mus. 11: 92, 1944 (Hattori
1944d). Bas.: Madotheca ulophylla Steph., Bull. Herb. Boissier 5 (2): 97, 1897
(Stephani 1897b).
** Porella undatorevoluta J.S.Lou, Acta Phytotax. Sin. 25 (6): 485, 1987 (Lou 1987).
** Porella urceolata S.Hatt., J. Hattori Bot. Lab. 33: 66, 1970 (Hattori 1970).
** Porella urogea (C.Massal.) S.Hatt., J. Hattori Bot. Lab. 32: 349, 1969 (Hattori 1969).
Bas.: Madotheca urogea C.Massal., Hepat. Shen-si: 28, 1897 (Massalongo 1897).
** Porella vallis-gratiae (Gottsche) E.W.Jones, Trans. Brit. Bryol. Soc. 4 (3): 450,
1963 (Jones 1963). Bas.: Madotheca vallis-gratiae Gottsche, Sp. Hepat. (Stephani)
4: 261, 1910 (Stephani 1910b).
** Porella variabilis (Kashyap et R.S.Chopra) Parihar, Univ. Allahabad Stud.,
Bot. 1961-2: 17, 1962 (Parihar 1962). Bas.: Madotheca variabilis Kashyap et
R.S.Chopra, Liverworts W. Himal. 2: 33, 1932 (Kashyap and Chopra 1932).
** Porella vernicosa Lindb., Acta Soc. Sci. Fenn. 10: 223, 1872 [1873] (Lindberg 1872b).
** Porella viridissima (Mitt.) Grolle, J. Hattori Bot. Lab. 36: 83, 1972 [1973] (Grolle
and Schultze-Motel 1972). Bas.: Madotheca viridissima Mitt., Fl. vit.: 411, 1871
[1873] (Mitten 1871).
** Porella wataugensis (Sull.) Underw. ex M.Howe, Bull. Torrey Bot. Club 24 (11):
519, 1897 (Howe 1897b). Bas.: Madotheca wataugensis Sull., Musc. Hepat. U.S.:
700, 1856 (Sullivant 1856).
Radulineae R.M.Schust.
***
Radulaceae Müll.Frib.
by M.A.M. Renner
he treatment of Radulaceae follows Devos et al. (2011). Taxonomic and nomenclatural notes can also be found in Renner et al. (2013b, 2014).
World checklist of hornworts and liverworts
425
*** Radula Dumort., Commentat. Bot. (Dumortier): 112, 1822 (Dumortier 1822)
nom. conserv.
*** subg. Amentuloradula Devos, M.A.M.Renner, Gradst., A.J.Shaw et Vanderp.,
Taxon 60 (6): 1630, 2011 (Devos et al. 2011).
*** Radula amentulosa Mitt., Bonplandia 9 (24): 367, 1861 (Mitten 1861).
*** Radula aneurismalis (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat.
2: 262, 1845 (Gottsche et al. 1845a). Bas.: Jungermannia aneurismalis Hook.f. et
Taylor, London J. Bot. 4: 86, 1845 (Hooker and Taylor 1845).
** Radula ceylanica K.Yamada, J. Jap. Bot. 50 (12): 373, 1975 (Yamada 1975b).
*** Radula issifolia Steph., Sp. Hepat. (Stephani) 6: 507, 1924 (Stephani 1924).
*** Radula formosa (C.F.W.Meissn. ex Spreng.) Nees, Syn. Hepat. 2: 258, 1845
(Gottsche et al. 1845a). Bas.: Jungermannia formosa C.F.W.Meissn. ex Spreng.
Syst. Veg. (ed. 16) [Sprengel] 4 (2): 325, 1827 (Sprengel 1827b).
*** Radula helix (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 2: 260,
1845 (Gottsche et al. 1845a). Bas.: Jungermannia helix Hook.f. et Taylor, London
J. Bot. 3: 475, 1844 (Hooker and Taylor 1844b).
*** Radula hicksiae K.Yamada, Cryptog. Bryol. Lichénol. 5 (1/2): 191, 1984 (Yamada
1984a).
*** Radula iwatsukii K.Yamada, J. Hattori Bot. Lab. 45: 275, 1979 (Yamada 1979b).
** Radula morobeana K.Yamada et Piippo, Ann. Bot. Fenn. 26 (4): 358, 1989 (Yamada and Piippo 1989).
** Radula multiamentula E.A.Hodgs., Rec. Domin. Mus. 4 (11): 122, 1962 (Hodgson 1962a).
*** Radula ornata E.A.Br. et Pócs, Telopea 9 (3): 436, 2001 (Brown and Pócs 2001).
*** Radula physoloba Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 255, 1843 (Montagne 1843).
*** Radula pseudoscripta M.A.M.Renner, New Zealand J. Bot. 44 (3): 340, 2006
(Renner 2006).
** Radula queenslandica K.Yamada, J. Hattori Bot. Lab. 62: 192, 1987 (Yamada 1987).
*** Radula scariosa Mitt., Bonplandia 9 (24): 367, 1861 (Mitten 1861).
*** Radula splendida M.A.M.Renner et Devos, Nova Hedwigia 90 (1/2): 113, 2010
(Renner et al. 2010a).
* Radula squarrosa K.Yamada, J. Jap. Bot. 65 (1): 1, 1990 (Yamada 1990). 378
*** Radula thiersiae K.Yamada, J. Hattori Bot. Lab. 62: 198, 1987 (Yamada 1987).
*** Radula uvifera (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 2:
258, 1845 (Gottsche et al. 1845a). Bas.: Jungermannia uvifera Hook.f. et Taylor,
London J. Bot. 3: 292 [392], 1844 (Hooker and Taylor 1844a).
* Radula vagans Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9): 85,
1911 (Stephani 1911b).
** Radula verrucosa K.Yamada, J. Hattori Bot. Lab. 45: 277, 1979 (Yamada 1979b).
378 Radula squarrosa is probably conspeciic with Radula morobeana.
426
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** subg. Cladoradula Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 315, 1885
(Spruce 1885).
** Radula auriculata Steph., Bull. Herb. Boissier 5 (2): 105, 1897 (Stephani 1897b).
* Radula bipinnata Mitt., J. Proc. Linn. Soc., Bot. 7 (27): 166, 1863 (Mitten 1863). 379
*** Radula boryana (F.Weber) Nees ex Mont., Ann. Sci. Nat. Bot. (sér. 2) 18: 13,
1842 (Montagne 1842b). Bas.: Jungermannia boryana F.Weber, Hist. Musc.
Hepat. Prodr.: 58, 1815 (Weber 1815).
*** Radula campanigera Mont., London J. Bot. 3: 634, 1844 (Montagne 1844a).
*** Radula campanigera subsp. obiensis (S.Hatt.) K.Yamada, J. Hattori Bot. Lab. 45:
309, 1979 (Yamada 1979b). Bas.: Radula obiensis S.Hatt., Bull. Tokyo Sci. Mus.
11: 83, 1944 (Hattori 1944d).
** Radula chinensis Steph., Nuovo Giorn. Bot. Ital. (n.ser.) 13 (4): 355, 1906 (Levier
1906).
*** Radula gottscheana Taylor, London J. Bot. 5: 374, 1846 (Taylor 1846b).
*** Radula perrottetii Gottsche, Hedwigia 23 (10): 154, 1884 (Stephani 1884a).
** Radula tenax Lindb., Acta Soc. Sci. Fenn. 10: 492, 1875 (Lindberg 1875).
*** subg. Dactyloradula Devos, M.A.M.Renner, Gradst., A.J.Shaw et Vanderp.,
Taxon 60 (6): 1630, 2011 (Devos et al. 2011).
*** Radula brunnea Steph., Sp. Hepat. (Stephani) 4: 232, 1910 (Stephani 1910b).
*** subg. Metaradula R.M.Schust., Phytologia 56 (2): 69, 1984 (Schuster 1984).
** Radula acuminata Steph., Sp. Hepat. (Stephani) 4: 230, 1910 (Stephani 1910b). 380
** Radula aguirrei R.M.Schust., Phytotaxa 202 (1): 70, 2015 (Söderström et al.
2015c). Based on: Radula aguirrei R.M.Schust., J. Hattori Bot. Lab. 70: 56, 1991
(Schuster 1991a), nom. inval.
** Radula anisotoma M.A.M.Renner, PhytoKeys 27: 30, 2013 (Renner et al. 2013a).
*** Radula assamica Steph., Hedwigia 23 (10): 151, 1884 (Stephani 1884a).
*** Radula australiana K.Yamada, J. Hattori Bot. Lab. 51: 323, 1982 (Yamada 1982a).
*** Radula buccinifera (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat.
2: 261, 1845 (Gottsche et al. 1845a). Bas.: Jungermannia buccinifera Hook.f. et
Taylor, London J. Bot. 3: 580, 1844 (Hooker and Taylor 1844c).
*** Radula demissa M.A.M.Renner, PhytoKeys 27: 53, 2013 (Renner et al. 2013a).
** Radula evansii Castle, Ann. Bryol. 11: 37, 1938 (Castle 1938).
** Radula laccida Lindenb. et Gottsche, Syn. Hepat. 5: 726, 1847 (Gottsche et al. 1847).
** Radula laccida var. brachycalyx Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:
321, 1885 (Spruce 1885).
*** Radula foricata M.A.M.Renner, Austral. Syst. Bot. 26 (4): 307, 2013 (Renner et
al. 2013c).
** Radula grevilleana Taylor, Ann. Mag. Nat. Hist. 20 (135): 380, 1847 (Taylor 1847a).
379 Radula bipinnata is distinct from Radula boryana according to molecular evidence (Devos et al. 2011).
380 Radula acuminata may be conspeciic with Radula tjibodensis.
World checklist of hornworts and liverworts
427
*** Radula imposita M.A.M.Renner, PhytoKeys 27: 65, 2013 (Renner et al. 2013a).
** Radula jovetiana K.Yamada, Cryptog. Bryol. Lichénol. 5 (1/2): 193, 1984 (Yamada
1984a).
** Radula kilgourii M.A.M.Renner, Austral. Syst. Bot. 26 (4): 313, 2013 (Renner et
al. 2013c).
** Radula loriana Castle, J. Hattori Bot. Lab. 21: 6, 1959 (Castle 1959).
*** Radula mammosa Spruce, Mem. Torrey Bot. Club 1 (3): 127, 1890 (Spruce 1890).
*** Radula mittenii Steph., Hedwigia 23 (10): 148, 1884 (Stephani 1884a).
*** Radula myriopoda M.A.M.Renner, Austral. Syst. Bot. 26 (4): 323, 2013 (Renner
et al. 2013c).
*** Radula notabilis M.A.M.Renner, PhytoKeys 27: 77, 2013 (Renner et al. 2013a).
** Radula nymannii Steph., Sp. Hepat. (Stephani) 4: 229, 1910 (Stephani 1910b).
** Radula protensa Lindenb., Bot. Zeitung (Berlin) 6 (25): 462, 1848 (Meissner 1848).
** Radula protensa var. erectilobula Schifn., Nova Acta Acad. Caes. Leop.-Carol. German.
Nat. Cur. 60 (2): 247, 1893 (Schifner 1893a).
** Radula pseudolaccida E.W.Jones, J. Bryol. 9 (4): 501, 1977 [1978] (Jones 1977).
** Radula psychosis M.A.M.Renner, Austral. Syst. Bot. 26 (4): 328, 2013 (Renner et
al. 2013c).
*** Radula ratkowskiana K.Yamada, J. Jap. Bot. 59 (3): 94, 1984 (Yamada 1984b).
*** Radula robinsonii Steph., Sp. Hepat. (Stephani) 4: 214, 1910 (Stephani 1910b).
** Radula stenocalyx Mont., Ann. Sci. Nat. Bot. (sér. 4) 3 (5): 315, 1855 (Montagne
1855).
*** Radula strangulata Hook.f. et Taylor, London J. Bot. 5: 377, 1846 (Taylor 1846b).
* Radula tjibodensis K.I.Goebel, Ann. Jard. Bot. Buitenzorg 7 (1): 53, 1888 (Goebel
1888). 381
*** Radula ventricosa Steph., Sp. Hepat. (Stephani) 4: 187, 1910 (Stephani 1910b).
*** Radula yanoella R.M.Schust., Phytologia 56 (2): 72, 1984 (Schuster 1984).
subg. Odontoradula K.Yamada, J. Hattori Bot. Lab. 45: 209, 1979 (Yamada 1979b).
Radula acuta Mitt., Fl. vit.: 410, 1871 [1873] (Mitten 1871).
Radula allisonii Castle, Rev. Bryol. Lichénol. 31 (3/4): 148, 1962 [1963] (Castle 1962).
Radula amoena Herzog, Mitt. Inst. Allg. Bot. Hamburg 7 (3): 192, 1931 (Herzog
1931a).
** Radula anceps Sande Lac., Ned. Kruidk. Arch. 3: 419, 1854 [1855] (Sande Lacoste
1854).
* Radula crenulata Schifn., Leberm., Forschungsr. Gazelle 4 (4): 21, 1890 (Schifner
1890). 382
*** Radula cuspidata Steph., Sp. Hepat. (Stephani) 4: 156, 1910 (Stephani 1910b).
*** Radula decora Gottsche, Hedwigia 23 (10): 145, 1884 (Stephani 1884a).
***
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**
381 Radula tjibodensis is a doubtful taxon. he type specimen has not been found.
382 Radula crenulata is conspeciic with Radula acuta in So (2006) and possibly conspeciic with Radula
apiculata in Yamada and Piippo (1989), but difers morphologically from both and needs to be further studied.
428
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Radula emarginata K.Yamada et Piippo, Ann. Bot. Fenn. 26 (4): 352, 1989 (Yamada and Piippo 1989).
** Radula kojana Steph., Bull. Herb. Boissier 5 (2): 105, 1897 (Stephani 1897b).
*** Radula lacerata Steph., Rev. Bryol. 35 (2): 33, 1908 (Stephani 1908l).
*** Radula novae-hollandiae Hampe, Nov. Stirp. Pug. 7: 24, 1838 (Lehmann 1838).
*** Radula ocellata K.Yamada, J. Hattori Bot. Lab. 45: 209, 1979 (Yamada 1979b).
*** Radula plicata Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 154, 1854 (Mitten 1854).
** Radula pugioniformis M.A.M.Renner, PhytoKeys 27: 84, 2013 (Renner et al. 2013a).
*** Radula pulchella Mitt., Hedwigia 23 (10): 149, 1884 (Stephani 1884a).
*** Radula retrolexa Taylor, London J. Bot. 5: 378, 1846 (Taylor 1846b).
*** Radula tasmanica Steph., Sp. Hepat. (Stephani) 4: 212, 1910 (Stephani 1910b).
*** Radula weymouthiana Steph., Sp. Hepat. (Stephani) 4: 190, 1910 (Stephani 1910b).
***
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subg. Radula
Radula acutiloba Steph., Hedwigia 28 (4): 271, 1889 (Stephani 1889c). 383
Radula appressa Mitt., Philos. Trans. 168: 397, 1879 (Mitten 1879).
Radula aquilegia (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 2:
260, 1845 (Gottsche et al. 1845a). Bas.: Jungermannia aquilegia Hook.f. et Taylor,
London J. Bot. 3: 291 [391], 1844 (Hooker and Taylor 1844a).
Radula australis Austin, Bot. Bull. (Hanover) 1 (7): 32, 1876 (Austin 1876b).
Radula borneensis Steph., Sp. Hepat. (Stephani) 4: 209, 1910 (Stephani 1910b).
Radula caduca K.Yamada, J. Hattori Bot. Lab. 45: 225, 1979 (Yamada 1979b).
Radula carringtonii J.B.Jack, Flora 64 (25): 385, 1881 (Jack 1881).
Radula complanata (L.) Dumort., Syll. Jungerm. Europ.: 38, 1831 (Dumortier
1831). Bas.: Jungermannia complanata L., Sp. Pl. 1: 1133, 1753 (Linnaeus 1753).
Radula constricta Steph., Sp. Hepat. (Stephani) 6: 506, 1924 (Stephani 1924).
Radula evelynae K.Yamada, J. Jap. Bot. 50 (4): 115, 1975 (Yamada 1975a).
Radula fendleri Gottsche, Hedwigia 23 (10): 146, 1884 (Stephani 1884a). 384
Radula grandis Steph., J. Linn. Soc., Bot. 29 (201): 271, 1892 (Stephani 1892b).
Radula japonica Gottsche, Hedwigia 23 (10): 152, 1884 (Stephani 1884a).
Radula javanica Gottsche, Syn. Hepat. 2: 257, 1845 (Gottsche et al. 1845a).
Radula jonesii Bouman, Dirkse et K.Yamada, J. Bryol. 15 (1): 161, 1988 (Bouman
et al. 1988).
Radula lindenbergiana Gottsche ex C.Hartm., Handb. Skand. l. (ed.9): 98, 1864
(Hartman 1864).
Radula madagascariensis Gottsche, Abh. Naturwiss. Vereins Bremen 7: 349, 1882
(Gottsche 1882).
Radula marojezica E.W.Jones, J. Bryol. 17 (2): 307, 1992 (Jones 1992).
383 Radula acutiloba is tentatively placed in subg. Radula. he specimen labelled Radula acutiloba in
Devos et al. (2011) was misidentiied and should be Radula australiana.
384 Radula fendleri may be conspeciic with Radula madagascariensis.
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429
Radula multilora Gottsche ex Schifn., Leberm., Forschungsr. Gazelle 4 (4): 20,
1890 (Schifner 1890). 385
Radula novoguineensis K.Yamada et Piippo, Ann. Bot. Fenn. 26 (4): 360, 1989
(Yamada and Piippo 1989).
Radula obconica Sull., Manual (Gray): 688, 1848 (Gray 1848).
Radula obtusiloba Steph., Bull. Herb. Boissier 5 (2): 105, 1897 (Stephani 1897b).
Radula obtusiloba subsp. polyclada (A.Evans) S.Hatt., J. Hattori Bot. Lab. 29: 275,
1966 (Hattori 1966d). Bas.: Radula polyclada A.Evans, Bull. Torrey Bot. Club 41
(12): 607, 1914 [1915] (Evans 1914a).
Radula oceania Castle, Rev. Bryol. Lichénol. 33 (3/4): 390, 1965 (Castle 1965). 386
Radula oreopsis M.A.M.Renner, Telopea 17: 123, 2014 (Renner 2014).
Radula portoricensis Steph., Hedwigia 27 (11/12): 298, 1888 (Stephani 1888c).
Radula prolifera Arnell, Ark. Bot. 13 (2): 12, 1913 (Arnell 1913).
Radula quadrata Gottsche, Syn. Hepat. 2: 255, 1845 (Gottsche et al. 1845a).
Radula relexa Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 255, 1843 (Montagne
1843).
Radula sharpii K.Yamada, J. Jap. Bot. 60 (9): 260, 1985 (Yamada 1985a).
Radula sumatrana Steph., Sp. Hepat. (Stephani) 4: 204, 1910 (Stephani 1910b).
Radula tokiensis Steph., Hedwigia 23 (10): 150, 1884 (Stephani 1884a).
Radula van-zantenii K.Yamada, J. Hattori Bot. Lab. 45: 260, 1979 (Yamada 1979b).
Radula varilobula Castle, J. Hattori Bot. Lab. 21: 19, 1959 (Castle 1959).
Radula wichurae Steph., Sp. Hepat. (Stephani) 4: 168, 1910 (Stephani 1910b).
*** subg. Volutoradula Devos, M.A.M.Renner, Gradst., A.J.Shaw et Vanderp.,
Taxon 60 (6): 1629, 2011 (Devos et al. 2011).
** Radula ankeinensis Gottsche, Hedwigia 23 (10): 152, 1884 (Stephani 1884a).
** Radula antilleana Castle, J. Hattori Bot. Lab. 21: 48, 1959 (Castle 1959).
** Radula comorensis Steph., Hedwigia 23 (9): 132, 1884 (Stephani 1884c).
** Radula cubensis K.Yamada, J. Hattori Bot. Lab. 54: 241, 1983 (Yamada 1983).
** Radula diversifolia Steph., Sp. Hepat. (Stephani) 4: 212, 1910 (Stephani 1910b).
*** Radula eggersii K.Yamada, J. Hattori Bot. Lab. 82: 339, 1997 (Yamada 1997).
** Radula episcia Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 318, 1885 (Spruce
1885).
** Radula loridana Castle, Rev. Bryol. Lichénol. 36 (1/2): 1, 1968 [1969] (Castle 1968).
** Radula fulvifolia (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 2:
261, 1845 (Gottsche et al. 1845a). Bas.: Jungermannia fulvifolia Hook.f. et Taylor,
London J. Bot. 4: 85, 1845 (Hooker and Taylor 1845).
*** Radula hastata Steph., Sp. Hepat. (Stephani) 4: 163, 1910 (Stephani 1910b).
* Radula holstiana Steph., Bot. Jahrb. Syst. 20 (3): 320, 1895 (Stephani 1895a).
385 Radula multilora is similar to, and may be conspeciic with Radula relexa, but it was accepted by
Renner and de Lange (2009).
386 Radula oceania was considered conspeciic with Radula javanica by So (2006), but molecular data
does not support it (Renner 2014).
430
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Radula holtii Spruce, J. Bot. 25: 209, 1887 (Spruce 1887b).
Radula husnotii Castle, J. Hattori Bot. Lab. 21: 45, 1959 (Castle 1959).
Radula inlexa Gottsche, Hedwigia 23 (10): 148, 1884 (Stephani 1884a).
Radula kegelii Gottsche ex Steph., Hedwigia 23 (10): 152, 1884 (Stephani 1884a).
Radula macroloba Steph., Bull. Soc. Roy. Bot. Belgique 31: 121, 1892 (Stephani
1892c).
Radula mazarunensis K.Yamada, Trop. Bryol. 1: 38, 1989 (Gradstein and
Florschütz-de Waard 1989).
Radula mexicana Lindenb. et Gottsche, Mexik. Leverm.: 150, 1863 (Gottsche 1863).
Radula microloba Gottsche, Syn. Hepat. 2: 259, 1845 (Gottsche et al. 1845a).
Radula neotropica Castle, J. Hattori Bot. Lab. 21: 31, 1959 (Castle 1959).
Radula nudicaulis Steph., Sp. Hepat. (Stephani) 4: 174, 1910 (Stephani 1910b).
Radula nudicaulis var. delicatula P.Allorge et V.Allorge, Rev. Bryol. Lichénol. 19
(1/2): 106, 1950 (Allorge and Allorge 1950).
Radula pocsii K.Yamada, J. Hattori Bot. Lab. 54: 245, 1983 (Yamada 1983).
Radula recubans Taylor, London J. Bot. 5: 376, 1846 (Taylor 1846b).
Radula saccatiloba Steph., Hedwigia 23 (8): 129, 1884 (Stephani 1884b).
Radula schaefer-verwimpii K.Yamada, J. Jap. Bot. 65 (1): 3, 1990 (Yamada 1990).
Radula schoieldiana K.Yamada, J. Hattori Bot. Lab. 82: 337, 1997 (Yamada 1997).
Radula stipatilora Steph., Sp. Hepat. (Stephani) 4: 159, 1910 (Stephani 1910b).
Radula striata Mitt., Hedwigia 23 (10): 155, 1884 (Stephani 1884a).
Radula subinlata Lindenb. et Gottsche, Syn. Hepat. 5: 724, 1847 (Gottsche et al. 1847).
Radula sullivantii Austin, Hepat. bor.-amer.: 22, 1873 (Austin 1873).
Radula tenera Mitt., Hedwigia 23 (10): 149, 1884 (Stephani 1884a).
Radula voluta Taylor, Syn. Hepat. 2: 255, 1845 (Gottsche et al. 1845a).
Incertae sedis
Radula acutangula Steph., Bull. Herb. Boissier 5 (10): 848, 1897 (Stephani 1897c).
Radula angulata Steph., Hedwigia 23 (8): 114, 1884 (Stephani 1884b).
Radula bogotensis Steph., Hedwigia 23 (8): 115, 1884 (Stephani 1884b).
Radula bolanderi Gottsche, Hedwigia 23 (10): 145, 1884 (Stephani 1884a).
Radula boninensis Furuki et K.Yamada, J. Jap. Bot. 61 (10): 312, 1986 (Furuki
and Yamada 1986).
Radula brasilica K.Yamada, J. Hattori Bot. Lab. 74: 35, 1993 (Yamada 1993).
Radula caespitosa Steph., Hedwigia 27 (3/4): 107, 1888 (Stephani 1888d).
Radula campanulata Lindenb. et Gottsche, Syn. Hepat. 2: 256, 1845 (Gottsche et
al. 1845a).
Radula castlei Grolle, Bryologist 73 (4): 662, 1970 (Grolle 1970a).
Radula cavifolia Hampe ex Gottsche, Lindenb. et Nees, Syn. Hepat. 2: 259, 1845
(Gottsche et al. 1845a).
Radula cochabambaensis K.Yamada, J. Hattori Bot. Lab. 74: 37, 1993 (Yamada 1993).
Radula conferta Lindenb. et Gottsche, Syn. Hepat. 5: 729, 1847 (Gottsche et al. 1847).
Radula cordata Mitt., Fl. vit.: 410, 1871 [1873] (Mitten 1871).
World checklist of hornworts and liverworts
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431
Radula costaricensis Gottsche, J. Bot. 15: 226, 1877 (Polakowski 1877).
Radula curvilobula M.L.So, J. Hattori Bot. Lab. 98: 176, 2005 (So 2005a).
Radula decurrens Mitt., Fl. vit.: 410, 1871 [1873] (Mitten 1871). 387
Radula densifolia Castle, Rev. Bryol. Lichénol. 33 (3/4): 385, 1965 (Castle 1965).
Radula diaphana K.I.Goebel, Organogr. Pl., ed. 2, 2 (1): 677, 1915 (Goebel 1915).
Radula dolabrata K.Yamada, J. Jap. Bot. 60 (9): 257, 1985 (Yamada 1985a).
Radula elliottii Castle, J. Hattori Bot. Lab. 21: 12, 1959 (Castle 1959).
Radula falcata Steph., Hedwigia 23 (8): 115, 1884 (Stephani 1884b).
Radula fauriana Steph., Sp. Hepat. (Stephani) 4: 207, 1910 (Stephani 1910b).
Radula fernandezana Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46
(9): 84, 1911 (Stephani 1911b).
Radula lavifolia (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat. 2:
259, 1845 (Gottsche et al. 1845a). Bas.: Jungermannia lavifolia Hook.f. et Taylor,
London J. Bot. 3: 476, 1844 (Hooker and Taylor 1844b).
Radula fujitae Furuki, Bryol. Res. 9 (5): 143, 2007 (Furuki 2007).
Radula galapagona Steph., Sp. Hepat. (Stephani) 4: 176, 1910 (Stephani 1910b).
Radula gedena Gottsche, Hedwigia 23 (10): 146, 1884 (Stephani 1884a).
Radula gracilis Mitt., Hedwigia 23 (10): 147, 1884 (Stephani 1884a).
Radula gradsteinii K.Yamada, Trop. Bryol. 1: 37, 1989 (Gradstein and Florschützde Waard 1989).
Radula grandifolia Steph., Sp. Hepat. (Stephani) 4: 184, 1910 (Stephani 1910b).
Radula grollei K.Yamada et Piippo, Ann. Bot. Fenn. 26 (4): 379, 1989 (Yamada
and Piippo 1989).
Radula guyanensis K.Yamada, Trop. Bryol. 1: 38, 1989 (Gradstein and Florschützde Waard 1989).
Radula hattorii K.Yamada, J. Jap. Bot. 60 (9): 259, 1985 (Yamada 1985a).
Radula hawaiica M.L.So, J. Hattori Bot. Lab. 98: 177, 2005 (So 2005a).
Radula hedingeri K.I.Goebel, Ann. Jard. Bot. Buitenzorg 7 (1): 51, 1888 (Goebel 1888).
Radula inouei K.Yamada, J. Hattori Bot. Lab. 45: 262, 1979 (Yamada 1979b).
Radula involvens Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 325, 1885
(Spruce 1885).
Radula iwatsukiana K.Yamada, J. Hattori Bot. Lab. 58: 114, 1985 (Yamada 1985b).
Radula jamaicensis Pearson, Ann. Bryol. 4: 103, 1931 (Pearson 1931b).
Radula jamesonii Taylor, London J. Bot. 5: 375, 1846 (Taylor 1846b).
Radula kinabaluensis K.Yamada, Misc. Bryol. Lichenol. 6 (6): 97, 1973 (Yamada
1973b).
Radula kitagawae K.Yamada, J. Hattori Bot. Lab. 58: 116, 1985 (Yamada 1985b).
Radula koponenii K.Yamada et Piippo, Ann. Bot. Fenn. 26 (4): 364, 1989 (Yamada and Piippo 1989).
Radula kurzii Steph., Hedwigia 23 (10): 153, 1884 (Stephani 1884a).
Radula laxiramea Steph., Sp. Hepat. (Stephani) 4: 178, 1910 (Stephani 1910b).
387 Radula decurrens may be conspeciic with Radula relexa.
432
**
**
***
***
**
*
***
***
**
**
**
***
**
**
**
***
***
**
**
**
**
**
**
**
*
***
***
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Radula leiboldii Steph., Hedwigia 23 (8): 116, 1884 (Stephani 1884b).
Radula lewisii K.Yamada, J. Hattori Bot. Lab. 74: 39, 1993 (Yamada 1993).
Radula ligula Steph., Sp. Hepat. (Stephani) 4: 228, 1910 (Stephani 1910b).
Radula lingulata Gottsche, Syn. Hepat. 2: 260, 1845 (Gottsche et al. 1845a).
Radula longiloba K.Yamada, J. Hattori Bot. Lab. 54: 243, 1983 (Yamada 1983).
Radula longispica Steph., Sp. Hepat. (Stephani) 4: 183, 1910 (Stephani 1910b). 388
Radula marginata Gottsche, Lindenb. et Nees, Syn. Hepat. 2: 261, 1845 (Gottsche
et al. 1845a). Nom. nov. pro Jungermannia marginata Hook.f. et Taylor, London
J. Bot. 3: 566, 1844 (Hooker and Taylor 1844a), nom. illeg.
Radula mauiensis M.L.So, J. Hattori Bot. Lab. 98: 178, 2005 (So 2005a).
Radula microlobula Castle, J. Hattori Bot. Lab. 21: 35, 1959 (Castle 1959).
Radula minutilobula K.Yamada et Piippo, Ann. Bot. Fenn. 26 (4): 377, 1989 (Yamada and Piippo 1989).
Radula mizutanii K.Yamada, J. Jap. Bot. 48 (5): 134, 1973 (Yamada 1973a).
Radula nigra Pearson, J. Linn. Soc., Bot. 46 (305): 31, 1922 (Pearson 1922b).
Radula nilgiriensis Udar et D.Kumar, J. Indian Bot. Soc. 61: 177, 1982 (Udar and
Kumar 1982b).
Radula norrisii K.Yamada et Piippo, Ann. Bot. Fenn. 26 (4): 374, 1989 (Yamada
and Piippo 1989).
Radula novivrieseana K.Yamada, J. Hattori Bot. Lab. 51: 326, 1982 (Yamada 1982a).
Radula novocaledonica Hürl. et K.Yamada, J. Jap. Bot. 54 (8): 238, 1979 (Hürlimann and Yamada 1979).
Radula novocaledoniensis K.Yamada, J. Hattori Bot. Lab. 58: 120, 1985 (Yamada
1985b).
Radula obovata Castle, J. Hattori Bot. Lab. 21: 16, 1959 (Castle 1959).
Radula obscura Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 107, 1860 [1861] (Mitten
1860c).
Radula okamurana Steph., Sp. Hepat. (Stephani) 4: 209, 1910 (Stephani 1910b).
Radula onraedtii K.Yamada, Misc. Bryol. Lichenol. 8 (6): 113, 1979 (Yamada 1979a).
Radula opaciuscula (Spruce) Castle, J. Hattori Bot. Lab. 21: 22, 1959 (Castle
1959). Bas.: Radula episcia var. opaciuscula Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 319, 1885 (Spruce 1885).
Radula ovalilobula K.Yamada, J. Hattori Bot. Lab. 45: 257, 1979 (Yamada 1979b).
Radula oyamensis Steph., Hedwigia 23 (10): 149, 1884 (Stephani 1884a).
Radula paganii Castle, J. Hattori Bot. Lab. 21: 33, 1959 (Castle 1959). 389
Radula pallens (Sw.) Nees ex Mont., Voy. Amér. Mérid. 7 (2): 71, 1839 (Montagne 1839a). Bas.: Jungermannia pallens Sw., Prodr. (Swartz): 143, 1788 (Swartz
1788).
Radula pandei Udar et Dh.Kumar, Lindbergia 9 (2): 133, 1983 (Udar and Kumar
1983b).
388 Radula longispica may be conspeciic with Radula javanica.
389 Radula paganii is doubtfully distinct from Radula neotropica.
World checklist of hornworts and liverworts
433
** Radula patens K.Yamada, Cryptog. Bryol. Lichénol. 5 (1/2): 197, 1984 (Yamada
1984a).
** Radula peruviana K.Yamada, Beih. Nova Hedwigia 88: 79, 1987 (Schultze-Motel
and Menzel 1987).
** Radula philippinensis K.Yamada, J. Hattori Bot. Lab. 45: 299, 1979 (Yamada 1979b).
* Radula pinnulata Mitt., Fl. vit.: 410, 1871 [1873] (Mitten 1871). 390
** Radula pseudostachya Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 319, 1885
(Spruce 1885).
** Radula punctata Steph., Hedwigia 23 (8): 135, 1884 (Stephani 1884b).
** Radula pusilla Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 320, 1885 (Spruce
1885).
*** Radula rhombiloba Steph., Sp. Hepat. (Stephani) 4: 204, 1910 (Stephani 1910b).
* Radula rupicola K.Yamada, J. Hattori Bot. Lab. 58: 124, 1985 (Yamada 1985b). 391
** Radula santacruziana K.Yamada et Gradst., Trop. Bryol. 4: 67, 1991 (Yamada and
Gradstein 1991).
** Radula silvestris Gottsche, Abh. Naturwiss. Vereins Bremen 7: 349, 1882 (Gottsche
1882).
** Radula sinskeana K.Yamada, J. Hattori Bot. Lab. 74: 41, 1993 (Yamada 1993).
** Radula sinuata Gottsche ex Steph., Sp. Hepat. (Stephani) 4: 161, 1910 (Stephani
1910b).
* Radula socorana Gerola, Lav. Bot. Ist. Bot. Univ. Padova 12: 475, 1947 (Gerola 1947).
** Radula sonsonensis Steph., Sp. Hepat. (Stephani) 4: 201, 1910 (Stephani 1910b).
** Radula stellatogemmipara C.Gao et Y.H.Wu, Nova Hedwigia 80 (1/2): 239, 2005
(Gao and Wu 2005).
** Radula subsimplex Steph., Hedwigia 23 (8): 130, 1884 (Stephani 1884b).
** Radula subsquarrosa S.W.Arnell, Ark. Bot. (n.ser.) 4 (1): 15, 1957 (Arnell 1957b).
** Radula tabularis Steph., Hedwigia 23 (9): 131, 1884 (Stephani 1884c).
** Radula taylorii Steph., Hedwigia 23 (9): 133, 1884 (Stephani 1884c).
** Radula tectiloba Steph., Hedwigia 27 (11/12): 298, 1888 (Stephani 1888c).
** Radula tenuis K.Yamada, J. Hattori Bot. Lab. 54: 247, 1983 (Yamada 1983).
** Radula underwoodii Castle, J. Hattori Bot. Lab. 21: 37, 1959 (Castle 1959).
** Radula venezuelensis K.Yamada, Misc. Bryol. Lichenol. 9 (6): 122, 1982 (Yamada
1982b).
*** Radula vieillardii Gottsche, Hedwigia 23 (10): 150, 1884 (Stephani 1884a).
** Radula visianica C.Massal., Ann. Bot. (Rome) 1 (4): 298, 1904 (Massalongo
1904).
** Radula vrieseana Sande Lac., Ann. Mus. Bot. Lugduno-Batavi 1: 305, 1864 (Sande
Lacoste 1864).
** Radula wrightii Castle, J. Hattori Bot. Lab. 21: 15, 1959 (Castle 1959).
390 Radula pinnulata was accepted by Yamada (1979b) and, indirectly through comparison with Radula
norrisii, by Yamada and Piippo (1989). It is conspeciic with Radula javanica according to So (2006).
391 Radula rupicola may be conspeciic with Radula vieillardii (So 2006).
434
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Radula xalapensis Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 5: 56, 1836 (Nees
and Montagne 1836).
Ptilidiales Schljakov
**
Herzogianthaceae Stotler et Crand.-Stotl.
** Herzogianthus R.M.Schust., J. Hattori Bot. Lab. 23: 71, 1960 [1961] (Schuster
1960b).
* Herzogianthus sanguineus R.M.Schust., Phytologia 56 (7): 457, 1985 (Schuster
1985c). 392
*** Herzogianthus vaginatus (Herzog) R.M.Schust., J. Hattori Bot. Lab. 23: 71, 1960
[1961] (Schuster 1960b). Bas.: Blepharostoma vaginatum Herzog, Trans. & Proc.
Roy. Soc. New Zealand 65 (3): 355, 1936 (Herzog 1936b).
**
Neotrichocoleaceae Inoue
** Neotrichocolea S.Hatt., J. Hattori Bot. Lab. 2: 9, 1947 [1948] (Hattori 1947b).
** Neotrichocolea bissetii (Mitt.) S.Hatt., J. Hattori Bot. Lab. 2: 10, 1947 [1948]
(Hattori 1947b). Bas.: Mastigophora bissetii Mitt., Trans. Linn. Soc. London, Bot.
3 (3): 200, 1891 (Mitten 1891).
** Trichocoleopsis S.Okamura, Bot. Mag. (Tokyo) 25 (293): 159, 1911 (Okamura 1911).
** Trichocoleopsis sacculata (Mitt.) S.Okamura, Bot. Mag. (Tokyo) 25 (293): 159,
1911 (Okamura 1911). Bas.: Blepharozia sacculata Mitt., Trans. Linn. Soc. London, Bot. 3 (3): 200, 1891 (Mitten 1891).
***
Ptilidiaceae H.Klinggr.
by L. Söderström
Ptilidiaceae was recently studied molecularly by Kreier et al. (2010) showing the occurrence of a possibly undescribed cryptic species from the Himalayas.
*** Ptilidium Nees, Naturgesch. Eur. Leberm. 1: 95, 1833 (Nees 1833c).
*** Ptilidium californicum (Austin) Pearson, List. Canad. Hepat.: 7, 1890 (Pearson
1890). Bas.: Lepidozia californica Austin, Bull. Torrey Bot. Club 6 (3): 19, 1875
(Austin 1875b).
392 Herzogianthus sanguineus is possibly conspeciic with Herzogianthus vaginatus.
World checklist of hornworts and liverworts
435
*** Ptilidium ciliare (L.) Hampe, Prod. l. hercyn.: 76, 1836 (Hampe 1836). Bas.:
Jungermannia ciliaris L., Sp. Pl. 1: 1134, 1753 (Linnaeus 1753).
*** Ptilidium pulcherrimum (Weber) Vain., Meddel. Soc. Fauna Fl. Fenn. 3: 88, 1878
(Vainio 1878). Bas.: Jungermannia pulcherrima Weber, Spic. Fl. Goett.: 150, 1778
(Weber 1778).
Metzgeriidae Barthol.-Began
Metzgeriales Chalaud
***
Aneuraceae H.Klinggr.
by M. Nebel
A molecular phylogeny of Aneuraceae was recently published by Preussing et al. (2010).
heir study revealed Verdoorniaceae to be nested within Aneuraceae and Cryptothallus within
Aneura (see also Wickett and Goinet 2008). Nomenclatural and taxonomic notes can also
be found in Söderström et al. (2010a), Söderström et al. (2012a) and Nebel et al. (2013).
*** Aneura Dumort., Commentat. Bot. (Dumortier): 115, 1822 (Dumortier 1822). 393
* Aneura amboinensis Steph., Bull. Herb. Boissier 7 (9): 678 (219), 1899 (Stephani
1899e).
* Aneura augustae Steph., Sp. Hepat. (Stephani) 6: 430, 1923 (Stephani 1923).
* Aneura bilora Colenso, Trans. & Proc. New Zealand Inst. 17: 262, 1885 (Colenso 1885).
*** Aneura blasioides (Horik.) Furuki, J. Hattori Bot. Lab. 70: 311, 1991 (Furuki
1991). Bas.: Riccardia blasioides Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2,
Bot. 1: 197, 1933 (Horikawa 1933).
** Aneura brasiliensis (Ångstr.) Steph., Hedwigia 32 (3): 137, 1893 (Stephani 1893b).
Bas.: Pseudoneura brasiliensis Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 33 (7):
91, 1876 [1877] (Ångström 1876).
* Aneura brevissima Steph., Sp. Hepat. (Stephani) 6: 21, 1917 (Stephani 1917a).
** Aneura cerebrata Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 94 (2): 185,
1970 (Hewson 1970b).
*** Aneura crateriformis Furuki et D.G.Long, J. Bryol. 18 (2): 281, 1994 (Furuki and
Long 1994).
393 Aneura includes also Acrostolia and Pseudoneura, but a few taxa have never been transferred nor synonymized. hey are listed in the section “Names in genera not currently accepted” below. he types of
Aneura augustae, Aneura hunsteinii, Aneura latemultiida, Aneura ledermannii, Aneura subledermannii and
Aneura subtenerrima were destroyed in B and their identity is doubtful (Grolle and Piippo 1984).
436
*
**
*
*
**
**
**
**
*
**
**
*
***
*
**
**
**
*
**
*
**
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***
***
**
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Aneura crinita C.Massal., Bull. Soc. Bot. Ital. 1917 (8/9): 81, 1917 (Massalongo
1917). 394
Aneura crumii L.Söderstr., A.Hagborg et von Konrat, Phytotaxa 65: 43, 2012 (Söderström et al. 2012a). Nom. nov. pro Cryptothallus hirsutus H.A.Crum, Bryologist
99 (4): 438, 1996 (Crum and Bruce 1996).
Aneura densa Steph., Sp. Hepat. (Stephani) 6: 24, 1917 (Stephani 1917a).
Aneura denticulata Mitt. ex hurn, Timehri 5: 222, 1886 (hurn 1886). 395
Aneura eachamensis Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 94 (2):
184, 1970 (Hewson 1970b).
Aneura erronea Steph., Sp. Hepat. (Stephani) 6: 20, 1917 (Stephani 1917a).
Aneura eskuchei Hässel, Veröf. Geobot. Inst. ETH Stiftung Rübel Zürich 91: 294,
1986 (Hässel 1986b).
Aneura gemmifera Furuki, J. Hattori Bot. Lab. 70: 309, 1991 (Furuki 1991).
Aneura giangena Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 94 (2): 190,
1970 (Hewson 1970b).
Aneura gibbsiana Steph., Sp. Hepat. (Stephani) 6: 28, 1917 (Stephani 1917a).
Aneura glaucescens Steph., Sp. Hepat. (Stephani) 6: 28, 1917 (Stephani 1917a).
Aneura goebeliana Steph., Sp. Hepat. (Stephani) 6: 28, 1917 (Stephani 1917a).
Aneura hirsuta Furuki, J. Hattori Bot. Lab. 70: 317, 1991 (Furuki 1991).
Aneura hunsteinii Steph., Sp. Hepat. (Stephani) 6: 431, 1923 (Stephani 1923).
Aneura imbricata Colenso, Trans. & Proc. New Zealand Inst. 16: 359, 1884 (Colenso 1884).
Aneura kaguaensis Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 94 (2):
190, 1970 (Hewson 1970b).
Aneura keniae Gola, Mem. Reale Accad. Sci. Torino (ser. 2) 65 (1): 3, 1916 (Gola
1916).
Aneura latemultiida Steph., Sp. Hepat. (Stephani) 6: 431, 1923 (Stephani 1923).
Aneura latissima Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 544, 1885
(Spruce 1885).
Aneura ledermannii Steph., Sp. Hepat. (Stephani) 6: 431, 1923 (Stephani 1923).
Aneura macrostachya Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 550, 1885
(Spruce 1885).
Aneura marianensis Furuki, Bryologist 97 (1): 87, 1994 (Furuki 1994b).
Aneura maxima (Schifn.) Steph., Bull. Herb. Boissier 7 (10): 760 (270), 1899
(Stephani 1899f). Bas.: Riccardia maxima Schifn., Denkschr. Kaiserl. Akad.
Wiss., Math.-Naturwiss. Kl. 67: 177, 1898 (Schifner 1898a).
Aneura mirabilis (Malmb.) Wickett et Goinet, Bot. J. Linn. Soc. 156 (1): 11,
2008 (Wickett and Goinet 2008). Bas.: Cryptothallus mirabilis Malmb., Ann.
Bryol. 6: 122, 1933 (von Malmborg 1933).
Aneura novaecaledoniae R.M.Schust., Phytologia 56 (7): 451, 1985 (Schuster 1985c).
394 Aneura crinita (type from Europe) has neither been recognized in any recent European treatment nor
synonymized.
395 Aneura denticulata is possibly conspeciic with Riccardia ciliolata (Hässel 2006b).
World checklist of hornworts and liverworts
437
*** Aneura novaguineensis Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 94 (2):
189, 1970 (Hewson 1970b).
* Aneura nymannii Steph., Sp. Hepat. (Stephani) 6: 35, 1917 (Stephani 1917a).
** Aneura pellucida Colenso, Trans. & Proc. New Zealand Inst. 18: 252, 1886 (Colenso 1886b).
*** Aneura pinguis (L.) Dumort., Syll. Jungerm. Europ.: 86, 1831 (Dumortier 1831).
Bas.: Jungermannia pinguis L., Sp. Pl. 1: 1136, 1753 (Linnaeus 1753).
** Aneura polyantha Colenso, Trans. & Proc. New Zealand Inst. 17: 262, 1885 (Colenso 1885).
** Aneura punctata Colenso, Trans. & Proc. New Zealand Inst. 18: 254, 1886 (Colenso 1886b).
** Aneura rodwayi Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 94 (2): 188,
1970 (Hewson 1970b).
* Aneura roraimensis Steph., Trans. Linn. Soc. London, Bot. 6 (1): 94, 1901 (Stephani
1901e).
** Aneura rotangicola Steph., Sp. Hepat. (Stephani) 6: 432, 1923 (Stephani 1923).
* Aneura serrulata Gottsche ex Steph., Sp. Hepat. (Stephani) 6: 42, 1917 (Stephani 1917a).
** Aneura sharpii Inoue et N.G.Mill., Bull. Natl. Sci. Mus. Tokyo, B 11 (3): 96,
1985 (Inoue and Miller 1985). 396
* Aneura singalangana (Schifn.) Steph., Bull. Herb. Boissier 7 (10): 751 (261),
1899 (Stephani 1899f). Bas.: Riccardia singalangana Schifn., Denkschr. Kaiserl.
Akad. Wiss., Math.-Naturwiss. Kl. 67: 174, 1898 (Schifner 1898a). 397
** Aneura subcanaliculata R.M.Schust., J. Hattori Bot. Lab. 67: 60, 1989 (Schuster
1989).
* Aneura subledermannii Steph., Sp. Hepat. (Stephani) 6: 432, 1923 (Stephani 1923).
* Aneura subtenerrima Steph., Sp. Hepat. (Stephani) 6: 432, 1923 (Stephani 1923).
* Aneura vincentina Steph., Sp. Hepat. (Stephani) 6: 45, 1917 (Stephani 1917a).
** Lobatiriccardia (Mizut. et S.Hatt.) Furuki, J. Hattori Bot. Lab. 70: 319, 1991
(Furuki 1991). Bas.: Riccardia subg. Lobatiriccardia Mizut. et S.Hatt., J. Hattori
Bot. Lab. 18: 38, 1957 (Mizutani and Hattori 1957).
*** Lobatiriccardia alterniloba (Hook.f. et Taylor) Furuki, J. Hattori Bot. Lab. 70:
319, 1991 (Furuki 1991). Bas.: Jungermannia alterniloba Hook.f. et Taylor, London J. Bot. 3: 572, 1844 (Hooker and Taylor 1844d).
** Lobatiriccardia alterniloba var. gigantea (Steph.) Nebel, Phytotaxa 81 (1): 10, 2013
(Nebel et al. 2013). Bas.: Aneura gigantea Steph., J. & Proc. Roy. Soc. New South
Wales 48 (1/2): 95, 1914 (Stephani and Watts 1914).
** Lobatiriccardia alterniloba var. robusta (Rodway) Nebel, Phytotaxa 81 (1): 10,
2013 (Nebel et al. 2013). Bas.: Aneura alterniloba f. robusta Rodway, Tasm. Bryoph.: 12, 1917 (Rodway 1917b).
396 Aneura sharpii is conspeciic with Aneura maxima in Schuster (1992b), but the taxa are not the same
(D.G. Long, pers. comm.).
397 Aneura singalangana may be a form of Riccardia subexalata (Stephani 1899f ).
438
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Lobatiriccardia athertonensis (Hewson) Furuki, J. Hattori Bot. Lab. 70: 319, 1991
(Furuki 1991). Bas.: Aneura athertonensis Hewson, Proc. Linn. Soc. New South
Wales (ser. 2) 94 (2): 188, 1970 (Hewson 1970b).
*** Lobatiriccardia coronopus (De Not.) Furuki, J. Hattori Bot. Lab. 100: 90, 2006
(Furuki 2006a). Bas.: Aneura coronopus De Not., Hedwigia 32 (1): 19, 1893
(Stephani 1893a).
** Lobatiriccardia coronopus subsp. australis (R.M.Schust.) Nebel, Preussing, Schäf.Verw. et D.Quandt, Taxon 59 (5): 1434, 2010 (Preussing et al. 2010). Bas.: Aneura
lobata subsp. australis R.M.Schust., Phytologia 56 (7): 451, 1985 (Schuster 1985c).
** Lobatiriccardia oberwinkleri Nebel, Preussing, Schäf.-Verw. et D.Quandt, Taxon
59 (5): 1435, 2010 (Preussing et al. 2010).
** Lobatiriccardia subaquatica (R.M.Schust.) Nebel, Phytotaxa 81 (1): 10, 2013 (Nebel et al. 2013). Bas.: Aneura subaquatica R.M.Schust., Phytologia 56 (7): 450,
1985 (Schuster 1985c).
** Lobatiriccardia verdoornioides Nebel, Preussing, Schäf.-Verw. et D.Quandt, Taxon
59 (5): 1437, 2010 (Preussing et al. 2010).
*** Lobatiriccardia yakusimensis (S.Hatt.) Furuki, J. Hattori Bot. Lab. 70: 321, 1991
(Furuki 1991). Bas.: Riccardia lobata var. yakusimensis S.Hatt., J. Hattori Bot. Lab.
6: 10, 1951 [1952] (Hattori 1951a).
*** Lobatiriccardia yunnanensis Furuki et D.G.Long, J. Bryol. 29 (3): 161, 2007 (Furuki and Long 2007).
*** Riccardia Gray, Nat. Arr. Brit. Pl. 1: 679, 1821 (Gray 1821) nom. conserv. 398
** Riccardia pectinata var. fasciculata (Steph.) Hürl., Bauhinia 5 (4): 208, 1976 (Hürlimann 1976). Bas.: Aneura fasciculata Steph., Sp. Hepat. (Stephani) 6: 25, 1917
(Stephani 1917a).
** subg. Arceoneura Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 204, 1972 (Hässel 1972a).
** Riccardia marionensis R.M.Schust., J. Hattori Bot. Lab. 67: 65, 1989 (Schuster 1989).
*** Riccardia prehensilis (Hook.f. et Taylor) C.Massal., Nuovo Giorn. Bot. Ital. 17 (3):
255, 1885 (Massalongo 1885). Bas.: Jungermannia prehensilis Hook.f. et Taylor,
London J. Bot. 3: 480, 1844 (Hooker and Taylor 1844b).
** subg. Corioneura Furuki, J. Hattori Bot. Lab. 70: 394, 1991 (Furuki 1991).
** Riccardia argentolimbata Hewson et Grolle, J. Hattori Bot. Lab. 29: 70, 1966
(Grolle 1966i).
*** Riccardia hattorii Furuki, J. Hattori Bot. Lab. 75: 257, 1994 (Furuki 1994a).
** subg. Hyaloneura R.M.Schust., Phytologia 56 (7): 452, 1985 (Schuster 1985c).
398 Riccardia includes Acrostolia, but a few taxa have neither been transferred nor synonymized. hey are
listed under “Names in genera not currently accepted” below.
World checklist of hornworts and liverworts
439
*
Riccardia albomarginata (Steph.) Schifn., Consp. Hepat. Arch. Ind.: 53, 1898
(Schifner 1898b). Bas.: Aneura albomarginata Steph., Hedwigia 32 (1): 18, 1893
(Stephani 1893a). 399
** Riccardia canaliculata (Nees) Kuntze, Revis. Gen. Pl. 2: 838, 1891 (Kuntze 1891).
Bas.: Jungermannia canaliculata Nees, Enum. Pl. Crypt. Javae: 10, 1830 (Nees 1830).
** Riccardia pindensis Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95 (1): 87,
1970 (Hewson 1970a).
** subg. Lophoneura Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 218, 1972 (Hässel 1972a).
** Riccardia fuegiensis C.Massal., Nuovo Giorn. Bot. Ital. 17 (3): 255, 1885 (Massalongo 1885).
** subg. Neoneura Furuki, J. Hattori Bot. Lab. 70: 385, 1991 (Furuki 1991).
*** Riccardia spongiosa Furuki, J. Hattori Bot. Lab. 70: 385, 1991 (Furuki 1991).
** subg. Phycaneura R.M.Schust., J. Hattori Bot. Lab. 26: 294, 1963 (Schuster 1963b).
** Riccardia aequicellularis (Steph.) Hewson, Proc. Linn. Soc. New South Wales
(ser. 2) 95 (1): 79, 1970 (Hewson 1970a). Bas.: Aneura aequicellularis Steph., J.
& Proc. Roy. Soc. New South Wales 48 (1/2): 95, 1914 (Stephani and Watts
1914).
** Riccardia asperulata R.M.Schust., J. Hattori Bot. Lab. 27: 209, 1964 (Schuster 1964a).
**
***
***
***
***
***
***
***
***
**
subg. Riccardia
Riccardia aeruginosa Furuki, J. Hattori Bot. Lab. 70: 345, 1991 (Furuki 1991).
Riccardia arcuata Furuki, J. Hattori Bot. Lab. 70: 361, 1991 (Furuki 1991).
Riccardia chamedryfolia (With.) Grolle, Trans. Brit. Bryol. Soc. 5 (4): 772, 1969
(Grolle 1969b). Bas.: Jungermannia chamedryfolia With., Bot. arr. veg. Gr. Brit. 2:
699, 1776 (Withering 1776).
Riccardia cochleata (Hook.f. et Taylor) Kuntze, Revis. Gen. Pl. 2: 838, 1891
(Kuntze 1891). Bas.: Riccia cochleata Hook.f. et Taylor, London J. Bot. 4: 96,
1845 (Hooker and Taylor 1845).
Riccardia eriocaula (Hook.) C.Massal., Nuovo Giorn. Bot. Ital. 17 (3): 256, 1885
(Massalongo 1885). Bas.: Jungermannia eriocaula Hook., Musci Exot. 1: tab. 72,
1818 (Hooker 1818).
Riccardia lavovirens Furuki, J. Hattori Bot. Lab. 70: 333, 1991 (Furuki 1991).
Riccardia fruticosa (Steph.) Furuki, Nat. Hist. Res. 5 (1): 1, 1998 (Furuki 1998).
Bas.: Aneura fruticosa Steph., Sp. Hepat. (Stephani) 6: 27, 1917 (Stephani 1917a).
Riccardia glauca Furuki, J. Hattori Bot. Lab. 70: 352, 1991 (Furuki 1991).
Riccardia kodamae Mizut. et S.Hatt., J. Hattori Bot. Lab. 18: 57, 1957 (Mizutani
and Hattori 1957).
399 Riccardia albomarginata is possibly conspeciic with Riccardia canaliculata (Söderström et al. 2010a).
440
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Riccardia latifrons (Lindb.) Lindb., Acta Soc. Sci. Fenn. 10: 513, 1875 (Lindberg
1875). Bas.: Aneura latifrons Lindb., Bot. Not. 26: 62, 1873 (Anonymous 1873).
** Riccardia latifrons subsp. arctica R.M.Schust. et Damsh., J. Hattori Bot. Lab. 62:
303, 1987 (Schuster 1987d).
** Riccardia latifrons var. miyakeana (Schifn.) Furuki, J. Hattori Bot. Lab. 70: 375,
1991 (Furuki 1991). Bas.: Riccardia miyakeana Schifn., Österr. Bot. Z. 49 (11):
388, 1899 (Schifner 1899c).
** Riccardia nagasakiensis (Steph.) S.Hatt., Bull. Tokyo Sci. Mus. 11: 164, 1944
(Hattori 1944d). Bas.: Aneura nagasakiensis Steph., Sp. Hepat. (Stephani) 6: 34,
1917 (Stephani 1917a).
*** Riccardia palmata (Hedw.) Carruth., J. Bot. 3: 302, 1865 (Carruthers 1865). Bas.: Jungermannia palmata Hedw., heoria generat.: 87, 1784 (Hedwig 1784), nom. conserv.
** Riccardia planilora (Steph.) S.Hatt., Bull. Tokyo Sci. Mus. 11: 164, 1944 (Hattori 1944d). Bas.: Aneura planilora Steph., Sp. Hepat. (Stephani) 6: 38, 1917
(Stephani 1917a).
** Riccardia planilora var. aequatorialis Furuki, Nat. Hist. Res. 4 (2): 77, 1997 (Furuki
1997).
** Riccardia pseudodendroceros R.M.Schust., Phytologia 56 (7): 452, 1985 (Schuster
1985c).
*** Riccardia pumila Furuki, J. Hattori Bot. Lab. 70: 361, 1991 (Furuki 1991).
** Riccardia pusilla Grolle, J. Jap. Bot. 41 (8): 231, 1966 (Grolle 1966d). Nom. nov.
pro Riccardia nana Mizut. et S.Hatt., J. Hattori Bot. Lab. 18: 53, 1957 (Mizutani
and Hattori 1957), nom. illeg.
*** Riccardia subalpina Furuki, J. Hattori Bot. Lab. 70: 367, 1991 (Furuki 1991).
** Riccardia tamariscina (Steph.) Schifn., Consp. Hepat. Arch. Ind.: 58, 1898
(Schifner 1898b). Bas.: Aneura tamariscina Steph., Hedwigia 32 (1): 27, 1893
(Stephani 1893a).
*** Riccardia vitrea Furuki, J. Hattori Bot. Lab. 70: 327, 1991 (Furuki 1991).
** sect. Alcicornia Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 17, 1972 (Hässel 1972a).
** Riccardia alcicornis (Hook.f. et Taylor) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser.
3), C. Sci. Mat. 4 (13): 431, 1877 (Trevisan 1877). Bas.: Jungermannia alcicornis
Hook.f. et Taylor, London J. Bot. 3: 479, 1844 (Hooker and Taylor 1844b).
** Riccardia conimitra (Steph.) A.Evans, Trans. Connecticut Acad. Arts 25 (2): 156,
1921 (Evans 1921a). Bas.: Aneura conimitra Steph., Bull. Herb. Boissier 7 (10):
749 (259), 1899 (Stephani 1899f).
** Riccardia corralensis (Steph.) A.Evans, Trans. Connecticut Acad. Arts 25 (2): 143,
1921 (Evans 1921a). Bas.: Aneura corralensis Steph., Bull. Herb. Boissier 7 (10):
742 (252), 1899 (Stephani 1899f).
*** Riccardia furtiva E.A.Br. et Braggins, J. Hattori Bot. Lab. 66: 35, 1989 (Brown
and Braggins 1989).
World checklist of hornworts and liverworts
441
** Riccardia fuscobrunnea (Steph.) A.Evans, Trans. Connecticut Acad. Arts 25 (2):
152, 1921 (Evans 1921a). Bas.: Aneura fuscobrunnea Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9): 7, 1911 (Stephani 1911b).
** Riccardia longioleata Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 30, 1972 (Hässel 1972a).
*** Riccardia multicorpora E.A.Br., J. Hattori Bot. Lab. 66: 40, 1989 (Brown and
Braggins 1989).
* Riccardia umida E.A.Br., J. Hattori Bot. Lab. 66: 38, 1989 (Brown and Braggins 1989).
** sect. Crassantia Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 41, 1972 (Hässel 1972a).
*** Riccardia aequitexta (Steph.) E.A.Br., J. Hattori Bot. Lab. 66: 78, 1989 (Brown
and Braggins 1989). Bas.: Aneura aequitexta Steph., J. Linn. Soc., Bot. 29 (201):
263, 1892 (Stephani 1892b).
*** Riccardia alba (Colenso) E.A.Br., J. Hattori Bot. Lab. 66: 66, 1989 (Brown and
Braggins 1989). Bas.: Aneura alba Colenso, Trans. & Proc. New Zealand Inst. 16:
357, 1884 (Colenso 1884).
** Riccardia amnicola Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 92, 1972 (Hässel 1972a).
** Riccardia bipinnatiida (Colenso) Hewson, Proc. Linn. Soc. New South Wales
(ser. 2) 95 (1): 93, 1970 (Hewson 1970a). Bas.: Aneura bipinnatiida Colenso,
Trans. & Proc. New Zealand Inst. 16: 358, 1884 (Colenso 1884).
** Riccardia calva (Schifn.) A.Evans, Trans. Connecticut Acad. Arts 25 (2): 134,
1921 (Evans 1921a). Bas.: Aneura calva Schifn., Leberm., Forschungsr. Gazelle 4
(4): 42, 1890 (Schifner 1890).
*** Riccardia colensoi (Steph.) W.Martin, Trans. & Proc. Roy. Soc. New Zealand 78
(4): 499, 1950 (Martin 1950). Bas.: Aneura colensoi Steph., J. Linn. Soc., Bot. 29
(201): 264, 1892 (Stephani 1892b).
*** Riccardia crassa (Schwägr.) C.Massal., Nuovo Giorn. Bot. Ital. 17 (3): 254, 1885
(Massalongo 1885). Bas.: Jungermannia crassa Schwägr., Hist. Musc. Hepat.
Prodr.: 31, 1814 (Schwägrichen 1814).
** Riccardia crassicrispa (Steph.) A.Evans, Trans. Connecticut Acad. Arts 25 (2):
194, 1921 (Evans 1921a). Bas.: Aneura crassicrispa Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9): 6, 1911 (Stephani 1911b).
** Riccardia diderma Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 56, 1972 (Hässel 1972a).
** Riccardia diversilora A.Evans, Trans. Connecticut Acad. Arts 25 (2): 167, 1921
(Evans 1921a).
** Riccardia diversilora subsp. paucigyna R.M.Schust., J. Hattori Bot. Lab. 67: 97,
1989 (Schuster 1989).
** Riccardia duriuscula Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 82, 1972 (Hässel 1972a).
442
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Riccardia falsiloribunda Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 119, 1972 (Hässel 1972a).
** Riccardia loribunda (Steph.) A.Evans, Trans. Connecticut Acad. Arts 25 (2): 182,
1921 (Evans 1921a). Bas.: Aneura loribunda Steph., Bull. Herb. Boissier 7 (10):
749 (259), 1899 (Stephani 1899f).
** Riccardia luvigena Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 73, 1972 (Hässel 1972a).
** Riccardia georgiensis (Steph.) Hässel, Lindbergia 1 (1/2): 80, 1971 [1972] (Grolle
1971a). Bas.: Aneura georgiensis Steph., Wiss. Ergebn. Schwed. Südpolar-Exped.
[1901–1903] 4 (1): 2, 1905 (Stephani 1905e).
** Riccardia georgiensis subsp. sympodea R.M.Schust., J. Hattori Bot. Lab. 67: 83,
1989 (Schuster 1989).
*** Riccardia graefei (Steph.) Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95
(1): 118, 1970 (Hewson 1970a). Bas.: Aneura graefei Steph., Hedwigia 32 (1): 21,
1893 (Stephani 1893a).
*** Riccardia marginata (Colenso) Pearson, Univ. Calif. Publ. Bot. 10 (4): 309, 1923
(Pearson 1923). Bas.: Aneura marginata Colenso, Trans. & Proc. New Zealand
Inst. 18: 253, 1886 (Colenso 1886b).
*** Riccardia marginata var. paciica Furuki, J. Hattori Bot. Lab. 70: 380, 1991 (Furuki 1991).
** Riccardia mycophora A.Evans, Trans. Connecticut Acad. Arts 25 (2): 175, 1921
(Evans 1921a).
** Riccardia negeri (Steph.) A.Evans, Trans. Connecticut Acad. Arts 25 (2): 172,
1921 (Evans 1921a). Bas.: Aneura negeri Steph., Bull. Herb. Boissier 7 (10): 747
(257), 1899 (Stephani 1899f).
** Riccardia nitida (Colenso) E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot. 3
(4): 92, 1965 (Hodgson 1965). Bas.: Aneura nitida Colenso, Trans. & Proc. New
Zealand Inst. 18: 253, 1886 (Colenso 1886b).
** Riccardia papillosa (C.Massal. et Steph.) Hässel, Revista Mus. Argent. Ci. Nat.,
Bernardino Rivadavia Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 60, 1972 (Hässel
1972a). Bas.: Aneura papillosa C.Massal. et Steph., Atti Reale Ist. Veneto Sci. Lett.
Arti 87 (2): 241, 1928 (Massalongo 1928).
** Riccardia regularis (Steph.) Kühnem., Revista Centro Estud. Doct. Ci. Nat. 1:
171, 1937 (Kühnemann 1937). Bas.: Aneura regularis Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9): 9, 1911 (Stephani 1911b).
** Riccardia rivularis Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 52, 1972 (Hässel 1972a).
** Riccardia saxicola Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 137, 1972 (Hässel 1972a).
** Riccardia spectabilis (Steph.) A.Evans, Trans. Connecticut Acad. Arts 25 (2): 140,
1921 (Evans 1921a). Bas.: Aneura spectabilis Steph., Bull. Herb. Boissier 7 (10):
746 (256), 1899 (Stephani 1899f).
World checklist of hornworts and liverworts
443
** Riccardia spegazziniana C.Massal., Nuovo Giorn. Bot. Ital. 17 (3): 254, 1885
(Massalongo 1885).
** Riccardia tenax (Steph.) A.Evans, Trans. Connecticut Acad. Arts 25 (2): 186, 1921
(Evans 1921a). Bas.: Aneura tenax Steph., Bull. Herb. Boissier 7 (10): 755 (265),
1899 (Stephani 1899f).
** Riccardia tenerrima (Steph.) A.Evans, Trans. Connecticut Acad. Arts 25 (2):
164, 1921 (Evans 1921a). Bas.: Aneura tenerrima Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46 (9): 9, 1911 (Stephani 1911b).
** Riccardia theliophora Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 96, 1972 (Hässel 1972a).
** sect. Pallidevirida Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 142, 1972 (Hässel 1972a).
** Riccardia granulata (Steph.) A.Evans, Trans. Connecticut Acad. Arts 25 (2): 192,
1921 (Evans 1921a). Bas.: Aneura granulata Steph., Hedwigia 32 (1): 21, 1893
(Stephani 1893a).
** Riccardia intercellula E.A.Br., J. Hattori Bot. Lab. 66: 58, 1989 (Brown and Braggins 1989).
** Riccardia opuntiiformis S.W.Arnell, Arch. Soc. Zool. Bot. Fenn. “Vanamo” 9: 52,
1954 (Arnell 1954b).
** Riccardia pallidevirens (Steph.) A.Evans, Trans. Connecticut Acad. Arts 25 (2):
189, 1921 (Evans 1921a). Bas.: Aneura pallidevirens Steph., Bull. Herb. Boissier 7
(10): 762 (272), 1899 (Stephani 1899f).
*** Riccardia pennata E.A.Br., J. Hattori Bot. Lab. 66: 53, 1989 (Brown and Braggins
1989).
*** Riccardia perspicua E.A.Br., J. Hattori Bot. Lab. 66: 55, 1989 (Brown and Braggins 1989).
** Riccardia xylophila Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 156, 1972 (Hässel 1972a).
** sect. Riccardia
*** Riccardia australis (Lehm.) Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95
(1): 110, 1970 (Hewson 1970a). Bas.: Sarcomitrium australe Lehm., Nov. Stirp.
Pug. 10: 19, 1857 (Lehmann 1857).
** Riccardia autoica (Steph.) A.Evans, Trans. Connecticut Acad. Arts 25 (2): 159,
1921 (Evans 1921a). Bas.: Aneura autoica Steph., Bull. Herb. Boissier 7 (9): 691
(232), 1899 (Stephani 1899e).
** Riccardia breviala E.A.Br., J. Hattori Bot. Lab. 66: 46, 1989 (Brown and Braggins
1989).
** Riccardia breviramosa (Steph.) A.Evans, Nat. Hist. Juan Fernandez (Botany) 2
(20): 559, 1930 (Evans 1930a). Bas.: Aneura breviramosa Steph., Kungl. Svenska
Vetensk.-Akad. Handl. (n.ser.) 46 (9): 6, 1911 (Stephani 1911b).
444
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Riccardia ilicina (Colenso) E.A.Hodgs., Rec. Domin. Mus. 4 (11): 129, 1962
(Hodgson 1962a). Bas.: Aneura ilicina Colenso, Trans. & Proc. New Zealand
Inst. 16: 358, 1884 (Colenso 1884).
** Riccardia leptostachya A.Evans, Nat. Hist. Juan Fernandez (Botany) 2 (20): 570,
1930 (Evans 1930a).
** Riccardia lobulata (Colenso) E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot.
3 (4): 92, 1965 (Hodgson 1965). Bas.: Zoopsis lobulata Colenso, Trans. & Proc.
New Zealand Inst. 18: 250, 1886 (Colenso 1886b).
** Riccardia mejlandii S.W.Arnell, Results Norweg. Sci. Exped. Tristan da Cunha 42:
30, 1958 (Arnell 1958b).
*** Riccardia multiida (L.) Gray, Nat. Arr. Brit. Pl. 1: 684, 1821 (Gray 1821). Bas.:
Jungermannia multiida L., Sp. Pl. 1: 1136, 1753 (Linnaeus 1753).
*** Riccardia multiida subsp. decrescens (Steph.) Furuki, J. Hattori Bot. Lab. 70: 341,
1991 (Furuki 1991). Bas.: Aneura decrescens Steph., Bull. Herb. Boissier 7 (9): 686
(227), 1899 (Stephani 1899e).
** Riccardia multiida subsp. synoica R.M.Schust., J. Hattori Bot. Lab. 62: 319, 1987
(Schuster 1987d).
** Riccardia multioleata Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 195, 1972 (Hässel 1972a).
*** Riccardia papulosa (Steph.) E.A.Br., J. Hattori Bot. Lab. 66: 49, 1989 (Brown
and Braggins 1989). Bas.: Aneura papulosa Steph., Hedwigia 32 (1): 25, 1893
(Stephani 1893a).
** Riccardia patens Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia Inst.
Nac. Invest. Ci. Nat. Bot. 4 (1): 165, 1972 (Hässel 1972a).
** Riccardia polyclada (Mitt.) Hässel, Bryologist 109 (1): 34, 2006 (Hässel 2006b).
Bas.: Aneura polyclada Mitt., Trans. Linn. Soc. London, Bot. 2 (13): 297, 1887
(Mitten 1887).
** Riccardia statensis Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 177, 1972 (Hässel 1972a).
** Riccardia thaxteri A.Evans, Trans. Connecticut Acad. Arts 25 (2): 126, 1921 (Evans 1921a).
** Riccardia tristaniana S.W.Arnell, Results Norweg. Sci. Exped. Tristan da Cunha
42: 31, 1958 (Arnell 1958b).
** subg. Spinella (Schifn.) Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 214, 1972 (Hässel 1972a). Bas.:
Spinella Schifn., Leberm., Forschungsr. Gazelle 4 (4): 41, 1890 (Schifner 1890).
** Riccardia spinulifera C.Massal., Nuovo Giorn. Bot. Ital. 17 (3): 254, 1885 (Massalongo 1885).
** subg. hornoneura Furuki, J. Hattori Bot. Lab. 70: 382, 1991 (Furuki 1991).
World checklist of hornworts and liverworts
445
* Riccardia baumannii Hürl., Bauhinia 5 (4): 203, 1976 (Hürlimann 1976). 400
** Riccardia deguchii Furuki et K.T.Yong, Hikobia 16 (3): 285, 2013 (Furuki et al. 2013).
*** Riccardia grossitexta (Steph.) Furuki, J. Hattori Bot. Lab. 70: 382, 1991 (Furuki
1991). Bas.: Aneura grossitexta Steph., Sp. Hepat. (Stephani) 6: 29, 1917 (Stephani
1917a).
** Riccardia inconspicua (Steph.) Reeb et Bardat, Cryptog. Bryol. 35 (1): 61, 2014
(Reeb and Bardat 2014). Bas.: Aneura inconspicua Steph., Hedwigia 32 (1): 23,
1893 (Stephani 1893a).
** subg. Trichothallia Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 224, 1972 (Hässel 1972a).
** Riccardia hyalitricha Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia
Inst. Nac. Invest. Ci. Nat. Bot. 4 (1): 225, 1972 (Hässel 1972a).
Incertae sedis
** Riccardia aberrans (Steph.) Gradst., J. Hattori Bot. Lab. 45: 129, 1979 (Gradstein
and Hekking 1979). Bas.: Aneura aberrans Steph., Hedwigia 32 (1): 18, 1893
(Stephani 1893a).
** Riccardia agumana Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95 (1): 82,
1970 (Hewson 1970a).
** Riccardia algoides (Taylor) Meenks, J. Hattori Bot. Lab. 62: 168, 1987 (Meenks
1987). Bas.: Metzgeria algoides Taylor, London J. Bot. 5: 410, 1846 (Taylor
1846b).
*** Riccardia amazonica (Spruce) Schifn. ex Gradst. et Hekking, J. Hattori Bot. Lab.
45: 129, 1979 (Gradstein and Hekking 1979). Bas.: Aneura amazonica Spruce,
Trans. & Proc. Bot. Soc. Edinburgh 15: 545, 1885 (Spruce 1885).
*** Riccardia andina (Spruce) Herzog, Svensk Bot. Tidskr. 46 (1): 65, 1952 (Herzog
1952e). Bas.: Aneura andina Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 548,
1885 (Spruce 1885).
** Riccardia angustata Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 126,
1934 (Horikawa 1934).
** Riccardia angustealata (Steph.) Hewson, Proc. Linn. Soc. New South Wales (ser. 2)
95 (1): 111, 1970 (Hewson 1970a). Bas.: Aneura angustealata Steph., Sp. Hepat.
(Stephani) 6: 20, 1917 (Stephani 1917a).
** Riccardia angustissima (Steph.) H.A.Mill., Phytologia 47 (4): 323, 1981 (Miller 1981). Bas.: Aneura angustissima Steph., Sp. Hepat. (Stephani) 6: 20, 1917
(Stephani 1917a).
** Riccardia aspera (Steph.) Grolle, J. Hattori Bot. Lab. 30: 117, 1967 (Grolle 1967b).
Bas.: Aneura aspera Steph., Sp. Hepat. (Stephani) 6: 21, 1917 (Stephani 1917a).
400 Riccardia baumannii is possibly conspeciic with Riccardia grossitexta or Riccardia tenuicostata (Furuki
1991, Söderström et al. 2010a).
446
*
**
**
**
**
**
**
**
**
**
**
***
**
**
**
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Riccardia baldwinii (Steph.) H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 527, 1963 (Miller
1963). Bas.: Aneura baldwinii Steph., Bull. Herb. Boissier 7 (10): 743 (253), 1899
(Stephani 1899f). 401
Riccardia barbilora (Steph.) Piippo, J. Hattori Bot. Lab. 68: 134, 1990 (Piippo
1990). Bas.: Aneura barbilora Steph., Mém. Soc. Nat. Sci. Nat. Math. Cherbourg
29: 209, 1894 (Stephani 1894b).
Riccardia bogotensis (Gottsche) Pagán, Bryologist 42 (1): 6, 1939 (Pagán 1939a).
Bas.: Pseudoneura bogotensis Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 184, 1864
(Gottsche 1864).
Riccardia boliviensis (Steph.) Meenks, J. Hattori Bot. Lab. 62: 170, 1987 (Meenks
1987). Bas.: Aneura boliviensis Steph., Biblioth. Bot. 87 (2): 176, 1916 (Stephani
1916a).
Riccardia bongeriana Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95 (1):
101, 1970 (Hewson 1970a).
Riccardia brunnea (Steph.) S.Hatt., Bull. Tokyo Sci. Mus. 11: 164, 1944 (Hattori
1944d). Bas.: Aneura brunnea Steph., Sp. Hepat. (Stephani) 6: 21, 1917 (Stephani
1917a).
Riccardia calcarea (Steph.) Meenks, J. Hattori Bot. Lab. 62: 170, 1987 (Meenks
1987). Bas.: Aneura calcarea Steph., Bull. Herb. Boissier 7 (10): 756 (266), 1899
(Stephani 1899f).
Riccardia capillacea (Steph.) Meenks et C.De Jong, Cryptog. Bryol. Lichénol. 6
(1): 6, 1985 (Meenks and de Jong 1985). Bas.: Aneura capillacea Steph., Biblioth.
Bot. 87 (2): 174, 1916 (Stephani 1916a).
Riccardia capillacea var. dentata Meenks, J. Hattori Bot. Lab. 62: 170, 1987
(Meenks 1987).
Riccardia cardotii (Steph.) Pandé et S.C.Srivast., Biol. Mem. 1 (1/2): 131, 1976
(Srivastava and Udar 1976). Bas.: Aneura cardotii Steph., Bull. Soc. Roy. Bot.
Belgique, Mém. 41 (1): 118, 1904 [1905] (Stephani 1904d).
Riccardia cataractarum (Spruce) Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl.,
Denkschr. 111: 18, 1964 (Schifner and Arnell 1964). Bas.: Aneura cataractarum
Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: cxcv, 1889 [1890] (Spruce 1889).
Riccardia cervicornis (Spruce) Herzog ex Gradst. et Hekking, J. Hattori Bot. Lab.
45: 129, 1979 (Gradstein and Hekking 1979). Bas.: Aneura cervicornis Spruce,
Trans. & Proc. Bot. Soc. Edinburgh 15: 550, 1885 (Spruce 1885).
Riccardia changbaishanensis C.Gao, Fl. Hepat. Chin. Boreali-Orient.: 209, 1981
(Gao and Chang 1981).
Riccardia chinensis C.Gao, Fl. Hepat. Chin. Boreali-Orient.: 209, 1981 (Gao and
Chang 1981).
Riccardia ciliolata (Spruce) Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot.
1: 198, 1933 (Horikawa 1933). Bas.: Aneura ciliolata Spruce, Trans. & Proc. Bot.
Soc. Edinburgh 15: 547, 1885 (Spruce 1885).
401 Riccardia baldwinii is possibly conspeciic with Riccardia laccida (Miller et al. 1983).
World checklist of hornworts and liverworts
447
** Riccardia columbica (Steph.) Hässel ex Gradst. et Hekking, J. Hattori Bot. Lab.
45: 130, 1979 (Gradstein and Hekking 1979). Bas.: Aneura columbica Steph., Sp.
Hepat. (Stephani) 6: 22, 1917 (Stephani 1917a).
** Riccardia comata (Steph.) H.A.Mill., Phytologia 47 (4): 323, 1981 (Miller 1981).
Bas.: Aneura comata Steph., Sp. Hepat. (Stephani) 6: 22, 1917 (Stephani 1917a).
** Riccardia compacta (Steph.) S.W.Arnell, Bot. Not. 105: 141, 1952 (Arnell 1952b).
Bas.: Aneura compacta Steph., Hedwigia 32 (1): 19, 1893 (Stephani 1893a).
** Riccardia comptonii (Pearson) H.A.Mill., Phytologia 47 (4): 323, 1981 (Miller
1981). Bas.: Aneura comptonii Pearson, J. Linn. Soc., Bot. 46 (305): 17, 1922
(Pearson 1922b).
** Riccardia costata (Steph.) Hürl., Bauhinia 5 (4): 206, 1976 (Hürlimann 1976).
Bas.: Aneura costata Steph., Sp. Hepat. (Stephani) 6: 23, 1917 (Stephani 1917a).
*** Riccardia crassicaulis (Steph.) Meenks et C.De Jong, Cryptog. Bryol. Lichénol. 6
(1): 8, 1985 (Meenks and de Jong 1985). Bas.: Aneura crassicaulis Steph., Biblioth.
Bot. 87 (2): 174, 1916 (Stephani 1916a).
** Riccardia crassiretis Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl.
67: 173, 1898 (Schifner 1898a).
** Riccardia crenulata Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl.
67: 173, 1898 (Schifner 1898a).
** Riccardia crenuliformis R.M.Schust., J. Hattori Bot. Lab. 67: 72, 1989 (Schuster 1989).
* Riccardia decolyana Schifn., J. Indian Bot. Soc. 38 (4): 538, 1959 [1960] (Schifner
et al. 1959). 402
** Riccardia densiramea (Steph.) S.Hatt., Bull. Tokyo Sci. Mus. 11: 164, 1944 (Hattori 1944d). Bas.: Aneura densiramea Steph., Sp. Hepat. (Stephani) 6: 24, 1917
(Stephani 1917a).
** Riccardia devexa Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl., Denkschr.
111: 12, 1964 (Schifner and Arnell 1964).
** Riccardia diablotina (Spruce) Pagán, Bryologist 45 (4): 80, 1942 (Pagán 1942b).
Bas.: Aneura diablotina Spruce, J. Linn. Soc., Bot. 30 (210): 366, 1895 (Gepp
1895b).
*** Riccardia digitiloba (Spruce) Pagán, Bryologist 42 (1): 6, 1939 (Pagán 1939a).
Bas.: Aneura digitiloba Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: cci, 1889
[1890] (Spruce 1889).
** Riccardia dilatata (Spruce) Schäf.-Verw. et Pócs, Cryptog. Bryol. 31 (4): 389,
2010 (Schäfer-Verwimp 2010). Bas.: Aneura dilatata Spruce, J. Linn. Soc., Bot.
30 (210): 368, 1895 (Gepp 1895b).
* Riccardia diminuta Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl.
67: 170, 1898 (Schifner 1898a). 403
* Riccardia diminuta var. thermarum Schifn., Denkschr. Kaiserl. Akad. Wiss.,
Math.-Naturwiss. Kl. 67: 172, 1898 (Schifner 1898a).
402 Riccardia decolyana is possibly conspeciic with Riccardia multiida (Srivastava and Udar 1976).
403 Riccardia diminuta is possibly conspeciic with Riccardia elata (Söderström et al. 2010a).
448
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Riccardia distans (Spruce) Pagán, Bryologist 45 (4): 80, 1942 (Pagán 1942b). Bas.:
Aneura distans Spruce, J. Linn. Soc., Bot. 30 (210): 367, 1895 (Gepp 1895b).
** Riccardia elata (Steph.) Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 67: 169, 1898 (Schifner 1898a). Bas.: Aneura elata Steph., Hedwigia 32
(1): 19, 1893 (Stephani 1893a).
* Riccardia elata var. laccida Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 67: 170, 1898 (Schifner 1898a).
* Riccardia elata var. intercedens Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.Naturwiss. Kl. 67: 170, 1898 (Schifner 1898a).
** Riccardia elegans (Steph.) Hürl., Bauhinia 5 (4): 196, 1976 (Hürlimann 1976).
Bas.: Aneura elegans Steph., Sp. Hepat. (Stephani) 6: 25, 1917 (Stephani 1917a).
** Riccardia elisabethae houvenot et Reeb, Telopea 17: 229, 2014 (houvenot and
Reeb 2014).
*** Riccardia emarginata (Steph.) K.G.Hell, Bol. Fac. Filos. Univ. São Paulo, Bot.
25: 100, 1969 (Hell 1969). Bas.: Aneura emarginata Steph., Hedwigia 32 (1): 20,
1893 (Stephani 1893a).
** Riccardia erosa (Steph.) E.W.Jones, Trans. Brit. Bryol. Soc. 3 (1): 83, 1956 (Jones
1956). Bas.: Aneura erosa Steph., Hedwigia 30 (6): 269, 1891 (Stephani 1891c).
** Riccardia fastigiata (Lehm.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 431, 1877 (Trevisan 1877). Bas.: Jungermannia fastigiata Lehm.,
Linnaea 4: 370, 1829 (Lehmann 1829).
** Riccardia fendleri (Steph.) Pagán, Bryologist 45 (4): 80, 1942 (Pagán 1942b). Bas.:
Aneura fendleri Steph., Hedwigia 32 (1): 20, 1893 (Stephani 1893a).
** Riccardia laccida (Steph.) S.Hatt., Bull. Tokyo Sci. Mus. 11: 164, 1944 (Hattori
1944d). Bas.: Aneura laccida Steph., Sp. Hepat. (Stephani) 6: 26, 1917 (Stephani
1917a).
** Riccardia laccidissima Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss.
Kl. 67: 167, 1898 (Schifner 1898a).
** Riccardia lagellaris (A.Gepp) H.A.Mill., Phytologia 47 (4): 323, 1981 (Miller
1981). Bas.: Aneura lagellaris A.Gepp, J. Linn. Soc., Bot. 39 (270): 194, 1909
(Gibbs 1909).
** Riccardia lagellifrons C.Gao, Fl. Hepat. Chin. Boreali-Orient.: 209, 1981 (Gao
and Chang 1981).
** Riccardia leischeri (Steph.) H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 527, 1963 (Miller 1963).
Bas.: Aneura leischeri Steph., Sp. Hepat. (Stephani) 6: 26, 1917 (Stephani 1917a).
** Riccardia foliacea Meenks et C.De Jong, Cryptog. Bryol. Lichénol. 6 (1): 8, 1985
(Meenks and de Jong 1985).
** Riccardia formosensis (Steph.) Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2,
Bot. 2: 125, 1934 (Horikawa 1934). Bas.: Aneura formosensis Steph., Sp. Hepat.
(Stephani) 6: 27, 1917 (Stephani 1917a).
*** Riccardia fucoidea (Sw.) C.Massal., Nuovo Giorn. Bot. Ital. 17 (3): 256, 1885
(Massalongo 1885). Bas.: Jungermannia fucoidea Sw., Prodr. (Swartz): 145, 1788
(Swartz 1788).
World checklist of hornworts and liverworts
449
** Riccardia geniana Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95 (1): 76,
1970 (Hewson 1970a).
*** Riccardia glaziovii (Spruce) Meenks, J. Hattori Bot. Lab. 62: 173, 1987 (Meenks
1987). Bas.: Aneura glaziovii Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: cci,
1889 [1890] (Spruce 1889).
** Riccardia gogolensis (Steph.) Hewson, Proc. Linn. Soc. New South Wales (ser. 2)
95 (1): 102, 1970 (Hewson 1970a). Bas.: Aneura gogolensis Steph., Bull. Herb.
Boissier 7 (9): 689 (230), 1899 (Stephani 1899e).
** Riccardia gracilis (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 295, 1963 (Schuster 1963b). Bas.: Aneura gracilis Steph., Bull. Herb. Boissier 7 (10): 752 (262),
1899 (Stephani 1899f).
** Riccardia grandilora (Steph.) H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 528, 1963 (Miller
1963). Bas.: Aneura grandilora Steph., Sp. Hepat. (Stephani) 6: 29, 1917 (Stephani
1917a).
*** Riccardia grollei Furuki, Haussknechtia, Beih. 9: 139, 1999 (Furuki 1999).
** Riccardia grossidens (Steph.) Pagán, Bryologist 45 (4): 80, 1942 (Pagán 1942b).
Bas.: Aneura grossidens Steph., Hedwigia 32 (1): 23, 1893 (Stephani 1893a).
** Riccardia gunniana (Steph.) H.A.Mill., Phytologia 47 (4): 323, 1981 (Miller
1981). Bas.: Aneura gunniana Steph., J. & Proc. Roy. Soc. New South Wales 48
(1/2): 96, 1914 (Stephani and Watts 1914).
** Riccardia hamatilora (Steph.) H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 526, 1963 (Miller 1963). Bas.: Aneura hamatilora Steph., Bull. Herb. Boissier 5 (10): 844, 1897
(Stephani 1897c).
*** Riccardia hans-meyeri (Steph.) Meenks et C.De Jong, Cryptog. Bryol. Lichénol. 6
(1): 12, 1985 (Meenks and de Jong 1985). Bas.: Aneura hans-meyeri Steph., Sp.
Hepat. (Stephani) 6: 29, 1917 (Stephani 1917a).
** Riccardia hans-meyeri var. dentata Meenks, J. Hattori Bot. Lab. 62: 173, 1987
(Meenks 1987).
** Riccardia hawaica (Steph.) H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 528, 1963 (Miller 1963).
Bas.: Aneura hawaica Steph., Sp. Hepat. (Stephani) 6: 30, 1917 (Stephani 1917a).
** Riccardia hebridensis (Steph.) H.A.Mill., Phytologia 47 (4): 323, 1981 (Miller
1981). Bas.: Aneura hebridensis Steph., J. & Proc. Roy. Soc. New South Wales 48
(1/2): 96, 1914 (Stephani and Watts 1914).
** Riccardia herzogiana (Steph.) Meenks et C.De Jong, Cryptog. Bryol. Lichénol. 6
(1): 12, 1985 (Meenks and de Jong 1985). Bas.: Aneura herzogiana Steph., Biblioth. Bot. 87 (2): 175, 1916 (Stephani 1916a).
** Riccardia heteroclada Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss.
Kl. 67: 175, 1898 (Schifner 1898a).
* Riccardia hirtilora (Steph.) Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl.,
Denkschr. 111: 19, 1964 (Schifner and Arnell 1964). Bas.: Aneura hirtilora
Steph., Arch. Mus. Nac. Rio de Janeiro 13: 116, 1905 (Stephani 1905c). 404
404 Riccardia hirtilora is possibly conspeciic with Riccardia emarginata (Gradstein and Costa 2003).
450
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Riccardia humilis (Gottsche) O.Yano, J. Hattori Bot. Lab. 56: 530, 1984 (Yano 1984).
Bas.: Pseudoneura humilis Gottsche, Mexik. Leverm.: 260, 1863 (Gottsche 1863).
** Riccardia hyalina (Steph.) H.A.Mill., Phytologia 47 (4): 324, 1981 (Miller 1981).
Bas.: Aneura hyalina Steph., Sp. Hepat. (Stephani) 6: 31, 1917 (Stephani 1917a).
** Riccardia hydra Hürl., Bauhinia 5 (4): 210, 1976 (Hürlimann 1976).
** Riccardia hymenophylloides Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 67: 175, 1898 (Schifner 1898a).
* Riccardia hymenophylloides var. laccida Schifn., Denkschr. Kaiserl. Akad. Wiss.,
Math.-Naturwiss. Kl. 67: 176, 1898 (Schifner 1898a).
*** Riccardia hymenophytoides (Spruce) Meenks, Beih. Nova Hedwigia 88: 101, 1987
(Schultze-Motel and Menzel 1987). Bas.: Aneura hymenophytoides Spruce, Trans.
& Proc. Bot. Soc. Edinburgh 15: 549, 1885 (Spruce 1885).
** Riccardia hypipamensis Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95 (1):
97, 1970 (Hewson 1970a).
** Riccardia ibana Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95 (1): 78,
1970 (Hewson 1970a).
*** Riccardia incurvata Lindb., Helsingf. Dagbl. 1878 (315, 18 Nov.): 2, 1878 (Lindberg 1878).
** Riccardia innovans (Steph.) Pagán, Bryologist 45 (4): 80, 1942 (Pagán 1942b).
Bas.: Aneura innovans Steph., Symb. Antill. 2: 470, 1901 (Stephani 1901f).
** Riccardia insularis Schifn., Deutsche Südpolar-Exped. 1901-1903, 8 (bot.) 1: 66,
1906 (Schifner 1906a).
** Riccardia intricata (Steph.) H.A.Mill., Phytologia 47 (4): 324, 1981 (Miller 1981).
Bas.: Aneura intricata Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 301, 1896 (Stephani 1896a).
** Riccardia jackii Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 67:
165, 1898 (Schifner 1898a).
* Riccardia jackii var. densa Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 67: 165, 1898 (Schifner 1898a).
** Riccardia judithae Meenks et C.De Jong, Cryptog. Bryol. Lichénol. 6 (1): 12,
1985 (Meenks and de Jong 1985).
*** Riccardia jugata R.M.Schust., J. Hattori Bot. Lab. 62: 305, 1987 (Schuster 1987d).
* Riccardia karstenii (Steph.) Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.Naturwiss. Kl. 67: 177, 1898 (Schifner 1898a). Bas.: Aneura karstenii Steph.,
Hedwigia 32 (1): 23, 1893 (Stephani 1893a).
** Riccardia laticostata (Spruce) Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl.,
Denkschr. 111: 12, 1964 (Schifner and Arnell 1964). Bas.: Aneura laticostata
Spruce, J. Linn. Soc., Bot. 30 (210): 367, 1895 (Gepp 1895b).
** Riccardia latifrondoides Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 67: 168, 1898 (Schifner 1898a).
** Riccardia lepidomitra (Spruce) Gradst., J. Hattori Bot. Lab. 45: 130, 1979 (Gradstein and Hekking 1979). Bas.: Aneura lepidomitra Spruce, Trans. & Proc. Bot.
Soc. Edinburgh 15: 549, 1885 (Spruce 1885).
World checklist of hornworts and liverworts
451
** Riccardia leptophylla (Spruce) Herzog, Svensk Bot. Tidskr. 46 (1): 65, 1952 (Herzog 1952e). Bas.: Aneura leptophylla Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 544, 1885 (Spruce 1885).
** Riccardia leptothallus R.M.Schust., J. Hattori Bot. Lab. 67: 87, 1989 (Schuster 1989).
** Riccardia levieri Schifn., Österr. Bot. Z. 49 (4): 130, 1899 (Schifner 1899b).
** Riccardia lichenoides (Steph.) H.A.Mill., Phytologia 47 (4): 324, 1981 (Miller
1981). Bas.: Aneura lichenoides Steph., Bot. Jahrb. Syst. 23 (1/2, 3): 301, 1896
(Stephani 1896a).
** Riccardia ligulata (Steph.) Pócs et Schäf.-Verw., Cryptog. Bryol. 31 (4): 390, 2010
(Schäfer-Verwimp 2010). Bas.: Aneura ligulata Steph., Sp. Hepat. (Stephani) 6:
32, 1917 (Stephani 1917a).
** Riccardia lilliena (Steph.) H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 527, 1963 (Miller 1963).
Bas.: Aneura lilliena Steph., Sp. Hepat. (Stephani) 6: 33, 1917 (Stephani 1917a).
** Riccardia limbata (Steph.) E.W.Jones, Trans. Brit. Bryol. Soc. 3 (1): 79, 1956
(Jones 1956). Bas.: Aneura limbata Steph., Hedwigia 30 (5): 203, 1891 (Stephani 1891a).
** Riccardia loefgrenii Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl., Denkschr.
111: 14, 1964 (Schifner and Arnell 1964).
** Riccardia longilora (Steph.) Hewson, Proc. Linn. Soc. New South Wales (ser. 2)
95 (1): 88, 1970 (Hewson 1970a). Bas.: Aneura longilora Steph., Bull. Herb. Boissier 7 (10): 746 (256), 1899 (Stephani 1899f).
** Riccardia longispica (Steph.) Pearson, Forh. Vidensk.-Selsk. Kristiania 1892 (14):
4, 1893 (Pearson 1893). Bas.: Aneura longispica Steph., Bot. Gaz. 15 (11): 281,
1890 (Stephani 1890c).
** Riccardia loriana (Steph.) H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 528, 1963 (Miller
1963). Bas.: Aneura loriana Steph., Bull. Herb. Boissier 7 (10): 733 (243), 1899
(Stephani 1899f).
** Riccardia macdonaldiana Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95
(1): 93, 1970 (Hewson 1970a).
** Riccardia macrantha (Pearson) H.A.Mill., Phytologia 47 (4): 324, 1981 (Miller
1981). Bas.: Aneura macrantha Pearson, J. Linn. Soc., Bot. 46 (305): 17, 1922
(Pearson 1922b).
*** Riccardia magnicellularis Furuki, J. Hattori Bot. Lab. 100: 93, 2006 (Furuki 2006a).
*** Riccardia metzgeriiformis (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 295, 1963
(Schuster 1963b). Bas.: Aneura metzgeriiformis Steph., Bull. Herb. Boissier 7 (10):
753 (263), 1899 (Stephani 1899f).
** Riccardia microscopica (Nees) Kuntze, Revis. Gen. Pl. 2: 838, 1891 (Kuntze
1891). Bas.: Aneura microscopica Nees, Syn. Hepat. 4: 500, 1846 (Gottsche et
al. 1846).
** Riccardia minuta (Steph.) W.Martin, Trans. & Proc. Roy. Soc. New Zealand 78
(4): 499, 1950 (Martin 1950). Bas.: Aneura minuta Steph., Sp. Hepat. (Stephani)
6: 34, 1917 (Stephani 1917a).
452
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**
**
**
**
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**
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**
*
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Riccardia multiidoides Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss.
Kl. 67: 166, 1898 (Schifner 1898a). 405
Riccardia multispica (Steph.) S.Hatt., Bull. Tokyo Sci. Mus. 11: 164, 1944 (Hattori 1944d). Bas.: Aneura multispica Steph., Sp. Hepat. (Stephani) 6: 34, 1917
(Stephani 1917a).
Riccardia nadeaudii (Steph.) Hürl., Bauhinia 5 (4): 201, 1976 (Hürlimann 1976).
Bas.: Aneura nadeaudii Steph., Bull. Herb. Boissier 7 (10): 750 (260), 1899
(Stephani 1899f).
Riccardia newellana (Steph.) H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 528, 1963 (Miller
1963). Bas.: Aneura newellana Steph., Sp. Hepat. (Stephani) 6: 35, 1917 (Stephani
1917a).
Riccardia nigra (Steph.) H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 527, 1963 (Miller 1963).
Bas.: Aneura nigra Steph., Sp. Hepat. (Stephani) 6: 35, 1917 (Stephani 1917a).
Riccardia nobilis (Steph.) Schifn., Consp. Hepat. Arch. Ind.: 56, 1898 (Schifner
1898b). Bas.: Aneura nobilis Steph., Hedwigia 32 (1): 24, 1893 (Stephani 1893a).
Riccardia novo-amstelodamensis Schifn., Deutsche Südpolar-Exped. 1901-1903, 8
(bot.) 1: 65, 1906 (Schifner 1906a).
Riccardia nudilora (Steph.) Grolle, Bryophyt. Biblioth. 48: 130, 1995 (Grolle 1995).
Bas.: Aneura nudilora Steph., Bot. Gaz. 15 (11): 282, 1890 (Stephani 1890c).
Riccardia obtusa S.W.Arnell, Bot. Not. 105: 142, 1952 (Arnell 1952b).
Riccardia obtusifrons (Steph.) H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 527, 1963 (Miller
1963). Bas.: Aneura obtusifrons Steph., Sp. Hepat. (Stephani) 6: 36, 1917 (Stephani
1917a).
Riccardia omkaliensis Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95 (1):
97, 1970 (Hewson 1970a).
Riccardia pallida (Spruce) Meenks et C.De Jong, Cryptog. Bryol. Lichénol. 6 (1):
15, 1985 (Meenks and de Jong 1985). Bas.: Aneura pallida Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 547, 1885 (Spruce 1885).
Riccardia palmatiida (Steph.) H.A.Mill., Phytologia 47 (4): 324, 1981 (Miller 1981). Bas.: Aneura palmatiida Steph., Sp. Hepat. (Stephani) 6: 36, 1917
(Stephani 1917a).
Riccardia palmatiformis Schifn., J. Indian Bot. Soc. 38 (4): 538, 1959 [1960]
(Schifner et al. 1959). 406
Riccardia papillata (Gottsche) Hässel ex Gradst. et Hekking, J. Hattori Bot.
Lab. 45: 130, 1979 (Gradstein and Hekking 1979). Bas.: Pseudoneura papillata
Gottsche, Ann. Sci. Nat. Bot. (sér. 5) 1: 185, 1864 (Gottsche 1864).
Riccardia paramorum Meenks, J. Hattori Bot. Lab. 62: 176, 1987 (Meenks 1987).
405 Riccardia multiidoides is conspeciic with Riccardia multiida in Meijer (1959), but that was not
based on examination of type material (Söderström et al. 2010a).
406 Riccardia palmatiformis was treated as a doubtful taxon with some ainity with Riccardia levieri by
Srivastava and Udar (1976).
World checklist of hornworts and liverworts
453
*** Riccardia parasitans (Steph.) Meenks et C.De Jong, Cryptog. Bryol. Lichénol. 6
(1): 17, 1985 (Meenks and de Jong 1985). Bas.: Aneura parasitans Steph., Biblioth. Bot. 87 (2): 175, 1916 (Stephani 1916a).
** Riccardia parvula Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl.
67: 172, 1898 (Schifner 1898a).
** Riccardia pauciramea (Steph.) H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 527, 1963 (Miller 1963). Bas.: Aneura pauciramea Steph., Bull. Herb. Boissier 5 (10): 845, 1897
(Stephani 1897c).
** Riccardia paulensis Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl., Denkschr.
111: 16, 1964 (Schifner and Arnell 1964).
** Riccardia pectinata (Austin) H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 526, 1963 (Miller
1963). Bas.: Aneura pectinata Austin, Bull. Torrey Bot. Club 5 (3): 15, 1874
(Austin 1874).
** Riccardia pellucida Piippo, J. Hattori Bot. Lab. 68: 134, 1990 (Piippo 1990). Nom.
nov. pro Aneura pellucida Steph., Sp. Hepat. (Stephani) 6: 37, 1917 (Stephani
1917a), nom. illeg.
** Riccardia pembaiensis (Steph.) Hürl., Bauhinia 5 (4): 205, 1976 (Hürlimann 1976).
Bas.: Aneura pembaiensis Steph., Sp. Hepat. (Stephani) 6: 37, 1917 (Stephani 1917a).
** Riccardia pengagensis Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95 (1):
108, 1970 (Hewson 1970a).
** Riccardia perssonii S.C.Srivast. et Udar, Lindbergia 4 (1/2): 127, 1977 (Srivastava
and Udar 1977).
*** Riccardia philippinensis Furuki, J. Hattori Bot. Lab. 100: 94, 2006 (Furuki 2006a).
** Riccardia phleganiana Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95 (1):
80, 1970 (Hewson 1970a).
** Riccardia plana (Steph.) Hürl., Bauhinia 5 (4): 205, 1976 (Hürlimann 1976).
Bas.: Aneura plana Steph., Sp. Hepat. (Stephani) 6: 38, 1917 (Stephani 1917a).
* Riccardia plana var. minor (Pearson) Hürl. ex H.A.Mill., H.Whittier et B.Whittier,
Bryophyt. Biblioth. 25: 303, 1983 (Miller et al. 1983). Bas.: Aneura plana var. minor Pearson, J. Linn. Soc., Bot. 46 (305): 16, 1922 (Pearson 1922b).
** Riccardia planifrons (Spruce) Pagán, Bryologist 45 (4): 80, 1942 (Pagán 1942b). Bas.:
Aneura planifrons Spruce, J. Linn. Soc., Bot. 30 (210): 368, 1895 (Gepp 1895b).
*** Riccardia plumiformis (Spruce) Hässel ex Meenks, Beih. Nova Hedwigia 88: 101,
1987 (Schultze-Motel and Menzel 1987). Bas.: Aneura plumiformis Spruce, Trans.
& Proc. Bot. Soc. Edinburgh 15: 548, 1885 (Spruce 1885).
** Riccardia plumosa (Mitt.) E.O.Campb., J. Roy. Soc. New Zealand 1 (1): 24, 1971
(Campbell 1971). Bas.: Sarcomitrium plumosum Mitt., Bonplandia 10 (2): 19,
1862 (Mitten 1862).
*** Riccardia poeppigiana (Lehm. et Lindenb.) Hässel ex Meenks et C.De Jong,
Cryptog. Bryol. Lichénol. 6 (1): 17, 1985 (Meenks and de Jong 1985). Bas.:
Jungermannia poeppigiana Lehm. et Lindenb., Nov. Stirp. Pug. 6: 23, 1834
(Lehmann 1834).
454
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Riccardia porcina (Hewson) L.Söderstr., Phytotaxa 202 (1): 70, 2015 (Söderström
et al. 2015c). Bas.: Riccardia bliklika var. porcina Hewson, Proc. Linn. Soc. New
South Wales (ser. 2) 95 (1): 84, 1970 (Hewson 1970a).
** Riccardia portoricensis (Steph.) Pagán, Bryologist 42 (1): 7, 1939 (Pagán 1939a).
Bas.: Aneura portoricensis Steph., Bull. Herb. Boissier 7 (10): 739 (249), 1899
(Stephani 1899f).
** Riccardia punahuina (Steph.) H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 526, 1963 (Miller
1963). Bas.: Aneura punahuina Steph., Sp. Hepat. (Stephani) 6: 39, 1917 (Stephani
1917a).
** Riccardia ramosissima (Steph.) Grolle, Bryophyt. Biblioth. 48: 130, 1995 (Grolle
1995). Bas.: Aneura ramosissima Steph., Bull. Soc. Roy. Bot. Belgique, Compt.
Rend. 30 (2): 196, 1891 [1892] (Stephani 1891b).
** Riccardia regina Meenks et C.De Jong, Cryptog. Bryol. Lichénol. 6 (1): 17, 1985
(Meenks and de Jong 1985).
*** Riccardia regnellii (Ångstr.) K.G.Hell, Bol. Fac. Filos. Univ. São Paulo, Bot. 25:
110, 1969 (Hell 1969). Bas.: Pseudoneura regnellii Ångstr., Öfvers. Kongl. Vetensk.-Akad. Förh. 33 (7): 90, 1876 [1877] (Ångström 1876).
** Riccardia reyesiana Meenks, Acta Bot. Hung. 32 (1/4): 207, 1986 (Meenks 1986).
** Riccardia riccioides Pearson, Univ. Calif. Publ. Bot. 10 (4): 310, 1923 (Pearson 1923).
* Riccardia rigida Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl.
67: 172, 1898 (Schifner 1898a).
** Riccardia robbinsii Hewson et Grolle, J. Hattori Bot. Lab. 29: 72, 1966 (Grolle 1966i).
** Riccardia robusta (Steph.) H.A.Mill., Phytologia 47 (4): 324, 1981 (Miller 1981).
Bas.: Aneura robusta Steph., Sp. Hepat. (Stephani) 6: 40, 1917 (Stephani 1917a).
** Riccardia rockii (Steph.) H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 527, 1963 (Miller 1963).
Bas.: Aneura rockii Steph., Sp. Hepat. (Stephani) 6: 41, 1917 (Stephani 1917a).
** Riccardia rupicola (Steph.) Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95
(1): 99, 1970 (Hewson 1970a). Bas.: Aneura rupicola Steph., Sp. Hepat. (Stephani) 6: 41, 1917 (Stephani 1917a).
** Riccardia russellii R.M.Schust., J. Hattori Bot. Lab. 67: 93, 1989 (Schuster 1989).
** Riccardia saccatilora (Steph.) S.W.Arnell, Bot. Not. 105: 144, 1952 (Arnell 1952b).
Bas.: Aneura saccatilora Steph., Bot. Gaz. 15 (11): 282, 1890 (Stephani 1890c).
** Riccardia santapaui Udar et S.C.Srivast., Rev. Bryol. Lichénol. 39 (1): 155, 1973
(Udar and Srivastava 1973).
*** Riccardia schwaneckei (Steph.) Pagán, Bryologist 42 (1): 7, 1939 (Pagán 1939a).
Bas.: Aneura schwaneckei Steph., Hedwigia 27 (11/12): 278, 1888 (Stephani 1888c).
** Riccardia singapurensis Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss.
Kl. 67: 165, 1898 (Schifner 1898a).
** Riccardia smaragdina Meenks et C.De Jong, Cryptog. Bryol. Lichénol. 6 (1): 20,
1985 (Meenks and de Jong 1985).
*** Riccardia sprucei (Steph.) Meenks et C.De Jong, Cryptog. Bryol. Lichénol. 6 (1):
22, 1985 (Meenks and de Jong 1985). Bas.: Aneura sprucei Steph., Bull. Herb.
Boissier 5 (10): 844, 1897 (Stephani 1897c).
World checklist of hornworts and liverworts
455
** Riccardia squamifera Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl., Denkschr. 111: 19, 1964 (Schifner and Arnell 1964).
* Riccardia stipatilora (Steph.) Pagán, Bryologist 45 (4): 81, 1942 (Pagán 1942b).
Bas.: Aneura stipatilora Steph., Hedwigia 32 (1): 27, 1893 (Stephani 1893a).
** Riccardia stricta R.M.Schust., J. Hattori Bot. Lab. 62: 326, 1987 (Schuster 1987d).
** Riccardia subantarctica Grolle et L.Söderstr., Phytotaxa 202 (1): 70, 2015 (Söderström et al. 2015c). Nom. nov. pro Riccardia pauciramea R.M.Schust., J. Hattori
Bot. Lab. 67: 102, 1989 (Schuster 1989), nom. illeg.
** Riccardia subexalata Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss.
Kl. 67: 163, 1898 (Schifner 1898a).
* Riccardia subexalata var. procera Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.Naturwiss. Kl. 67: 164, 1898 (Schifner 1898a).
* Riccardia submersa (Hook.f. et Taylor) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser.
3), C. Sci. Mat. 4 (13): 431, 1877 (Trevisan 1877). Bas.: Jungermannia multiida var.
submersa Hook.f. et Taylor, London J. Bot. 4: 94, 1845 (Hooker and Taylor 1845).
** Riccardia submultiida Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 128,
1934 (Horikawa 1934).
** Riccardia subpalmata (Steph.) Hürl., Bauhinia 5 (4): 194, 1976 (Hürlimann 1976).
Bas.: Aneura subpalmata Steph., Sp. Hepat. (Stephani) 6: 43, 1917 (Stephani 1917a).
** Riccardia subsimplex (Steph.) Pagán, Bryologist 45 (4): 81, 1942 (Pagán 1942b).
Bas.: Aneura subsimplex Steph., Hedwigia 32 (1): 27, 1893 (Stephani 1893a).
** Riccardia sumatrana Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss.
Kl. 67: 173, 1898 (Schifner 1898a).
** Riccardia tahitensis (Steph.) Hürl., Bauhinia 5 (4): 201, 1976 (Hürlimann 1976).
Bas.: Aneura tahitensis Steph., Bull. Herb. Boissier 7 (10): 728 (238), 1899
(Stephani 1899f).
* Riccardia tenella Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95 (1): 95,
1970 (Hewson 1970a).
*** Riccardia tenuicula (Spruce) Meenks, Beih. Nova Hedwigia 88: 101, 1987
(Schultze-Motel and Menzel 1987). Bas.: Aneura tenuicula Spruce, Trans. & Proc.
Bot. Soc. Edinburgh 15: 545, 1885 (Spruce 1885).
** Riccardia tenuis (Steph.) Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 67: 172, 1898 (Schifner 1898a). Bas.: Aneura tenuis Steph., Hedwigia 32
(1): 28, 1893 (Stephani 1893a).
* Riccardia tjibodensis Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss.
Kl. 67: 165, 1898 (Schifner 1898a).
*** Riccardia trichomanoides (Spruce) Hässel ex Meenks, Beih. Nova Hedwigia 88:
101, 1987 (Schultze-Motel and Menzel 1987). Bas.: Aneura trichomanoides
Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 547, 1885 (Spruce 1885).
** Riccardia trukensis H.A.Mill. et Bonner, Beih. Nova Hedwigia 11: 70, 1963 (Miller et al. 1963).
** Riccardia tumbareriensis Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95
(1): 87, 1970 (Hewson 1970a).
456
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Riccardia upoluna (Steph.) Grolle, J. Hattori Bot. Lab. 36: 77, 1972 [1973] (Grolle
and Schultze-Motel 1972). Bas.: Aneura upoluna Steph., Denkschr. Kaiserl. Akad.
Wiss., Math.-Naturwiss. Kl. 91: 165, 1915 (Stephani 1915a).
** Riccardia valida (Steph.) J.J.Engel, Bryologist 78 (3): 362, 1975 (Engel 1975).
Bas.: Aneura valida Steph., Sp. Hepat. (Stephani) 6: 44, 1917 (Stephani 1917a).
** Riccardia venosa (Steph.) Hürl., Bauhinia 5 (4): 208, 1976 (Hürlimann 1976).
Bas.: Aneura venosa Steph., Sp. Hepat. (Stephani) 6: 45, 1917 (Stephani 1917a).
* Riccardia villosa (Steph.) Pandé et S.C.Srivast., Biol. Mem. 1 (1/2): 129, 1976
(Srivastava and Udar 1976). Bas.: Aneura villosa Steph., Sp. Hepat. (Stephani)
6: 45, 1917 (Stephani 1917a).
** Riccardia virens (Steph.) Hürl., Bauhinia 5 (4): 209, 1976 (Hürlimann 1976).
Bas.: Aneura virens Steph., Sp. Hepat. (Stephani) 6: 45, 1917 (Stephani 1917a).
** Riccardia virgata (Gottsche) Pagán, Bryologist 42 (1): 7, 1939 (Pagán 1939a).
Bas.: Aneura virgata Gottsche, Hedwigia 27 (11/12): 277, 1888 (Stephani
1888c).
** Riccardia wallisii (Steph.) Gradst., J. Hattori Bot. Lab. 45: 130, 1979 (Gradstein and Hekking 1979). Bas.: Aneura wallisii Steph., Hedwigia 32 (1): 28, 1893
(Stephani 1893a).
** Riccardia wettsteinii Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss.
Kl. 67: 162, 1898 (Schifner 1898a).
* Riccardia wettsteinii var. angustilimbia Schifn., Denkschr. Kaiserl. Akad. Wiss.,
Math.-Naturwiss. Kl. 67: 163, 1898 (Schifner 1898a).
* Riccardia wettsteinii var. crassa Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.Naturwiss. Kl. 67: 163, 1898 (Schifner 1898a).
* Riccardia wettsteinii var. procera Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.Naturwiss. Kl. 67: 162, 1898 (Schifner 1898a).
* Riccardia wettsteinii var. tenuiretis Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.Naturwiss. Kl. 67: 162, 1898 (Schifner 1898a).
** Riccardia womersleyana Hewson, Proc. Linn. Soc. New South Wales (ser. 2) 95
(1): 98, 1970 (Hewson 1970a).
** Verdoornia R.M.Schust., J. Hattori Bot. Lab. 26: 291, 1963 (Schuster 1963b).
*** Verdoornia succulenta R.M.Schust., J. Hattori Bot. Lab. 26: 291, 1963 (Schuster
1963b).
***
Metzgeriaceae H.Klinggr.
by D. P. Costa
he treatment of Metzgeriaceae follows what was outlined in Hewson (1982), Kuwahara (1986), Crandall-Stotler and Stotler (2000), and Costa (2008).
World checklist of hornworts and liverworts
457
*** Metzgeria Raddi, Jungermanniogr. Etrusca: 34, 1818 (Raddi 1818a). 407
*** Metzgeria acuminata Steph., Bull. Herb. Boissier 7 (12): 934 (282), 1899 (Stephani 1899g).
*** Metzgeria adscendens Steph. ex K.I.Goebel, Flora 77: 427, 1893 (Goebel 1893a).
*** Metzgeria agnewiae Kuwah., Bryologist 76 (4): 569, 1973 (Kuwahara 1973b). 408
*** Metzgeria albinea Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: cci, 1889
[1890] (Spruce 1889).
*** Metzgeria albinea var. aberrans Schifn., Österr. Akad. Wiss., Math.-Naturwiss.
Kl., Denkschr. 111: 27, 1964 (Schifner and Arnell 1964).
*** Metzgeria albinea var. angusta (Steph.) D.P.Costa et Gradst., Bryologist 103 (4):
757, 2000 [2001] (Costa and Gradstein 2000). Bas.: Metzgeria angusta Steph.,
Bull. Herb. Boissier 7 (12): 944 (292), 1899 (Stephani 1899g).
*** Metzgeria allionii Steph., Sp. Hepat. (Stephani) 6: 47, 1917 (Stephani 1917a).
*** Metzgeria alpina R.M.Schust. et J.J.Engel, Brittonia 40 (2): 203, 1988 (Engel and
Schuster 1988).
*** Metzgeria americana Masuzaki, Hikobia 15 (4): 441, 2010 (Masuzaki et al. 2010a).
*** Metzgeria attenuata Steph., Biblioth. Bot. 87 (2): 177, 1916 (Stephani 1916a).
*** Metzgeria aurantiaca Steph., Bull. Herb. Boissier 7 (12): 938 (286), 1899 (Stephani 1899g).
*** Metzgeria auriculata Grolle et Kuwah., Bryophyt. Biblioth. 28: 194, 1986 (Kuwahara 1986).
*** Metzgeria bahiensis Schifn., Österr. Bot. Z. 61 (7/8): 262, 1911 (Schifner 1911).
*** Metzgeria bartlettii Kuwah., Mem. New York Bot. Gard. 45: 561, 1987 (Kuwahara 1987).
*** Metzgeria bischlerae Kuwah., J. Hattori Bot. Lab. 40: 264, 1976 (Kuwahara
1976a).
*** Metzgeria bracteata Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 553, 1885
(Spruce 1885).
*** Metzgeria brasiliensis Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl., Denkschr. 111: 22, 1964 (Schifner and Arnell 1964).
*** Metzgeria chilensis Steph., Bull. Herb. Boissier 7 (12): 937 (285), 1899 (Stephani
1899g).
*** Metzgeria ciliata Raddi, Critt. Brasil.: 17, 1822 (Raddi 1822). 409
*** Metzgeria clavilora Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 556, 1885
(Spruce 1885).
407 Metzgeria was divided into many subdivisions proposed by Kuwahara in a series of publications, but
they are artiicial and not accepted here.
408 Metzgeria agnewiae is conspeciic with Metzgeria consanguinea in So (2004), but it was accepted by
Costa (2008).
409 Metzgeria ciliata is conspeciic with Metzgeria furcata in Grolle (2002) and with Metzgeria dichotoma
in Gradstein and Costa (2003), but it was accepted by Costa (2008).
458
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Metzgeria cleeii Kuwah., Proc. Kon. Ned. Akad. Wetensch. C 85 (3): 360, 1982
(Kuwahara 1982).
** Metzgeria comata Steph., Bull. Herb. Boissier 7 (12): 939 (287), 1899 (Stephani
1899g).
*** Metzgeria conjugata Lindb., Acta Soc. Sci. Fenn. 10: 495, 1875 (Lindberg 1875). 410
*** Metzgeria consanguinea Schifn., Nova Acta Acad. Caes. Leop.-Carol. German.
Nat. Cur. 60 (2): 271, 1893 (Schifner 1893a).
*** Metzgeria convoluta Steph., Bull. Herb. Boissier 7 (12): 940 (288), 1899 (Stephani
1899g).
** Metzgeria coorgensis S.C.Srivast. et S.Srivast., Phytotaxonomy 4: 81, 2004 (Srivastava
and Srivastava 2004).
** Metzgeria corralensis Steph., Bull. Herb. Boissier 7 (12): 933 (281), 1899 (Stephani
1899g).
*** Metzgeria crassipilis (Lindb.) A.Evans, Rhodora 11 (130): 188, 1909 (Evans 1909).
Bas.: Metzgeria furcata subsp. crassipilis Lindb., Acta Soc. Fauna Fl. Fenn. 1 (2):
42, 1877 (Lindberg 1877b).
*** Metzgeria cratoneura Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl., Denkschr.
111: 24, 1964 (Schifner and Arnell 1964).
*** Metzgeria cylindra Kuwah., Bryologist 81 (3): 405, 1978 (Kuwahara 1978).
** Metzgeria decrescens Steph., Bull. Herb. Boissier 7 (12): 932 (280), 1899 (Stephani
1899g).
*** Metzgeria dichotoma (Sw.) Nees, Syn. Hepat. 4: 504, 1846 (Gottsche et al. 1846).
Bas.: Jungermannia dichotoma Sw., Prodr. (Swartz): 145, 1788 (Swartz 1788).
** Metzgeria divaricata A.Evans, Proc. Amer. Acad. Arts 58 (7): 288, 1923 (Evans
1923a).
*** Metzgeria dorsipara (Herzog) Kuwah., J. Hattori Bot. Lab. 40: 269, 1976 (Kuwahara 1976a). Bas.: Metzgeria violacea var. dorsipara Herzog, Svensk Bot. Tidskr. 51
(1): 187, 1957 (Herzog 1957a).
*** Metzgeria duricosta Steph., Sp. Hepat. (Stephani) 6: 50, 1917 (Stephani 1917a).
** Metzgeria engelii Kuwah., Hikobia 8 (3/4): 275, 1980 (Kuwahara 1980a).
** Metzgeria epiphylla A.Evans, Proc. Amer. Acad. Arts 58 (7): 303, 1923 (Evans 1923a).
*** Metzgeria ilicina Mitt., Hooker’s J. Bot. Kew Gard. Misc. 3: 361, 1851 (Mitten 1851).
** Metzgeria lavovirens Colenso, Trans. & Proc. New Zealand Inst. 21: 79, 1889
(Colenso 1889).
* Metzgeria foliicola Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl.
67: 181, 1898 (Schifner 1898a). 411
*** Metzgeria francana Steph., Sp. Hepat. (Stephani) 6: 51, 1917 (Stephani 1917a).
410 Metzgeria conjugata is a complex taxon. Fuselier et al. (2009) demonstrated the occurrence of two
clearly separated lineages, a northern North American lineage (possibly corresponding to Metzgeria
conjugata s.str.) and a southern North American and European lineage (possibly corresponding to
Metzgeria simplex).
411 Metzgeria foliicola is conspeciic with Metzgeria albinea in Kuwahara (1986), but it was accepted by
So (2002c).
World checklist of hornworts and liverworts
459
** Metzgeria frontipilis Lindb., Acta Soc. Fauna Fl. Fenn. 1 (2): 14, 1877 (Lindberg 1877b).
*** Metzgeria fruticola Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 554, 1885
(Spruce 1885).
*** Metzgeria furcata (L.) Corda, Gen. hepat.: 654, 1829 (Corda 1829). Bas.: Jungermannia furcata L., Sp. Pl. 1: 1136, 1753 (Linnaeus 1753). 412
* Metzgeria furcata var. expansa Douin, Rev. Bryol. 30 (3): 47, 1903 (Douin 1903).
* Metzgeria furcata var. paciica Brinkm., Bryologist 34 (2): 15, 1931 (Brinkman 1931).
*** Metzgeria grandilora A.Evans, Torreya 16 (3): 68, 1916 (Evans 1916).
** Metzgeria hasselii Kuwah., J. Hattori Bot. Lab. 40: 509, 1976 (Kuwahara 1976c).
** Metzgeria hebridensis Steph., Sp. Hepat. (Stephani) 6: 51, 1917 (Stephani 1917a).
*** Metzgeria hegewaldii Kuwah., Nova Hedwigia 34: 784, 1981 (Kuwahara 1981).
*** Metzgeria herminieri Schifn., Österr. Bot. Z. 61 (7/8): 261, 1911 (Schifner 1911).
* Metzgeria heteroramea Steph., Biblioth. Bot. 87 (2): 178, 1916 (Stephani 1916a).
** Metzgeria imberbis J.B.Jack et Steph., Hedwigia 34 (6): 316, 1895 (Jack and
Stephani 1895).
*** Metzgeria inlata Steph., Bull. Herb. Boissier 7 (12): 936 (284), 1899 (Stephani
1899g).
*** Metzgeria jamesonii Kuwah., Bryophyt. Biblioth. 28: 157, 1986 (Kuwahara 1986).
** Metzgeria kanaii Kuwah., Fl. E. Himalaya 2: 240, 1971 (Hattori 1971a).
*** Metzgeria kinabaluensis Masuzaki, Hikobia 16 (1): 59, 2011 (Masuzaki 2011).
Based on: Apometzgeria pubescens var. kinabaluensis Kuwah., J. Hattori Bot. Lab.
28: 166, 1965 (Kuwahara 1965), nom. inval.
** Metzgeria kuwaharae Piippo, Acta Bot. Fenn. 143: 8, 1991 (Piippo 1991).
** Metzgeria laciniata Kuwah., Bryologist 81 (3): 406, 1978 (Kuwahara 1978).
*** Metzgeria lechleri Steph., Bull. Herb. Boissier 7 (12): 942 (290), 1899 (Stephani
1899g).
*** Metzgeria leptoneura Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 555, 1885
(Spruce 1885).
** Metzgeria leptoneura var. breviseta (Schifn.) O.Yano, J. Hattori Bot. Lab. 56: 532,
1984 (Yano 1984). Bas.: Metzgeria hamata var. breviseta Schifn., Österr. Akad.
Wiss., Math.-Naturwiss. Kl., Denkschr. 111: 27, 1964 (Schifner and Arnell 1964).
** Metzgeria leptoneura var. polychaeta R.M.Schust., Phytotaxa 202 (1): 70, 2015
(Söderström et al. 2015c). Based on: Metzgeria leptoneura var. polychaeta
R.M.Schust., J. Hattori Bot. Lab. 70: 150, 1991 (Schuster 1991b), nom. inval.
*** Metzgeria liebmaniana Lindenb. et Gottsche, Syn. Hepat. 4: 505, 1846 (Gottsche
et al. 1846).
*** Metzgeria lindbergii Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss.
Kl. 67: 182, 1898 (Schifner 1898a).
** Metzgeria litoralis J.J.Engel et Kuwah., Bryologist 76 (2): 293, 1973 (Engel and
Kuwahara 1973).
412 Metzgeria furcata includes several cryptic species as demonstrated by Fuselier et al. (2009).
460
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Metzgeria longitexta Steph., Bull. Herb. Boissier 7 (12): 940 (288), 1899 (Stephani
1899g).
** Metzgeria macrospora Kuwah., J. Hattori Bot. Lab. 32: 17, 1969 (Kuwahara
1969a).
** Metzgeria macveanii Kuwah., Rev. Bryol. Lichénol. 36 (3/4): 539, 1969 [1970]
(Kuwahara 1969b).
*** Metzgeria maegdefraui Kuwah., Hikobia 8 (3/4): 269, 1980 (Kuwahara 1980b).
** Metzgeria magellanica Schifn., Leberm., Forschungsr. Gazelle 4 (4): 43, 1890
(Schifner 1890).
*** Metzgeria metaensis Kuwah., Proc. Kon. Ned. Akad. Wetensch. C 85 (3): 375,
1982 (Kuwahara 1982).
*** Metzgeria mexicana Steph., Sp. Hepat. (Stephani) 6: 55, 1917 (Stephani 1917a).
413
** Metzgeria monoica Kuwah. et J.J.Engel, Hikobia 8 (3/4): 284, 1980 (Kuwahara
1980a).
*** Metzgeria myriopoda Lindb., Acta Soc. Fauna Fl. Fenn. 1 (2): 22, 1877 (Lindberg
1877b).
*** Metzgeria neotropica Kuwah., Nova Hedwigia 34: 779, 1981 (Kuwahara 1981).
*** Metzgeria nudifrons Steph., Hedwigia 31 (3): 126, 1892 (Stephani 1892d).
*** Metzgeria parviinvolucrata Kuwah., Nova Hedwigia 34: 774, 1981 (Kuwahara
1981).
** Metzgeria patagonica Steph., Bull. Herb. Boissier 7 (12): 940 (288), 1899 (Stephani 1899g).
*** Metzgeria polytricha Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 553, 1885
(Spruce 1885).
*** Metzgeria procera Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 166, 1855 (Mitten
1855).
*** Metzgeria psilocraspeda Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl., Denkschr. 111: 25, 1964 (Schifner and Arnell 1964).
*** Metzgeria pubescens (Schrank) Raddi, Jungermanniogr. Etrusca: 35, 1818 (Raddi 1818a). Bas.: Jungermannia pubescens Schrank, Prim. Fl. Salisb.: 231, 1792
(Schrank 1792).
*** Metzgeria pulvinata Steph., Biblioth. Bot. 87 (2): 179, 1916 (Stephani 1916a).
** Metzgeria quadrifaria Steph., Bull. Herb. Boissier 7 (12): 953 (301), 1899 (Stephani 1899g).
** Metzgeria raoi S.C.Srivast. et S.Srivast., Phytotaxonomy 4: 83, 2004 (Srivastava
and Srivastava 2004).
** Metzgeria rigida Lindb., Acta Soc. Fauna Fl. Fenn. 1 (2): 43, 1877 (Lindberg
1877b).
** Metzgeria robinsonii Steph., Sp. Hepat. (Stephani) 6: 60, 1917 (Stephani 1917a).
413 Metzgeria mexicana is conspeciic with Metzgeria subundulata in Schuster (1992b), but it was accepted
by Costa (2008).
World checklist of hornworts and liverworts
461
** Metzgeria roivainenii Kuwah., J. Hattori Bot. Lab. 40: 516, 1976 (Kuwahara
1976c).
*** Metzgeria rufula Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 555, 1885
(Spruce 1885).
** Metzgeria saccata Mitt., J. Linn. Soc., Bot. 22 (145): 241, 1886 (Mitten 1886a).
** Metzgeria saxbyi Pearson, Ann. Cryptog. Exot. 4 (2): 70, 1931 (Pearson 1931a).
** Metzgeria scobina Mitt., J. Linn. Soc., Bot. 22 (145): 243, 1886 (Mitten 1886a).
*** Metzgeria scyphigera A.Evans, Trans. Connecticut Acad. Arts 18 (5): 299, 1914
(Evans 1914c).
*** Metzgeria senjoana Masuzaki, Hikobia 15 (4): 445, 2010 (Masuzaki et al. 2010a).
** Metzgeria setigera R.M.Schust. ex Crand.-Stotl. et L.Söderstr., Phytotaxa 202
(1): 69, 2015 (Söderström et al. 2015c). Based on: Metzgeria furcata var. setigera
R.M.Schust., J. Hattori Bot. Lab. 70: 149, 1991 (Schuster 1991b), nom. inval.
** Metzgeria sikkimensis S.C.Srivast. et K.K.Rawat, Geophytology 31 (1/2): 71, 2001
[2003] (Srivastava and Rawat 2001).
* Metzgeria simplex Lorb. ex Müll.Frib., Hedwigia 80 (1/2): 115, 1941 (Müller
1941).
*** Metzgeria sinuata Loitl., Diagn. pl. nov.: 25, 1894 (Loitlesberger 1894).
** Metzgeria sparrei Kuwah., Hikobia 8 (3/4): 278, 1980 (Kuwahara 1980a).
*** Metzgeria spindleri Steph., Biblioth. Bot. 87 (2): 179, 1916 (Stephani 1916a).
*** Metzgeria subaneura Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl., Denkschr.
111: 22, 1964 (Schifner and Arnell 1964).
** Metzgeria submarginata M.L.So, New Zealand J. Bot. 40 (2): 201, 2002 (So 2002b).
** Metzgeria subundulata (Lindb.) Kuwah., Bryologist 86 (3): 276, 1983 [1984] (Kuwahara 1983). Bas.: Metzgeria furcata subsp. subundulata Lindb., Acta Soc. Fauna Fl.
Fenn. 1 (2): 42, 1877 (Lindberg 1877b).
** Metzgeria temperata Kuwah., J. Hattori Bot. Lab. 40: 219, 1976 (Kuwahara 1976b).
*** Metzgeria uncigera A.Evans, Ann. Bot. (Oxford) 24 (2): 276, 1910 (Evans 1910).
*** Metzgeria undulata Kuwah., Nova Hedwigia 34: 792, 1981 (Kuwahara 1981).
*** Metzgeria violacea (Ach.) Dumort., Recueil Observ. Jungerm.: 26, 1835 (Dumortier 1835). Bas.: Jungermannia violacea Ach., Beitr. Naturk. (Weber & Mohr) 1:
77, 1805 (Acharius 1805).
** Steereella Kuwah., Amer. J. Bot. 60 (6): 602, 1973 (Kuwahara 1973a).
*** Steereella lilliana (Steph.) Kuwah., Bryophyt. Biblioth. 28: 179, 1986 (Kuwahara
1986). Bas.: Metzgeria lilliana Steph., Sp. Hepat. (Stephani) 6: 53, 1917 (Stephani
1917a).
*** Steereella linearis (Sw.) Kuwah., Amer. J. Bot. 60 (6): 604, 1973 (Kuwahara 1973a).
Bas.: Jungermannia linearis Sw., Prodr. (Swartz): 145, 1788 (Swartz 1788).
** Vandiemenia Hewson, J. Hattori Bot. Lab. 52: 163, 1982 (Hewson 1982).
** Vandiemenia ratkowskiana Hewson, J. Hattori Bot. Lab. 52: 163, 1982 (Hewson
1982).
462
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Pleuroziales Schljakov
***
Pleuroziaceae Müll.Frib.
by B. hiers
*** Pleurozia Dumort., Recueil Observ. Jungerm.: 15, 1835 (Dumortier 1835).
** subg. Constantifolia B.M.hiers, Bryologist 96 (4): 526, 1993 (hiers 1993).
*** Pleurozia conchifolia (Hook. et Arn.) Austin, Bull. Torrey Bot. Club 5 (3): 16,
1874 (Austin 1874). Bas.: Jungermannia conchifolia Hook. et Arn., Bot. Beechey
Voy. 3: 110, 1832 (Hooker and Arnott 1832).
** Pleurozia conchifolia var. papillosa B.M.hiers, Bryologist 96 (4): 528, 1993 (hiers
1993).
*** Pleurozia purpurea Lindb., Acta Soc. Fauna Fl. Fenn. 1 (2): 27, 1877 (Lindberg
1877b). Based on: Jungermannia purpurea Lightf., Fl. Scot. 2: 778, 1777 (Lightfoot 1777), nom. illeg.
** subg. Diversifolia B.M.hiers, Bryologist 96 (4): 531, 1993 (hiers 1993).
*** Pleurozia acinosa (Mitt.) Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci.
Mat. 4 (13): 412, 1877 (Trevisan 1877). Bas.: Physiotium acinosum Mitt., J. Proc.
Linn. Soc., Bot. 5 (18): 102, 1860 [1861] (Mitten 1860c).
*** Pleurozia articulata (Lindb.) Lindb. et Lackström, Hepat. Scand. Exsicc.: 5 (adnot.), 1874 (Lindberg and Lackström 1874). Bas.: Physiotium articulatum Lindb.,
Öfvers. Förh. Finska Vetensk.-Soc. 12 (2): 78, 1870 (Lindberg 1870).
*** Pleurozia caledonica (Gottsche) Steph., Rev. Bryol. 33 (2): 29 (Paris 1906a). Bas.:
Physiotium caledonicum Gottsche, Hedwigia 25 (2/3): 81, 1886 (Jack 1886).
*** Pleurozia curiosa B.M.hiers, Bryologist 96 (4): 537, 1993 (hiers 1993).
*** Pleurozia heterophylla Steph. ex Fulford, Mem. New York Bot. Gard. 23: 842,
1972 (Fulford 1972).
*** Pleurozia johannis-winkleri Herzog, Mitt. Inst. Allg. Bot. Hamburg 7 (3): 195,
1931 (Herzog 1931a).
*** Pleurozia paradoxa (J.B.Jack) Schifn., Hepat. (Engl.-Prantl): 115, 1893 (Schifner 1893b). Bas.: Physiotium paradoxum J.B.Jack, Hedwigia 25 (2/3): 85, 1886
(Jack 1886).
*** Pleurozia subinlata (Austin) Austin, Bull. Torrey Bot. Club 5 (3): 16, 1874 (Austin
1874). Bas.: Physiotium subinlatum Austin, Proc. Acad. Nat. Sci. Philadelphia 21:
224, 1869 (Austin 1869).
** subg. Pleurozia
*** Pleurozia gigantea (F.Weber) Lindb., Hepat. Scand. Exsicc.: no. 5, 1874 (Lindberg
and Lackström 1874). Bas.: Jungermannia gigantea F.Weber, Hist. Musc. Hepat.
Prodr.: 57, 1815 (Weber 1815).
World checklist of hornworts and liverworts
463
Incertae sedis
** Pleurozia pocsii Frank Müll., Polish Bot. J. 58 (1): 50, 2013 (Müller 2013).
Pelliidae He-Nygrén, Juslén, Ahonen, Glenny et Piippo
Fossombroniales Schljakov
Calyculariineae He-Nygrén, Juslén, Ahonen, Glenny et Piippo
***
Calyculariaceae He-Nygrén, Juslén, Ahonen, Glenny et Piippo
*** Calycularia Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 122, 1860 [1861] (Mitten 1860c).
*** Calycularia crispula Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 122, 1860 [1861] (Mitten 1860c).
*** Calycularia laxa Lindb. et Arnell, Kongl. Svenska Vetensk.-Akad. Handl. (n.ser.)
23 (5): 66, 1889 (Lindberg and Arnell 1889).
Fossombroniineae R.M.Schust. ex Stotler et Crand.-Stotl.
**
Allisoniaceae Schljakov
*** Allisonia Herzog, Hedwigia 80 (1/2): 77, 1941 (Herzog 1941a).
*** Allisonia cockaynei (K.I.Goebel) R.M.Schust., J. Hattori Bot. Lab. 26: 294, 1963 (Schuster
1963b). Bas.: Moerckia cockaynei K.I.Goebel, Flora 96: 190, 1906 (Goebel 1906).
***
Fossombroniaceae Hazsl. nom. conserv.
by R. Stotler, B. J. Crandall-Stotler and D. C. Cargill
*** Fossombronia Raddi, Jungermanniogr. Etrusca: 29, 1818 (Raddi 1818a).
*** Fossombronia alaskana Steere et Inoue, Bryologist 77 (1): 66, 1974 (Steere and
Inoue 1974).
*** Fossombronia alata G.A.M.Scott et D.C.Pike, J. Hattori Bot. Lab. 56: 340, 1984
(Scott and Pike 1984).
*** Fossombronia altilamellosa G.A.M.Scott et D.C.Pike, Fl. Austral. Suppl. 21:
114, 2003 (McCarthy 2003). Based on: Fossombronia altilamellosa G.A.M.Scott
et D.C.Pike, J. Hattori Bot. Lab. 62: 367, 1987 (Scott and Pike 1987), nom.
inval.
*** Fossombronia angulifolia Perold, Bothalia 28 (2): 159, 1998 (Perold 1998b).
*** Fossombronia angulosa (Dicks.) Raddi, Jungermanniogr. Etrusca: 29, 1818 (Raddi
1818a). Bas.: Jungermannia angulosa Dicks., Fasc. Pl. Crypt. Brit. 1: 7, 1785
(Dickson 1785).
464
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Fossombronia areolata G.A.M.Scott et D.C.Pike, J. Hattori Bot. Lab. 62: 368,
1987 (Scott and Pike 1987).
** Fossombronia auricolor G.A.M.Scott et D.C.Pike, Fl. Austral. Suppl. 21: 114, 2003
(McCarthy 2003). Based on: Fossombronia auricolor G.A.M.Scott et D.C.Pike, J.
Hattori Bot. Lab. 62: 370, 1987 (Scott and Pike 1987), nom. inval.
*** Fossombronia australis Mitt., J. Linn. Soc., Bot. 15 (82): 73, 1876 (Mitten 1876a).
*** Fossombronia caespitiformis (Raddi) De Not. ex Rabenh., Hepat. Eur., Leberm.
13-14: no. 123, 1860 [1861] (Rabenhorst 1860). Bas.: Fossombronia angulosa var.
caespitiformis Raddi, Jungermanniogr. Etrusca: 30, 1818 (Raddi 1818a).
*** Fossombronia caespitiformis subsp. multispira (Schifn.) J.R.Bray et Cargill, Bryologist 106 (1): 131, 2003 (Stotler et al. 2003). Bas.: Fossombronia caespitiformis var.
multispira Schifn., Österr. Bot. Z. 67 (4/5): 152, 1918 (Schifner 1918).
*** Fossombronia caledonica Steph., Sp. Hepat. (Stephani) 6: 71, 1917 (Stephani 1917a).
*** Fossombronia cederbergensis Perold, Bothalia 28 (1): 1, 1998 (Perold 1998c).
*** Fossombronia cerebriformis G.A.M.Scott et D.C.Pike, J. Hattori Bot. Lab. 56: 343,
1984 (Scott and Pike 1984).
*** Fossombronia crassifolia Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 527,
1885 (Spruce 1885).
*** Fossombronia crispa Nees, Syn. Hepat. 4: 469, 1846 (Gottsche et al. 1846).
* Fossombronia crispula (Brot.) R.M.Schust., Hepat. Anthocerotae N. Amer. 5: 383,
1992 (Schuster 1992b). Bas.: Jungermannia crispula Brot., Fl. lusit. 2: 422, 1804
[1805] (Brotero 1804). 414
*** Fossombronia cristula Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 228, 1869
(Austin 1869).
*** Fossombronia cultriformis G.A.M.Scott et D.C.Pike, J. Hattori Bot. Lab. 62: 371,
1987 (Scott and Pike 1987).
*** Fossombronia densa G.A.M.Scott et D.C.Pike, J. Hattori Bot. Lab. 62: 372, 1987
(Scott and Pike 1987).
*** Fossombronia densilamellata S.W.Arnell, Bot. Not. 105: 317, 1952 (Arnell 1952a).
*** Fossombronia echinata Macvicar, Rev. Bryol. 38 (4): 73, 1911 (Macvicar 1911).
*** Fossombronia elsieae Perold, Bothalia 29 (1): 25, 1999 (Perold 1999c).
*** Fossombronia fernandeziensis Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.
ser.) 46 (9): 15, 1911 (Stephani 1911b).
*** Fossombronia imbriata Paton, J. Bryol. 8 (1): 1, 1974 (Paton 1974).
*** Fossombronia leischeri Osterwald ex Loeske, Verh. Bot. Vereins Prov. Brandenburg 70: 125, 1928 (Loeske 1928).
*** Fossombronia foveolata Lindb., Helsingf. Dagbl. 1873 (353, 28 Dec): 2, 1873
(Lindberg 1873b).
** Fossombronia fuhreri G.A.M.Scott et D.C.Pike, J. Hattori Bot. Lab. 62: 374, 1987
(Scott and Pike 1987).
414 Fossombronia crispula is based on the description most likely Fossombronia pusilla, but no material has
been found.
World checklist of hornworts and liverworts
465
*** Fossombronia gemmifera Perold, Bothalia 27 (1): 19, 1997 (Perold 1997a).
*** Fossombronia glenii Perold, Bothalia 27 (1): 20, 1997 (Perold 1997a).
** Fossombronia grandis Steph., Mém. Herb. Boissier 16: 29 (383), 1900 (Stephani
1900a).
* Fossombronia gregaria Colenso, Trans. & Proc. New Zealand Inst. 20: 252, 1888
(Colenso 1888). 415
* Fossombronia grossepapillata Steph., J. & Proc. Roy. Soc. New South Wales 48
(1/2): 105, 1914 (Stephani and Watts 1914). 416
*** Fossombronia hamatohirta Steph., Hedwigia 33 (1): 8, 1894 (Stephani 1894a).
** Fossombronia hewsoniae G.A.M.Scott et D.C.Pike, J. Hattori Bot. Lab. 62: 375,
1987 (Scott and Pike 1987).
*** Fossombronia himalayensis Kashyap, New Phytol. 14 (1): 4, 1915 (Kashyap 1915).
*** Fossombronia hyalorhiza Perold, Bothalia 29 (1): 83, 1999 (Perold 1999d).
*** Fossombronia incurva Lindb., Helsingf. Dagbl. 1873 (273, 7 Oct): 2, 1873 (Lindberg 1873c).
*** Fossombronia indica Steph., Sp. Hepat. (Stephani) 6: 73, 1917 (Stephani 1917a).
* Fossombronia integerrima Steph., Mém. Herb. Boissier 16: 40 (394), 1900 (Stephani
1900a). 417
* Fossombronia integrifolia Steph., Sp. Hepat. (Stephani) 6: 73, 1917 (Stephani 1917a). 418
*** Fossombronia intestinalis Taylor, London J. Bot. 5: 408, 1846 (Taylor 1846b).
*** Fossombronia japonica Schifn., Österr. Bot. Z. 49 (11): 389, 1899 (Schifner
1899c).
** Fossombronia laciniata G.A.M.Scott et D.C.Pike, J. Hattori Bot. Lab. 62: 375,
1987 (Scott and Pike 1987).
*** Fossombronia lamellata Steph., Hedwigia 33 (1): 9, 1894 (Stephani 1894a).
*** Fossombronia leucoxantha (Lehm.) Lehm. et Lindenb., Syn. Hepat. 4: 469, 1846
(Gottsche et al. 1846). Bas.: Jungermannia leucoxantha Lehm., Linnaea 4: 368,
1829 (Lehmann 1829).
*** Fossombronia longiseta (Austin) Austin, Hepat. bor.-amer.: 30, 1873 (Austin
1873). Bas.: Androcryphia longiseta Austin, Proc. Acad. Nat. Sci. Philadelphia 21:
228, 1869 (Austin 1869).
*** Fossombronia lophoclada Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 529,
1885 (Spruce 1885).
** Fossombronia lophoscypha Hässel, Beih. Nova Hedwigia 131: 16, 2007 (Hässel and
Villagrán 2007).
*** Fossombronia luetzelburgiana K.I.Goebel, Flora 105: 55, 1912 (Goebel 1912).
*** Fossombronia macrocalyx Steph., Sp. Hepat. (Stephani) 6: 74, 1917 (Stephani 1917a).
415 Fossombronia gregaria is possibly conspeciic with Fossombronia australis (Scott and Pike 1988b).
416 Fossombronia grossepapillata may be conspeciic with Fossombronia papillata, but the type specimen is
too immature to allow determination (Scott and Pike 1984).
417 Fossombronia integerrima is possibly conspeciic with Fossombronia australis.
418 Fossombronia integrifolia is possibly conspeciic with Fossombronia foveolata (Scott and Pike 1984).
466
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Fossombronia macrophylla Colenso, Trans. & Proc. New Zealand Inst. 18: 285,
1886 (Colenso 1886a).
Fossombronia magnaspora G.A.M.Scott et D.C.Pike, Fl. Austral. Suppl. 21: 114,
2003 (McCarthy 2003). Based on: Fossombronia magnaspora G.A.M.Scott et
D.C.Pike, J. Hattori Bot. Lab. 62: 377, 1987 (Scott and Pike 1987), nom. inval.
Fossombronia marindae Perold, Bothalia 29 (1): 86, 1999 (Perold 1999d).
Fossombronia maritima (Paton) Paton, J. Bryol. 18 (2): 367, 1994 (Paton 1994).
Bas.: Fossombronia pusilla var. maritima Paton, J. Bryol. 7 (3): 244, 1973 (Paton
1973).
Fossombronia marshii J.R.Bray et Stotler, Phytologia 92 (2): 230, 2010 (Stotler et
al. 2010).
Fossombronia microlamellata G.A.M.Scott et D.C.Pike, J. Hattori Bot. Lab. 56:
347, 1984 (Scott and Pike 1984).
Fossombronia mittenii Tind., J. Bot. 36: 44, 1898 (Tindall 1898).
Fossombronia montaguensis S.W.Arnell, Bot. Not. 105: 316, 1952 (Arnell 1952a).
Fossombronia monticola Perold, Bothalia 29 (1): 87, 1999 (Perold 1999d).
Fossombronia mylioides Inoue, J. Hattori Bot. Lab. 37: 296, 1973 (Inoue 1973).
Fossombronia nigricaulis Colenso, Trans. & Proc. New Zealand Inst. 18: 248,
1886 (Colenso 1886b). 419
Fossombronia nyikaensis Perold, Bothalia 31 (1): 48, 2001 (Perold 2001c).
Fossombronia papillata Steph., Hedwigia 28 (3): 157, 1889 (Stephani 1889d).
Fossombronia paranapanemae Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl.,
Denkschr. 111: 34, 1964 (Schifner and Arnell 1964).
Fossombronia peruviana Gottsche et Hampe, Linnaea 27 (5): 555, 1854 (Hampe
1854).
Fossombronia porphyrorhiza (Nees) Prosk., Bryologist 58 (3): 197, 1955 (Proskauer 1955). Bas.: Jungermannia porphyrorhiza Nees, Fl. Bras. (Martius) 1 (1): 343,
1833 (Nees 1833a).
Fossombronia pulvinata Steph., Wiss. Ergebn. Deut. Zentr.-Afr. Exped. (190708), Bot. 2: 113, 1911 (Stephani 1911a).
Fossombronia punctata G.A.M.Scott et D.C.Pike, J. Hattori Bot. Lab. 56: 341,
1984 (Scott and Pike 1984).
Fossombronia purpureospora G.A.M.Scott et D.C.Pike, Fl. Austral. Suppl. 21: 114,
2003 (McCarthy 2003). Based on: Fossombronia purpureospora G.A.M.Scott et
D.C.Pike, J. Hattori Bot. Lab. 62: 379, 1987 (Scott and Pike 1987), nom. inval.
Fossombronia pusilla (L.) Nees, Naturgesch. Eur. Leberm. 3: 319, 1838 (Nees
1838b). Bas.: Jungermannia pusilla L., Sp. Pl. 1: 1136, 1753 (Linnaeus 1753).
Fossombronia renateae Perold, Bothalia 29 (1): 77, 1999 (Perold 1999b).
Fossombronia reticulata Steph., Hedwigia 33 (1): 9, 1894 (Stephani 1894a).
419 Fossombronia nigricaulis is possibly conspeciic with Fossombronia australis (Scott and Pike 1984).
World checklist of hornworts and liverworts
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467
Fossombronia rosulata Colenso, Trans. & Proc. New Zealand Inst. 18: 248, 1886
(Colenso 1886b).
Fossombronia rudis G.A.M.Scott et D.C.Pike, Beih. Nova Hedwigia 90: 110, 1988
(Scott and Pike 1988a).
Fossombronia ruminata Cargill, Phytotaxa 65: 45, 2012 (Cargill et al. 2012). Nom.
nov. pro Fossombronia maritima G.A.M.Scott et D.C.Pike, Fl. Austral. Suppl. 21:
114, 2003 (McCarthy 2003), nom. illeg.
Fossombronia rupestris G.A.M.Scott et D.C.Pike, Fl. Austral. Suppl. 21: 114, 2003
(McCarthy 2003). Based on: Fossombronia rupestris G.A.M.Scott et D.C.Pike, J.
Hattori Bot. Lab. 62: 381, 1987 (Scott and Pike 1987), nom. inval.
Fossombronia rwandaensis Perold, Bothalia 28 (1): 45, 1998 (Perold 1998a).
Fossombronia scrobiculata G.A.M.Scott et D.C.Pike, J. Hattori Bot. Lab. 56: 341,
1984 (Scott and Pike 1984).
Fossombronia spinifolia Steph., Mém. Herb. Boissier 16: 35 (389), 1900 (Stephani
1900a).
Fossombronia spinosa Perold, Bothalia 29 (1): 29, 1999 (Perold 1999c).
Fossombronia stephanii Schifn. ex Steph., Mém. Herb. Boissier 16: 27 (381), 1900
(Stephani 1900a).
Fossombronia straussiana Perold, Bothalia 27 (1): 24, 1997 (Perold 1997a).
Fossombronia subsaccata Steph., Sp. Hepat. (Stephani) 6: 75, 1917 (Stephani
1917a). 420
Fossombronia swaziensis Perold, Bothalia 28 (2): 162, 1998 (Perold 1998b).
Fossombronia tesselata G.A.M.Scott et D.C.Pike, J. Hattori Bot. Lab. 62: 382,
1987 (Scott and Pike 1987).
Fossombronia texana Lindb., Acta Soc. Sci. Fenn. 10: 533, 1875 (Lindberg 1875).
Fossombronia truncata G.A.M.Scott et D.C.Pike, Fl. Austral. Suppl. 21: 114, 2003
(McCarthy 2003). Based on: Fossombronia truncata G.A.M.Scott et D.C.Pike, J.
Hattori Bot. Lab. 62: 383, 1987 (Scott and Pike 1987), nom. inval.
Fossombronia tumida Mitt., J. Linn. Soc., Bot. 16 (91): 193, 1877 (Mitten 1877).
Fossombronia valparaisiana Hässel, Beih. Nova Hedwigia 131: 14, 2007 (Hässel
and Villagrán 2007).
Fossombronia vermiculata G.A.M.Scott et D.C.Pike, J. Hattori Bot. Lab. 56: 345,
1984 (Scott and Pike 1984).
Fossombronia wattsii Steph., Sp. Hepat. (Stephani) 6: 75, 1917 (Stephani 1917a).
Fossombronia wondraczekii (Corda) Dumort. ex Lindb., Helsingf. Dagbl. 1873
(273, 7 Oct): 2, 1873 (Lindberg 1873c). Bas.: Jungermannia wondraczekii Corda,
Deutschl. Fl. (Sturm), Abt. 2, Cryptog.: 30, 1830 (Corda 1830).
Fossombronia wrightii Austin, Bot. Bull. (Hanover) 1 (8): 36, 1876 (Austin 1876a).
Fossombronia zuurbergensis Perold, Bothalia 31 (1): 25, 2001 (Perold 2001b).
420 Fossombronia subsaccata has a sterile and unidentiiable type specimen (Scott and Pike 1988b).
468
***
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Petalophyllaceae Stotler et Crand.-Stotl.
by R. Stotler and B.J. Crandall-Stotler
*** Petalophyllum Nees et Gottsche, Nov. Stirp. Pug. 8: 29, 1844 (Lehmann 1844).
*** Petalophyllum americanum C.H.Ford et Crand.-Stotl., Novon 12 (3): 335, 2002
(Crandall-Stotler et al. 2002).
*** Petalophyllum hodgsoniae C.H.Ford et Crand.-Stotl., Novon 12 (3): 336, 2002
(Crandall-Stotler et al. 2002).
*** Petalophyllum indicum Kashyap, J. Indian Bot. Soc. 7: 14, 1928 (Kashyap 1928).
*** Petalophyllum preissii Gottsche, Nov. Stirp. Pug. 8: 30, 1844 (Lehmann 1844).
*** Petalophyllum ralfsii (Wilson) Nees et Gottsche, Nov. Stirp. Pug. 8: 30, 1844
(Lehmann 1844). Bas.: Jungermannia ralfsii Wilson, Suppl. Engl. Bot. 4: tab.
2874, 1849 (Borrer et al. 1849).
*** Sewardiella Kashyap, New Phytol. 14 (1): 5, 1915 (Kashyap 1915).
*** Sewardiella tuberifera Kashyap, New Phytol. 14 (1): 5, 1915 (Kashyap 1915).
Makinoiineae He-Nygrén, Juslén, Ahonen, Glenny et Piippo
***
Makinoaceae Nakai
by R. Stotler and B.J. Crandall-Stotler
*** Makinoa Miyake, Bot. Mag. (Tokyo) 13 (144): 23, 1899 (Miyake 1899).
*** Makinoa crispata (Steph.) Miyake, Bot. Mag. (Tokyo) 13 (144): 23, 1899 (Miyake 1899).
Bas.: Pellia crispata Steph., Bull. Herb. Boissier 5 (2): 103, 1897 (Stephani 1897b).
Pallaviciniales W.Frey et M.Stech
Pallaviciniineae R.M.Schust.
***
Hymenophytaceae R.M.Schust.
by R. Stotler and B.J. Crandall-Stotler
*** Hymenophyton Dumort., Recueil Observ. Jungerm.: 25, 1835 (Dumortier 1835).
*** Hymenophyton labellatum (Labill.) Dumort. ex Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci. Mat. 4 (13): 430, 1877 (Trevisan 1877). Bas.: Jungermannia labellata Labill., Nov. Holl. Pl. 2: 109, 1806 (Labillardière 1806).
*** Hymenophyton leptopodum (Hook.f. et Taylor) A.Evans, Trans. Connecticut Acad.
Arts 8 (16): 274, 1893 (Evans 1893). Bas.: Jungermannia leptopoda Hook.f. et
Taylor, London J. Bot. 3: 571, 1844 (Hooker and Taylor 1844d).
World checklist of hornworts and liverworts
469
*** Hymenophyton pedicellatum Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.
ser.) 46 (9): 11, 1911 (Stephani 1911b).
***
Moerckiaceae K.I.Goebel ex Stotler et Crand.-Stotl.
by B.J. Crandall-Stotler and R. Stotler
Mamontov et al. (2015) showed that the family Moerckiaceae is heterogeneous and
they moved Hattorianthus and Moerckia lotoviana to their new family Cordaeaceae.
However, the type of Moerckiaceae (Moerckia hibernica) was not included in the study
and only provisionally left in Moerckia. We prefer to follow Crandall-Stotler & Stotler
(2007) until M. hibernica is further studied.
*** Hattorianthus R.M.Schust. et Inoue, Bull. Natl. Sci. Mus. Tokyo, B 1 (3): 103,
1975 (Schuster and Inoue 1975).
*** Hattorianthus erimonus (Steph.) R.M.Schust. et Inoue, Bull. Natl. Sci. Mus. Tokyo, B 1 (3): 106, 1975 (Schuster and Inoue 1975). Bas.: Pallavicinia erimona
Steph., Bull. Herb. Boissier 5 (2): 102, 1897 (Stephani 1897b).
*** Moerckia Gottsche, Hepat. Eur., Leberm. 13-14: no. 121, 1860 [1861] (Rabenhorst 1860).
*** Moerckia blyttii (Mørch) Brockm., Arch. Vereins Freunde Naturgesch. Mecklenburg 17: 190, 1863 (Brockmüller 1863). Bas.: Jungermannia blyttii Mørch, Fl.
Danica 12: 6, 1830 (Hornemann 1830).
*** Moerckia lotoviana (Nees) Schifn., Österr. Bot. Z. 51 (2): 43, 1901 (Schifner
1901). Bas.: Cordaea lotoviana Nees, Flora 16 (26): 405, 1833 (Nees 1833b).
*** Moerckia hibernica (Hook.) Gottsche, Hepat. Eur., Leberm. 13-14: no. 121, 1860
[1861] (Rabenhorst 1860). Bas.: Jungermannia hibernica Hook., Brit. Jungermann.: tab. 78, 1816 (Hooker 1816a).
***
Pallaviciniaceae Mig.
by R. Stotler and B.J. Crandall-Stotler
**
Pallavicinioideae Mig. ex Grolle
*** Jensenia Lindb., Not. Sällsk. Fauna Fl. Fenn. Förh. 9: 13, 1868 (Lindberg 1868a).
*** Jensenia connivens (Colenso) Grolle, Rev. Bryol. Lichénol. 33 (1/2): 228, 1964
[1965] (Grolle 1964j). Bas.: Symphyogyna connivens Colenso, Trans. & Proc. New
Zealand Inst. 20: 254, 1888 (Colenso 1888).
470
*
***
***
***
***
***
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Jensenia crassifrons (Steph.) S.Schuette et Stotler, J. Hattori Bot. Lab. 97: 300,
2005 (Schuette and Stotler 2005). Bas.: Pallavicinia crassifrons Steph., Mém.
Herb. Boissier 11: 21 (325), 1900 (Stephani 1900c). 421
Jensenia decipiens (Mitt.) Grolle, Rev. Bryol. Lichénol. 33 (1/2): 228, 1964 [1965]
(Grolle 1964j). Bas.: Steetzia decipiens Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 123,
1860 [1861] (Mitten 1860c).
Jensenia diformis (Nees) Grolle, Rev. Bryol. Lichénol. 33 (1/2): 228, 1964 [1965]
(Grolle 1964j). Bas.: Jungermannia diformis Nees, Fl. Bras. (Martius) 1 (1): 329,
1833 (Nees 1833a).
Jensenia lorschuetzii Gronde, Proc. Kon. Ned. Akad. Wetensch. C 83 (3): 273,
1980 (Van der Gronde 1980).
Jensenia spinosa (Lindenb. et Gottsche) Grolle, Acta Bot. Fenn. 133: 65, 1986
(Grolle and Piippo 1986). Bas.: Symphyogyna spinosa Lindenb. et Gottsche, Syn.
Hepat. 5: 786, 1847 (Gottsche et al. 1847).
Jensenia wallisii (J.B.Jack et Steph.) Grolle, Rev. Bryol. Lichénol. 33 (1/2): 228,
1964 [1965] (Grolle 1964j). Bas.: Pallavicinia wallisii J.B.Jack et Steph., Hedwigia
31 (1): 23, 1892 (Jack and Stephani 1892).
*** Pallavicinia Gray, Nat. Arr. Brit. Pl. 1: 775, 1821 (Gray 1821) nom. conserv.
*** Pallavicinia ambigua (Mitt.) Steph., Mém. Herb. Boissier 11: 8 (312), 1900 (Stephani 1900a). Bas.: Steetzia ambigua Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 123,
1860 [1861] (Mitten 1860c).
** Pallavicinia baldwinii (Austin) A.Evans, Trans. Connecticut Acad. Arts 8 (15):
259, 1891 (Evans 1891). Bas.: Steetzia baldwinii Austin, Bull. Torrey Bot. Club 6
(52): 303, 1879 (Austin 1879).
* Pallavicinia bipinnata Steph., Sp. Hepat. (Stephani) 6: 62, 1917 (Stephani 1917a).
** Pallavicinia camisassai Gola, Mem. Reale Accad. Sci. Torino (ser. 2) 65 (1): 4,
1916 (Gola 1916).
** Pallavicinia cylindrica (Austin) A.Evans, Trans. Connecticut Acad. Arts 8 (15):
259, 1891 (Evans 1891). Bas.: Steetzia cylindrica Austin, Bull. Torrey Bot. Club 5
(3): 17, 1874 (Austin 1874).
** Pallavicinia himalayensis Schifn., Mém. Herb. Boissier 11: 13 (317), 1900
(Stephani 1900c).
*** Pallavicinia indica Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl.
67: 183, 1898 (Schifner 1898a).
*** Pallavicinia levieri Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl.
67: 184, 1898 (Schifner 1898a).
*** Pallavicinia lyellii (Hook.) Gray, Nat. Arr. Brit. Pl. 1: 775, 1821 (Gray 1821). Bas.:
Jungermannia lyellii Hook., Brit. Jungermann.: tab. 77, 1816 (Hooker 1816a).
** Pallavicinia pilifera Steph., Hedwigia 30 (6): 271, 1891 (Stephani 1891c).
** Pallavicinia purpurea Steph., Sp. Hepat. (Stephani) 6: 64, 1917 (Stephani 1917a).
421 Jensenia crassifrons is possibly conspeciic with Jensenia diformis.
World checklist of hornworts and liverworts
471
** Pallavicinia ridleyi Steph., Sp. Hepat. (Stephani) 6: 64, 1917 (Stephani 1917a).
*** Pallavicinia rubristipa Schifn., Österr. Bot. Z. 56 (1): 24, 1906 (Schifner 1906b).
*** Pallavicinia subciliata (Austin) Steph., Mém. Herb. Boissier 11: 9 (313), 1900
(Stephani 1900c). Bas.: Steetzia subciliata Austin, Bull. Torrey Bot. Club 6 (52):
303, 1879 (Austin 1879).
*** Pallavicinia xiphoides (Hook.f. et Taylor) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 427, 1877 (Trevisan 1877). Bas.: Jungermannia xiphoides
Hook.f. et Taylor, London J. Bot. 3: 569, 1844 (Hooker and Taylor 1844d).
*** Podomitrium Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 164, 1855 (Mitten 1855).
*** Podomitrium malaccense (Steph.) Campb., Amer. J. Bot. 2 (5): 199, 1915 (Campbell 1915). Bas.: Hymenophyton malaccense Steph., Hedwigia 34 (2): 46, 1895
(Stephani 1895c).
*** Podomitrium marginatum (Steph.) Hürl., Bauhinia 4 (1): 78, 1968 [1969] (Hürlimann 1968). Bas.: Hymenophyton marginatum Steph., Sp. Hepat. (Stephani) 6:
61, 1917 (Stephani 1917a).
*** Podomitrium phyllanthus (Hook.) Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2):
164, 1855 (Mitten 1855). Bas.: Jungermannia phyllanthus Hook., Musci Exot. 1:
tab. 95, 1818 (Hooker 1818).
**
Symphyogynoideae R.M.Schust. ex Grolle
** Greeneothallus Hässel, J. Bryol. 11 (1): 115, 1980 (Hässel 1980).
*** Greeneothallus gemmiparus Hässel, J. Bryol. 11 (1): 115, 1980 (Hässel 1980).
*** Seppeltia Grolle, J. Hattori Bot. Lab. 60: 276, 1986 (Grolle and Seppelt 1986).
*** Seppeltia succuba Grolle, J. Hattori Bot. Lab. 60: 276, 1986 (Grolle and Seppelt 1986).
*** Symphyogyna Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 5: 66, 1836 (Nees and
Montagne 1836).
*** Symphyogyna apiculispina Steph., Biblioth. Bot. 87 (2): 180, 1916 (Stephani 1916a).
*** Symphyogyna aspera Steph. ex F.A.McCormick, Bot. Gaz. 58 (5): 403, 1914 (McCormick 1914).
** Symphyogyna atronervia Taylor, London J. Bot. 5: 409, 1846 (Taylor 1846b).
** Symphyogyna bogotensis Steph., Mém. Herb. Boissier 11: 42 (346), 1900 (Stephani
1900c). Based on: Symphyogyna hymenophyllum var. bogotensis Gottsche, Ann. Sci.
Nat. Bot. (sér. 5) 1: 181, 1864 (Gottsche 1864), nom. inval.
* Symphyogyna boliviensis Steph., Biblioth. Bot. 87 (2): 180, 1916 (Stephani 1916a).
*** Symphyogyna brasiliensis Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 5: 67, 1836
(Nees and Montagne 1836). Nom. nov. pro Jungermannia brasiliensis Nees, Enum.
Pl. Crypt. Javae: 11, 1830 (Nees 1830), nom. illeg.
** Symphyogyna brasiliensis var. angustior (Gottsche, Lindenb. et Nees) Gottsche, Ann.
Sci. Nat. Bot. (sér. 5) 1: 183, 1864 (Gottsche 1864). Bas.: Symphyogyna brasiliensis
472
**
***
***
***
**
***
***
**
**
**
**
**
***
**
***
***
***
**
*
***
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
β angustior Gottsche, Lindenb. et Nees, Syn. Hepat. 4: 484, 1846 (Gottsche et al.
1846).
Symphyogyna brasiliensis var. subsinuata Schifn., Österr. Akad. Wiss., Math.Naturwiss. Kl., Denkschr. 111: 32, 1964 (Schifner and Arnell 1964).
Symphyogyna brongniartii Mont., Ann. Sci. Nat. Bot. (sér. 2) 19: 265, 1843 (Montagne 1843).
Symphyogyna circinata Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 5: 69, 1836
(Nees and Montagne 1836).
Symphyogyna digitisquama Steph., Mém. Herb. Boissier 11: 31 (335), 1900
(Stephani 1900c).
Symphyogyna fuscovirens A.Evans, Trans. Connecticut Acad. Arts 28 (6): 320,
1927 (Evans 1927).
Symphyogyna hochstetteri Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 5: 68, 1836
(Nees and Montagne 1836).
Symphyogyna hymenophyllum (Hook.) Nees et Mont., Ann. Sci. Nat. Bot. (sér.
2) 5: 68, 1836 (Nees and Montagne 1836). Bas.: Jungermannia hymenophyllum
Hook., Musci Exot. 1: tab. 14, 1818 (Hooker 1818).
Symphyogyna hymenophyllum var. heterogenum Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 536, 1885 (Spruce 1885).
Symphyogyna ignambiensis Hürl., Bauhinia 4 (1): 79, 1968 [1969] (Hürlimann 1968).
Symphyogyna interrupta Carrington et Pearson, Proc. Linn. Soc. New South Wales
(ser. 2) 2 (4): 1053, 1888 (Carrington and Pearson 1888a).
Symphyogyna irregularis Steph., Mém. Herb. Boissier 11: 29 (333), 1900 (Stephani
1900c).
Symphyogyna lacerosquama Steph., Sp. Hepat. (Stephani) 6: 67, 1917 (Stephani
1917a).
Symphyogyna leptothelia Taylor, London J. Bot. 5: 408, 1846 (Taylor 1846b).
Symphyogyna lindmanii A.Evans, Trans. Connecticut Acad. Arts 28 (6): 316, 1927
(Evans 1927).
Symphyogyna luetzelburgii Herzog, Repert. Spec. Nov. Regni Veg. 21 (1/7): 22,
1925 (Herzog 1925a).
Symphyogyna marginata Steph., Mém. Herb. Boissier 11: 30 (334), 1900 (Stephani 1900c).
Symphyogyna mexicana Steph., Rev. Bryol. 36 (6): 140, 1909 (Stephani 1909b).
Symphyogyna multilora Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2):
133, 1914 (Stephani and Watts 1914).
Symphyogyna paucidens Steph., Kungl. Svenska Vetensk.-Akad. Handl. (n.ser.) 46
(9): 13, 1911 (Stephani 1911b). 422
Symphyogyna podophylla (hunb.) Nees et Mont., Flora 29 (9): 135, 1846 (Krauss
1846). Bas.: Jungermannia podophylla hunb., Prodr. Pl. Cap. 2: 174, 1800 (hunberg
1800).
422 Symphyogyna paucidens is a doubtful taxon. Hässel and Rubies (2009) could not ind the type specimen.
World checklist of hornworts and liverworts
*
***
**
***
***
**
**
***
***
***
***
*
***
**
473
Symphyogyna purpureolimbata E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot. 3
(4): 95, 1965 (Hodgson 1965).
Symphyogyna rectidens Grolle, Acta Bot. Fenn. 133: 68, 1986 (Grolle and Piippo 1986).
Symphyogyna rhodina (Hook.f. et Taylor) Gottsche, Lindenb. et Nees, Syn. Hepat.
4: 487, 1846 (Gottsche et al. 1846). Bas.: Jungermannia rhodina Hook.f. et Taylor, London J. Bot. 4: 93, 1845 (Hooker and Taylor 1845).
Symphyogyna rubescens Steph., Mém. Herb. Boissier 11: 29 (333), 1900 (Stephani
1900c).
Symphyogyna rubritincta A.Evans, Trans. Connecticut Acad. Arts 27 (1): 38, 1925
(Evans 1925).
Symphyogyna semi-involucrata Austin, Bull. Torrey Bot. Club 5 (3): 15, 1874 (Austin
1874).
Symphyogyna similis Grolle, Acta Bot. Fenn. 133: 70, 1986 (Grolle and Piippo 1986).
Symphyogyna sinuata (Sw.) Nees et Mont., Voy. Amér. Mérid., Bot. 7 (1): 61,
1839 (Montagne 1839b). Bas.: Jungermannia sinuata Sw., Prodr. (Swartz): 145,
1788 (Swartz 1788).
Symphyogyna subsimplex Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 166, 1855
(Mitten 1855).
Symphyogyna tenuinervis (Hook.f. et Taylor) Grolle, J. Hattori Bot. Lab. 61: 253,
1986 [1987] (Grolle 1986a). Bas.: Jungermannia tenuinervis Hook.f. et Taylor,
London J. Bot. 3: 570, 1844 (Hooker and Taylor 1844d).
Symphyogyna trivittata Spruce, J. Linn. Soc., Bot. 30 (210): 365, 1895 (Gepp 1895b).
Symphyogyna ulvoides (Reinw., Blume et Nees) Nees, Syn. Hepat. 4: 487, 1846
(Gottsche et al. 1846). Bas.: Jungermannia ulvoides Reinw., Blume et Nees, Nova
Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12 (1): 196, 1824 [1825]
(Reinwardt et al. 1824a). 423
Symphyogyna undulata Colenso, Trans. & Proc. New Zealand Inst. 16: 356, 1884
(Colenso 1884).
Symphyogyna volkensii Steph., Mém. Herb. Boissier 11: 35 (339), 1900 (Stephani
1900c).
** Symphyogynopsis Grolle, Acta Bot. Fenn. 133: 72, 1986 (Grolle and Piippo 1986).
*** Symphyogynopsis gottscheana (Mont. et Nees) Grolle, J. Hattori Bot. Lab. 63: 441,
1987 (Grolle 1987). Bas.: Symphyogyna gottscheana Mont. et Nees, Syn. Hepat. 4:
484, 1846 (Gottsche et al. 1846).
** Xenothallus R.M.Schust., J. Hattori Bot. Lab. 26: 293, 1963 (Schuster 1963b).
*** Xenothallus vulcanicola R.M.Schust., J. Hattori Bot. Lab. 26: 293, 1963 (Schuster
1963b).
423 Symphyogyna ulvoides is probably a ilmy fern (Schifner 1900a), but he did not see any type material.
474
**
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Sandeothallaceae R.M.Schust.
** Sandeothallus R.M.Schust., Nova Hedwigia 36: 10, 1982 (Schuster 1982).
** Sandeothallus japonicus (Inoue) Crand.-Stotl. et Stotler, Beih. Nova Hedwigia
131: 58, 2007 (Crandall-Stotler and Stotler 2007). Bas.: Moerckia japonica Inoue,
Bull. Natl. Sci. Mus. Tokyo, B 11 (1): 8, 1985 (Inoue 1985).
** Sandeothallus radiculosus (Schifn.) R.M.Schust., Nova Hedwigia 36: 11, 1982
(Schuster 1982). Bas.: Moerckia radiculosa Schifn., Österr. Bot. Z. 51 (2): 48,
1901 (Schifner 1901).
Phyllothalliineae R.M.Schust.
***
Phyllothalliaceae E.A.Hodgs. ex T.Katag.
*** Phyllothallia E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot. 2 (19): 247, 1964
(Hodgson 1964).
*** Phyllothallia fuegiana R.M.Schust., Trans. Brit. Bryol. Soc. 5 (2): 284, 1967
(Schuster 1967d).
*** Phyllothallia nivicola E.A.Hodgs., Trans. Roy. Soc. New Zealand, Bot. 2 (19):
247, 1964 (Hodgson 1964).
Pelliales He-Nygrén, Juslén, Ahonen, Glenny et Piippo
***
Noterocladaceae W.Frey et M.Stech
Frey and Stech (2005a) proposed Noterocladaceae as a monogeneric family based on
molecular and morphological evidence, which was later corroborated by CrandallStotler et al. (2010).
*** Noteroclada Taylor ex Hook.f. et Wilson, London J. Bot. 3: 166, 1844 (Hooker
and Wilson 1844).
*** Noteroclada conluens Taylor, London J. Bot. 3: 166, 1844 (Hooker and Wilson
1844).
*
Excluded from the genus
Noteroclada longiuscula Colenso, Trans. & Proc. New Zealand Inst. 19: 299, 1887
(Colenso 1887). 424
424 Noteroclada longiuscula is neither a Noteroclada nor a Fossombronia species. he type specimen could
not be found (Crandall-Stotler et al. 2010).
World checklist of hornworts and liverworts
***
475
Pelliaceae H.Klinggr.
Frey and Stech (2005b) recognized Pelliaceae as a monogeneric family, which was later
supported by Crandall-Stotler et al. (2010).
** Pellia Raddi, Jungermanniogr. Etrusca: 38, 1818 (Raddi 1818a) nom. conserv.
*** subg. Apopellia Grolle, J. Bryol. 12 (3): 427, 1983 (Grolle 1983a).
** Pellia alpicola R.M.Schust. ex L.Söderstr., A.Hagborg et von Konrat, Phytotaxa 76
(3): 39, 2013 (Söderström et al. 2013d). Based on: Pellia endiviifolia subsp. alpicola
R.M.Schust., J. Hattori Bot. Lab. 70: 145, 1991 (Schuster 1991b), nom. inval.
*** Pellia endiviifolia (Dicks.) Dumort., Recueil Observ. Jungerm.: 27, 1835 (Dumortier 1835). Bas.: Jungermannia endiviifolia Dicks., Fasc. Pl. Crypt. Brit. 4: 19,
1801 (Dickson 1801).
** Pellia megaspora R.M.Schust., J. Bryol. 11 (3): 419, 1981 (Schuster 1981b).
*** subg. Pellia
** Pellia appalachiana R.M.Schust., Phytotaxa 76 (3): 39, 2013 (Söderström et al.
2013d). Based on: Pellia appalachiana R.M.Schust., J. Hattori Bot. Lab. 70: 145,
1991 (Schuster 1991b), nom. inval.
*** Pellia epiphylla (L.) Corda, Gen. hepat.: 654, 1829 (Corda 1829). Bas.: Jungermannia epiphylla L., Sp. Pl. 1: 1135, 1753 (Linnaeus 1753).
** Pellia epiphylla subsp. borealis (Lorb.) Messe, Bull. Soc. Roy. Bot. Belgique 114
(1): 13, 1981 (Messe 1981). Bas.: Pellia borealis Lorb., Jahrb. Wiss. Bot. 80: 697,
1934 (Lorbeer 1934).
*** Pellia neesiana (Gottsche) Limpr., Hedwigia 15 (2): 18, 1876 (Limpricht 1876).
Bas.: Pellia epiphylla f. neesiana Gottsche, Hedwigia 6 (5): 69, 1867 (Gottsche 1867).
*
*
*
*
*
Incertae sedis
Pellia cordaeana Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C. Sci. Mat. 4
(13): 433, 1877 (Trevisan 1877).
Pellia crispa P.Kumm., Führer Leberm.: 60, 1875 (Kummer 1875).
Pellia gottscheana Kreh, Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur.
90 (4): 237, 1909 (Kreh 1909).
Pellia longifolia P.Kumm., Führer Leberm.: 60, 1875 (Kummer 1875).
Pellia undulata P.Kumm., Führer Leberm.: 60, 1875 (Kummer 1875).
476
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
MARCHANTIOPSIDA Cronquist, Takht. et W.Zimm.
Blasiidae He-Nygrén, Juslén, Ahonen, Glenny et Piippo
Blasiales Stotler et Crand.-Stotl.
***
Blasiaceae H.Klinggr.
by L. Söderström
*** Blasia L., Sp. Pl. 1: 1138, 1753 (Linnaeus 1753).
*** Blasia pusilla L., Sp. Pl. 1: 1138, 1753 (Linnaeus 1753).
*** Cavicularia Steph., Bull. Herb. Boissier 5 (2): 87, 1897 (Stephani 1897b).
*** Cavicularia densa Steph., Bull. Herb. Boissier 5 (2): 87, 1897 (Stephani 1897b).
Marchantiidae Engl.
Lunulariales H.Klinggr.
***
Lunulariaceae H.Klinggr.
by D.G. Long
*** Lunularia Adans., Fam. Pl. (Adanson) 2: 15, 1763 (Adanson 1763).
*** Lunularia cruciata (L.) Dumort. ex Lindb., Not. Sällsk. Fauna Fl. Fenn. Förh. 9:
298, 1868 (Lindberg 1868b). Bas.: Marchantia cruciata L., Sp. Pl. 1: 1137, 1753
(Linnaeus 1753).
* Lunularia cruciata subsp. thaxteri (A.Evans et Herzog) R.M.Schust., Hepat.
Anthocerotae N. Amer. 6: 91, 1992 (Schuster 1992d). Bas.: Lunularia thaxteri
A.Evans et Herzog, Arch. Esc. Fárm. Fac. Ci. Méd. Córdoba 7: 5, 1938 (Herzog
and Hosseus 1938). 425
Marchantiales Limpr.
***
Aytoniaceae Cavers
by D.G. Long
he genera of Aytoniaceae follow the treatment of Bischler-Causse et al. (2005) with
the exception of Asterella and Mannia which were re-deined by Schill et al. (2010).
425 Lunularia cruciata subsp. thaxteri was synonymized with subsp. cruciata by Boisselier-Dubayle et al.
(1995), but their subsp. thaxteri specimen was identiied with some doubt and originated far from
the known range.
World checklist of hornworts and liverworts
477
*** Asterella P.Beauv., Dict. Sci. Nat. [F. Cuvier] 3: 257, 1805 (Palisot de Beauvois
1805a) nom. conserv. 426
** subg. Asterella
** sect. Asterella
*** Asterella tenella (L.) P.Beauv., Dict. Sci. Nat. [F. Cuvier] 3: 258, 1805 (Palisot de
Beauvois 1805a). Bas.: Marchantia tenella L., Sp. Pl. 1: 1137, 1753 (Linnaeus 1753).
** sect. Brachyblepharis (Nees) D.G.Long, J. Bryol. 22 (2): 113, 2000 (Grolle and
Long 2000). Bas.: Fimbraria subg. Brachyblepharis Nees, Syn. Hepat. 4: 569, 1846
(Gottsche et al. 1846).
*** Asterella abyssinica (Gottsche) Grolle, Explor. Hydrobiol. Lac Bangweolo Luapula:
170, 1972 (Vanden Berghen 1972b). Bas.: Fimbraria abyssinica Gottsche, Syn.
Hepat. 4: 569, 1846 (Gottsche et al. 1846).
*** Asterella africana (Mont.) Underw. ex A.Evans, Contr. U.S. Natl. Herb. 20: 250,
1920 (Evans 1920). Bas.: Fimbraria africana Mont., Hist. Nat. Îles Canaries 3 (2):
61, 1840 (Montagne 1840b).
*** Asterella blumeana (Nees) Kachroo, J. Gauhati India Univ. 3: 130, 1952 (Kachroo
1952). Bas.: Fimbraria blumeana Nees, Syn. Hepat. 4: 564, 1846 (Gottsche et al. 1846).
*** Asterella chilensis (Nees et Mont.) A.Evans, Bull. Torrey Bot. Club 46 (12): 469,
1919 (Evans 1919b). Bas.: Fimbraria chilensis Nees et Mont., Ann. Sci. Nat. Bot.
(sér. 2) 9: 41, 1838 (Montagne 1838).
*** Asterella cruciata (Steph.) Horik., Hikobia 1 (2): 79, 1951 (Horikawa 1951c). Bas.:
Fimbraria cruciata Steph., Sp. Hepat. (Stephani) 6: 12, 1917 (Stephani 1917a).
*** Asterella dissoluta (Steph.) Grolle, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.Naturwiss. Reihe 38 (2): 237, 1989 (Grolle 1989b). Bas.: Fimbraria dissoluta
Steph., Planzenw. Ost-Afrikas C: 62, 1895 (Stephani 1895d).
*** Asterella dominicensis S.W.Arnell, Bryologist 61 (2): 140, 1958 (Arnell 1958c).
*** Asterella khasyana (Grif.) Grolle, Khumbu Himal 1 (4): 267, 1966 (Grolle 1966k).
Bas.: Octokepos khasyanus Grif., Not. pl. asiat. 2: 343, 1849 (Griith 1849).
*** Asterella leptophylla (Mont.) Grolle, Feddes Repert. 87 (3/4): 246, 1976 (Grolle
1976a). Bas.: Fimbraria leptophylla Mont., Ann. Sci. Nat. Bot. (sér. 2) 18: 16,
1842 (Montagne 1842b).
*** Asterella limbata D.G.Long et Grolle, J. Bryol. 18 (2): 287, 1994 (Long and Grolle
1994).
** Asterella shimizuana Inoue, Bull. Natl. Sci. Mus. Tokyo (n.ser.) 10 (3): 361, 1967
(Inoue 1967b). 427
426 Asterella includes Fimbraria and Hypenantron, but a few taxa have neither been transferred nor
synonymized. hey are listed in the “Names in genera not currently accepted” section below.
427 Asterella shimizuana is conspeciic with Asterella khasyana in Grolle and Piippo (1984), but it was
accepted by Long (2006).
478
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Asterella tenera (Mitt.) R.M.Schust., J. Hattori Bot. Lab. 26: 298, 1963 (Schuster
1963b). Bas.: Fimbraria tenera Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 170,
1855 (Mitten 1855).
* Asterella tenerrima (Steph.) H.A.Mill., Phytologia 47 (4): 320, 1981 (Miller 1981).
Bas.: Fimbraria tenerrima Steph., Sp. Hepat. (Stephani) 6: 17, 1917 (Stephani
1917a). 428
*** Asterella venosa (Lehm. et Lindenb.) A.Evans, Contr. U.S. Natl. Herb. 20: 286,
1920 (Evans 1920). Bas.: Fimbraria venosa Lehm. et Lindenb., Nov. Stirp. Pug. 4:
29, 1832 (Lehmann 1832).
** subg. Phragmoblepharis Grolle, Feddes Repert. 87 (3/4): 246, 1976 (Grolle 1976a).
*** Asterella australis (Hook.f. et Taylor) Verd., Ann. Bryol. 5: 126, 1932 (Verdoorn
1932c). Bas.: Fimbraria australis Hook.f. et Taylor, London J. Bot. 3: 573, 1844
(Hooker and Taylor 1844d).
*** Asterella bachmannii (Steph.) S.W.Arnell, Hepat. South Africa: 62, 1963 (Arnell 1963b).
Bas.: Fimbraria bachmannii Steph., Hedwigia 33 (1): 7, 1894 (Stephani 1894a).
*** Asterella bolanderi (Austin) Underw., Bot. Gaz. 20 (2): 61, 1895 (Underwood
1895). Bas.: Fimbraria bolanderi Austin, Proc. Acad. Nat. Sci. Philadelphia 21:
230, 1869 (Austin 1869).
* Asterella caucasica (Steph.) H.Buch, A.Evans et Verd., Ann. Bryol. 10: 8, 1937
[1938] (Buch et al. 1937). Bas.: Fimbraria caucasica Steph., Bull. Herb. Boissier 7
(3): 206 (132), 1899 (Stephani 1899b). 429
*** Asterella conocephala (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 298, 1963
(Schuster 1963b). Bas.: Fimbraria conocephala Steph., Bull. Herb. Boissier 7 (3):
205 (131), 1899 (Stephani 1899b).
** Asterella coronata (Steph.) H.A.Mill., Phytologia 47 (4): 319, 1981 (Miller 1981).
Bas.: Fimbraria coronata Steph., Sp. Hepat. (Stephani) 6: 12, 1917 (Stephani
1917a).
** Asterella dioica (Steph.) H.A.Mill., Phytologia 47 (4): 319, 1981 (Miller 1981).
Bas.: Fimbraria dioica Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2):
104, 1914 (Stephani and Watts 1914).
* Asterella dognyensis H.A.Mill., Phytologia 47 (4): 319, 1981 (Miller 1981). Nom.
nov. pro Fimbraria umbonata Steph., Sp. Hepat. (Stephani) 6: 17, 1917 (Stephani
1917a), nom. illeg. 430
*** Asterella drummondii (Taylor) R.M.Schust. ex D.G.Long, J. Bryol. 21 (1): 76,
1999 (Long 1999b). Bas.: Fimbraria drummondii Taylor, London J. Bot. 5: 412,
1846 (Taylor 1846b).
*** Asterella echinella (Gottsche) Underw., Bot. Gaz. 20 (2): 62, 1895 (Underwood 1895).
Bas.: Fimbraria echinella Gottsche, Mexik. Leverm.: 271, 1863 (Gottsche 1863).
428 Asterella tenerrima is probably conspeciic with Asterella tenera.
429 Asterella caucasica is probably conspeciic with Asterella whiteleggeana (Grolle 1975c, Scott and Bradshaw
1985).
430 Asterella dognyensis is possibly conspeciic with Asterella heterolora.
World checklist of hornworts and liverworts
479
*** Asterella elegans (Spreng.) Trevis., Rendiconti Reale Ist. Lombardo Sci. (ser. 2) 7:
785, 1874 (Trevisan 1874). Bas.: Fimbraria elegans Spreng. Syst. Veg. (ed. 16)
[Sprengel] 4 (1): 235, 1827 (Sprengel 1827a).
*** Asterella heterolora (Steph.) H.A.Mill., Phytologia 47 (4): 320, 1981 (Miller 1981).
Bas.: Fimbraria heterolora Steph., Sp. Hepat. (Stephani) 6: 14, 1917 (Stephani
1917a).
*** Asterella innovans (Austin) H.A.Mill., Ark. Bot. (n.ser.) 5 (2): 529, 1963 (Miller
1963). Bas.: Marchantia innovans Austin, Bull. Torrey Bot. Club 5 (3): 14, 1874
(Austin 1874).
*** Asterella lateralis M.Howe, Bull. Torrey Bot. Club 25 (4): 189, 1898 (Howe
1898b).
*** Asterella lindenbergiana (Corda ex Nees) Lindb. ex Arnell, Lebermoosstud. nördl.
Norwegen: 2, 1892 (Arnell 1892). Bas.: Fimbraria lindenbergiana Corda ex Nees,
Naturgesch. Eur. Leberm. 4: 266, 1838 (Nees 1838a).
*** Asterella linearis (Steph.) M.Howe, Bull. Torrey Bot. Club 25 (4): 191, 1898
(Howe 1898b). Bas.: Fimbraria linearis Steph., Bot. Jahrb. Syst. 20 (3): 302, 1895
(Stephani 1895a).
*** Asterella longebarbata (Steph.) H.A.Mill., Phytologia 47 (4): 320, 1981 (Miller
1981). Bas.: Fimbraria longebarbata Steph., Hedwigia 28 (3): 156, 1889 (Stephani
1889d).
*** Asterella macropoda (Spruce) A.Evans, Bull. Torrey Bot. Club 46 (12): 472, 1919
(Evans 1919b). Bas.: Fimbraria macropoda Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 564, 1885 (Spruce 1885).
*** Asterella marginata (Nees) S.W.Arnell, Hepat. South Africa: 63, 1963 (Arnell
1963b). Bas.: Fimbraria marginata Nees, Horae Phys. Berol.: 44, 1820 (Nees
1820).
** Asterella muelleri (Gottsche) R.M.Schust., J. Hattori Bot. Lab. 26: 298, 1963
(Schuster 1963b). Bas.: Fimbraria muelleri Gottsche, Bull. Herb. Boissier 7 (3):
203 (129), 1899 (Stephani 1899b).
*** Asterella multilora (Steph.) Kachroo, J. Hattori Bot. Lab. 19: 3, 1958 (Kachroo
1958). Bas.: Fimbraria multilora Steph., Bull. Herb. Boissier 7 (3): 198 (124),
1899 (Stephani 1899b).
*** Asterella mussuriensis (Kashyap) Verd., Ann. Bryol. 8: 156, 1935 (Verdoorn 1935).
Bas.: Fimbraria mussuriensis Kashyap, J. Bombay Nat. Hist. Soc. 24 (2): 345, 1916
(Kashyap 1916).
*** Asterella mussuriensis subsp. crassa (Shimizu et S.Hatt.) D.G.Long, Lindbergia 26
(1): 44, 2001 (Long 2001). Bas.: Asterella crassa Shimizu et S.Hatt., J. Hattori Bot.
Lab. 8: 48, 1952 (Shimizu and Hattori 1952).
*** Asterella pappii (Gola) Grolle, Feddes Repert. 87 (3/4): 246, 1976 (Grolle 1976a).
Bas.: Fimbraria pappii Gola, Ann. Bot. (Rome) 13 (1): 65, 1914 (Gola 1914a).
*** Asterella persica (Steph.) M.Howe, Bull. Torrey Bot. Club 25 (4): 191, 1898
(Howe 1898b). Bas.: Fimbraria persica Steph., Hedwigia 33 (1): 7, 1894 (Stephani
1894a).
480
*
**
***
**
***
***
***
***
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Asterella preussii (Schifn.) M.Howe, Bull. Torrey Bot. Club 25 (4): 191, 1898
(Howe 1898b). Bas.: Fimbraria preussii Schifn., Bot. Jahrb. Syst. 20 (3): 303,
1895 (Stephani 1895a). 431
Asterella setisquama (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 298, 1963
(Schuster 1963b). Bas.: Fimbraria setisquama Steph., Hedwigia 28 (3): 156, 1889
(Stephani 1889d).
Asterella syngenesica (Bory) Grolle, Lindbergia 2 (3/4): 230, 1974 (Grolle and
Onraedt 1974). Bas.: Marchantia syngenesica Bory, Voy. îles Afrique 2: 95, 1804
(Bory 1804).
Asterella tasmanica (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 298, 1963
(Schuster 1963b). Bas.: Fimbraria tasmanica Steph., Bull. Herb. Boissier 7 (3):
206 (132), 1899 (Stephani 1899b).
Asterella versicolor A.Evans, Contr. U.S. Natl. Herb. 20: 307, 1920 (Evans 1920).
Asterella vulcanica (Schifn.) Kachroo et Bapna, J. Indian Bot. Soc. 56 (1): 75,
1977 (Kachroo et al. 1977). Bas.: Hypenantron vulcanicum Schifn., Denkschr.
Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 67: 155, 1898 (Schifner 1898a).
Asterella whiteleggeana (Steph.) R.M.Schust., J. Hattori Bot. Lab. 26: 298, 1963
(Schuster 1963b). Bas.: Fimbraria whiteleggeana Steph., Hedwigia 28 (3): 155,
1889 (Stephani 1889d).
Asterella wilmsii (Steph.) S.W.Arnell, Hepat. South Africa: 62, 1963 (Arnell 1963b).
Bas.: Fimbraria wilmsii Steph., Hedwigia 31 (3): 122, 1892 (Stephani 1892d).
** subg. Saccatae (Grolle) D.G.Long, J. Bryol. 22 (2): 113, 2000 (Grolle and Long
2000). Bas.: Asterella subg. Phragmoblepharis sect. Saccatae Grolle, Feddes Repert.
87 (3/4): 346, 1976 (Grolle 1976a).
*** Asterella alpina (Steph.) D.G.Long, J. Hattori Bot. Lab. 93: 9, 2003 (Gradstein
et al. 2003). Bas.: Fimbraria alpina Steph., Bull. Herb. Boissier 7 (3): 211 (137),
1899 (Stephani 1899b).
*** Asterella grollei D.G.Long, Bryologist 102 (2): 169, 1999 (Long 1999a).
*** Asterella muscicola (Steph.) S.W.Arnell, Mitt. Bot. Staatssamml. München 2 (16):
263, 1957 (Arnell 1957c). Bas.: Fimbraria muscicola Steph., Hedwigia 31 (3): 121,
1892 (Stephani 1892d).
*** Asterella palmeri (Austin) Underw., Bot. Gaz. 20 (2): 63, 1895 (Underwood 1895).
Bas.: Fimbraria palmeri Austin, Bull. Torrey Bot. Club 6 (7): 47, 1875 (Austin 1875c).
*** Asterella pringlei Underw., Bot. Gaz. 20 (2): 64, 1895 (Underwood 1895).
*** Asterella rugosa A.Evans, Contr. U.S. Natl. Herb. 20: 289, 1920 (Evans 1920).
*** Asterella saccata (Wahlenb.) A.Evans, Contr. U.S. Natl. Herb. 20: 276, 1920 (Evans
1920). Bas.: Marchantia saccata Wahlenb., Mag. Neuesten Entdeck. Gesammten
Naturk. Ges. Naturf. Freunde Berlin 5: 296, 1811 (Wahlenberg 1811).
431 Asterella preussii was provisionally placed in synonymy with Asterella bachmannii (Wigginton and
Grolle 1996).
World checklist of hornworts and liverworts
481
** subg. Wallichianae D.G.Long, Lindbergia 26 (1): 43, 2001 (Long 2001).
** sect. Californicae D.G.Long, J. Hattori Bot. Lab. 97: 257, 2005 (Long 2005).
*** Asterella californica (Hampe ex Austin) Underw., Bot. Gaz. 20 (2): 60, 1895 (Underwood 1895). Bas.: Fimbraria californica Hampe ex Austin, Hepat. bor.-amer.:
33, 1873 (Austin 1873).
** sect. Wallichianae D.G.Long, Phytotaxa 173 (1): 87, 2014 (Long et al. 2014).
*** Asterella wallichiana (Lehm. et Lindenb.) Grolle, Khumbu Himal 1 (4): 262, 1966
(Grolle 1966k). Bas.: Fimbraria wallichiana Lehm. et Lindenb., Nov. Stirp. Pug.
4: 4, 1832 (Lehmann 1832).
** Cryptomitrium Austin ex Underw., Bull. Illinois State Lab. Nat. Hist. 2 (1): 36,
1884 (Underwood 1884).
*** Cryptomitrium himalayense Kashyap, New Phytol. 14 (1): 2, 1915 (Kashyap 1915).
*** Cryptomitrium oreades Perold, Bothalia 24 (2): 149, 1994 (Perold 1994).
*** Cryptomitrium tenerum (Hook.) Austin ex Underw., Bull. Illinois State Lab. Nat.
Hist. 2 (1): 36, 1884 (Underwood 1884). Bas.: Marchantia tenera Hook., Syn. Pl.
(Kunth) 1: 45, 1822 (Kunth 1822).
*** Mannia Corda, Gen. hepat.: 646, 1829 (Corda 1829) nom. conserv.
** subg. Mannia
*** Mannia androgyna (L.) A.Evans, Chron. Bot. 4: 224, 1938 (Evans 1938a). Bas.:
Marchantia androgyna L., Sp. Pl. 1: 1138, 1753 (Linnaeus 1753).
*** Mannia californica (Gottsche) L.C.Wheeler, Bryologist 37 (5): 88, 1934 [1935]
(Wheeler 1934). Bas.: Grimaldia californica Underw., Bot. Gaz. 13 (5): 114, 1888
(Underwood 1888).
*** Mannia controversa (Meyl.) D.B.Schill, Edinburgh J. Bot. 65 (1): 36, 2008 (Schill
et al. 2008). Bas.: Grimaldia controversa Meyl., Beitr. Kryptogamenl. Schweiz 6
(4): 87, 1924 (Meylan 1924).
*** Mannia controversa subsp. asiatica D.B.Schill et D.G.Long, Edinburgh J. Bot. 65
(1): 45, 2008 (Schill et al. 2008).
*** Mannia fragrans (Balb.) Frye et L.Clark, Univ. Wash. Publ. Biol. 6 (1): 62, 1937
(Frye and Clark 1937). Bas.: Marchantia fragrans Balb., Mem. Acad. Sci. Turin,
Sci. Phys. 15: 76, 1804 (Balbis 1804).
** Mannia fragrans subsp. orientalis R.M.Schust., Hepat. Anthocerotae N. Amer. 6:
201, 1992 (Schuster 1992d). Nom. nov. pro Mannia barbifrons Shimizu et S.Hatt.,
J. Hattori Bot. Lab. 10: 49, 1953 (Shimizu and Hattori 1953b).
* Mannia perssonii Udar et V.Chandra, Canad. J. Bot. 43 (1): 150, 1965 (Udar and
Chandra 1965). 432
432 Mannia perssonii is possibly conspeciic with Mannia sibirica.
482
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Mannia sibirica (Müll.Frib.) Frye et L.Clark, Univ. Wash. Publ. Biol. 6 (1): 66,
1937 (Frye and Clark 1937). Bas.: Grimaldia pilosa var. sibirica Müll.Frib., Lebermoose 1 (5): 265, 1907 (Müller 1907a).
** subg. Neesiella (Schifn.) D.B.Schill et D.G.Long, Bryologist 113 (1): 175,
2010 (Schill et al. 2010). Bas.: Neesiella Schifn., Hepat. (Engl.-Prantl): 32, 1893
(Schifner 1893b).
*** Mannia gracilis (F.Weber) D.B.Schill et D.G.Long, Bryologist 113 (1): 173, 2010
(Schill et al. 2010). Bas.: Marchantia gracilis F.Weber, Hist. Musc. Hepat. Prodr.:
105, 1815 (Weber 1815).
* Mannia hegewaldii Bischl., Fl. Neotrop. Monogr. 97: 184, 2005 (Bischler-Causse
et al. 2005). 433
*** Mannia pilosa (Hornem.) Frye et L.Clark, Univ. Wash. Publ. Biol. 6 (1): 64, 1937
(Frye and Clark 1937). Bas.: Marchantia pilosa Hornem., Fl. Danica 8 (24): 7, tab.
1426, 1810 (Hornemann 1810).
*** Mannia triandra (Scop.) Grolle, J. Bryol. 8 (4): 487, 1975 (Grolle 1975d). Bas.:
Marchantia triandra Scop., Fl. Carniol. (ed. 2) 2: 354, 1772 (Scopoli 1772).
*
Incertae sedis
Mannia paradoxa R.M.Schust., Phytologia 57 (6): 410, 1985 (Schuster 1985b). 434
*** Plagiochasma Lehm., Nov. Stirp. Pug. 4: 13, 1832 (Lehmann 1832) nom. conserv.
* Plagiochasma megacarpon (Grif.) Steph., Bull. Herb. Boissier 6 (10): 789 (86),
1898 (Stephani 1898c). Bas.: Antrocephalus megacarpon Grif., Not. pl. asiat. 2:
338, 1849 (Griith 1849). 435
** subg. Micropylum Bischl., Rev. Bryol. Lichénol. 43 (1): 103, 1977 (Bischler 1977).
*** Plagiochasma rupestre (J.R.Forst. et G.Forst.) Steph., Bull. Herb. Boissier 6 (10):
783 (80), 1898 (Stephani 1898c). Bas.: Aytonia rupestris J.R.Forst. et G.Forst.,
Char. gen. pl., ed. 2: 148, 1776 (Forster and Forster 1776).
*** Plagiochasma rupestre var. volkii Bischl., Rev. Bryol. Lichénol. 44 (3): 289, 1978
(Bischler 1978).
** subg. Plagiochasma
*** Plagiochasma appendiculatum Lehm. et Lindenb., Nov. Stirp. Pug. 4: 14, 1832
(Lehmann 1832).
*** Plagiochasma argentinicum Bischl., Rev. Bryol. Lichénol. 45 (3): 301, 1979
(Bischler 1979a).
433 Mannia hegewaldii is possibly conspeciic with Mannia triandra.
434 Mannia paradoxa is possibly a Reboulia species.
435 Plagiochasma megacarpon may not be a Plagiochasma species. Bischler (1979b) could not ind any
specimen and did not know where to refer it.
World checklist of hornworts and liverworts
483
*** Plagiochasma beccarianum Steph., Bull. Herb. Boissier 6 (10): 781 (78), 1898
(Stephani 1898c).
*** Plagiochasma cordatum Lehm. et Lindenb., Nov. Stirp. Pug. 4: 13, 1832 (Lehmann
1832).
*** Plagiochasma crenulatum Gottsche, Mexik. Leverm.: 266, 1863 (Gottsche 1863).
*** Plagiochasma cuneatum A.Evans, Amer. J. Bot. 19 (7): 627, 1932 (Evans 1932a).
*** Plagiochasma eximium (Schifn.) Steph., Bull. Herb. Boissier 6 (10): 781 (78),
1898 (Stephani 1898c). Bas.: Aytonia eximia Schifn., Bot. Jahrb. Syst. 20 (3):
300, 1895 (Stephani 1895a).
*** Plagiochasma intermedium Lindenb. et Gottsche, Syn. Hepat. 4: 513, 1846
(Gottsche et al. 1846).
*** Plagiochasma jamaicense (Haynes) A.Evans, Bull. Torrey Bot. Club 42 (5): 292,
1915 (Evans 1915). Bas.: Aytonia jamaicensis Haynes, Bull. Torrey Bot. Club 34
(2): 58, 1907 (Haynes 1907).
*** Plagiochasma japonicum (Steph.) C.Massal., Hepat. Shen-si: 47, 1897 (Massalongo 1897). Bas.: Aytonia japonica Steph., Bull. Herb. Boissier 5 (2): 84, 1897
(Stephani 1897b).
*** Plagiochasma landii A.Evans, Bull. Torrey Bot. Club 42 (5): 298, 1915 (Evans 1915).
*** Plagiochasma microcephalum (Steph.) Steph., Bull. Herb. Boissier 6 (10): 781 (78),
1898 (Stephani 1898c). Bas.: Aytonia microcephala Steph., Bot. Jahrb. Syst. 20 (3):
301, 1895 (Stephani 1895a).
*** Plagiochasma microcephalum var. tunesicum Bischl., Rev. Bryol. Lichénol. 44 (3):
247, 1978 (Bischler 1978).
*** Plagiochasma muenchianum Steph., Sp. Hepat. (Stephani) 6: 9, 1917 (Stephani 1917a).
*** Plagiochasma pterospermum C.Massal., Hepat. Shen-si: 46, 1897 (Massalongo 1897).
*** Plagiochasma wrightii Sull., Musc. Hepat. U.S.: 688, 1856 (Sullivant 1856).
Incertae sedis
** Plagiochasma udarii A.Alam et S.C.Srivast., Indian J. Forest. 32 (4): 631, 2009
(Alam and Srivastava 2009).
*** Reboulia Raddi, Opusc. Sci. 2 (6): 357, 1818 (Raddi 1818b) nom. conserv.
*** Reboulia hemisphaerica (L.) Raddi, Opusc. Sci. 2 (6): 357, 1818 (Raddi 1818b).
Bas.: Marchantia hemisphaerica L., Sp. Pl. 1: 1138, 1753 (Linnaeus 1753).
** Reboulia hemisphaerica subsp. acrogyna (R.M.Schust.) R.M.Schust., Hepat. Anthocerotae N. Amer. 6: 168, 1992 (Schuster 1992d). Bas.: Asterella bolanderi subsp.
acrogyna R.M.Schust., Phytologia 57 (6): 410, 1985 (Schuster 1985b).
** Reboulia hemisphaerica subsp. australis R.M.Schust., Phytologia 56 (7): 460, 1985
(Schuster 1985c).
** Reboulia hemisphaerica subsp. dioica R.M.Schust., Phytologia 56 (7): 462, 1985
(Schuster 1985c).
** Reboulia hemisphaerica var. issisquama Herzog, Symb. Sin. 5: 5, 1930 (Nicholson
et al. 1930).
484
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Reboulia hemisphaerica subsp. orientalis R.M.Schust., Phytologia 56 (7): 461, 1985
(Schuster 1985c).
** Reboulia hemisphaerica var. turkestanica C.E.O.Jensen ex Herzog, Symb. Sin. 5: 5,
1930 (Nicholson et al. 1930).
***
Cleveaceae Cavers
by D.G. Long
he genera of Cleveaceae were re-deined by Rubasinghe et al. (2011a) based on molecular evidence.
*** Athalamia Falc., Ann. Mag. Nat. Hist. (ser. 2) 1 (5): 375, 1848 (Anonymous 1848).
* Athalamia dioica Kashyap, J. Bombay Nat. Hist. Soc. 24 (2): 348, 1916 (Kashyap 1916).
*** Athalamia pinguis Falc., Ann. Mag. Nat. Hist. (ser. 2) 1 (5): 375, 1848 (Anonymous
1848).
* Athalamia pulcherrima (Steph.) S.Hatt., J. Hattori Bot. Lab. 12: 54, 1954 (Shimizu and Hattori 1954). Bas.: Clevea pulcherrima Steph., Bot. Jahrb. Syst. 20 (3):
303, 1895 (Stephani 1895a).
*** Clevea Lindb., Not. Sällsk. Fauna Fl. Fenn. Förh. 9: 289, 1868 (Lindberg 1868b).
*** Clevea hyalina (Sommerf.) Lindb., Not. Sällsk. Fauna Fl. Fenn. Förh. 9: 291,
1868 (Lindberg 1868b). Bas.: Marchantia hyalina Sommerf., Mag. Naturvidensk.
11 (2): 234, 1833 (Sommerfeldt 1833).
* Clevea hyalina var. californica M.Howe, Mem. Torrey Bot. Club 7: 38, 1899
(Howe 1899).
* Clevea pedicellata (Grif.) Lindb., Acta Soc. Fauna Fl. Fenn. 2 (3): 11, 1882 (Lindberg 1882). Bas.: Plagiochasma pedicellatum Grif., Not. pl. asiat. 2: 331, 1849
(Griith 1849).
*** Clevea pusilla (Steph.) Rubas. et D.G.Long, J. Bryol. 33 (2): 167, 2011 (Rubasinghe et al. 2011b). Bas.: Gollaniella pusilla Steph., Hedwigia 44 (2): 74, 1905
(Stephani 1905h).
*** Clevea spathysii (Lindenb.) Müll.Frib., Hedwigia 79 (1/2): 75, 1940 (Müller 1940).
Bas.: Marchantia spathysii Lindenb., Syn. hepat. eur: 104, 1829 (Lindenberg 1829).
*** Peltolepis Lindb., Morgonbladet (Helsinki) 1876 (106, 9 May): 1, 1876 (Elfving
1876).
** Peltolepis japonica (Shimizu et S.Hatt.) S.Hatt., J. Hattori Bot. Lab. 14: 103, 1955
(Shimizu and Hattori 1955). Bas.: Peltolepis quadrata var. japonica Shimizu et
S.Hatt., J. Hattori Bot. Lab. 12: 69, 1954 (Shimizu and Hattori 1954).
*** Peltolepis quadrata (Saut.) Müll.Frib., Hedwigia 79 (1/2): 74, 1940 (Müller 1940).
Bas.: Sauteria quadrata Saut., Flora 43 (22): 351, 1860 (Sauter 1860).
World checklist of hornworts and liverworts
485
*** Sauteria Nees, Naturgesch. Eur. Leberm. 4: 139, 1838 (Nees 1838a).
** sect. Sauchia (Kashyap) R.M.Schust., Phytologia 57 (6): 411, 1985 (Schuster
1985b). Bas.: Sauchia Kashyap, J. Bombay Nat. Hist. Soc. 24 (2): 347, 1916
(Kashyap 1916).
* Sauteria japonica (Shimizu et S.Hatt.) S.Hatt., J. Hattori Bot. Lab. 12: 62, 1954
(Shimizu and Hattori 1954). Bas.: Sauchia japonica Shimizu et S.Hatt., J. Hattori
Bot. Lab. 9: 32, 1953 (Shimizu and Hattori 1953a). 436
*** Sauteria spongiosa (Kashyap) S.Hatt., J. Hattori Bot. Lab. 12: 62, 1954 (Shimizu
and Hattori 1954). Bas.: Sauchia spongiosa Kashyap, J. Bombay Nat. Hist. Soc. 24
(2): 347, 1916 (Kashyap 1916).
** sect. Sauteria
*** Sauteria alpina (Nees) Nees, Naturgesch. Eur. Leberm. 4: 143, 1838 (Nees 1838a).
Bas.: Lunularia alpina Nees, Flora 13 (25): 399, 1830 (Nees and Bischof 1830).
* Sauteria inlata C.Gao et K.C.Chang, Acta Bot. Yunnan. 3 (4): 389, 1981 (Gao et
al. 1981). 437
*
Incertae sedis
Sauteria chilensis (Lindenb.) Grolle, J. Hattori Bot. Lab. 58: 200, 1985 (Grolle
1985b). Bas.: Grimaldia chilensis Lindenb., Voy. Amér. Mérid. 7 (2): 53, 1839
(Montagne 1839a).
Sauteria crassipes Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 229, 1869 (Austin
1869).
Sauteria nyikaensis Perold, Bothalia 33 (2): 167, 2003 (Perold 2003).
***
Conocephalaceae Müll.Frib. ex Grolle
*
*
by D.G. Long
*** Conocephalum Hill, Gener. Nat. Hist. 2 Hist. pl. (ed. 2): 118, 1773 (Hill 1773)
nom. conserv.
** subg. Conocephalum
*** Conocephalum conicum (L.) Dumort., Commentat. Bot. (Dumortier): 115, 1822
(Dumortier 1822). Bas.: Marchantia conica L., Sp. Pl. 1: 1138, 1753 (Linnaeus 1753).
*** Conocephalum salebrosum Szweyk., Buczk. et Odrzyk., Pl. Syst. Evol. 253 (1/4):
146, 2005 (Szweykowski et al. 2005).
436 Sauteria japonica is possibly conspeciic with Sauteria spongiosa.
437 Sauteria inlata is possibly conspeciic with Sauteria alpina (Schuster 1992d).
486
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** subg. Sandea (Lindb.) Inoue, Ill. Jap. Hep. 2: 192, 1976 (Inoue 1976a). Bas.:
Sandea Lindb., Acta Soc. Fauna Fl. Fenn. 2 (5): 3, 1884 (Lindberg 1884).
*** Conocephalum japonicum (hunb.) Grolle, J. Hattori Bot. Lab. 55: 501, 1984
(Grolle 1984a). Bas.: Lichen japonicus hunb., Fl. Jap. (hunberg): 344, 1784
(hunberg 1784).
**
Corsiniaceae Engl.
by D.G. Long
**
Corsinioideae Schiffn.
*** Corsinia Raddi, Opusc. Sci. 2 (6): 354, 1818 (Raddi 1818b).
*** Corsinia coriandrina (Spreng.) Lindb., Hepaticol. Utveckl.: 30, 1877 (Lindberg
1877c). Bas.: Riccia coriandrina Spreng. Anleit. Kenntn. Gew. 3: 320, 1804
(Sprengel 1804).
***
Cronisioideae R.M.Schust.
*** Cronisia Berk., Introd. crypt. bot.: 434, 1857 (Berkeley 1857).
*** Cronisia imbriata (Nees) Whittem. et Bischl., Cryptog. Bryol. 22 (3): 170, 2001
(Bischler and Whittemore 2001). Bas.: Riccia imbriata Nees, Fl. Bras. (Martius) 1
(1): 301, 1833 (Nees 1833a).
*** Cronisia weddellii (Mont.) Grolle, J. Bryol. 9 (4): 532, 1977 [1978] (Grolle 1977a).
Bas.: Boschia weddellii Mont., Ann. Sci. Nat. Bot. (sér. 4) 5: 352, 1856 (Montagne
1856c).
***
Cyathodiaceae Stotler et Crand.-Stotl.
by D.G. Long
*** Cyathodium Kunze, Nov. Stirp. Pug. 6: 17, 1834 (Lehmann 1834).
*** Cyathodium aureonitens (Grif.) Mitten, J. Linn. Soc., Bot. 22 (146): 327 (Mitten
1886b). Bas.: Synhymenium aureonitens Grif., Not. pl. asiat. 2: 344, 1849 (Grifith 1849).
*** Cyathodium bischlerianum N.Salazar, Bryologist 104 (1): 141, 2001 (Salazar 2001).
*** Cyathodium cavernarum Kunze, Nov. Stirp. Pug. 6: 18, 1834 (Lehmann 1834).
** Cyathodium denticulatum Udar et S.C.Srivast., Geophytology 1 (2): 166, 1971
(Udar and Srivastava 1971).
World checklist of hornworts and liverworts
487
*** Cyathodium foetidissimum Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 67: 154, 1898 (Schifner 1898a).
*** Cyathodium indicum Udar et D.K.Singh, J. Bryol. 10 (2): 139, 1978 [1979] (Udar
and Singh 1978).
*** Cyathodium mehranum D.K.Singh, Misc. Bryol. Lichenol. 9 (8): 173, 1983 (Singh
1983b).
*** Cyathodium smaragdinum Schifn., Ann. Jard. Bot. Buitenzorg, suppl. 3: 480,
1910 (Schifner 1910c).
*** Cyathodium spruceanum Prosk., Bryologist 54 (4): 243, 1951 [1952] (Proskauer
1951b).
* Cyathodium spurium (Dicks.) Lindb. ex Braithw., J. Bot. 16: 55, 1878 (Braithwaite
1878). Bas.: Riccia spuria Dicks., Fasc. Pl. Crypt. Brit. 4: 20, 1801 (Dickson 1801). 438
*** Cyathodium steerei Hässel, Rev. Bryol. Lichénol. 30 (3/4): 223, 1961 (Hässel 1961).
*** Cyathodium tuberculatum Udar et D.K.Singh, Bryologist 79 (2): 235, 1976 (Udar
and Singh 1976).
*** Cyathodium tuberosum Kashyap, New Phytol. 13 (6/7): 210, 1914 (Kashyap
1914a).
***
Dumortieraceae D.G.Long
by D.G. Long
*** Dumortiera Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12
(1): 410, 1824 [1825] (Reinwardt et al. 1824b).
*** Dumortiera hirsuta (Sw.) Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol.
Nat. Cur. 12 (1): 410, 1824 [1825] (Reinwardt et al. 1824b). Bas.: Marchantia
hirsuta Sw., Prodr. (Swartz): 145, 1788 (Swartz 1788). 439
* Dumortiera hirsuta subsp. nepalensis (Taylor) R.M.Schust., Hepat. Anthocerotae N. Amer. 6: 386, 1992 (Schuster 1992d). Bas.: Hygrophila nepalensis Taylor,
Trans. Linn. Soc. London 17 (3): 392, 1836 (Taylor 1836b).
* Dumortiera hirsuta subsp. tatunoi Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2,
Bot. 6: 38, 1951 (Horikawa 1951a).
***
Exormothecaceae Müll.Frib. ex Grolle
by D.G. Long
*** Aitchisoniella Kashyap, New Phytol. 13 (6/7): 219, 1914 (Kashyap 1914a).
438 Cyathodium spurium may be conspeciic with Cyathodium cavernarum (Braithwaite 1878). he type
specimen is from Scotland, but it is probably mislabelled as no Cyathodium is known from boreal Europe.
439 Dumortiera hirsuta is a species complex (Forrest et al. 2011).
488
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Aitchisoniella himalayensis Kashyap, New Phytol. 13 (6/7): 219, 1914 (Kashyap
1914a).
*** Exormotheca Mitt., Nat. hist. Azores: 325, 1870 (Mitten 1870).
** subg. Corbierella (Douin et Trab.) Schifn., Hedwigia 81 (1/2): 71, 1942
(Schifner 1942). Bas.: Corbierella Douin et Trab., Rev. Gén. Bot. 31: 326, 1919
(Douin and Trabut 1919).
*** Exormotheca bischlerae Furuki et Higuchi, Cryptog. Bryol. 27 (1): 98, 2006 (Furuki
and Higuchi 2006).
*** Exormotheca holstii Steph., Bull. Herb. Boissier 7 (3): 219 (145), 1899 (Stephani
1899b).
*** Exormotheca welwitschii Steph., Bull. Herb. Boissier 7 (3): 220 (146), 1899
(Stephani 1899b).
** subg. Exormotheca
*** Exormotheca pustulosa Mitt., Nat. hist. Azores: 326, 1870 (Mitten 1870).
Incertae sedis
*** Exormotheca bulbigena Bornefeld, O.H.Volk et R.Wolf, Bothalia 26 (2): 159,
1996 (Bornefeld et al. 1996).
*** Exormotheca ceylonensis Meijer, J. Hattori Bot. Lab. 16: 72, 1956 (Meijer 1956).
* Exormotheca gollanii Steph., Sp. Hepat. (Stephani) 6: 18, 1917 (Stephani 1917a).
*** Exormotheca tuberifera Kashyap, New Phytol. 13 (9): 309, 1914 (Kashyap
1914b).
*** Stephensoniella Kashyap, New Phytol. 13 (9): 312, 1914 (Kashyap 1914b).
*** Stephensoniella brevipedunculata Kashyap, New Phytol. 13 (9): 312, 1914 (Kashyap
1914b).
***
Marchantiaceae Lindl.
by D.G. Long
he treatment of Marchantiaceae is mainly following Bischler (1984, 1989b) and
Bischler-Causse (1993).
**
Bucegioideae R.M.Schust.
*** Bucegia Radian, Bull. Herb. Inst. Bot. Bucarest 3-4: 3, 1903 (Radian 1903).
*** Bucegia romanica Radian, Bull. Herb. Inst. Bot. Bucarest 3-4: 4, 1903 (Radian 1903).
World checklist of hornworts and liverworts
**
489
Marchantioideae Schiffn.
*** Marchantia L., Sp. Pl. 1: 1137, 1753 (Linnaeus 1753).
** subg. Chlamidium (Corda) Bischl., Cryptog. Bryol. Lichénol. 3 (4): 362, 1982
(Bischler 1982). Bas.: Chlamidium Corda, Gen. hepat.: 647, 1829 (Corda 1829).
*** Marchantia breviloba A.Evans, Trans. Connecticut Acad. Arts 21 (3): 265, 1917
(Evans 1917a).
*** Marchantia inlexa Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 9: 43, 1838 (Montagne 1838).
** sect. Chlamidium (Corda) Nees, Naturgesch. Eur. Leberm. 4: 101, 1838 (Nees
1838a). Bas.: Chlamidium Corda, Gen. hepat.: 647, 1829 (Corda 1829).
*** Marchantia chenopoda L., Sp. Pl. 1: 1137, 1753 (Linnaeus 1753).
*** Marchantia crenata Austin, Bull. Torrey Bot. Club 5 (3): 14, 1874 (Austin 1874).
*** Marchantia foliacea Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 168, 1855 (Mitten
1855).
*** Marchantia formosana Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 121,
1934 (Horikawa 1934).
*** Marchantia globosa Brid., Hist. Musc. Hepat. Prodr.: 102, 1815 (Weber 1815).
*** Marchantia hexaptera Reichardt, Verh. K.K. Zool.-Bot. Ges. Wien 16: 957, 1866
(Reichardt 1866).
*** Marchantia linearis Lehm. et Lindenb., Nov. Stirp. Pug. 4: 8, 1832 (Lehmann 1832).
*** Marchantia miqueliana Lehm., Nov. Stirp. Pug. 10: 20, 1857 (Lehmann 1857).
*** Marchantia novoguineensis Bischl., Bryophyt. Biblioth. 38: 130, 1989 (Bischler
1989b).
*** Marchantia pappeana Lehm., Nov. Stirp. Pug. 10: 21, 1857 (Lehmann 1857).
** Marchantia pappeana subsp. robusta (Steph.) Bischl., Bryophyt. Biblioth. 45: 91,
1993 (Bischler-Causse 1993). Bas.: Marchantia robusta Steph., Candollea 14: 111,
1953 (Bonner 1953b).
*** Marchantia pileata Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 169, 1855 (Mitten
1855).
*** Marchantia pinnata Steph., Candollea 14: 109, 1953 (Bonner 1953b).
*** Marchantia rubribarba Steph., Bull. Herb. Boissier 7 (5): 400 (172), 1899 (Stephani
1899c).
*** Marchantia vitiensis Steph., Bull. Herb. Boissier 7 (7): 520 (182), 1899 (Stephani
1899d).
** sect. Paleaceae Bischl., Bryophyt. Biblioth. 38: 90, 1989 (Bischler 1989b).
*** Marchantia paleacea Bertol., Opusc. Sci. 1: 242, 1817 (Bertoloni 1817).
*** Marchantia paleacea subsp. diptera (Nees et Mont.) Inoue, J. Jap. Bot. 64 (7): 194,
1989 (Inoue 1989c). Bas.: Marchantia diptera Nees et Mont., Ann. Sci. Nat. Bot.
(sér. 2) 19: 243, 1843 (Montagne 1843).
490
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** sect. Papillatae Bischl., Cryptog. Bryol. Lichénol. 10 (1): 69, 1989 (Bischler
1989a).
*** Marchantia debilis K.I.Goebel, Organogr. Pl., ed. 2, 2 (1): 901, 1915 (Goebel 1915).
*** Marchantia emarginata Reinw., Blume et Nees, Nova Acta Phys.-Med. Acad.
Caes. Leop.-Carol. Nat. Cur. 12 (1): 192, 1824 [1825] (Reinwardt et al. 1824a).
*** Marchantia emarginata subsp. lecordiana (Steph.) Bischl., Cryptog. Bryol. Lichénol. 10 (1): 78, 1989 (Bischler 1989a). Bas.: Marchantia lecordiana Steph., Bull.
Herb. Boissier 7 (7): 525 (187), 1899 (Stephani 1899d).
*** Marchantia emarginata subsp. tosana (Steph.) Bischl., Cryptog. Bryol. Lichénol.
10 (1): 77, 1989 (Bischler 1989a). Bas.: Marchantia tosana Steph., Bull. Herb.
Boissier 5 (2): 99, 1897 (Stephani 1897b).
*** Marchantia papillata Raddi, Critt. Brasil.: 20, 1822 (Raddi 1822).
*** Marchantia papillata subsp. grossibarba (Steph.) Bischl., Cryptog. Bryol. Lichénol.
10 (1): 78, 1989 (Bischler 1989a). Bas.: Marchantia grossibarba Steph., Mém. Soc.
Nat. Sci. Nat. Math. Cherbourg 29: 221, 1894 (Stephani 1894b).
** subg. Marchantia
*** Marchantia plicata Nees et Mont., Ann. Sci. Nat. Bot. (sér. 2) 9: 43, 1838 (Montagne 1838).
** sect. Berteroanae R.M.Schust., Hepat. Anthocerotae N. Amer. 6: 310, 1992
(Schuster 1992d).
*** Marchantia berteroana Lehm. et Lindenb., Nov. Stirp. Pug. 6: 21, 1834 (Lehmann
1834).
** sect. Marchantia
*** Marchantia polymorpha L., Sp. Pl. 1: 1137, 1753 (Linnaeus 1753).
*** Marchantia polymorpha subsp. montivagans Bischl. et Boissel.-Dub., J. Bryol. 16
(3): 364, 1991 (Bischler-Causse and Boisselier-Dubayle 1991).
*** Marchantia polymorpha subsp. ruderalis Bischl. et Boissel.-Dub., J. Bryol. 16 (3):
364, 1991 (Bischler-Causse and Boisselier-Dubayle 1991).
** subg. Protomarchantia R.M.Schust., Phytologia 57 (6): 410, 1985 (Schuster 1985b).
** sect. Protomarchantia (R.M.Schust.) L.Söderstr., Phytotaxa 202 (1): 69, 2015
(Söderström et al. 2015c). Bas.: Marchantia subg. Protomarchantia R.M.Schust.,
Phytologia 57 (6): 410, 1985 (Schuster 1985b).
*** Marchantia acaulis Steph., Bull. Herb. Boissier 7 (7): 533 (195), 1899 (Stephani
1899d).
*** Marchantia antiqua Steph., Candollea 14: 103, 1953 (Bonner 1953b).
*** Marchantia carrii Bischl., Bryophyt. Biblioth. 38: 256, 1989 (Bischler 1989b).
*** Marchantia geminata Reinw., Blume et Nees, Nova Acta Phys.-Med. Acad. Caes.
Leop.-Carol. Nat. Cur. 12 (1): 194, 1824 [1825] (Reinwardt et al. 1824a).
World checklist of hornworts and liverworts
491
*** Marchantia hartlessiana Steph., Candollea 14: 107, 1953 (Bonner 1953b).
*** Marchantia macropora Mitt., Bot. antarct. voy. II (Fl. Nov.-Zel. 2): 169, 1855
(Mitten 1855).
*** Marchantia philippinensis Bischl., Bryophyt. Biblioth. 38: 245, 1989 (Bischler 1989b).
*** Marchantia solomonensis Bischl., Bryophyt. Biblioth. 38: 281, 1989 (Bischler 1989b).
*** Marchantia streimannii Bischl., Bryophyt. Biblioth. 38: 250, 1989 (Bischler 1989b).
*** Marchantia subintegra Mitt., J. Proc. Linn. Soc., Bot. 5 (18): 125, 1860 [1861]
(Mitten 1860c).
*** Marchantia treubii Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl.
67: 160, 1898 (Schifner 1898a).
*** Marchantia wallisii J.B.Jack et Steph., Bull. Herb. Boissier 7 (7): 520 (182), 1899
(Stephani 1899d).
** sect. Subgeminatae Bischl., Bryophyt. Biblioth. 38: 219, 1989 (Bischler 1989b).
*** Marchantia subgeminata Steph., Bull. Herb. Boissier 7 (7): 530 (192), 1899
(Stephani 1899d).
*
*
*
*
*
*
*
*
Incertae sedis
Marchantia assamica Grif., Not. pl. asiat. 2: 327, 1849 (Griith 1849). 440
Marchantia balboi Gola, Mem. Reale Accad. Sci. Torino (ser. 2) 65 (1): 2, 1916
(Gola 1916). 441
Marchantia balboi var. acutisquamata Gerola, Lav. Bot. Ist. Bot. Univ. Padova 12:
472, 1947 (Gerola 1947). 442
Marchantia cagnii Gola, Ann. Bot. (Rome) 6 (2): 271, 1907 (Gola 1907). 443
Marchantia cengiana Gerola, Lav. Bot. Ist. Bot. Univ. Padova 12: 473, 1947
(Gerola 1947). 444
Marchantia friedrichsthaliana Trevis., Mem. Reale Ist. Lombardo Sci. (Ser. 3), C.
Sci. Mat. 4 (13): 438, 1877 (Trevisan 1877).
Marchantia keniae Gola, Mem. Reale Accad. Sci. Torino (ser. 2) 65 (1): 3, 1916
(Gola 1916). 445
Marchantia papyracea Gola, Ann. Bot. (Rome) 6 (2): 271, 1907 (Gola 1907). 446
440 Marchantia assamica is a doubtful taxon. Bischler (1989b) could not ind the type specimen.
441 Marchantia balboi is a doubtful taxon. Bischler-Causse (1993) could not ind the type specimen.
442 Marchantia balboi var. acutisquamata is a doubtful taxon, Bischler-Causse (1993) could not ind the
type specimen and did not know what it is.
443 Marchantia cagnii is a doubtful taxon. Bischler-Causse (1993) could not ind the type specimen, but
the description points to Marchantia paleacea.
444 Marchantia cengiana is a doubtful taxon. Bischler-Causse (1993) could not ind the type specimen,
but the description points to Marchantia pappeana.
445 Marchantia keniae is a doubtful taxon. Bischler-Causse (1993) could not ind the type specimen, but
the description points to Marchantia pappeana.
446 Marchantia papyracea is a doubtful taxon. Bischler-Causse (1993) could not ind the type specimen,
but the description points to Marchantia pappeana.
492
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
* Marchantia quadriloba Steph., Candollea 14: 110, 1953 (Bonner 1953b).
* Marchantia sellae Gola, Ann. Bot. (Rome) 6 (2): 271, 1907 (Gola 1907). 447
** Marchantia stoloniscyphulus (C.Gao et K.C.Chang) Piippo, J. Hattori Bot. Lab.
68: 134, 1990 (Piippo 1990). Bas.: Marchantiopsis stoloniscyphulus C.Gao et
K.C.Chang, Bull. Bot. Res., Harbin 2 (4): 114, 1982 (Gao and Chang 1982).
* Marchantia trilocularis Roth, Tent. Fl. Germ. 1: 487, 1788 (Roth 1788). 448
* Marchantia tusui Gola, Mem. Reale Accad. Sci. Torino (ser. 2) 65 (1): 3, 1916
(Gola 1916). 449
** Preissia Corda, Gen. hepat.: 647, 1829 (Corda 1829).
*** Preissia quadrata (Scop.) Nees, Naturgesch. Eur. Leberm. 4: 135, 1838 (Nees
1838a). Bas.: Marchantia quadrata Scop., Fl. Carniol. (ed. 2) 2: 355, 1772 (Scopoli 1772).
** Preissia quadrata subsp. hyperborea R.M.Schust., Phytologia 57 (6): 410, 1985
(Schuster 1985b).
***
Monocleaceae A.B.Frank
by D.G. Long
***
***
***
***
Monoclea Hook., Musci Exot. 2: tab. clxxiv, 1820 (Hooker 1820).
Monoclea forsteri Hook., Musci Exot. 2: tab. clxxiv, 1820 (Hooker 1820).
Monoclea gottschei Lindb., Rev. Bryol. 13 (6): 102, 1886 (Lindberg 1886).
Monoclea gottschei subsp. elongata Gradst. et Mues, Pl. Syst. Evol. 180 (1/2): 133,
1992 (Gradstein et al. 1992).
***
Monosoleniaceae Inoue
by D.G.Long
*** Monosolenium Grif., Not. pl. asiat. 2: 341, 1849 (Griith 1849).
*** Monosolenium tenerum Grif., Not. pl. asiat. 2: 341, 1849 (Griith 1849).
447 Marchantia sellae is a doubtful taxon. Bischler-Causse (1993) could not ind the type specimen, but
the description points to Marchantia pappeana or Marchantia paleacea.
448 Marchantia trilocularis is a doubtful taxon which Bischler-Causse (1993) could not place. he name
may have priority once its identity is determined.
449 Marchantia tusui is a doubtful taxon. Bischler-Causse (1993) could not ind the type specimen, but
the description points to Marchantia pappeana.
World checklist of hornworts and liverworts
***
493
Oxymitraceae Müll.Frib. ex Grolle
by D.G. Long
*** Oxymitra Bisch. ex Lindenb., Syn. hepat. eur: 124, 1829 (Lindenberg 1829).
*** Oxymitra cristata Garside, Bothalia 23 (2): 211, 1993 (Perold 1993). Based on: Oxymitra cristata Garside, J. S. African Bot. 24: 83, 1958 (Garside 1958), nom. inval.
*** Oxymitra incrassata (Brot.) Sérgio et Sim-Sim, J. Bryol. 15 (4): 662, 1989 (Sérgio
and Sim-Sim 1989). Bas.: Riccia incrassata Brot., Fl. lusit. 2: 428, 1804 [1805]
(Brotero 1804).
***
Ricciaceae Rchb.
by R. Stotler, B.J. Crandall-Stotler and D.C. Cargill
** Riccia L., Sp. Pl. 1: 1138, 1753 (Linnaeus 1753) nom. conserv.
** subg. Chartaceae Perold, Bothalia 16 (1): 29, 1986 (Volk and Perold 1986c).
*** Riccia schelpei O.H.Volk et Perold, Bothalia 16 (1): 29, 1986 (Volk and Perold
1986c).
** subg. Leptoriccia R.M.Schust., Phytologia 56 (2): 72, 1984 (Schuster 1984).
*** Riccia membranacea Gottsche et Lindenb., Syn. Hepat. 4: 608, 1846 (Gottsche et
al. 1846).
** subg. Riccia
*** Riccia albida Sull. ex Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 231, 1869
(Austin 1869).
*** Riccia albopunctata Jovet-Ast, Cryptog. Bryol. Lichénol. 12 (3): 237, 1991 (JovetAst 1991).
*** Riccia australis Steph., Bull. Herb. Boissier 6 (4): 337 (29), 1898 (Stephani 1898a).
*** Riccia boliviensis Jovet-Ast, Cryptog. Bryol. Lichénol. 12 (3): 242, 1991 (Jovet-Ast
1991).
*** Riccia brasiliensis Schifn., Österr. Akad. Wiss., Math.-Naturwiss. Kl., Denkschr.
111: 6, 1964 (Schifner and Arnell 1964).
*** Riccia breutelii Hampe, Bull. Herb. Boissier 6 (4): 325 (17), 1898 (Stephani 1898a).
*** Riccia brittonii M.Howe, Ann. Missouri Bot. Gard. 2 (1/2): 50, 1915 (Britton 1915).
* Riccia chudoana Steph., Sp. Hepat. (Stephani) 6: 1, 1917 (Stephani 1917a).
*** Riccia coracina Jovet-Ast, Cryptog. Bryol. 24 (3): 212, 2003 (Jovet-Ast 2003).
*** Riccia corrugata Jovet-Ast, Cryptog. Bryol. 21 (4): 308, 2000 (Jovet-Ast 2000).
*** Riccia crassivenia Jovet-Ast, Cryptog. Bryol. 21 (4): 312, 2000 (Jovet-Ast 2000).
*** Riccia cubensis S.W.Arnell, Bryologist 61 (2): 142, 1958 (Arnell 1958c).
494
***
***
***
***
***
***
***
***
***
**
***
***
***
*
***
***
***
***
**
***
**
***
***
***
***
***
***
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Riccia discolor Lehm. et Lindenb., Nov. Stirp. Pug. 4: 1, 1832 (Lehmann 1832).
Riccia ekmanii S.W.Arnell, Bryologist 61 (2): 143, 1958 (Arnell 1958c).
Riccia elliottii Steph., Bull. Herb. Boissier 6 (4): 324 (16), 1898 (Stephani 1898a).
Riccia enyae Jovet-Ast, Cryptog. Bryol. Lichénol. 12 (3): 230, 1991 (Jovet-Ast 1991).
Riccia erythrocarpa Jovet-Ast, Cryptog. Bryol. Lichénol. 12 (3): 257, 1991 (JovetAst 1991).
Riccia fruchartii Steph., Bull. Herb. Boissier 6 (4): 330 (22), 1898 (Stephani
1898a).
Riccia gangetica Ahmad ex L.Söderstr., A.Hagborg et von Konrat, Phytotaxa 65:
57, 2012 (Söderström et al. 2012f). Based on: Riccia gangetica Ahmad, Curr. Sci.
11 (11): 433, 1942 (Ahmad 1942), nom. inval.
Riccia grandis Nees, Fl. Bras. (Martius) 1 (1): 300, 1833 (Nees 1833a).
Riccia helenae Jovet-Ast, J. Hattori Bot. Lab. 74: 96, 1993 (Jovet-Ast 1993a).
Riccia hirta (Austin) Underw., Bot. Gaz. 19 (7): 274, 1894 (Underwood 1894).
Bas.: Riccia arvensis var. hirta Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 232,
1869 (Austin 1869).
Riccia horrida Jovet-Ast, Cryptog. Bryol. Lichénol. 12 (3): 226, 1991 (Jovet-Ast 1991).
Riccia hortorum Bory, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur.
18 (1): 435, 1836 [1837] (Lindenberg 1836).
Riccia howellii M.Howe, Proc. Calif. Acad. Sci. (ser. 4) 21 (17): 202, 1934 (Howe
1934).
Riccia ianthina Jovet-Ast, Rev. Bryol. Lichénol. 44 (4): 418, 1978 (Jovet-Ast
1978). 450
Riccia inlexa Taylor, London J. Bot. 5: 417, 1846 (Taylor 1846b).
Riccia iodocheila M.Howe, Proc. Calif. Acad. Sci. (ser. 4) 21 (17): 200, 1934
(Howe 1934).
Riccia lanceolata Steph., Hedwigia 27 (3/4): 110, 1888 (Stephani 1888d).
Riccia lindmanii Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 23 (III, 2):
29, 1897 (Stephani 1897a).
Riccia macallisteri M.Howe, Bryologist 20 (3): 35, 1917 (Howe 1917).
Riccia macrospora Steph., Bull. Herb. Boissier 6 (4): 328 (20), 1898 (Stephani 1898a).
Riccia mamillata Trab. ex Steph., Rev. Bryol. 16 (5): 65, 1889 (Stephani 1889b).
Riccia mauryana Steph., Bull. Herb. Boissier 6 (4): 327 (19), 1898 (Stephani 1898a).
Riccia olgensis Na-halang, Brunonia 3 (1): 100, 1980 (Na-halang 1980).
Riccia planobiconvexa Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 23 (III,
2): 29, 1897 (Stephani 1897a).
Riccia ridleyi A.Gepp, J. Linn. Soc., Bot. 27 (181): 74, 1890 (Gepp 1890).
Riccia sanguineisporis Jovet-Ast, Cryptog. Bryol. Lichénol. 12 (3): 253, 1991
(Jovet-Ast 1991).
Riccia squamata Nees, Fl. Bras. (Martius) 1 (1): 302, 1833 (Nees 1833a).
450 Riccia ianthina seems identical to glabrous phases of Riccia atromarginata (Schuster 1992d). It is related to Riccia iodocheila and Riccia violacea (Bischler-Causse et al. 2005), but it is only known from
the type specimen.
World checklist of hornworts and liverworts
495
*** Riccia subdepilata Jovet-Ast, Cryptog. Bryol. Lichénol. 12 (3): 228, 1991 (Jovet-Ast 1991).
*** Riccia subplana Steph., Symb. Antill. (Urban) 3 (2): 275, 1902 (Stephani 1902e).
*** Riccia taeniiformis Jovet-Ast, Cryptog. Bryol. Lichénol. 12 (3): 270, 1991 (Jovet-Ast 1991).
*** Riccia viannae Jovet-Ast, Cryptog. Bryol. Lichénol. 12 (3): 261, 1991 (Jovet-Ast 1991).
*** Riccia vitalii Jovet-Ast, Mem. New York Bot. Gard. 45: 285, 1987 (Jovet-Ast 1987).
*** Riccia weinionis Steph., Bull. Herb. Boissier 6 (4): 326 (18), 1898 (Stephani 1898a).
** sect. Pilifer O.H.Volk, Mitt. Bot. Staatssamml. München 19: 455, 1983 (Volk 1983).
*** Riccia alatospora O.H.Volk et Perold, Bothalia 15 (3/4): 534, 1985 (Volk and
Perold 1985).
*** Riccia albomarginata Bisch. ex C.Krauss, Flora 29 (9): 135, 1846 (Krauss 1846).
*** Riccia albovestita O.H.Volk, Mitt. Bot. Staatssamml. München 17: 245, 1981
(Volk 1981).
*** Riccia ampullacea Perold, Bothalia 20 (2): 168, 1990 (Perold 1990e).
*** Riccia concava Bisch. ex C.Krauss, Flora 29 (9): 135, 1846 (Krauss 1846).
*** Riccia elongata Perold, Bothalia 20 (2): 167, 1990 (Perold 1990e).
*** Riccia furfuracea Perold, Bothalia 20 (2): 176, 1990 (Perold 1990b).
*** Riccia hantamensis Perold, Bothalia 19 (2): 157, 1989 (Perold 1989b).
*** Riccia hirsuta O.H.Volk et Perold, Bothalia 16 (2): 187, 1986 (Volk and Perold 1986a).
*** Riccia namaquensis Perold, Bothalia 20 (2): 180, 1990 (Perold 1990b).
*** Riccia parvoareolata O.H.Volk et Perold, Bothalia 15 (1/2): 117, 1984 (Volk and
Perold 1984).
*** Riccia pulveracea Perold, Bothalia 20 (2): 185, 1990 (Perold 1990c).
*** Riccia radicosa Pearson, Natuurw. Tijdschr. 4 (5/6): 142, 1922 (Pearson 1922a).
*** Riccia simii Perold, Bothalia 20 (1): 36, 1990 (Perold 1990a).
*** Riccia trachyglossa Perold, Bothalia 20 (2): 172, 1990 (Perold 1990e).
*** Riccia villosa Steph., Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr. 88: 724,
1913 (Stephani 1913b).
*** Riccia vitrea Perold, Bothalia 20 (2): 178, 1990 (Perold 1990b).
** sect. Riccia
*** Riccia albolimbata S.W.Arnell, Mitt. Bot. Staatssamml. München 2 (16): 264,
1957 (Arnell 1957c).
*** Riccia alboporosa Perold, Bothalia 19 (1): 12, 1989 (Perold 1989c).
*** Riccia albornata O.H.Volk et Perold, Bothalia 18 (2): 160, 1988 (Volk et al.
1988).
*** Riccia angolensis Steph., Bull. Herb. Boissier 6 (4): 323 (15), 1898 (Stephani 1898a).
*** Riccia argenteolimbata O.H.Volk et Perold, Bothalia 18 (2): 155, 1988 (Volk et al.
1988).
** Riccia atlantica Sérgio et Perold, J. Bryol. 17 (1): 127, 1992 (Sérgio and Perold 1992).
*** Riccia atromarginata Levier, Nuovo Giorn. Bot. Ital. 21 (2): 291, 1889 (Martelli 1889).
496
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Riccia atromarginata var. jovet-astiae Rauh et Buchloh, Rev. Bryol. Lichénol. 30
(1/2): 77, 1961 (Rauh and Buchloh 1961).
*** Riccia atropurpurea Sim, Trans. Roy. Soc. South Africa 15 (1): 11, 1926 (Sim 1926).
*** Riccia beyrichiana Hampe, Nov. Stirp. Pug. 7: 1, 1838 (Lehmann 1838).
*** Riccia bicarinata Lindb., Rev. Bryol. 4 (3): 41, 1877 (Lindberg 1877d).
*** Riccia bicolorata Perold, Bothalia 20 (2): 188, 1990 (Perold 1990c).
*** Riccia bifurca Hofm., Deutschl. Fl., heil 2 (Hofm.): 95, 1795 [1796] (Hofmann
1795).
*** Riccia billardierei Mont. et Nees, Syn. Hepat. 4: 602, 1846 (Gottsche et al. 1846).
*** Riccia breidleri Jur. ex Steph., Hedwigia 24 (1): 6, 1885 (Stephani 1885e).
*** Riccia californica Austin, Bull. Torrey Bot. Club 6 (7): 46, 1875 (Austin 1875c).
*** Riccia campbelliana M.Howe, Mem. Torrey Bot. Club 7: 26, 1899 (Howe 1899).
*** Riccia ciliata Hofm., Deutschl. Fl., heil 2 (Hofm.): 95, 1795 [1796] (Hofmann
1795).
*** Riccia ciliifera Link, Syn. hepat. eur: 119, 1829 (Lindenberg 1829).
* Riccia congoana Steph., Bull. Herb. Boissier 6 (4): 328 (20), 1898 (Stephani 1898a). 451
*** Riccia crinita Taylor, London J. Bot. 5: 415, 1846 (Taylor 1846b).
*** Riccia crozalsii Levier, Rev. Bryol. 29 (4): 73, 1902 (Levier 1902).
* Riccia crustata Trab., Bull. Soc. Hist. Nat. Afrique N. 7: 87, 1916 (Trabut 1916). 452
** Riccia dictyospora M.Howe, Bull. Torrey Bot. Club 28 (3): 163, 1901 (Howe 1901a).
*** Riccia glauca L., Sp. Pl. 1: 1139, 1753 (Linnaeus 1753).
** Riccia glauca var. ciliaris Warnst., Verh. Bot. Vereins Prov. Brandenburg 27 (1):
87, 1886 (Warnstorf 1886).
** Riccia gothica Damsh. et Hallingb., Lindbergia 12 (2/3): 100, 1986 [1987] (Damsholt and Hallingbäck 1986).
*** Riccia gougetiana Durieu et Mont., Ann. Sci. Nat. Bot. (sér. 3) 11: 35, 1849 (Montagne 1849).
** Riccia gougetiana var. armatissima Levier ex Müll.Frib., Lebermoose 1 (3): 161,
1907 (Müller 1907b).
*** Riccia lamellosa Raddi, Opusc. Sci. 2 (6): 351, 1818 (Raddi 1818b).
*** Riccia ligula Steph., Bull. Herb. Boissier 6 (4): 315 (7), 1898 (Stephani 1898a).
*** Riccia limbata Bisch. ex C.Krauss, Flora 29 (9): 135, 1846 (Krauss 1846).
*** Riccia macrocarpa Levier, Bull. Soc. Bot. Ital. 1894: 114, 1894 (Levier 1894).
*** Riccia mammifera O.H.Volk et Perold, Bothalia 16 (2): 176, 1986 (Volk and Perold
1986b).
*** Riccia melitensis C.Massal., Bull. Soc. Bot. Ital. 1913 (2/3): 52, 1913 (Massalongo
1913).
*** Riccia michelii Raddi, Opusc. Sci. 2 (6): 352, 1818 (Raddi 1818b).
*** Riccia microciliata O.H.Volk et Perold, Bothalia 16 (2): 173, 1986 (Volk and
Perold 1986b).
451 Riccia congoana is possibly conspeciic with Riccia billardierei (Perold 1989a).
452 Riccia crustata may be conspeciic with Riccia albida (Jovet-Ast 1986).
World checklist of hornworts and liverworts
497
*** Riccia montana Perold, Bothalia 19 (1): 9, 1989 (Perold 1989c).
*** Riccia natalensis Sim, Trans. Roy. Soc. South Africa 15 (1): 9, 1926 (Sim 1926).
*** Riccia nigrella DC., Fl. Franç. (DC. & Lamarck), 5 (6): 193, 1815 (De Candolle
and Lamarck 1815).
*** Riccia okahandjana S.W.Arnell, Mitt. Bot. Staatssamml. München 2 (16): 268,
1957 (Arnell 1957c).
** Riccia ozarkiana McGregor, Bryologist 63 (1): 30, 1960 (McGregor 1960).
*** Riccia papillosa Moris, Stirp. Sard. Elench.: 18, 1829 (Moris 1829).
*** Riccia pottsiana Sim, Trans. Roy. Soc. South Africa 15 (1): 10, 1926 (Sim 1926).
*** Riccia rosea O.H.Volk et Perold, Bothalia 16 (2): 181, 1986 (Volk and Perold 1986d).
* Riccia runssorensis Steph., Bull. Herb. Boissier 6 (4): 330 (22), 1898 (Stephani
1898a). 453
*** Riccia sommieri Levier, Isola Giglio: 119, 1900 (Bottini et al. 1900).
*** Riccia sorocarpa Bisch., Bem. Leberm.: 145, 1835 (Bischof 1835).
** Riccia sorocarpa var. heegii Schifn., Hedwigia 53 (1/2): 36, 1912 (Schifner 1912a).
*** Riccia subbifurca Warnst. ex Croz., Rev. Bryol. 30 (4): 62, 1903 (Crozals 1903b).
** Riccia tenella D.L.Jacobs, Bryologist 52 (4): 168, 1949 [1950] (Jacobs 1949).
*** Riccia trabutiana Steph., Rev. Bryol. 16 (5): 65, 1889 (Stephani 1889b).
*** Riccia violacea M.Howe, Ann. Missouri Bot. Gard. 2 (1/2): 51, 1915 (Britton 1915).
*** Riccia violacea var. laevis Jovet-Ast, Cryptog. Bryol. Lichénol. 10 (2): 100, 1989
(Jovet-Ast 1989).
*** Riccia warnstorii Limpr. ex Warnst., Verh. Bot. Vereins Prov. Brandenburg 27
(1): 85, 1886 (Warnstorf 1886).
** subg. Ricciella (A.Braun) Boulay, Musc. France 2: 198, 1904 (Boulay 1904).
Bas.: Ricciella A.Braun, Flora 4 (2): 756, 1821 (Braun 1821).
*** Riccia cancellata Taylor, London J. Bot. 5: 414, 1846 (Taylor 1846b).
*** Riccia cincta Jovet-Ast, Cryptog. Bryol. 21 (4): 303, 2000 (Jovet-Ast 2000).
*** Riccia cruciata Kashyap, J. Bombay Nat. Hist. Soc. 24 (2): 349, 1916 (Kashyap 1916).
*** Riccia eburnea Jovet-Ast, Cryptog. Bryol. 21 (4): 300, 2000 (Jovet-Ast 2000).
*** Riccia hasskarliana Steph., Bull. Herb. Boissier 6 (5): 374 (49), 1898 (Stephani 1898b).
*** Riccia junghuhniana Nees et Lindenb., Syn. Hepat. 4: 609, 1846 (Gottsche et al.
1846).
** Riccia junghuhniana var. simplex Schifn., Hep. Fl. Buitenzorg: 14, 1900 (Schifner
1900a).
*** Riccia mangalorica Ahmad ex Jovet-Ast, Cryptog. Bryol. 24 (3): 223, 2003 (JovetAst 2003). Based on: Riccia mangalorica Ahmad, Curr. Sci. 11 (11): 433, 1942
(Ahmad 1942), nom. inval.
*** Riccia multiida (Steph.) Steph., Bull. Herb. Boissier 6 (5): 365 (40), 1898 (Stephani
1898b). Bas.: Ricciella multiida Steph., Hedwigia 28 (4): 273, 1889 (Stephani 1889c).
453 Riccia runssorensis is possibly conspeciic with Riccia macrospora (Seppelt 1998).
498
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
** Riccia polycarpa (Trab.) Jelenc, Bull. Trimestriel Geogr. Archeol. Oran 73 (228):
88, 1950 (Jelenc 1950). Bas.: Ricciella polycarpa Trab., Mém. Soc. Hist. Nat. Afrique N. 3: 36, 1933 (Maire 1933).
*** Riccia porosa Taylor, London J. Bot. 5: 416, 1846 (Taylor 1846b).
*** Riccia pullulans Jovet-Ast, Cryptog. Bryol. Lichénol. 18 (3): 183, 1997 (Jovet-Ast 1997).
** sect. Ricciella (A.Braun) Bisch., Bem. Leberm.: 160, 1835 (Bischof 1835). Bas.:
Ricciella A.Braun, Flora 4 (2): 756, 1821 (Braun 1821).
* Riccia bahiensis Steph., Bull. Herb. Boissier 6 (5): 375 (50), 1898 (Stephani 1898b). 454
*** Riccia canaliculata Hofm., Deutschl. Fl., heil 2 (Hofm.): 96, 1795 [1796]
(Hofmann 1795).
*** Riccia chiapasensis Jovet-Ast, Cryptog. Bryol. Lichénol. 14 (3): 235, 1993 (Jovet-Ast 1993b).
*** Riccia crassifrons Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 570, 1885
(Spruce 1885).
*** Riccia duplex Lorb. ex Müll.Frib., Hedwigia 80 (1/2): 100, 1941 (Müller 1941).
** Riccia duplex var. megaspora Na-halang, Brunonia 3 (1): 128, 1980 (Na-halang
1980).
*** Riccia dussiana Steph., Symb. Antill. (Urban) 3 (2): 275, 1902 (Stephani 1902e).
*** Riccia luitans L., Sp. Pl. 1: 1139, 1753 (Linnaeus 1753).
*** Riccia frostii Austin, Bull. Torrey Bot. Club 6 (3): 17, 1875 (Austin 1875b).
* Riccia frostii var. crystallinoides Schifn., Ann. K. K. Naturhist. Hofmus. 27: 503,
1913 (Schifner 1913).
*** Riccia geissleriana Jovet-Ast, Cryptog. Bryol. Lichénol. 14 (3): 236, 1993 (JovetAst 1993b).
*** Riccia hegewaldiana Jovet-Ast, Cryptog. Bryol. Lichénol. 14 (3): 238, 1993 (JovetAst 1993b).
*** Riccia huebeneriana Lindenb., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol.
Nat. Cur. 18 (1): 504d, 1836 [1837] (Lindenberg 1836).
* Riccia huebeneriana subsp. sullivantii (Austin) R.M.Schust., Hepat. Anthocerotae
N. Amer. 6: 457, 1992 (Schuster 1992d). Bas.: Riccia sullivantii Austin, Proc.
Acad. Nat. Sci. Philadelphia 21: 233, 1869 (Austin 1869).
*** Riccia jovet-astiae E.Vianna, Bol. Inst. Bioci. Univ. Fed. Rio Grande do Sul 38:
165, 1985 (Vianna 1985).
*** Riccia limicola Jovet-Ast, Rev. Bryol. Lichénol. 44 (4): 422, 1978 (Jovet-Ast 1978).
*** Riccia paraguayensis Spruce, Bull. Soc. Bot. France (Congr. Bot.) 36: cxcvi, 1889
[1890] (Spruce 1889).
*** Riccia paranaensis Hässel, Opera Lilloana 7: 228, 1962 [1963] (Hässel 1962). 455
*** Riccia perennis Steph., Bull. Herb. Boissier 6 (5): 372 (47), 1898 (Stephani 1898b).
454 Riccia bahiensis is conspeciic with Riccia cavernosa in Jovet-Ast (1965), but it was accepted by
Bischler-Causse et al. (2005). It is only known from the type specimen.
455 Riccia paranaensis may be conspeciic with Riccia huebeneriana (Schuster 1992d), but it was accepted
by Hässel and Rubies (2009).
World checklist of hornworts and liverworts
***
**
**
***
499
Riccia purpurascens Lehm., Linnaea 4: 371, 1829 (Lehmann 1829).
Riccia rhenana Lorb. ex Müll.Frib., Hedwigia 80 (1/2): 94, 1941 (Müller 1941).
Riccia rhenana var. violacea M.F.Boiko, Chornom. Bot. J. 7: 93, 2011 (Boiko 2011).
Riccia stricta (Lindenb.) Perold, Bothalia 20 (2): 197, 1990 (Perold 1990d). Bas.:
Riccia luitans var. stricta Lindenb., Nova Acta Phys.-Med. Acad. Caes. Leop.Carol. Nat. Cur. 18 (1): 444, 1836 [1837] (Lindenberg 1836).
** sect. Spongodes Nees, Naturgesch. Eur. Leberm. 4: 391, 1838 (Nees 1838a).
*** Riccia bullosa Link, Syn. hepat. eur: 119, 1829 (Lindenberg 1829).
*** Riccia cavernosa Hofm., Deutschl. Fl., heil 2 (Hofm.): 95, 1795 [1796] (Hofmann 1795).
*** Riccia crystallina L., Sp. Pl. 1: 1138, 1753 (Linnaeus 1753).
*** Riccia cupulifera A.V.Duthie, Trans. Roy. Soc. South Africa 24 (2): 116, 1936
(Duthie and Garside 1936).
*** Riccia garsidei Sim, Trans. Roy. Soc. South Africa 15 (1): 13, 1926 (Sim 1926).
*** Riccia moenkemeyeri Steph., Bot. Jahrb. Syst. 8 (2): 95, 1886 (Stephani 1886d).
*** Riccia rubricollis Garside et A.V.Duthie ex Perold, Bothalia 21 (1): 51, 1991 (Perold
1991a).
*** Riccia volkii S.W.Arnell, Mitt. Bot. Staatssamml. München 2 (16): 271, 1957
(Arnell 1957c).
*** Riccia vulcanicola Eb.Fisch., Trop. Bryol. 8: 70, 1993 (Fischer 1993).
** subg. hallocarpus (Lindb.) Jovet-Ast, Cryptog. Bryol. Lichénol. 14 (3): 220,
1993 (Jovet-Ast 1993b). Bas.: hallocarpus Lindb., Not. Sällsk. Fauna Fl. Fenn.
Förh. 13: 377, 1874 (Lindberg 1874a).
*** Riccia curtisii (Austin) Austin, Bull. Torrey Bot. Club 6 (52): 305, 1879 (Austin
1879). Bas.: Cryptocarpus curtisii Austin, Proc. Acad. Nat. Sci. Philadelphia 21:
231, 1869 (Austin 1869).
** Riccia leptothallus R.M.Schust., J. Hattori Bot. Lab. 71: 271, 1992 (Schuster 1992c).
*** Riccia perssonii Sultan Khan, Svensk Bot. Tidskr. 49 (3): 433, 1955 (Kahn 1955).
** subg. Triseriata Jovet-Ast, Cryptog. Bryol. Lichénol. 17 (2): 132, 1996 (JovetAst 1996).
*** Riccia singularis Jovet-Ast, Cryptog. Bryol. Lichénol. 17 (2): 127, 1996 (Jovet-Ast
1996).
Incertae sedis
** Riccia abuensis Bapna, Trans. Brit. Bryol. Soc. 4 (2): 249, 1962 (Bapna 1962).
** Riccia acutisulca Steph., Sp. Hepat. (Stephani) 6: 1, 1917 (Stephani 1917a).
* Riccia amboinensis Schifn., Leberm., Forschungsr. Gazelle 4 (4): 44, 1890 (Schifner
1890).
** Riccia aravalliensis Pandé et Udar, J. Indian Bot. Soc. 36 (3): 249, 1957 (Pandé
and Udar 1957).
500
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
* Riccia arnellii Sultan Khan, Bryologist 60 (1): 29, 1957 (Khan 1957).
** Riccia asprella Carrington et Pearson, Proc. Linn. Soc. New South Wales (ser. 2) 2
(4): 1059, 1888 (Carrington and Pearson 1888a).
* Riccia asservanda De Not. ex Lamothe, Rech. Anat. Taxinom. Gamét. Marchantiales: 138, 1919 (Lamothe 1919).
** Riccia attenuata Pandé, Proc. Natl. Inst. Sci. India B 25 (2): 92, 1959 (Pandé and
Udar 1959).
* Riccia balansae Steph., Bull. Herb. Boissier 6 (5): 370 (45), 1898 (Stephani 1898b).
* Riccia bialbistrata Hässel, Opera Lilloana 7: 243, 1962 [1963] (Hässel 1962). 456
** Riccia biokoensis Perold, Nova Hedwigia 64 (1/2): 244, 1997 (Perold 1997b).
** Riccia blackii Na-halang, Brunonia 3 (1): 81, 1980 (Na-halang 1980).
*** Riccia caroliniana Na-halang, Brunonia 3 (1): 72, 1980 (Na-halang 1980).
** Riccia cartilaginosa Steph., Hedwigia 28 (4): 272, 1889 (Stephani 1889c).
* Riccia chartacea K.I.Goebel, Organogr. Pl., ed. 2, 2 (1): 630, 1915 (Goebel 1915).
** Riccia chinensis Herzog, Symb. Sin. 5: 1, 1930 (Nicholson et al. 1930).
** Riccia collata Na-halang, Brunonia 3 (1): 122, 1980 (Na-halang 1980).
** Riccia compacta Garside, Trans. Roy. Soc. South Africa 27 (1): 17, 1939 (Duthie
and Garside 1939).
** Riccia convexa Steph., Sp. Hepat. (Stephani) 6: 2, 1917 (Stephani 1917a).
* Riccia coronata Sim, Trans. Roy. Soc. South Africa 15 (1): 9, 1926 (Sim 1926). 457
** Riccia crassa Steph., Bull. Herb. Boissier 6 (5): 376 (51), 1898 (Stephani 1898b).
** Riccia crenatodentata O.H.Volk, Nova Hedwigia 46 (1/2): 27, 1988 (Volk 1988).
** Riccia delavayi Steph., Bull. Herb. Boissier 6 (5): 367 (42), 1898 (Stephani 1898b).
** Riccia deserticola Steph., Bull. Herb. Boissier 6 (5): 373 (48), 1898 (Stephani
1898b).
*** Riccia erubescens Perold, J. Bryol. 16 (3): 371, 1991 (Perold 1991b).
** Riccia esulcata Steph., Sp. Hepat. (Stephani) 6: 2, 1917 (Stephani 1917a).
** Riccia fertilissima Steph., Sp. Hepat. (Stephani) 6: 2, 1917 (Stephani 1917a).
* Riccia gemmifera O.H.Volk, Nova Hedwigia 39: 131, 1984 (Volk 1984). 458
** Riccia grollei Udar, Curr. Sci. 34 (4): 126, 1965 (Udar 1965). Nom. nov. pro Riccia
tuberculata Pandé et Udar, Proc. Natl. Inst. Sci. India B 24 (2): 83, 1958 (Pandé
and Udar 1958), nom. illeg.
** Riccia handelii Schifn., Symb. sin. 2: 81, 1937 (Schifner 1937).
** Riccia hawaiiensis Hürl., Phytologia 61 (5): 339, 1986 (Hürlimann 1986).
** Riccia indica Udar et A.Gupta, Proc. V Indian Geophytol. Conf., Special Publ.:
307, 1984 (Udar and Gupta 1984).
** Riccia indira-gandhiensis Dabhade et A.Hasan, J. Bombay Nat. Hist. Soc. 83 (2):
400, 1986 (Dabhade and Hasan 1986).
456 Riccia bialbistrata is a doubtful taxon closely related to Riccia lindmanii (Bischler-Causse et al. 2005).
457 Riccia coronata is only known from the type specimen, which could not be traced. he description is
very brief and it is suspected that it refers to smaller plants of Riccia natalensis (Perold 1999a).
458 Riccia gemmifera is possibly conspeciic with Riccia atropurpurea (Perold 1999a).
World checklist of hornworts and liverworts
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501
Riccia intermedia Roum., Mém. Soc. Arts Sci. Carcassonne 5: 198, 1888 (Roumeguère 1888). 459
Riccia jodhpurensis Bapna, Bot. Not. 114 (2): 181, 1961 (Bapna 1961).
Riccia kirinensis C.Gao et K.C.Chang, Acta Phytotax. Sin. 16 (4): 117, 1978 (Gao
and Chang 1978).
Riccia laxisquamata (Steph.) Steph., Bull. Herb. Boissier 6 (5): 371 (46), 1898
(Stephani 1898b). Bas.: Ricciella laxisquamata Steph., Bot. Jahrb. Syst. 20 (3):
299, 1895 (Stephani 1895a).
Riccia liaoningensis C.Gao et K.C.Chang, Acta Phytotax. Sin. 16 (4): 113, 1978
(Gao and Chang 1978).
Riccia linearis (Schifn.) Steph., Bull. Herb. Boissier 6 (5): 371 (46), 1898 (Stephani 1898b). Bas.: Ricciella linearis Schifn., Leberm., Forschungsr. Gazelle 4 (4): 43,
1890 (Schifner 1890).
Riccia luticola Na-halang, Brunonia 3 (1): 123, 1980 (Na-halang 1980).
Riccia mamrensis Perold, Cryptog. Bryol. 26 (1): 68, 2005 (Perold 2005).
Riccia marginata Lindb., Meddel. Soc. Fauna Fl. Fenn. 1: 106, 1876 [1877] (Lindberg 1876b). 460
Riccia melanospora Kashyap, Liverworts W. Himal. 1: 94, 1929 (Kashyap 1929).
Riccia miyakeana Schifn., Österr. Bot. Z. 49 (11): 386, 1899 (Schifner 1899c).
Riccia muscicola Steph., Hedwigia 24 (1): 4, 1885 (Stephani 1885e).
Riccia nigerica E.W.Jones, Trans. Brit. Bryol. Soc. 3 (2): 225, 1957 (Jones 1957a).
Riccia nigrescens Mont., Voy. Amér. Mérid., Bot. 7 (1): 15, 1839 (Montagne 1839b). 461
Riccia nipponica S.Hatt., J. Hattori Bot. Lab. 9: 38, 1953 (Shimizu and Hattori 1953a).
Riccia novo-hannoverana Schifn., Leberm., Forschungsr. Gazelle 4 (4): 44, 1890
(Schifner 1890).
Riccia numeensis Steph., Bull. Herb. Boissier 6 (4): 343 (35), 1898 (Stephani 1898a).
Riccia obtusa Meijer, J. Hattori Bot. Lab. 20: 113, 1958 (Meijer 1958). 462
Riccia oerstediana Lindenb. et Hampe, Linnaea 24 (3): 304, 1851 [1852] (Hampe
1851b).
Riccia pandei Udar, J. Indian Bot. Soc. 38 (1): 149, 1959 (Udar 1959).
Riccia papillispora Steph., Bull. Herb. Boissier 6 (4): 334 (26), 1898 (Stephani 1898a).
Riccia papulosa (Steph.) Steph., Bull. Herb. Boissier 6 (5): 377 (52), 1898 (Stephani
1898b). Bas.: Ricciella papulosa Steph., Hedwigia 28 (4): 273, 1889 (Stephani 1889c).
Riccia papulosa var. variabilis Na-halang, Brunonia 3 (1): 112, 1980 (Na-halang
1980).
Riccia pathankotensis Kashyap, J. Bombay Nat. Hist. Soc. 24 (2): 349, 1916
(Kashyap 1916).
459 Riccia intermedia (type from France) has neither been recognized in any recent treatment nor synonymized. he name may have priority once its identity is determined.
460 Riccia marginata is possibly conspeciic with Riccia beyrichiana (Potemkin and Ahti 2012).
461 Riccia nigrescens is an Anthoceros species (Hässel and Rubies 2009).
462 Riccia obtusa may be conspeciic with Riccia gangetica (Söderström et al. 2010a).
502
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Riccia perthiana Steph. ex K.I.Goebel, Organogr. Pl., ed. 2, 2 (1): 630, 1915
(Goebel 1915).
Riccia prominens Meijer, J. Hattori Bot. Lab. 20: 111, 1958 (Meijer 1958). 463
Riccia pseudoluitans C.Gao et K.C.Chang, Acta Phytotax. Sin. 16 (4): 116, 1978
(Gao and Chang 1978).
Riccia pubescens S.Hatt., Nat. Sci. Mus. 14 (6): 141, 1943 (Hattori 1943a).
Riccia radiata Perold, Bothalia 34 (1): 23, 2004 (Perold 2004).
Riccia rechingeri Steph., Akad. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr. 81:
288, 1907 (Stephani 1907a).
Riccia reticulatula Udar, Bull. Bot. Soc. Univ. Saugar 13: 49, 1961 (Udar 1961).
Riccia rorida Na-halang, Brunonia 3 (1): 101, 1980 (Na-halang 1980).
Riccia saharensis Steph. ex Jovet-Ast, Rev. Bryol. Lichénol. 26 (1/2): 62, 1957
(Gillett and Jovet-Ast 1957). 464
Riccia satoi S.Hatt., Bot. Mag. (Tokyo) 62 (733/734): 109, 1949 (Hattori 1949).
Riccia schroederi Steph., 52 (5): 304, 1912 (Stephani 1912a).
Riccia schweinfurthii Steph., Bull. Herb. Boissier 6 (4): 339 (31), 1898 (Stephani
1898a).
Riccia sibayenii Perold, Bothalia 31 (1): 151, 2001 (Perold 2001a).
Riccia somaliensis Perold, J. Bryol. 16 (3): 367, 1991 (Perold 1991b).
Riccia spongiosula Na-halang, Brunonia 3 (1): 113, 1980 (Na-halang 1980).
Riccia subtilis (Steph.) Steph., Bull. Herb. Boissier 6 (5): 364 (39), 1898 (Stephani
1898b). Bas.: Ricciella subtilis Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl.
23 (III, 2): 31, 1897 (Stephani 1897a).
Riccia sumatrana Meijer, J. Hattori Bot. Lab. 20: 114, 1958 (Meijer 1958).
Riccia symoensii Vanden Berghen, Explor. Hydrobiol. Lac Bangweolo Luapula:
191, 1972 (Vanden Berghen 1972b).
Riccia tasmanica Steph. ex Rodway, Tasm. Bryoph.: 4, 1917 (Rodway 1917b).
Riccia tomentosa O.H.Volk et Perold, Bothalia 20 (1): 25, 1990 (Volk and Perold 1990).
Riccia treubiana Steph., Bull. Herb. Boissier 6 (4): 323 (15), 1898 (Stephani 1898a).
Riccia treubiana var. subrubescens Schifn., Hep. Fl. Buitenzorg: 16, 1900 (Schifner
1900a).
Riccia triangularis Steph., Bull. Mus. Natl. Hist. Nat. 18 (2): 116, 1912 (Corbière
1912).
Riccia tuberculata Poir., Encycl. (Lamarck) 6: 199, 1804 (Lamarck and Poiret 1804). 465
Riccia udarii Kanwal, J. Indian Bot. Soc. 58 (3): 282, 1979 (Kanwal 1979).
Riccia velenovskyi Kavina, Arch. Přír. Výzk. Čech 16 (2): 75, 1915 (Kavina 1915). 466
463 Riccia prominens may be conspeciic with Riccia junghuhniana (Söderström et al. 2010a).
464 Riccia saharensis is possibly conspeciic with Riccia argenteolimbata (Perold 1995).
465 Riccia tuberculata (type from France) has neither been recognized in any recent European treatment
nor synonymized. he name may have priority once its identity is determined.
466 Riccia velenovskyi (type from Czech Republic) has neither been recognized in any recent European
treatment nor synonymized. he name may have priority once its identity is determined.
World checklist of hornworts and liverworts
503
** Riccia velimalaiana A.E.D.Daniels et P.Daniel, Bull. Bot. Surv. India 44 (1/4):
139, 2002 [2003] (Daniels and Daniel 2002).
** Riccia victoriensis Steph., Bull. Herb. Boissier 6 (5): 370, 1898 (Stephani 1898b).
** Riccia weymouthiana Steph. ex Rodway, Tasm. Bryoph.: 5, 1917 (Rodway 1917b).
** Riccia wichurae Steph., Bull. Herb. Boissier 6 (4): 330 (22), 1898 (Stephani 1898a).
** Ricciocarpos Corda, Gen. hepat.: 651, 1829 (Corda 1829).
*** Ricciocarpos natans (L.) Corda, Gen. hepat.: 651, 1829 (Corda 1829). Bas.: Riccia
natans L., Syst. Nat., ed. 10., 2: 1339, 1759 (Linnaeus 1759).
***
Targioniaceae Dumort.
by D.G. Long
*** Targionia L., Sp. Pl. 1: 1136, 1753 (Linnaeus 1753).
** subg. Prototargionia R.M.Schust., Hepat. Anthocerotae N. Amer. 6: 69, 1992
(Schuster 1992d).
** Targionia stellaris (Müll.Frib.) Hässel, Opera Lilloana 7: 74, 1962 [1963] (Hässel
1962). Bas.: Grimaldia stellaris Müll.Frib., Feddes Repert. Spec. Nov. Regni Veg.
58: 61, 1955 (Müller 1955).
** subg. Targionia
*** Targionia hypophylla L., Sp. Pl. 1: 1136, 1753 (Linnaeus 1753).
** Targionia hypophylla subsp. linealis W.Frey et Kürschner, Nova Hedwigia 57
(1/2): 127, 1993 (Frey and Kürschner 1993).
** Targionia lorbeeriana Müll.Frib., Hedwigia 79 (1/2): 78, 1940 (Müller 1940).
*
*
*
*
*
Incertae sedis
Targionia dioica Schifn., Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl.
67: 154, 1898 (Schifner 1898a). 467
Targionia elongata Bisch., Syn. Hepat. 4: 576, 1846 (Gottsche et al. 1846).
Targionia iorii Gola, Ann. Bot. (Rome) 13 (1): 62, 1914 (Gola 1914a).
Targionia formosica Horik., J. Jap. Bot. 11: 499, 1935 (Horikawa 1935).
Targionia indica Udar et A.Gupta, Geophytology 13 (1): 83, 1983 (Udar and
Gupta 1983).
467 Targionia dioica is probably conspeciic with Targionia hypophylla (Söderström et al. 2010a).
504
***
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Wiesnerellaceae Inoue
by D.G.Long
*** Wiesnerella Schifn., Österr. Bot. Z. 46 (3): 86, 1896 (Schifner 1896b).
*** Wiesnerella denudata (Mitt.) Steph., Bull. Herb. Boissier 7 (5): 382 (154), 1899
(Stephani 1899c). Bas.: Dumortiera denudata Mitt., J. Proc. Linn. Soc., Bot. 5
(18): 125, 1860 [1861] (Mitten 1860c).
* Wiesnerella fasciaria C.Gao et K.C.Chang, Acta Bot. Yunnan. 3 (4): 391, 1981
(Gao et al. 1981).
Neohodgsoniales D.G.Long
***
Neohodgsoniaceae D.G.Long
by D.G. Long
*** Neohodgsonia Perss., Bot. Not. 107 (1): 40, 1954 (Persson 1954). Nom. nov. pro
Hodgsonia Perss., Hodgsonia Leal. Stockholm: 1, 1953 (Persson 1953).
*** Neohodgsonia mirabilis (Perss.) Perss., Bot. Not. 107 (1): 40, 1954 (Persson 1954).
Bas.: Hodgsonia mirabilis Perss., Hodgsonia Leal. Stockholm: 1, 1953 (Persson
1953).
Sphaerocarpales Cavers
***
Monocarpaceae D.J.Carr ex Schelpe
by D.G. Long
*** Monocarpus D.J.Carr, Austral. J. Bot. 4 (2): 176, 1956 (Carr 1956).
*** Monocarpus sphaerocarpus D.J.Carr, Austral. J. Bot. 4 (2): 176, 1956 (Carr 1956).
***
Riellaceae Engl.
by J. G. Segarra-Moragues and F. Puche
*** Austroriella Cargill et J.Milne, Polish Bot. J. 58 (1): 72, 2013 (Cargill and Milne
2013).
*** Austroriella salta J.Milne et Cargill, Polish Bot. J. 58 (1): 72, 2013 (Cargill and
Milne 2013).
World checklist of hornworts and liverworts
505
*** Riella Mont., Ann. Sci. Nat. Bot. (sér. 3) 18: 11, 1852 (Montagne 1852).
*** subg. Riella
*** Riella alatospora Wigglesw., J. Linn. Soc., Bot. 51 (339): 317, 1937 (Wigglesworth
1937).
*** Riella americana M.Howe et Underw., Bull. Torrey Bot. Club 30 (4): 218, 1903
(Howe and Underwood 1903).
* Riella battandieri Trab., Rev. Bryol. 13 (3): 35, 1886 (Trabut 1886).
*** Riella bialata Trab., Rev. Bryol. 35 (4): 96, 1908 (Trabut 1908).
*** Riella capensis Cavers, Rev. Bryol. 30 (5): 81, 1903 (Cavers 1903).
*** Riella choconensis Hässel, Symp. Biol. Hung. 35: 341, 1987 (Hässel 1987).
* Riella cyrenaica Maire, Bull. Soc. Hist. Nat. Afrique N. 30 (5): 312, 1939 (Maire
and Weiler 1939).
*** Riella echinospora Wigglesw., J. Linn. Soc., Bot. 51 (339): 321, 1937 (Wigglesworth 1937).
* Riella gallica Balansa ex Trab., Rev. Gén. Bot. 3 (35): 450, 1891 (Trabut 1891).
*** Riella halophila Banwell, Trans. Brit. Bryol. Soc. 1 (5): 475, 1951 (Banwell 1951).
*** Riella helicophylla (Bory et Mont.) Mont., Ann. Sci. Nat. Bot. (sér. 3) 18: 12, 1852
(Montagne 1852). Bas.: Duriaea helicophylla Bory et Mont., Ann. Sci. Nat. Bot.
(sér. 3) 1: 229, 1844 (Bory and Montagne 1844).
* Riella helicophylla var. macrocarpa P.Allorge, Sched. Br. Iber. (ser. 2): 4, 1929 (Allorge 1929).
* Riella indica Steph. ex Kashyap, J. Bombay Nat. Hist. Soc. 25 (2): 279, 1917
(Kashyap 1917).
*** Riella notarisii (Mont.) Mont., Ann. Sci. Nat. Bot. (sér. 3) 18: 12, 1852 (Montagne 1852). Bas.: Sphaerocarpos notarisii Mont., Ann. Sci. Nat. Bot. (sér. 2) 9: 39,
1838 (Montagne 1838). 468
** Riella numidica Trab., Bull. Soc. Hist. Nat. Afrique N. 25 (9): 391, 1934 [1935]
(Trabut 1934).
*** Riella pampae Hässel, Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia Inst.
Nac. Invest. Ci. Nat. Bot. 5 (9): 207, 1979 (Hässel 1979).
*** Riella parisii Gottsche, Hepat. Eur., Leberm. 38-39: no. 375, 1867 (Gottsche and
Rabenhorst 1867).
*** Riella purpureospora Wigglesw., J. Linn. Soc., Bot. 51 (339): 312, 1937 (Wigglesworth 1937).
* Riella reuteri Mont., Ann. Sci. Nat. Bot. (sér. 3) 18: 12, 1852 (Montagne 1852).
* Riella sersuensis Trab., Bull. Soc. Hist. Nat. Afrique N. 25 (9): 392, 1934 [1935]
(Trabut 1934).
** Riella spiculata J.Taylor, Kew Bull. 9 (1): 45, 1954 (Taylor 1954).
468 Riella notarisii is a species complex with poorly deined taxonomic boundaries including Riella
battandieri, Riella cyrenaica, Riella gallica, Riella indica, Riella reuteri and Riella sersuensis.
506
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
*** Riella trigonospora Segarra et Puche, S. African J. Bot. 94: 175, 2014 (Segarra-Moragues and Puche 2014).
*** Riella undulata Hässel, Symp. Biol. Hung. 35: 341, 1987 (Hässel 1987).
*** subg. Trabutiella Porsild, Bot. Tidsskr. 24 (3): 327, 1902 (Porsild 1902).
*** Riella ainis M.Howe et Underw., Bull. Torrey Bot. Club 30 (4): 221, 1903
(Howe and Underwood 1903).
*** Riella cossoniana Trab., Atlas l. Alger 1: 6, 1886 (Battandier and Trabut 1886).
*** Riella echinata (Müll.Frib.) Segarra, Puche et Sabovlj., Phytotaxa 159 (3): 165,
2014 (Segarra-Moragues et al. 2014). Bas.: Riella cossoniana var. echinata Müll.
Frib., Rev. Bryol. Lichénol. 22 (3/4): 132, 1953 [1954] (Müller 1953).
*** Riella gamundiae Hässel, Rev. Bryol. Lichénol. 38 (3/4): 580, 1972 [1973] (Hässel
1972b).
*** Riella heliospora Segarra, Puche et Sabovlj., Syst. Bot. 37 (2): 315, 2012 (Segarra-Moragues et al. 2012).
*** Riella mediterranea Segarra, Puche, Sabovlj., M.Infante et Heras, Phytotaxa 159
(3): 170, 2014 (Segarra-Moragues et al. 2014).
***
Sphaerocarpaceae Heeg
by D. Long
*** Geothallus Campb., Bot. Gaz. 21 (1): 13, 1896 (Campbell 1896).
*** Geothallus tuberosus Campb., Bot. Gaz. 21 (1): 13, 1896 (Campbell 1896).
*** Sphaerocarpos Boehm., Def. gen. pl., ed. 3: 501, 1760 (Ludwig 1760).
** subg. Austrosphaerocarpos R.M.Schust., Hepat. Anthocerotae N. Amer. 5: 813,
1992 (Schuster 1992b).
*** Sphaerocarpos stipitatus Bisch. ex Lindenb., Nova Acta Phys.-Med. Acad. Caes.
Leop.-Carol. Nat. Cur. 18 (1): 504i, 1836 [1837] (Lindenberg 1836).
** subg. Sphaerocarpos
*** Sphaerocarpos michelii Bellardi, App. l. pedem.: 52, 1792 (Bellardi 1792).
** Sphaerocarpos texanus Austin, Bull. Torrey Bot. Club 6 (30): 158, 1877 (Austin 1877).
Incertae sedis
*** Sphaerocarpos cristatus M.Howe, Mem. Torrey Bot. Club 7: 66, 1899 (Howe 1899).
*** Sphaerocarpos donnellii Austin, Bull. Torrey Bot. Club 6 (30): 157, 1877 (Austin 1877).
*** Sphaerocarpos drewiae Wigglesw., Univ. Calif. Publ. Bot. 16 (3): 129, 1929 (Wigglesworth 1929).
World checklist of hornworts and liverworts
507
*** Sphaerocarpos europaeus Lorb., Jahrb. Wiss. Bot. 80: 665, 1934 (Lorbeer 1934). 469
*** Sphaerocarpos hians Haynes, Bull. Torrey Bot. Club 37 (5): 225, 1910 (Haynes 1910).
** Sphaerocarpos muccilloi E.Vianna, Lindbergia 7 (1): 58, 1981 (Vianna 1981).
Names in genera not currently accepted
he following taxa in unsupported genera are all poorly understood. We list them here
rather than making new combinations for names we do not know the status of.
Acrostolia Dumort., Recueil Observ. Jungerm.: 26, 1835 (Dumortier 1835). 470
* Acrostolia alata (Gottsche et Rabenh.) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 431, 1877 (Trevisan 1877). Bas.: Pseudoneura alata
Gottsche et Rabenh., Hepat. Eur., Leberm. 56-57: no. 560, 1873 (Gottsche and
Rabenhorst 1873b).
* Acrostolia brevifolia (Gottsche et Rabenh.) Trevis., Mem. Reale Ist. Lombardo Sci.
(Ser. 3), C. Sci. Mat. 4 (13): 431, 1877 (Trevisan 1877). Bas.: Pseudoneura brevifolia Gottsche et Rabenh., Hepat. Eur., Leberm. 56-57: no. 560, 1873 (Gottsche
and Rabenhorst 1873b).
Aphanolejeunea A.Evans, Bull. Torrey Bot. Club 38 (6): 272, 1911 (Evans 1911).
* Aphanolejeunea lancifera R.M.Schust., Phytologia 45 (5): 434, 1980 (Schuster
1980b). 471
* Aphanolejeunea minima Tixier, Ann. Fac. Sci. Yaoundé 20: 7, 1975 (Tixier 1975b). 472
Aspiromitus Steph., Sp. Hepat. (Stephani) 5: 957, 1916 (Stephani 1916b). 473
* Aspiromitus asper Schifn., Arch. Hydrobiol., suppl. 21 (3/4): 402, 1955 (Schifner
1955).
* Aspiromitus bullosus Schifn., Arch. Hydrobiol., suppl. 21 (3/4): 403, 1955
(Schifner 1955).
* Aspiromitus crenatifrons Steph., Sp. Hepat. (Stephani) 5: 968, 1916 (Stephani 1916b).
* Aspiromitus lobatus Schifn., Arch. Hydrobiol., suppl. 21 (3/4): 405, 1955
(Schifner 1955).
* Aspiromitus squamulosus Schifn., Arch. Hydrobiol., suppl. 21 (3/4): 407, 1955
(Schifner 1955).
469
470
471
472
Sphaerocarpos europaeus is clearly distinct from Sphaerocarpos texanus (Bell et al. 2013).
Acrostolia is congeneric with Riccardia, but a few taxa have neither been transferred nor synonymized.
Aphanolejeunea lancifera is a Drepanolejeunea species (Pócs et al. 2014).
Aphanolejeunea minima seems to be close to Cololejeunea gracilis based on the drawing in the original
description, but the type specimen (not available for study) seems to be very fragmentary, having only
reduced leaves, so its identity is uncertain (Pócs and Bernecker 2009).
473 Aspiromitus is congeneric with Anthoceros, but some taxa have neither been transferred nor synonymized.
508
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Crossotolejeunea (Spruce) Schifn., Hepat. (Engl.-Prantl): 127, 1893 (Schifner
1893b). Bas.: Lejeunea subg. Crossotolejeunea Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 161, 1884 (Spruce 1884). 474
* Crossotolejeunea curvifolia Steph., Hedwigia 35 (3): 75, 1896 (Stephani 1896b). 475
Eulejeunea Steph., Hedwigia 27 (2): 60, 1888 (Stephani 1888a). 476
* Eulejeunea setulosa Steph., Sp. Hepat. (Stephani) 6: 421, 1923 (Stephani 1923).
* Eulejeunea subpililoba Steph., Sp. Hepat. (Stephani) 6: 420, 1923 (Stephani 1923). 477
Euosmolejeunea (Spruce) Steph., Hedwigia 28 (3): 170, 1889 (Stephani 1889d).
Bas.: Lejeunea subg. Euosmolejeunea Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 241, 1884 (Spruce 1884). 478
* Euosmolejeunea parvistipula (Lindenb. et Gottsche) Steph., Hedwigia 29 (1): 80,
1890 (Stephani 1890a). Bas.: Lejeunea parvistipula Lindenb. et Gottsche, Syn.
Hepat. 5: 761, 1847 (Gottsche et al. 1847).
* Euosmolejeunea tenerrima (Nees) Steph., Sp. Hepat. (Stephani) 5: 589, 1914
(Stephani 1914b). Bas.: Jungermannia sordida var. tenerrima Nees, Fl. Bras. (Martius) 1 (1): 363, 1833 (Nees 1833a). 479
Fimbraria Nees, Horae Phys. Berol.: 44, 1820 (Nees 1820) nom. illeg. 480
* Fimbraria gigantea Steph., Bull. Herb. Boissier 7 (2): 93 (106), 1899 (Stephani
1899a). 481
* Fimbraria incrassata Steph., Bull. Herb. Boissier 7 (2): 87 (100), 1899 (Stephani
1899a). 482
* Fimbraria kamerunensis Steph., Sp. Hepat. (Stephani) 6: 14, 1917 (Stephani
1917a). 483
* Fimbraria pirottae Gola, Ann. Bot. (Rome) 13 (1): 64, 1914 (Gola 1914a).
474 Crossotolejeunea is congeneric with Lejeunea, but one taxon has neither been transferred nor
synonymized.
475 Crossotolejeunea curvifolia has doubtful status, the type specimen could not be found in G (ReinerDrehwald and Goda 2000).
476 Eulejeunea is congeneric with Lejeunea subg. Lejeunea, but a few taxa have neither been transferred
nor synonymized.
477 Eulejeunea subpililoba is morphologically similar to Lejeunea spiniloba.
478 Euosmolejeunea is here treated as a subgenus of Cheilolejeunea, but a few taxa have neither been transferred nor synonymized.
479 Euosmolejeunea tenerrima is a doubtful taxon described with three syntypes, two from Brazil and one
from Java. he Java material is probably not the same species as the material from Brazil.
480 Fimbraria is congeneric with Asterella, but the following taxa have neither been transferred nor synonymized, and their status is doubtful.
481 Fimbraria gigantea belongs to Asterella subg. Asterella sect. Brachyblepharis.
482 Fimbraria incrassata was provisionally placed in synonymy with Asterella abyssinica (Wigginton and
Grolle 1996), and was listed as an unassigned Asterella species by Wigginton (2009).
483 Fimbraria kamerunensis was provisionally placed in synonymy with Asterella abyssinica (Wigginton
and Grolle 1996), and was listed as an unassigned Asterella species by Wigginton (2009).
World checklist of hornworts and liverworts
509
Hygrolejeunea (Spruce) Schifn., Hepat. (Engl.-Prantl): 124, 1893 (Schifner 1893b).
Bas.: Lejeunea subg. Hygrolejeunea Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 230, 1884 (Spruce 1884). 484
* Hygrolejeunea cubensis Steph., Sp. Hepat. (Stephani) 5: 533, 1914 (Stephani 1914b).
* Hygrolejeunea harpaphylla Steph., Sp. Hepat. (Stephani) 5: 552, 1914 (Stephani
1914b).
* Hygrolejeunea paciica Steph., Sp. Hepat. (Stephani) 6: 411, 1923 (Stephani 1923).
* Hygrolejeunea parvicalycina Steph., Hedwigia 35 (3): 103, 1896 (Stephani 1896b).
* Hygrolejeunea parvistipula Steph., Sp. Hepat. (Stephani) 5: 568, 1914 (Stephani 1914b).
* Hygrolejeunea patellirostris Steph., Hedwigia 35 (3): 103, 1896 (Stephani 1896b). 485
* Hygrolejeunea staudtiana Steph., Sp. Hepat. (Stephani) 5: 528, 1914 (Stephani
1914b). 486
Hypenantron Corda, Gen. hepat.: 648, 1829 (Corda 1829). 487
* Hypenantron brachypus Steph. ex Lamothe, Rech. Anat. Taxinom. Gamét.
Marchantiales: 107, 1919 (Lamothe 1919).
* Hypenantron brasiliense Steph. ex Lamothe, Rech. Anat. Taxinom. Gamét.
Marchantiales: 104, 1919 (Lamothe 1919).
Jamesoniella (Spruce) Carrington, Cat. Brit. Moss. Hepat.: 25, 1881 (Carrington
1881). Bas.: Jungermannia subg. Jamesoniella Spruce, J. Bot. 14: 230, 1876
(Spruce 1876a). 488
* Jamesoniella convoluta Steph., Sp. Hepat. (Stephani) 6: 433, 1924 (Stephani 1924).
Kingiolejeunea H.Rob., Bryologist 70 (1): 53, 1967 (Robinson 1967). 489
* Kingiolejeunea guayanensis H.Rob., Bol. Soc. Venez. Ci. Nat. 32 (132/133): 259,
1976 (Robinson 1976b).
484 Hygrolejeunea is congeneric with Lejeunea, but some taxa have neither been transferred nor synonymized.
485 Hygrolejeunea patellirostris is conspeciic with Lejeunea acuta in Tixier (1995b) and Wigginton and
Grolle (1996), but it was kept as an unassigned Lejeunea species by Wigginton (2009).
486 Hygrolejeunea staudtiana is likely to be either Lejeunea brenanii or Lejeunea isophylla (Wigginton and
Grolle 1996). It was kept as an unassigned Lejeunea species by Wigginton (2009).
487 Hypenantron is congeneric with Asterella, but a few taxa have neither been transferred nor synonymized.
488 Jamesoniella is congeneric with Syzygiella (Feldberg et al. 2010a), but one taxon has neither been transferred nor synonymized. he holotype was burned in B (Grolle 1971b) and it is not clear where it belongs.
489 Kingiolejeunea is congeneric with Lepidolejeunea, but one taxon has neither been transferred nor
synonymized.
510
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Leptocolea (Spruce) A.Evans, Bull. Torrey Bot. Club 38 (6): 261, 1911 (Evans
1911). Bas.: Lejeunea sect. Leptocolea Spruce, Trans. & Proc. Bot. Soc. Edinburgh
15: 294, 1884 (Spruce 1884). 490
* Leptocolea sumatrana Herzog, Ann. Bryol. 5: 96, 1932 (Herzog 1932a). 491
Mastigobryum (Nees) Lindenb. et Gottsche, Syn. Hepat. 2: 214, 1845 (Gottsche et
al. 1845a) nom. illeg. Bas.: Herpetium sect. Mastigobryum Nees, Naturgesch. Eur.
Leberm. 3: 43, 1838 (Nees 1838b). 492
* Mastigobryum aberrans Steph., Sp. Hepat. (Stephani) 6: 486, 1924 (Stephani 1924).
* Mastigobryum asperum Steph., J. & Proc. Roy. Soc. New South Wales 48 (1/2):
121, 1914 (Stephani and Watts 1914).
* Mastigobryum deningeri Herzog, Beih. Bot. Centralbl. 38 (2): 322, 1921 (Herzog 1921).
* Mastigobryum karstenii Steph., Bull. Herb. Boissier (sér. 2) 8 (12): 952 (502),
1908 (Stephani 1908b).
* Mastigobryum ledermannii Steph., Sp. Hepat. (Stephani) 6: 486, 1924 (Stephani 1924).
* Mastigobryum londbergii Steph., Sp. Hepat. (Stephani) 6: 487, 1924 (Stephani 1924).
* Mastigobryum longifolium Steph., Sp. Hepat. (Stephani) 6: 487, 1924 (Stephani 1924).
* Mastigobryum minutitextum Steph., Sp. Hepat. (Stephani) 6: 487, 1924 (Stephani
1924).
* Mastigobryum multidens Steph., Sp. Hepat. (Stephani) 6: 487, 1924 (Stephani 1924).
* Mastigobryum muscicola Steph., Sp. Hepat. (Stephani) 6: 486, 1924 (Stephani 1924).
* Mastigobryum nigricans Herzog, Beih. Bot. Centralbl. 38 (2): 322, 1921 (Herzog 1921).
* Mastigobryum nipuranum Steph., Sp. Hepat. (Stephani) 6: 485, 1924 (Stephani
1924).
* Mastigobryum palmicola Steph., Sp. Hepat. (Stephani) 6: 488, 1924 (Stephani 1924).
* Mastigobryum rajanum Herzog, Mitt. Inst. Allg. Bot. Hamburg 7 (3): 190, 1931
(Herzog 1931a).
* Mastigobryum ribehanum Steph., Sp. Hepat. (Stephani) 6: 488, 1924 (Stephani 1924).
* Mastigobryum ruicaule Beauverd, Sp. Hepat. (Stephani) 6: 485, 1924 (Stephani 1924).
* Mastigobryum schraderbergii Steph., Sp. Hepat. (Stephani) 6: 489, 1924 (Stephani 1924).
* Mastigobryum squamulistipum Steph., Sp. Hepat. (Stephani) 6: 480, 1924 (Stephani
1924).
* Mastigobryum subhyalinum Steph., Sp. Hepat. (Stephani) 6: 482, 1924 (Stephani 1924).
* Mastigobryum venezuelanum Molk., Syn. hepat. jav.: 104, 1856 [1857] (Sande
Lacoste 1856b).
* Mastigobryum vermiculare Herzog, Hedwigia 66 (6): 339, 1926 (Herzog 1926).
490 Leptocolea is congeneric with Cololejeunea, but one taxon has neither been transferred nor synonymized.
491 Leptocolea sumatrana is possibly conspeciic with Cololejeunea equialbi.
492 Mastigobryum is congeneric with Bazzania, but many species have not been studied recently and thus
not transferred. It is possible that some prove to be older names of existing taxa when their identities are known. Grolle and Piippo (1984) could not study Mastigobryum ledermannii, Mastigobryum
londbergii and Mastigobryum longifolium since the types were destroyed in B.
World checklist of hornworts and liverworts
511
Nemoursia Mérat, Ann. Agric. Franç. (ser. 4) 2 (7): 10, 1840 (Mérat 1840).
* Nemoursia tuberculata Mérat, Ann. Agric. Franç. (ser. 4) 2 (7): 10, 1840 (Mérat
1840). 493
Physocolea (Spruce) Steph., Sp. Hepat. (Stephani) 5: 863, 1916 (Stephani 1916b).
Bas.: Lejeunea sect. Physocolea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15:
292, 1884 (Spruce 1884). 494
* Physocolea tambillensis (Loitl.) Steph., Sp. Hepat. (Stephani) 5: 885, 1916 (Stephani
1916b). Bas.: Lejeunea tambillensis Loitl., Diagn. pl. nov.: 20, 1894 (Loitlesberger
1894). 495
Plectocolea (Mitt.) Mitt., Fl. vit.: 405, 1871 [1873] (Mitten 1871). Bas.: Solenostoma subg. Plectocolea Mitt., J. Linn. Soc., Bot. 8 (31): 156, 1864 [1865] (Mitten
1864a). 496
* Plectocolea subamoena S.Winkl., Rev. Bryol. Lichénol. 42 (3): 821, 1976 (Winkler
1976). 497
Polyotus Gottsche, Syn. Hepat. 2: 244, 1845 (Gottsche et al. 1845a) nom. illeg. 498
* Polyotus peckianus Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 224, 1869 (Austin
1869).
Schisma Dumort., Commentat. Bot. (Dumortier): 114, 1822 (Dumortier 1822)
nom. illeg. 499
* Schisma orizabense (Gottsche) Steph., Sp. Hepat. (Stephani) 4: 19, 1909 (Stephani 1909d). Bas.: Sendtnera orizabensis Gottsche, Mexik. Leverm.: 139, 1863
(Gottsche 1863).
* Schisma uleanum Steph., Hedwigia 44 (4): 225, 1905 (Stephani 1905a).
Strepsilejeunea (Spruce) Schifn., Hepat. (Engl.-Prantl): 127, 1893 (Schifner
1893b). Bas.: Lejeunea sect. Strepsilejeunea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 168, 1884 (Spruce 1884). 500
493
494
495
496
497
498
499
500
Nemoursia tuberculata is a Conocephalum species.
Physocolea is congeneric with Cololejeunea, but one taxon has neither been transferred nor synonymized.
Physocolea tambillensis is possibly conspeciic with Myriocoleopsis minutissima.
Plectocolea is now regarded a subgenus of Solenostoma, but one taxon has neither been transferred nor
synonymized.
Plectocolea subamoena is probably conspeciic with Solenostoma amoena.
Polyotus is a superluous name for Lepidolaena, but a few taxa have neither been transferred nor synonymized.
Schisma is congeneric with Herbertus, but a few taxa have neither been transferred nor synonymized.
Strepsilejeunea is congeneric with Cheilolejeunea, but some taxa have neither been transferred nor
synonymized.
512
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Strepsilejeunea apollinea (Gottsche) Steph., Sp. Hepat. (Stephani) 5: 291, 1913
(Stephani 1913a). Bas.: Lejeunea apollinea Gottsche, Fragm. (Mueller): 64, 1880
(Gottsche 1880).
Strepsilejeunea durelii Schifn., Österr. Bot. Z. 49 (6): 206, 1899 (Schifner 1899a).
Strepsilejeunea hamatifolia Steph., Sp. Hepat. (Stephani) 6: 396, 1923 (Stephani 1923).
Strepsilejeunea lanceolata (Gottsche) Steph., Sp. Hepat. (Stephani) 5: 283, 1913
(Stephani 1913a). Bas.: Lejeunea lanceolata Gottsche, Syn. Hepat. 3: 353, 1845
(Gottsche et al. 1845b).
Strepsilejeunea muscicola Herzog, Hedwigia 74 (2): 96, 1934 (Herzog 1934a).
Strepsilejeunea novae-guineae Steph., Sp. Hepat. (Stephani) 6: 397, 1923 (Stephani
1923).
Strepsilejeunea obtusistipula Steph., Biblioth. Bot. 87 (2): 258, 1916 (Stephani 1916a).
Strepsilejeunea papulifolia Steph., Biblioth. Bot. 87 (2): 259, 1916 (Stephani 1916a).
Strepsilejeunea pectiniformis (Gottsche) Steph., Sp. Hepat. (Stephani) 5: 285, 1913
(Stephani 1913a). Bas.: Lejeunea pectiniformis Gottsche, Ann. Sci. Nat. Bot. (sér.
5) 1: 156, 1864 (Gottsche 1864).
Strepsilejeunea renistipula Steph., Sp. Hepat. (Stephani) 5: 289, 1913 (Stephani 1913a).
Strepsilejeunea vatovae Gerola, Lav. Bot. Ist. Bot. Univ. Padova 12: 479, 1947
(Gerola 1947).
hysanolejeunea (Spruce) Steph., Hedwigia 31 (1): 20, 1892 (Jack and Stephani
1892). Bas.: Lejeunea subg. hysanolejeunea Spruce, Trans. & Proc. Bot. Soc.
Edinburgh 15: 105, 1885 (Spruce 1885).
* hysanolejeunea africana Sim, Trans. Roy. Soc. South Africa 15 (1): 50, 1926 (Sim
1926). 501
Trachylejeunea (Spruce) Steph., Hedwigia 28 (4): 262, 1889 (Stephani 1889c) nom.
rejic. Bas.: Lejeunea subg. Trachylejeunea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 180, 1884 (Spruce 1884).
* Trachylejeunea conifera Steph., Sp. Hepat. (Stephani) 5: 302, 1913 (Stephani 1913a).
* Trachylejeunea cristulilora Steph., Hedwigia 35 (3): 137, 1896 (Stephani 1896b).
* Trachylejeunea englishii Steph., Bull. Misc. Inform. Kew 1899 (151/152): 126,
1899 (MacGregor 1899). 502
* Trachylejeunea jamaicensis Pearson, Ann. Bryol. 4: 98, 1931 (Pearson 1931b).
* Trachylejeunea kusaiensis Inoue et H.A.Mill., Bull. Natl. Sci. Mus. Tokyo (n.ser.)
8 (2): 147, 1965 (Inoue and Miller 1965).
501 hysanolejeunea africana may be conspeciic with Caudalejeunea africana.
502 Trachylejeunea englishii is not a Trachylejeunea species (see Zhu and So 1999a). he status needs further study.
World checklist of hornworts and liverworts
513
Alphabetic list of accepted taxa
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Acanthocoleus aberrans (Lindenb. et Gottsche) Kruijt ..................................................................297
Acanthocoleus aberrans var. laevis Gradst. .....................................................................................297
Acanthocoleus chrysophyllus (Lehm.) Kruijt...................................................................................297
Acanthocoleus elgonensis Gyarmati et Pócs ....................................................................................297
Acanthocoleus gilvus (Gottsche) Kruijt .........................................................................................297
Acanthocoleus javanicus (Steph.) Kruijt ........................................................................................297
Acanthocoleus juddii Kruijt ..........................................................................................................297
Acanthocoleus madagascariensis (Steph.) Kruijt .............................................................................297
Acanthocoleus trigonus (Nees et Mont.) Gradst. ...........................................................................297
Acanthocoleus yoshinaganus (S.Hatt.) Kruijt .................................................................................298
Acrobolbus africanus (Pearson) Briscoe...........................................................................................93
Acrobolbus anisodontus (Hook.f. et Taylor) Briscoe........................................................................94
Acrobolbus antillanus R.M.Schust. .................................................................................................94
Acrobolbus azoricus (Grolle et Perss.) Briscoe .................................................................................94
Acrobolbus bispinosus (J.B.Jack et Steph.) Steph. ............................................................................96
Acrobolbus caducifolius R.M.Schust. ..............................................................................................94
Acrobolbus ciliatus (Mitt.) Schifn. .................................................................................................94
Acrobolbus cinerascens (Lehm. et Lindenb.) Bastow .......................................................................94
Acrobolbus concinnus (Mitt.) Grolle ...............................................................................................94
Acrobolbus cuneifolius (Steph.) Briscoe...........................................................................................94
Acrobolbus diversifolius R.M.Schust. ..............................................................................................94
Acrobolbus epiphytus (Colenso) Briscoe ..........................................................................................94
Acrobolbus lavicans (J.J.Engel et Grolle) Briscoe et J.J.Engel .........................................................94
Acrobolbus gradsteinii (Grolle) Briscoe ...........................................................................................94
Acrobolbus integrifolius (A.Evans) Briscoe ......................................................................................94
Acrobolbus knightii (Mitt.) Briscoe ................................................................................................94
Acrobolbus kunkelii (Hässel et Solari) Briscoe et J.J.Engel ..............................................................94
Acrobolbus laxus (Lehm. et Lindenb.) Briscoe ................................................................................94
Acrobolbus limbatus (Steph.) Briscoe et J.J.Engel ...........................................................................95
Acrobolbus lophocoleoides (Mitt.) Mitt. ...........................................................................................95
Acrobolbus madeirensis (Grolle et Perss.) Briscoe ............................................................................95
Acrobolbus mittenii Steph. .............................................................................................................95
Acrobolbus ochrophyllus (Hook.f. et Taylor) R.M.Schust................................................................95
Acrobolbus papillosus (J.J.Engel et Glenny) Briscoe ........................................................................95
Acrobolbus perpusillus (Colenso) Briscoe ........................................................................................95
Acrobolbus perpusillus var. denticulatus (J.J.Engel et Glenny) Briscoe .............................................95
Acrobolbus plagiochiloides (J.J.Engel et Glenny) Briscoe .................................................................95
Acrobolbus pseudosaccatus (Grolle) Briscoe .....................................................................................95
Acrobolbus renifolius (Hässel et Solari) Briscoe et J.J.Engel ............................................................95
Acrobolbus ruwenzorensis (S.W.Arnell) Briscoe ..............................................................................95
Acrobolbus saccatus (Hook.) Trevis. ...............................................................................................95
Acrobolbus setulosus (Mitt.) Briscoe ................................................................................................95
Acrobolbus spinifolius R.M.Schust..................................................................................................95
Acrobolbus sumatranus (Schifn.) Briscoe .......................................................................................96
Acrobolbus surculosus (Nees) Trevis. ...............................................................................................96
Acrobolbus tenellus (Taylor) Trevis.................................................................................................96
Acrobolbus tenellus var. diversifolius (E.A.Hodgs.) Briscoe ..............................................................96
Acrobolbus urvilleanus (Mont.) Trevis. ..........................................................................................96
Acrobolbus viridis (Mitt.) Briscoe et J.J.Engel ................................................................................96
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Acrobolbus wilsonii Nees ................................................................................................................96
Acrobolbus wilsonii var. andinus Spruce .........................................................................................96
Acrochila biserialis (Lehm. et Lindenb.) Grolle ............................................................................219
Acrochila caledonica (Steph.) Inoue .............................................................................................219
Acrolejeunea abnormis (Gottsche) Pearson ...................................................................................401
Acrolejeunea allisonii Gradst. .......................................................................................................400
Acrolejeunea arcuata (Nees) Grolle et Gradst. ..............................................................................399
Acrolejeunea arcuata subsp. gradsteinii M.A.M.Renner ................................................................399
Acrolejeunea aulacophora (Mont.) Steph. .....................................................................................400
Acrolejeunea comptonii Pearson....................................................................................................401
Acrolejeunea crassicaulis (Steph.) Jian Wang bis et Gradst. ...........................................................401
Acrolejeunea emergens (Mitt.) Steph. ............................................................................................399
Acrolejeunea emergens var. confertissima (Steph.) Gradst. ..............................................................399
Acrolejeunea fertilis (Reinw., Blume et Nees) Schifn. ..................................................................399
Acrolejeunea heterophylla (A.Evans) Grolle et Gradst. ..................................................................399
Acrolejeunea inlexa (Gottsche) Pearson .......................................................................................401
Acrolejeunea infuscata (Mitt.) Jian Wang bis et Gradst. ...............................................................401
Acrolejeunea meghalayensis (Ajit P.Singh et V.Nath) Jian Wang bis et Gradst. .............................401
Acrolejeunea mollis (Hook.f. et Taylor) Schifn. ...........................................................................400
Acrolejeunea parvula (Mizut.) Gradst. .........................................................................................399
Acrolejeunea pusilla (Steph.) Grolle et Gradst. .............................................................................400
Acrolejeunea pycnoclada (Taylor) Schifn......................................................................................399
Acrolejeunea pycnoclada subsp. latistipula Gradst. ........................................................................399
Acrolejeunea recurvata Gradst. .....................................................................................................400
Acrolejeunea sandvicensis (Gottsche) Steph. .................................................................................399
Acrolejeunea securifolia (Nees) Steph............................................................................................400
Acrolejeunea securifolia subsp. caledonica (Steph.) Gradst. ............................................................400
Acrolejeunea securifolia subsp. hartmannii (Steph.) Gradst. ..........................................................400
Acrolejeunea securifolia subsp. pallida (Ångstr.) Gradst. ...............................................................400
Acrolejeunea sikkimensis (Mizut.) Gradst. ....................................................................................400
Acrolejeunea sinensis (Jian Wang bis, R.L.Zhu et Gradst.) Jian Wang bis et Gradst. ....................401
Acrolejeunea tjibodensis (Verd.) Grolle et Gradst. .........................................................................400
Acrolejeunea torulosa (Lehm. et Lindenb.) Schifn. ......................................................................399
Acrolophozia fuegiana R.M.Schust. ..............................................................................................110
Acrolophozia pectinata R.M.Schust. .............................................................................................110
Acrolophozia sulcata Hässel ..........................................................................................................110
Acromastigum adaptatum Hürl. ...................................................................................................134
Acromastigum anisostomum (Lehm. et Lindenb.) A.Evans............................................................143
Acromastigum anisostomum var. minutum E.A.Hodgs. .................................................................144
Acromastigum aurescens A.Evans ..................................................................................................144
Acromastigum bancanum (Sande Lac.) A.Evans ...........................................................................144
Acromastigum brotheri (Steph.) A.Evans ......................................................................................144
Acromastigum caledonicum (Steph.) Grolle ..................................................................................143
Acromastigum capillare (Steph.) A.Evans .....................................................................................144
Acromastigum cavifolium R.M.Schust. .........................................................................................143
Acromastigum colensoanum (Mitt.) A.Evans ex Reimers ...............................................................144
Acromastigum cunninghamii (Steph.) A.Evans .............................................................................144
Acromastigum curtilobum A.Evans ...............................................................................................144
Acromastigum divaricatum (Nees) A.Evans ex Reimers ................................................................144
Acromastigum echinatiforme (De Not.) A.Evans...........................................................................144
Acromastigum echinatum (Gottsche) A.Evans ..............................................................................144
Acromastigum exiguum (Steph.) A.Evans .....................................................................................144
World checklist of hornworts and liverworts
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515
Acromastigum exile (Lindenb.) A.Evans .......................................................................................144
Acromastigum ilum (Steph.) A.Evans ..........................................................................................144
Acromastigum ilum var. papillosum N.Kitag. ...............................................................................144
Acromastigum imbriatum (Steph.) A.Evans .................................................................................144
Acromastigum fumosum E.A.Br. et M.A.M.Renner ......................................................................145
Acromastigum furcatifolium (Steph.) E.A.Br. ...............................................................................145
Acromastigum herzogii Grolle ......................................................................................................143
Acromastigum homodictyon (Herzog) Grolle ................................................................................143
Acromastigum inaequilaterum (Lehm. et Lindenb.) A.Evans ........................................................145
Acromastigum integrifolium (Austin) A.Evans ..............................................................................143
Acromastigum interstisiale E.A.Br. et M.A.M.Renner ...................................................................145
Acromastigum laetevirens (Sande Lac. ex Steph.) A.Evans.............................................................145
Acromastigum laevigatum A.Evans ...............................................................................................145
Acromastigum leptophyllum Herzog..............................................................................................145
Acromastigum linganum (De Not.) A.Evans ................................................................................145
Acromastigum lobuliferum A.Evans ..............................................................................................145
Acromastigum longirete Grolle......................................................................................................145
Acromastigum marginatum E.A.Hodgs. .......................................................................................145
Acromastigum microstictum A.Evans ............................................................................................145
Acromastigum mooreanum (Steph.) E.A.Hodgs. ...........................................................................145
Acromastigum moratii N.Kitag. ...................................................................................................145
Acromastigum obliquatum (Mitt.) A.Evans ..................................................................................145
Acromastigum prismaticale E.A.Br. et M.A.M.Renner..................................................................145
Acromastigum pusillum N.Kitag...................................................................................................145
Acromastigum rigidum R.M.Schust..............................................................................................145
Acromastigum stellare N.Kitag. ....................................................................................................143
Acromastigum stenophyllum R.M.Schust. .....................................................................................143
Acromastigum subechinatiforme Hürl. ..........................................................................................145
Acromastigum tenax (Steph.) A.Evans ..........................................................................................145
Acromastigum verticale (Steph.) E.A.Hodgs. ................................................................................143
Acroscyphella iwatsukii (N.Kitag.) N.Kitag. et Grolle .....................................................................99
Acroscyphella phoenicorhiza (Grolle) N.Kitag. et Grolle..................................................................99
Acroscyphella tjiwideiensis (Sande Lac.) N.Kitag. et Grolle .............................................................99
Acrostolia alata (Gottsche et Rabenh.) Trevis. .............................................................................507
Acrostolia brevifolia (Gottsche et Rabenh.) Trevis. .......................................................................507
Adelanthus aureomarginatus R.M.Schust. ......................................................................................44
Adelanthus carabayensis (Mont.) Grolle .........................................................................................44
Adelanthus falcatus (Hook.) Mitt...................................................................................................44
Adelanthus gemmiparus (R.M.Schust.) E.A.Hodgs.........................................................................44
Adelanthus integerrimus Grolle ......................................................................................................44
Adelanthus lindenbergianus (Lehm.) Mitt. .....................................................................................44
Adelanthus lingulatus J.J.Engel et Váňa..........................................................................................44
Adelanthus occlusus (Hook.f. et Taylor) Carrington .......................................................................44
Adelanthus pittieri (Steph.) Grolle .................................................................................................44
Adelanthus squarrosus Grolle..........................................................................................................44
Adelanthus tenuis J.J.Engel et Grolle..............................................................................................44
Aitchisoniella himalayensis Kashyap .............................................................................................488
Allisonia cockaynei (K.I.Goebel) R.M.Schust. ..............................................................................463
Allisoniella nigra (Rodway) R.M.Schust. .......................................................................................65
Allisoniella nigra var. acutiloba J.J.Engel ........................................................................................65
Allisoniella nigra subsp. novaezelandiae R.M.Schust. .....................................................................65
Allisoniella recurva R.M.Schust. ....................................................................................................65
516
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Allisoniella scottii (R.M.Schust.) R.M.Schust. ................................................................................65
Allisoniella subbipartita (C.Massal.) R.M.Schust. et J.J.Engel ........................................................65
Allisoniella tasmanica R.M.Schust. ................................................................................................65
Alobiella husnotii (Spruce) Schifn. ................................................................................................56
Alobiellopsis acroscypha (Spruce) R.M.Schust. ................................................................................56
Alobiellopsis dominicensis (Spruce) Fulford .....................................................................................56
Alobiellopsis heteromorpha (Lehm.) R.M.Schust. ............................................................................56
Alobiellopsis parvifolia (Steph.) R.M.Schust. ..................................................................................57
Alobiellopsis pillansii (Sim) R.M.Schust. ........................................................................................57
Amazoopsis diplopoda (Pócs) J.J.Engel et G.L.Merr. ....................................................................183
Amazoopsis dissotricha (Spruce) J.J.Engel et G.L.Merr. ................................................................183
Amazoopsis gracilis J.J.Engel et G.L.Merr. ...................................................................................183
Amphicephalozia africana Váňa et M.Wigginton ...........................................................................65
Amphicephalozia amplexicaulis R.M.Schust. ..................................................................................65
Amphicephalozia geisslerae Pócs et Váňa.........................................................................................65
Anastrepta orcadensis (Hook.) Schifn. ...........................................................................................50
Anastrophyllopsis involutifolia (Mont. ex Gottsche, Lindenb. et Nees) Váňa et L.Söderstr. .............65
Anastrophyllopsis revoluta (Steph.) Váňa et L.Söderstr. ...................................................................65
Anastrophyllopsis subcomplicata (Lehm. et Lindenb.) Váňa et L.Söderstr. .......................................66
Anastrophyllum alpinum Steph. .....................................................................................................50
Anastrophyllum assimile (Mitt.) Steph. ...........................................................................................50
Anastrophyllum auritum (Lehm.) Steph. ........................................................................................50
Anastrophyllum ciliatum Steph. .....................................................................................................50
Anastrophyllum donnianum (Hook.) Steph. ...................................................................................50
Anastrophyllum ellipticum Inoue ....................................................................................................50
Anastrophyllum esenbeckii (Mont.) Steph. ......................................................................................50
Anastrophyllum issum Steph. .........................................................................................................50
Anastrophyllum joergensenii Schifn................................................................................................50
Anastrophyllum lignicola D.B.Schill et D.G.Long ..........................................................................50
Anastrophyllum michauxii (F.Weber) H.Buch ...............................................................................50
Anastrophyllum nigrescens (Mitt.) Steph. ........................................................................................51
Anastrophyllum obtusum Herzog ....................................................................................................51
Anastrophyllum piligerum (Nees) Steph. ........................................................................................51
Anastrophyllum squarrosum Herzog ...............................................................................................51
Anastrophyllum stellatum R.M.Schust. ...........................................................................................51
Anastrophyllum tubulosum (Nees) Grolle .......................................................................................51
Andrewsianthus aberrans (Nees et Mont.) Grolle ...........................................................................77
Andrewsianthus bidens (Mitt. ex Steph.) R.M.Schust. ....................................................................77
Andrewsianthus bilobus (Mitt.) Grolle ...........................................................................................77
Andrewsianthus carinatus Grolle ....................................................................................................77
Andrewsianthus cavifolius Grolle et Váňa .......................................................................................77
Andrewsianthus chimbuensis R.M.Schust. ......................................................................................77
Andrewsianthus hodgsoniae (R.M.Schust.) R.M.Schust. ex J.J.Engel ..............................................78
Andrewsianthus kinabaluensis N.Kitag. ..........................................................................................77
Andrewsianthus koponenii Váňa et Piippo ......................................................................................77
Andrewsianthus marionensis (S.W.Arnell) Grolle ...........................................................................78
Andrewsianthus mizutanii N.Kitag. ...............................................................................................77
Andrewsianthus papillosus N.Kitag. ................................................................................................78
Andrewsianthus perigonialis (Hook.f. et Taylor) R.M.Schust. ........................................................78
Andrewsianthus puniceus (Nees) R.M.Schust. ex Grolle .................................................................78
Andrewsianthus recurvifolius (Nees) R.M.Schust. ...........................................................................78
Andrewsianthus scabrellus (C.Massal.) R.M.Schust. ex J.J.Engel ....................................................78
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Andrewsianthus sphenoloboides (R.M.Schust.) R.M.Schust. ex J.J.Engel .........................................78
Andrewsianthus sundaicus (Schifn.) R.M.Schust. ..........................................................................78
Andrewsianthus zantenii Váňa .......................................................................................................78
Aneura amboinensis Steph. ..........................................................................................................435
Aneura augustae Steph.................................................................................................................435
Aneura bilora Colenso ................................................................................................................435
Aneura blasioides (Horik.) Furuki ................................................................................................435
Aneura brasiliensis (Ångstr.) Steph...............................................................................................435
Aneura brevissima Steph. .............................................................................................................435
Aneura cerebrata Hewson ............................................................................................................435
Aneura crateriformis Furuki et D.G.Long ....................................................................................435
Aneura crinita C.Massal. .............................................................................................................436
Aneura crumii L.Söderstr., A.Hagborg et von Konrat ..................................................................436
Aneura densa Steph. ....................................................................................................................436
Aneura denticulata Mitt. ex hurn ..............................................................................................436
Aneura eachamensis Hewson........................................................................................................436
Aneura erronea Steph. .................................................................................................................436
Aneura eskuchei Hässel ................................................................................................................436
Aneura gemmifera Furuki ............................................................................................................436
Aneura giangena Hewson ............................................................................................................436
Aneura gibbsiana Steph. ..............................................................................................................436
Aneura glaucescens Steph. ............................................................................................................436
Aneura goebeliana Steph. .............................................................................................................436
Aneura hirsuta Furuki .................................................................................................................436
Aneura hunsteinii Steph. .............................................................................................................436
Aneura imbricata Colenso ...........................................................................................................436
Aneura kaguaensis Hewson ..........................................................................................................436
Aneura keniae Gola .....................................................................................................................436
Aneura latemultiida Steph. .........................................................................................................436
Aneura latissima Spruce...............................................................................................................436
Aneura ledermannii Steph. ..........................................................................................................436
Aneura macrostachya Spruce ........................................................................................................436
Aneura marianensis Furuki ..........................................................................................................436
Aneura maxima (Schifn.) Steph..................................................................................................436
Aneura mirabilis (Malmb.) Wickett et Goinet...........................................................................436
Aneura novaecaledoniae R.M.Schust. ...........................................................................................436
Aneura novaguineensis Hewson....................................................................................................437
Aneura nymannii Steph. ..............................................................................................................437
Aneura pellucida Colenso ............................................................................................................437
Aneura pinguis (L.) Dumort. .......................................................................................................437
Aneura polyantha Colenso ...........................................................................................................437
Aneura punctata Colenso ............................................................................................................437
Aneura rodwayi Hewson .............................................................................................................437
Aneura roraimensis Steph. ...........................................................................................................437
Aneura rotangicola Steph. ............................................................................................................437
Aneura serrulata Gottsche ex Steph. ............................................................................................437
Aneura sharpii Inoue et N.G.Mill. ..............................................................................................437
Aneura singalangana (Schifn.) Steph. .........................................................................................437
Aneura subcanaliculata R.M.Schust. ............................................................................................437
Aneura subledermannii Steph. .....................................................................................................437
Aneura subtenerrima Steph. .........................................................................................................437
Aneura vincentina Steph..............................................................................................................437
518
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Anoplolejeunea conferta (C.F.W.Meissn. ex Spreng.) A.Evans ......................................................361
Anthelia julacea (L.) Dumort. .......................................................................................................98
Anthelia juratzkana (Limpr.) Trevis. .............................................................................................98
Anthoceros adscendens Lehm. et Lindenb. ......................................................................................27
Anthoceros aethyopicus Gola ...........................................................................................................30
Anthoceros agrestis Paton nom. conserv.(p. 000)
Anthoceros alpinus Steph................................................................................................................27
Anthoceros angustifolius Gottsche, Lindenb. et Nees ......................................................................28
Anthoceros angustus Steph. .............................................................................................................28
Anthoceros bharadwajii Udar et A.K.Asthana.................................................................................28
Anthoceros brunnthaleri Steph. ......................................................................................................30
Anthoceros buettneri Steph. ............................................................................................................28
Anthoceros capricornii Cargill et G.A.M.Scott ................................................................................28
Anthoceros caucasicus Steph............................................................................................................28
Anthoceros cavernosus Steph. ..........................................................................................................28
Anthoceros chambensis Kashyap ......................................................................................................28
Anthoceros chungii Khanna ............................................................................................................28
Anthoceros crispatus Grif. ..............................................................................................................28
Anthoceros dimorphus Sim .............................................................................................................28
Anthoceros erectus Kashyap ............................................................................................................28
Anthoceros expansus (Steph.) J.C.Villarreal et Cargill .....................................................................28
Anthoceros ferdinandi-muelleri Steph. ............................................................................................28
Anthoceros loribundus Steph. ........................................................................................................30
Anthoceros fragilis Steph. ...............................................................................................................28
Anthoceros fusiformis Austin ...........................................................................................................28
Anthoceros fusiformis var. taiwanensis J.Haseg. ...............................................................................28
Anthoceros gasongorii Gola .............................................................................................................28
Anthoceros granulatus Gottsche......................................................................................................28
Anthoceros harrisanus (Steph.) Parihar ...........................................................................................28
Anthoceros helmsii Steph. ...............................................................................................................28
Anthoceros jamesonii Taylor ex Mitt. .............................................................................................28
Anthoceros javanicoides H.A.Mill. ..................................................................................................29
Anthoceros jungermannioides Schwein. ...........................................................................................29
Anthoceros kajumas (K.I.Goebel) Prosk. ........................................................................................29
Anthoceros koshii Khanna ..............................................................................................................29
Anthoceros lamellatus Steph. ..........................................................................................................29
Anthoceros laminifer Steph.............................................................................................................29
Anthoceros macounii M.Howe .......................................................................................................29
Anthoceros macrosporus Steph. .......................................................................................................29
Anthoceros maritimus Steph. ..........................................................................................................29
Anthoceros megasporus Meijer.........................................................................................................29
Anthoceros mildbraedii Steph. ........................................................................................................30
Anthoceros muscoides Colenso ........................................................................................................29
Anthoceros myriandroecius Steph. ...................................................................................................29
Anthoceros natalensis Steph. ...........................................................................................................29
Anthoceros neesii Prosk. .................................................................................................................29
Anthoceros niger Steph. ..................................................................................................................29
Anthoceros orizabensis (Steph.) Hässel ............................................................................................29
Anthoceros pandei Udar et A.K.Asthana .........................................................................................29
Anthoceros parvifrons Steph. ..........................................................................................................30
Anthoceros patagonicus Hässel ........................................................................................................29
Anthoceros patagonicus subsp. gremmenii J.C.Villarreal, J.J.Engel et Váňa ......................................29
World checklist of hornworts and liverworts
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519
Anthoceros peruvianus Steph. .........................................................................................................29
Anthoceros pinnatus Steph. ............................................................................................................29
Anthoceros pseudocostus Steph. .......................................................................................................30
Anthoceros punctatus L. ..................................................................................................................29
Anthoceros pusillus Colenso............................................................................................................30
Anthoceros rossoi Gola ....................................................................................................................30
Anthoceros rosulans J.Haseg. ..........................................................................................................30
Anthoceros sambesianus Steph. .......................................................................................................30
Anthoceros scariosus Austin.............................................................................................................30
Anthoceros schroederi Steph. ...........................................................................................................30
Anthoceros serratus Steph. ..............................................................................................................30
Anthoceros simulans M.Howe ........................................................................................................30
Anthoceros spongiosus Steph. ..........................................................................................................30
Anthoceros subtilis Steph. ...............................................................................................................30
Anthoceros telaganus Steph. ............................................................................................................30
Anthoceros tristanianus J.C.Villarreal, J.J.Engel et Váňa .................................................................30
Anthoceros tuberculatus Lehm. et Lindenb. ....................................................................................30
Anthoceros venosus Lindenb. et Gottsche........................................................................................30
Aphanolejeunea lancifera R.M.Schust. .........................................................................................507
Aphanolejeunea minima Tixier ....................................................................................................507
Aphanotropis saxicola Herzog .......................................................................................................317
Aponardia huerlimannii (Váňa et Grolle) Váňa ...........................................................................123
Apotreubia hortoniae Konstant. .....................................................................................................42
Apotreubia nana (S.Hatt. et Inoue) S.Hatt. et Mizut. ....................................................................42
Apotreubia pusilla (R.M.Schust.) Grolle ........................................................................................42
Apotreubia yunnanensis Higuchi ....................................................................................................43
Archeophylla paradoxa R.M.Schust. .............................................................................................253
Archeophylla pungens (Herzog) R.M.Schust. ................................................................................253
Archeophylla schusteri (E.A.Hodgs. et Allison) R.M.Schust. .........................................................253
Archilejeunea abbreviata (Mitt.) Vanden Berghen .......................................................................401
Archilejeunea africana Steph. .......................................................................................................401
Archilejeunea alata Steph.............................................................................................................401
Archilejeunea amakawana Inoue ..................................................................................................402
Archilejeunea auberiana (Mont.) Steph........................................................................................402
Archilejeunea autoica Vanden Berghen.........................................................................................402
Archilejeunea badia (Spruce) Steph..............................................................................................402
Archilejeunea bilabiata (Mitt.) Steph. ..........................................................................................402
Archilejeunea bischleriana Gradst. ................................................................................................402
Archilejeunea bongardii Steph. .....................................................................................................402
Archilejeunea brachyantha J.B.Jack et Steph.................................................................................402
Archilejeunea brevilobula Steph. ..................................................................................................402
Archilejeunea crispistipula (Spruce) Steph. ...................................................................................402
Archilejeunea eberhardtii Steph. ...................................................................................................402
Archilejeunea elobulata Steph.......................................................................................................402
Archilejeunea gradsteinii X.Q.Shi et R.L.Zhu ..............................................................................402
Archilejeunea incrassata Steph. .....................................................................................................402
Archilejeunea jonesii Vanden Berghen..........................................................................................402
Archilejeunea juliformis (Nees) Gradst. ........................................................................................402
Archilejeunea kiushiana (Horik.) Verd. ........................................................................................402
Archilejeunea ludoviciana (De Not.) P.Geissler et Gradst. ...........................................................402
Archilejeunea ludoviciana subsp. porelloides (Spruce) Gradst. .......................................................402
Archilejeunea nebeliana Gradst. et Schäf.-Verw. ..........................................................................403
520
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Archilejeunea negrensis Steph. ......................................................................................................403
Archilejeunea olivacea (Hook.f. et Taylor) Steph..........................................................................403
Archilejeunea ovata Herzog..........................................................................................................403
Archilejeunea parvilora (Nees) Steph. .........................................................................................403
Archilejeunea planifolia (Horik.) Mizut. ......................................................................................403
Archilejeunea planiuscula (Mitt.) Steph........................................................................................403
Arctoscyphus fuegiensis (C.Massal.) Hässel ....................................................................................123
Arctoscyphus ronsmithii Hässel .....................................................................................................123
Arnellia fennica (Gottsche) Lindb. ................................................................................................99
Ascidiota blepharophylla C.Massal. ...............................................................................................417
Ascidiota blepharophylla subsp. alaskana Steere et R.M.Schust. ....................................................417
Aspiromitus asper Schifn. ............................................................................................................507
Aspiromitus bullosus Schifn. ........................................................................................................507
Aspiromitus crenatifrons Steph. ....................................................................................................507
Aspiromitus lobatus Schifn. .........................................................................................................507
Aspiromitus squamulosus Schifn. .................................................................................................507
Asterella abyssinica (Gottsche) Grolle ...........................................................................................477
Asterella africana (Mont.) Underw. ex A.Evans ...........................................................................477
Asterella alpina (Steph.) D.G.Long..............................................................................................480
Asterella australis (Hook.f. et Taylor) Verd. .................................................................................478
Asterella bachmannii (Steph.) S.W.Arnell ....................................................................................478
Asterella blumeana (Nees) Kachroo..............................................................................................477
Asterella bolanderi (Austin) Underw. ...........................................................................................478
Asterella californica (Hampe ex Austin) Underw. .........................................................................481
Asterella caucasica (Steph.) H.Buch, A.Evans et Verd. .................................................................478
Asterella chilensis (Nees et Mont.) A.Evans ..................................................................................477
Asterella conocephala (Steph.) R.M.Schust. ..................................................................................478
Asterella coronata (Steph.) H.A.Mill. ...........................................................................................478
Asterella cruciata (Steph.) Horik. .................................................................................................477
Asterella dioica (Steph.) H.A.Mill. ...............................................................................................478
Asterella dissoluta (Steph.) Grolle .................................................................................................477
Asterella dognyensis H.A.Mill. ......................................................................................................478
Asterella dominicensis S.W.Arnell.................................................................................................477
Asterella drummondii (Taylor) R.M.Schust. ex D.G.Long ...........................................................478
Asterella echinella (Gottsche) Underw..........................................................................................478
Asterella elegans (Spreng.) Trevis. ................................................................................................479
Asterella grollei D.G.Long ...........................................................................................................480
Asterella heterolora (Steph.) H.A.Mill. ........................................................................................479
Asterella innovans (Austin) H.A.Mill. ..........................................................................................479
Asterella khasyana (Grif.) Grolle .................................................................................................477
Asterella lateralis M.Howe ...........................................................................................................479
Asterella leptophylla (Mont.) Grolle .............................................................................................477
Asterella limbata D.G.Long et Grolle ..........................................................................................477
Asterella lindenbergiana (Corda ex Nees) Lindb. ex Arnell ...........................................................479
Asterella linearis (Steph.) M.Howe ..............................................................................................479
Asterella longebarbata (Steph.) H.A.Mill. .....................................................................................479
Asterella macropoda (Spruce) A.Evans ..........................................................................................479
Asterella marginata (Nees) S.W.Arnell .........................................................................................479
Asterella muelleri (Gottsche) R.M.Schust.....................................................................................479
Asterella multilora (Steph.) Kachroo ...........................................................................................479
Asterella muscicola (Steph.) S.W.Arnell ........................................................................................480
Asterella mussuriensis (Kashyap) Verd. .........................................................................................479
World checklist of hornworts and liverworts
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521
Asterella mussuriensis subsp. crassa (Shimizu et S.Hatt.) D.G.Long ..............................................479
Asterella palmeri (Austin) Underw. ..............................................................................................480
Asterella pappii (Gola) Grolle ......................................................................................................479
Asterella persica (Steph.) M.Howe ...............................................................................................479
Asterella preussii (Schifn.) M.Howe ............................................................................................480
Asterella pringlei Underw.............................................................................................................480
Asterella rugosa A.Evans...............................................................................................................480
Asterella saccata (Wahlenb.) A.Evans ...........................................................................................480
Asterella setisquama (Steph.) R.M.Schust. ....................................................................................480
Asterella shimizuana Inoue ..........................................................................................................477
Asterella syngenesica (Bory) Grolle................................................................................................480
Asterella tasmanica (Steph.) R.M.Schust. .....................................................................................480
Asterella tenella (L.) P.Beauv........................................................................................................477
Asterella tenera (Mitt.) R.M.Schust. ............................................................................................478
Asterella tenerrima (Steph.) H.A.Mill. .........................................................................................478
Asterella venosa (Lehm. et Lindenb.) A.Evans ..............................................................................478
Asterella versicolor A.Evans ..........................................................................................................480
Asterella vulcanica (Schifn.) Kachroo et Bapna ...........................................................................480
Asterella wallichiana (Lehm. et Lindenb.) Grolle .........................................................................481
Asterella whiteleggeana (Steph.) R.M.Schust. ...............................................................................480
Asterella wilmsii (Steph.) S.W.Arnell ...........................................................................................480
Athalamia dioica Kashyap ...........................................................................................................484
Athalamia pinguis Falc. ...............................................................................................................484
Athalamia pulcherrima (Steph.) S.Hatt........................................................................................484
Aureolejeunea aurifera R.M.Schust. .............................................................................................304
Aureolejeunea lumae (Herzog) van Slageren .................................................................................304
Aureolejeunea paramicola (Herzog) R.M.Schust. .........................................................................304
Aureolejeunea quinquecarinata R.M.Schust. ................................................................................304
Aureolejeunea tonduzana (Steph.) Gradst.....................................................................................304
Austrolophozia andina R.M.Schust. ...............................................................................................96
Austrolophozia camensis (Steph.) Grolle ex Hässel et Solari ............................................................96
Austrolophozia paradoxa R.M.Schust. ............................................................................................96
Austroriella salta J.Milne et Cargill ..............................................................................................504
Balantiopsis asymmetrica (Herzog) J.J.Engel...................................................................................99
Balantiopsis bisbiida (Steph.) Steph. ...........................................................................................100
Balantiopsis brasiliensis Steph.......................................................................................................100
Balantiopsis cancellata (Nees) Steph. ............................................................................................100
Balantiopsis ciliaris S.Hatt. ..........................................................................................................100
Balantiopsis ciliaris subsp. novoguineensis S.Hatt. .........................................................................100
Balantiopsis convexiuscula Berggr. ................................................................................................100
Balantiopsis crocea Herzog ...........................................................................................................100
Balantiopsis diplophylla (Hook.f. et Taylor) Mitt. ........................................................................100
Balantiopsis diplophylla var. hockenii (Berggr.) J.J.Engel et G.L.Merr. ..........................................100
Balantiopsis erinacea (Hook.f. et Taylor) Mitt. ............................................................................100
Balantiopsis lingulata R.M.Schust................................................................................................100
Balantiopsis montana (Colenso) J.J.Engel et G.L.Merr. ...............................................................100
Balantiopsis neocaledonica Pearson ...............................................................................................100
Balantiopsis paucidens Steph. .......................................................................................................100
Balantiopsis purpurata Mitt. ........................................................................................................100
Balantiopsis rosea Berggr. .............................................................................................................100
Balantiopsis splendens (Steph.) J.J.Engel et G.L.Merr. ..................................................................100
Balantiopsis tumida Berggr. .........................................................................................................100
522
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Balantiopsis verrucosa J.J.Engel et G.L.Merr. ...............................................................................100
Barbilophozia barbata (Schmidel ex Schreb.) Loeske .....................................................................51
Barbilophozia hatcheri (A.Evans) Loeske........................................................................................51
Barbilophozia lycopodioides (Wallr.) Loeske ...................................................................................51
Barbilophozia rubescens (R.M.Schust. et Damsh.) Kartt. et L.Söderstr. ..........................................51
Barbilophozia sudetica (Nees ex Huebener) L.Söderstr., De Roo et Hedd. .....................................51
Bazzania acanthostipa Spruce ......................................................................................................146
Bazzania accreta (Lehm. et Lindenb.) Trevis. ..............................................................................146
Bazzania acinaciformis Steph.......................................................................................................146
Bazzania acuminata (Lindenb. et Gottsche) Trevis. ....................................................................146
Bazzania acutifolia (Steph.) Schifn. ............................................................................................146
Bazzania adnexa (Lehm. et Lindenb.) Trevis. .............................................................................146
Bazzania adnexa var. aucklandica (Lindenb. et Gottsche) J.J.Engel et G.L.Merr. ........................146
Bazzania ainis (Lindenb. et Gottsche) Trevis. ...........................................................................146
Bazzania albifolia Horik. ............................................................................................................146
Bazzania ambigua (Lindenb.) Trevis. ..........................................................................................146
Bazzania amblyphylla Meagher ...................................................................................................146
Bazzania aneityensis (Steph.) Tixier .............................................................................................146
Bazzania angusta (Steph.) Herzog ...............................................................................................146
Bazzania angustifalcata Herzog ...................................................................................................146
Bazzania angustifolia Horik. .......................................................................................................147
Bazzania angustisedens (Steph.) N.Kitag. .....................................................................................147
Bazzania angustistipula N.Kitag. .................................................................................................147
Bazzania appendiculata (Mitt.) S.Hatt. .......................................................................................147
Bazzania approximata Onr..........................................................................................................147
Bazzania arcuata (Lindenb. et Gottsche) Trevis. .........................................................................147
Bazzania arcuata var. mamillosa Gradst. et A.R.Benitez ..............................................................147
Bazzania armatistipula (Steph.) Fulford ......................................................................................147
Bazzania asperrima Steph............................................................................................................147
Bazzania asymmetrica (Steph.) N.Kitag. ......................................................................................147
Bazzania aterrima (Steph.) N.Kitag. ...........................................................................................147
Bazzania aurescens Spruce ...........................................................................................................147
Bazzania avia Meagher ...............................................................................................................147
Bazzania azorica H.Buch et Perss. ..............................................................................................147
Bazzania baldwinii Austin ..........................................................................................................147
Bazzania bernieri (Steph.) Inoue et H.A.Mill. .............................................................................147
Bazzania bescherellei Steph. .........................................................................................................147
Bazzania bhutanica N.Kitag. et Grolle ........................................................................................147
Bazzania bicrenata N.Kitag. ........................................................................................................147
Bazzania bidens (Gottsche et Lindenb.) Trevis. ...........................................................................148
Bazzania bidens var. heterodonta Spruce ......................................................................................148
Bazzania bidentula (Steph.) Yasuda.............................................................................................148
Bazzania bilobata N.Kitag. .........................................................................................................148
Bazzania borneensis N.Kitag........................................................................................................148
Bazzania brasiliensis (Gottsche et Lindenb.) Trevis. ....................................................................148
Bazzania brighamii (Austin) A.Evans ..........................................................................................148
Bazzania cadens N.Kitag. ............................................................................................................148
Bazzania calcarata (Sande Lac.) Schifn. .....................................................................................148
Bazzania callida (Sande Lac. ex Steph.) Abeyw. ..........................................................................148
Bazzania canelensis (Steph.) Fulford ............................................................................................148
Bazzania caudata (Steph.) Herzog ..............................................................................................148
Bazzania caudistipula (Steph.) Inoue et H.A.Mill. ......................................................................148
World checklist of hornworts and liverworts
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523
Bazzania ceylanica (Mitt.) Steph. ................................................................................................148
Bazzania chilensis (Steph.) Fulford ..............................................................................................148
Bazzania chimantensis Fulford.....................................................................................................148
Bazzania cincinnata (De Not.) Trevis. ........................................................................................149
Bazzania citharodes Meagher .......................................................................................................149
Bazzania combinata (J.B.Jack et Steph.) Steph. ...........................................................................149
Bazzania commutata (Lindenb. et Gottsche) Schifn. ..................................................................149
Bazzania comorensis Steph...........................................................................................................149
Bazzania confertifolia (Steph.) Herzog.........................................................................................149
Bazzania conistipula (Steph.) H.A.Mill. ......................................................................................149
Bazzania conophylla (Sande Lac.) Schifn. ...................................................................................149
Bazzania consanguinea (Hampe et Lindenb.) Trevis. ...................................................................149
Bazzania consociata (Steph.) H.A.Mill. .......................................................................................149
Bazzania corbieri (Steph.) Meagher .............................................................................................149
Bazzania crassidentata Fulford.....................................................................................................149
Bazzania crassitexta Steph. ..........................................................................................................149
Bazzania crenata (Trevis.) Trevis. ...............................................................................................149
Bazzania cubensis (Gottsche ex Steph.) Pagán .............................................................................149
Bazzania cucullata Onr. ..............................................................................................................150
Bazzania cuneistipula (Gottsche et Lindenb.) Trevis. ..................................................................150
Bazzania curvidens Steph. ...........................................................................................................150
Bazzania debilis N.Kitag. ............................................................................................................150
Bazzania deciduifolia Onr. ..........................................................................................................150
Bazzania decrescens (Lehm. et Lindenb.) Trevis. ..........................................................................150
Bazzania decrescens var. dentistipula Kiaer et Pearson...................................................................150
Bazzania decrescens subsp. molleri (Steph.) E.W.Jones .................................................................150
Bazzania decrescens subsp. pumila (Mitt.) Pócs ............................................................................150
Bazzania densa (Sande Lac.) Schifn............................................................................................150
Bazzania densa var. connata (Sande Lac.) Schifn. .......................................................................150
Bazzania denticulata (Lindenb. et Gottsche) Trevis. ...................................................................150
Bazzania denticulifera Mägd. ......................................................................................................150
Bazzania denudata (Lindenb. et Gottsche) Trevis. ......................................................................150
Bazzania deplanchei (Gottsche) Jovet-Ast ....................................................................................150
Bazzania deplanchei var. ilamentosa Tixier ..................................................................................150
Bazzania desciscens (Steph.) Meijer ..............................................................................................150
Bazzania didericiana (Gottsche ex Steph.) Steph. ........................................................................151
Bazzania diminuta Herzog..........................................................................................................151
Bazzania distans (Nees) Trevis. ...................................................................................................151
Bazzania diversicuspis Spruce ......................................................................................................151
Bazzania drepanophylla Herzog ...................................................................................................151
Bazzania dulitensis Herzog ..........................................................................................................151
Bazzania dulongensis L.P.Zhou et Li Zhang ................................................................................151
Bazzania eggersiana (Steph.) Pagán .............................................................................................151
Bazzania elmeri (Steph.) N.Kitag. ...............................................................................................151
Bazzania elongata Fulford ...........................................................................................................151
Bazzania emarginata (Steph.) C.M.Cooke ..................................................................................151
Bazzania engelii Glenny ..............................................................................................................151
Bazzania erosa (Reinw., Blume et Nees) Trevis. ..........................................................................151
Bazzania erosa var. pulopenangensis (Lindenb. et Gottsche) Schifn. ............................................151
Bazzania exempta J.J.Engel .........................................................................................................151
Bazzania falcata (Lindenb.) Trevis. .............................................................................................151
Bazzania falcifolia (Steph.) H.A.Mill. .........................................................................................151
524
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Bazzania fallax (Sande Lac.) Schifn. ..........................................................................................152
Bazzania fasciculata (Steph.) Meagher.........................................................................................152
Bazzania fauriana (Steph.) S.Hatt. .............................................................................................152
Bazzania iliformis Steph. ............................................................................................................152
Bazzania laccida (Dumort.) Grolle.............................................................................................152
Bazzania lavescens (Sande Lac. ex Steph.) Schifn. ......................................................................152
Bazzania leischeri (Steph.) Abeyw. .............................................................................................152
Bazzania francana (Steph.) N.Kitag. ...........................................................................................152
Bazzania friabilis N.Kitag. et T.Kodama.....................................................................................152
Bazzania fuhreri Meagher ...........................................................................................................152
Bazzania fuscescens A.Evans .........................................................................................................152
Bazzania gamscottii Meagher.......................................................................................................152
Bazzania gedeana (Steph.) Meijer................................................................................................152
Bazzania gracilis (Hampe et Gottsche) Steph. .............................................................................152
Bazzania grandiretis (Steph.) Herzog ..........................................................................................152
Bazzania griithiana (Steph.) Mizut. ..........................................................................................152
Bazzania gunniana (Steph.) H.A.Mill. ........................................................................................152
Bazzania hainanensis L.P.Zhou et Li Zhang................................................................................153
Bazzania halconiensis (Steph.) N.Kitag. .......................................................................................153
Bazzania hamatifolia (Steph.) H.A.Mill. .....................................................................................153
Bazzania harpago (De Not.) Schifn............................................................................................153
Bazzania hebridensis (Steph.) H.A.Mill. ......................................................................................153
Bazzania herminieri (Gottsche ex Steph.) Pagán .........................................................................153
Bazzania herzogiana Meijer .........................................................................................................153
Bazzania heterostipa (Steph.) Fulford ..........................................................................................153
Bazzania himalayana (Mitt.) Schifn. ..........................................................................................153
Bazzania hochstetteri (Reichardt) E.A.Hodgs. ..............................................................................153
Bazzania hookeri (Lindenb.) Trevis. ............................................................................................153
Bazzania horridula Schifn. .........................................................................................................153
Bazzania inaequabilis Steph. .......................................................................................................153
Bazzania inaequitexta Steph. .......................................................................................................153
Bazzania incrassata (Steph.) N.Kitag. ..........................................................................................153
Bazzania indica (Gottsche et Lindenb.) Trevis. ...........................................................................153
Bazzania indigenarum (Steph.) N.Kitag. .....................................................................................154
Bazzania insignis (De Not.) Trevis. .............................................................................................154
Bazzania intermedia (Gottsche et Lindenb.) Trevis. ....................................................................154
Bazzania intermedia var. sarawakiana (De Not.) Schifn. ............................................................154
Bazzania involuta (Mont.) Trevis................................................................................................154
Bazzania involuta var. submutica (Lindenb. et Gottsche) J.J.Engel et G.L.Merr. .........................154
Bazzania involutiformis (De Not.) Trevis. ...................................................................................154
Bazzania irregularis (Steph.) Schifn............................................................................................154
Bazzania jamaicensis (Lehm. et Lindenb.) Trevis. .......................................................................154
Bazzania japonica (Sande Lac.) Lindb. ........................................................................................154
Bazzania japonica var. sumatrana Herzog ....................................................................................154
Bazzania javanica (Sande Lac.) Schifn. ......................................................................................154
Bazzania kernii Steph. ................................................................................................................154
Bazzania kokawana N.Kitag. et T.Kodama .................................................................................154
Bazzania latifolia Steph...............................................................................................................154
Bazzania lehmanniana (Lindenb.) Trevis. ...................................................................................155
Bazzania leratii (Beauverd) H.A.Mill. .........................................................................................155
Bazzania lessonii (Steph.) H.A.Mill. ............................................................................................155
Bazzania levieri (Steph.) N.Kitag. ...............................................................................................155
World checklist of hornworts and liverworts
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525
Bazzania liebmaniana (Lindenb. et Gottsche) Trevis. .................................................................155
Bazzania linearis Herzog .............................................................................................................155
Bazzania linguiformis (Sande Lac.) Trevis. ..................................................................................155
Bazzania longa (Nees) Trevis. .....................................................................................................155
Bazzania longa var. papillata (Steph.) Fulford .............................................................................155
Bazzania longicaulis (Sande Lac.) Schifn. ...................................................................................155
Bazzania longicaulis var. latiareata Herzog ..................................................................................155
Bazzania longistipula (Lindenb.) Trevis. ......................................................................................155
Bazzania loricata (Reinw., Blume et Nees) Trevis. ......................................................................155
Bazzania lowii (Sande Lac. ex Steph.) Schifn. ............................................................................155
Bazzania luzonensis (Steph.) Del Ros. .........................................................................................156
Bazzania macgregorii Steph. ........................................................................................................156
Bazzania macrostipula Fulford ....................................................................................................156
Bazzania magna Horik................................................................................................................156
Bazzania magnistipula N.Kitag. ..................................................................................................156
Bazzania malaccensis (Steph.) Tixier ...........................................................................................156
Bazzania manillana (Gottsche ex Steph.) S.Hatt. ........................................................................156
Bazzania marginata (Steph.) N.Kitag. .........................................................................................156
Bazzania marginella (Herzog) N.Kitag. et T.Kodama .................................................................156
Bazzania mascarena (Steph.) Herzog ...........................................................................................156
Bazzania mayebarae S.Hatt. ........................................................................................................156
Bazzania menzelii E.D.Cooper ...................................................................................................156
Bazzania merrillana (Steph.) Inoue ex Bonner ............................................................................156
Bazzania minuta (Austin) A.Evans ..............................................................................................156
Bazzania minutidens (Steph.) Inoue et H.A.Mill. ........................................................................156
Bazzania minutiserra (Steph.) N.Kitag. .......................................................................................156
Bazzania missionum (Herzog) Jovet-Ast ......................................................................................157
Bazzania mittenii (Steph.) Steph. ................................................................................................157
Bazzania monilinervis (Lehm. et Lindenb.) Trevis.......................................................................157
Bazzania morokensis (Steph.) Grolle ............................................................................................157
Bazzania nitida (F.Weber) Grolle ...............................................................................................157
Bazzania novae-zelandiae (Mitt.) Besch. et C.Massal...................................................................157
Bazzania nudicaulis A.Evans .......................................................................................................157
Bazzania nuuanuensis C.M.Cooke ..............................................................................................157
Bazzania obcuneata (Steph.) H.A.Mill. .......................................................................................157
Bazzania obtusata (Mitt.) Abeyw. ...............................................................................................157
Bazzania okaritana Meagher et Glenny .......................................................................................157
Bazzania orbanii Pócs .................................................................................................................157
Bazzania ovistipula (Steph.) Abeyw. ............................................................................................157
Bazzania pallidevirens (Steph.) Fulford .......................................................................................157
Bazzania papillosa S.W.Arnell .....................................................................................................157
Bazzania paradoxa (Sande Lac.) Steph. .......................................................................................157
Bazzania parisii (Steph.) N.Kitag. ...............................................................................................157
Bazzania parvitexta Steph. ..........................................................................................................158
Bazzania patens (Mont.) Trevis. ..................................................................................................158
Bazzania patentistipa (Sande Lac.) Schifn...................................................................................158
Bazzania paucidens (Steph.) H.A.Mill. ........................................................................................158
Bazzania pearsonii Steph. ............................................................................................................158
Bazzania pectinata (Lindenb. et Gottsche) Schifn. .....................................................................158
Bazzania perfalcata N.Kitag. .......................................................................................................158
Bazzania perrotana E.W.Jones ....................................................................................................158
Bazzania peruviana (Nees) Trevis. ..............................................................................................158
526
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Bazzania phyllobola Spruce .........................................................................................................158
Bazzania placophylla (Taylor) Grolle ...........................................................................................158
Bazzania platycnema (Schwägr. ex Steph.) H.A.Mill. ..................................................................158
Bazzania pompeana (Sande Lac.) Mitt. .......................................................................................158
Bazzania praerupta (Reinw., Blume et Nees) Trevis. ...................................................................158
Bazzania praerupta var. obliquata (Nees) Schifn. ........................................................................158
Bazzania praerupta var. thermalis Schifn. ...................................................................................158
Bazzania pseudovittata N.Kitag. et T.Kodama ............................................................................159
Bazzania pusilla (Mitt.) Steph. ....................................................................................................159
Bazzania pycnophylla (Taylor) Trevis. .........................................................................................159
Bazzania quadratistipula H.A.Mill. .............................................................................................159
Bazzania rabenhorstii (Steph.) Abeyw. ........................................................................................159
Bazzania recurva (Mont.) Trevis. ................................................................................................159
Bazzania recurva var. major (Sande Lac.) Schifn. .......................................................................159
Bazzania relexa (Gottsche) Steph. ..............................................................................................159
Bazzania reinwardtii (Sande Lac.) Schifn. ..................................................................................159
Bazzania renistipula Steph. .........................................................................................................159
Bazzania revoluta (Steph.) N.Kitag. ............................................................................................159
Bazzania rimosa Meagher............................................................................................................159
Bazzania roccatii Gola.................................................................................................................159
Bazzania roraimensis (Steph.) Fulford .........................................................................................159
Bazzania sandvicensis (Gottsche ex Steph.) Steph. .......................................................................159
Bazzania sauropoda Meagher ......................................................................................................159
Bazzania scalaris Meagher ...........................................................................................................159
Bazzania schlimiana (Gottsche) Fulford ......................................................................................159
Bazzania schultze-motelii N.Kitag................................................................................................159
Bazzania schusterana N.Kitag. .....................................................................................................160
Bazzania schwaneckiana (Hampe et Gottsche) Trevis..................................................................160
Bazzania scutigera (Nees et Mont.) Trevis. ..................................................................................160
Bazzania semicordata (Lindenb. et Gottsche) Kuntze ..................................................................160
Bazzania serpentina (Nees) Trevis. ..............................................................................................160
Bazzania serrapiculata Inoue et H.A.Mill. ...................................................................................160
Bazzania serrata Fulford .............................................................................................................160
Bazzania serrulatoides Horik. ......................................................................................................160
Bazzania sikkimensis (Steph.) Herzog ..........................................................................................160
Bazzania spinosa S.Okamura .......................................................................................................160
Bazzania spiralis (Reinw., Blume et Nees) Meijer .......................................................................160
Bazzania spruceana Steph............................................................................................................160
Bazzania squarrosa (Steph.) H.A.Mill. ........................................................................................160
Bazzania stolonifera (Sw.) Trevis. ................................................................................................160
Bazzania stolonifera var. granatensis (Gottsche) Fulford ...............................................................160
Bazzania stresemannii (Herzog) N.Kitag. ....................................................................................160
Bazzania subacuta (Mitt.) Steph..................................................................................................161
Bazzania subaequitexta (Steph.) N.Kitag. ....................................................................................161
Bazzania subintegra (Steph.) L.Söderstr. et A.Hagborg ...............................................................161
Bazzania sublonga Fulford...........................................................................................................161
Bazzania subserrifolia (Beauverd) H.A.Mill. ................................................................................161
Bazzania subserrulata A.Evans .....................................................................................................161
Bazzania subtilis (Sande Lac.) Trevis. ..........................................................................................161
Bazzania succulenta N.Kitag. ......................................................................................................161
Bazzania sumatrana (Sande Lac. ex Steph.) Steph. ......................................................................161
Bazzania sumbavensis (Gottsche ex Steph.) Steph........................................................................161
World checklist of hornworts and liverworts
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527
Bazzania taleana (Gottsche) Fulford ...........................................................................................161
Bazzania tayloriana (Mitt.) Kuntze .............................................................................................161
Bazzania temariana (Steph.) H.A.Mill. .......................................................................................161
Bazzania tessellata Meagher .........................................................................................................161
Bazzania tiaoloensis Mizut. et K.C.Chang ...................................................................................161
Bazzania tricrenata (Wahlenb.) Lindb. .......................................................................................161
Bazzania tricrenata var. fulfordiae W.S.Hong ..............................................................................161
Bazzania tridens (Reinw., Blume et Nees) Trevis. .......................................................................162
Bazzania tridens var. assamica (Steph.) Pócs ................................................................................162
Bazzania tridens var. cornutistipula (Steph.) Pócs ........................................................................162
Bazzania trilobata (L.) Gray ........................................................................................................162
Bazzania trilobata var. depauperata (Müll.Frib.) Grolle ...............................................................162
Bazzania uncigera (Reinw., Blume et Nees) Trevis. .....................................................................162
Bazzania uncigera var. brevifolia Herzog .....................................................................................162
Bazzania uncigera var. gibba (Sande Lac.) Meijer ........................................................................162
Bazzania undulata Herzog ..........................................................................................................162
Bazzania vietnamica Pócs............................................................................................................162
Bazzania vitiana Mitt. ................................................................................................................162
Bazzania vittata (Gottsche) Trevis. .............................................................................................162
Bazzania vittata var. luxurians (De Not.) Schifn. .......................................................................162
Bazzania wallichiana (Lindenb.) Trevis. .....................................................................................162
Bazzania watanabei Inoue ..........................................................................................................162
Bazzania wattsiana (Steph.) Meagher..........................................................................................162
Bazzania wiltensii (Sande Lac. ex Steph.) Schifn. .......................................................................163
Bazzania wooroonooran Meagher ................................................................................................163
Bazzania wrightii (Gottsche ex Steph.) Steph..............................................................................163
Bazzania yoshinagana (Steph.) Yasuda.........................................................................................163
Bazzania zollingeri (Lindenb.) Trevis. .........................................................................................163
Bazzania zonulata Meagher ........................................................................................................163
Biantheridion undulifolium (Nees) Konstant. et Vilnet ..................................................................52
Blasia pusilla L. ...........................................................................................................................476
Blepharidophyllum densifolium (Hook.) Ångstr. ex C.Massal. ......................................................103
Blepharidophyllum vertebrale (Gottsche) Ångstr. ex C.Massal. .....................................................103
Blepharolejeunea chimantaensis van Slageren et Kruijt ..................................................................298
Blepharolejeunea fuegiana (Besch. et C.Massal.) Gradst. ..............................................................298
Blepharolejeunea incongrua (Lindenb. et Gottsche) van Slageren et Kruijt ...................................298
Blepharolejeunea saccata (Steph.) van Slageren et Kruijt ...............................................................298
Blepharolejeunea securifolia (Steph.) R.M.Schust. ........................................................................298
Blepharostoma arachnoideum M.Howe ........................................................................................137
Blepharostoma indicum G.Asthana, M.Saxena et Maurya.............................................................137
Blepharostoma minor Horik. ........................................................................................................137
Blepharostoma trichophyllum (L.) Dumort. ..................................................................................137
Blepharostoma trichophyllum subsp. brevirete (Bryhn et Kaal.) R.M.Schust. .................................137
Brachiolejeunea conduplicata (Steph.) Gradst. ..............................................................................298
Brachiolejeunea fernandeziana S.W.Arnell ...................................................................................298
Brachiolejeunea laxifolia (Taylor) Schifn. ....................................................................................298
Brachiolejeunea leiboldiana (Gottsche et Lindenb.) Schifn. .........................................................298
Brachiolejeunea phyllorhiza (Nees) Kruijt et Gradst. ....................................................................298
Brachiolejeunea spruceana (C.Massal.) Schifn. ............................................................................298
Bragginsella anomala R.M.Schust. ...............................................................................................189
Brevianthus lavus (Grolle) J.J.Engel et R.M.Schust.....................................................................137
Brevianthus lavus subsp. crenulatus J.J.Engel...............................................................................137
528
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Brevianthus hypocanthidium M.A.M.Renner et J.J.Engel .............................................................137
Bromeliophila helenae Gradst. ......................................................................................................350
Bromeliophila natans (Steph.) R.M.Schust...................................................................................351
Bryopteris difusa (Sw.) Nees ........................................................................................................403
Bryopteris ilicina (Sw.) Nees .......................................................................................................403
Bryopteris issiloba Steph. .............................................................................................................403
Bryopteris gaudichaudii Gottsche .................................................................................................403
Bucegia romanica Radian.............................................................................................................488
Calatholejeunea lamii Mizut. .......................................................................................................317
Calatholejeunea paradoxa (Schifn.) K.I.Goebel ...........................................................................317
Calycularia crispula Mitt. ............................................................................................................463
Calycularia laxa Lindb. et Arnell .................................................................................................463
Calypogeia aeruginosa Mitt. .........................................................................................................106
Calypogeia amazonica (Spruce) Steph. .........................................................................................106
Calypogeia andicola Bischl. ..........................................................................................................104
Calypogeia angusta Steph. ............................................................................................................106
Calypogeia annabonensis Steph. ...................................................................................................104
Calypogeia apiculata (Steph.) Steph. ............................................................................................106
Calypogeia arguta Nees et Mont. .................................................................................................104
Calypogeia asakawana S.Hatt. ex Inoue .......................................................................................106
Calypogeia azorica Bischl. ............................................................................................................104
Calypogeia azurea Stotler et Crotz ...............................................................................................104
Calypogeia bidentula (F.Weber) Nees ..........................................................................................104
Calypogeia ceylanica S.Hatt. et Mizut. .........................................................................................106
Calypogeia contracta Inoue ..........................................................................................................106
Calypogeia cuspidata (Steph.) Steph. ............................................................................................106
Calypogeia decurrens (Steph.) Steph. ............................................................................................106
Calypogeia falcata Bischl. .............................................................................................................104
Calypogeia issa (L.) Raddi ...........................................................................................................104
Calypogeia issa subsp. neogaea R.M.Schust..................................................................................104
Calypogeia formosana Horik. .......................................................................................................106
Calypogeia fujisana Inoue ............................................................................................................106
Calypogeia goebelii (Schifn.) Steph..............................................................................................104
Calypogeia goebelii var. siamensis N.Kitag. ...................................................................................104
Calypogeia grandistipula (Steph.) Steph. ......................................................................................104
Calypogeia granulata Inoue..........................................................................................................106
Calypogeia integristipula Steph.....................................................................................................104
Calypogeia japonica Steph............................................................................................................106
Calypogeia khasiana Ajit P.Singh et V.Nath ................................................................................106
Calypogeia latissima Steph. ..........................................................................................................106
Calypogeia laxa Gottsche et Lindenb. ..........................................................................................104
Calypogeia lechleri (Steph.) Steph. ...............................................................................................104
Calypogeia lechleri var. densifolia (Steph.) Bischl. .........................................................................104
Calypogeia longifolia Steph. .........................................................................................................104
Calypogeia lophocoleoides Steph. ...................................................................................................104
Calypogeia lunata Mitt. ...............................................................................................................106
Calypogeia marginella Mitt. .........................................................................................................107
Calypogeia mascarenensis Bischl. ..................................................................................................105
Calypogeia microstipula (Steph.) Steph. .......................................................................................105
Calypogeia miquelii Mont. ex Gottsche, Lindenb. et Nees ...........................................................105
Calypogeia muelleriana (Schifn.) Müll.Frib.................................................................................105
Calypogeia muelleriana subsp. blomquistii R.M.Schust. ...............................................................105
World checklist of hornworts and liverworts
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529
Calypogeia neesiana (C.Massal. et Carestia) Müll.Frib. ................................................................105
Calypogeia neesiana subsp. subalpina (Inoue) Inoue .....................................................................105
Calypogeia oblata Herzog ............................................................................................................105
Calypogeia obovata R.M.Schust. ..................................................................................................107
Calypogeia peruviana Nees et Mont. ............................................................................................105
Calypogeia rhombifolia (Spruce) Steph. ........................................................................................105
Calypogeia rhombifolia var. colombiana Bischl..............................................................................105
Calypogeia sphagnicola (Arnell et J.Perss.) Warnst. et Loeske .......................................................105
Calypogeia steyermarkii Fulford ...................................................................................................107
Calypogeia subintegra (Gottsche, Lindenb. et Nees) Bischl. .........................................................105
Calypogeia subintegra var. dussiana (Steph.) Bischl.......................................................................105
Calypogeia suecica (Arnell et J.Perss.) Müll.Frib. ..........................................................................106
Calypogeia sullivantii Austin ........................................................................................................104
Calypogeia tenax (Spruce) Steph. .................................................................................................106
Calypogeia tosana (Steph.) Steph. ................................................................................................107
Calypogeia udarii Sudipa Das et D.K.Singh ................................................................................107
Calypogeia uncinulatula Herzog...................................................................................................106
Capillolejeunea geisslerae (Pócs) R.L.Zhu, Qiong He, Y.M.Wei et Pócs .......................................353
Capillolejeunea mascarena S.W.Arnell..........................................................................................353
Castanoclobos julaceus (Hatcher ex J.J.Engel) J.J.Engel et Glenny ................................................254
Caudalejeunea acutifolia Gerola ...................................................................................................404
Caudalejeunea africana (Steph.) Schifn. .....................................................................................404
Caudalejeunea cristiloba (Steph.) Gradst. .....................................................................................404
Caudalejeunea cristiloba subsp. samoana (Steph.) Gradst. ............................................................404
Caudalejeunea dusenii Steph........................................................................................................404
Caudalejeunea grolleana Gradst. ..................................................................................................403
Caudalejeunea hanningtonii (Mitt.) Schifn. ................................................................................404
Caudalejeunea katangensis Vanden Berghen ................................................................................404
Caudalejeunea lehmanniana (Gottsche) A.Evans .........................................................................404
Caudalejeunea lewallei Vanden Berghen ......................................................................................404
Caudalejeunea mauritiana Horik.................................................................................................404
Caudalejeunea reniloba (Gottsche) Steph.....................................................................................404
Caudalejeunea streimannii Gyarmati ...........................................................................................404
Caudalejeunea tridentata R.L.Zhu, Y.M.Wei et Qiong He ..........................................................404
Caudalejeunea yangambiensis (Vanden Berghen) E.W.Jones ........................................................404
Cavicularia densa Steph. .............................................................................................................476
Cephalantholejeunea temnanthoides (R.M.Schust.) R.M.Schust. ..................................................405
Cephalojonesia incuba Grolle et Vanden Berghen ..........................................................................66
Cephalojonesia incuba subsp. mexicana Burghardt, Gradst. et Váňa ...............................................66
Cephalolejeunea parvilobula Mizut...............................................................................................405
Cephalomitrion aterrimum (Steph.) R.M.Schust. ...........................................................................66
Cephalozia acuminata (Herzog) Váňa............................................................................................57
Cephalozia acutiloba (Inoue) Váňa ................................................................................................57
Cephalozia albula Steph. ...............................................................................................................57
Cephalozia ambigua C.Massal. ......................................................................................................57
Cephalozia austrigena R.M.Schust. ex J.J.Engel .............................................................................57
Cephalozia badia (Gottsche) Steph................................................................................................57
Cephalozia bicuspidata (L.) Dumort. .............................................................................................57
Cephalozia bicuspidata subsp. lammersiana (Huebener) R.M.Schust. ............................................57
Cephalozia chilensis (J.J.Engel et R.M.Schust.) R.M.Schust. ..........................................................57
Cephalozia conchata (Grolle et Váňa) Váňa ...................................................................................57
Cephalozia crossii Spruce ...............................................................................................................57
530
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Cephalozia darjeelingensis Udar et D.Kumar..................................................................................57
Cephalozia drucei (R.M.Schust.) Váňa ..........................................................................................58
Cephalozia fuegiensis Váňa .............................................................................................................58
Cephalozia hamatiloba Steph. ........................................................................................................58
Cephalozia hamatiloba subsp. siamensis (N.Kitag.) Váňa ...............................................................58
Cephalozia hians Steph. .................................................................................................................59
Cephalozia indica Udar et D.Kumar .............................................................................................59
Cephalozia kodaikanalensis G.Asthana et Saumya Srivast. ..............................................................59
Cephalozia lacinulata (J.B.Jack ex Gottsche et Rabenh.) Spruce ....................................................58
Cephalozia lucens (A.Evans) Steph. ................................................................................................58
Cephalozia macgregorii (Steph.) Váňa ............................................................................................58
Cephalozia macounii (Austin) Austin.............................................................................................58
Cephalozia maxima Steph. ............................................................................................................58
Cephalozia mollusca (De Not.) Váňa .............................................................................................58
Cephalozia neesiana Steph. ............................................................................................................58
Cephalozia nishimurae (N.Kitag.) Váňa .........................................................................................58
Cephalozia pachygyna R.M.Schust. ex J.J.Engel .............................................................................58
Cephalozia parvifolia Arnell...........................................................................................................59
Cephalozia physocaula (Hook.f. et Taylor) Steph. ..........................................................................58
Cephalozia schusteriana J.J.Engel ...................................................................................................58
Cephalozia stolonacea (Herzog) Váňa .............................................................................................58
Cephalozia tricuspidata (Nees) Trevis. ...........................................................................................59
Cephalozia tubulata (Hook.f. et Taylor) Trevis. ............................................................................58
Cephaloziella acanthophora (S.Hatt.) Horik. ..................................................................................71
Cephaloziella aenigmatica R.M.Schust. ..........................................................................................66
Cephaloziella anthelioides S.W.Arnell ............................................................................................66
Cephaloziella antillana (Besch. et Spruce) Fulford .........................................................................70
Cephaloziella arctogena (R.M.Schust.) Konstant. ...........................................................................66
Cephaloziella arenaria (Steph.) R.M.Schust. ..................................................................................66
Cephaloziella aspericaulis Jørg. .......................................................................................................66
Cephaloziella baumgartneri Schifn. ...............................................................................................66
Cephaloziella biokoensis Váňa et Frank Müll. .................................................................................71
Cephaloziella breviperianthia C.Gao ..............................................................................................66
Cephaloziella brinkmannii Douin ..................................................................................................66
Cephaloziella calyculata (Durieu et Mont.) Müll.Frib. ...................................................................70
Cephaloziella capensis (Sim) S.W.Arnell.........................................................................................66
Cephaloziella capillaris (Steph.) Douin ..........................................................................................67
Cephaloziella crassigyna (R.M.Schust.) R.M.Schust. ......................................................................67
Cephaloziella crispata N.Kitag. ......................................................................................................67
Cephaloziella densifolia R.M.Schust. ..............................................................................................67
Cephaloziella densifolia var. dubia R.M.Schust. .............................................................................67
Cephaloziella dentata (Raddi) Steph. .............................................................................................70
Cephaloziella dentifolia Udar et Ad.Kumar ....................................................................................71
Cephaloziella divaricata (Sm.) Schifn. ..........................................................................................67
Cephaloziella divaricata var. scabra (M.Howe) Haynes ..................................................................67
Cephaloziella dusenii Steph. ...........................................................................................................67
Cephaloziella elachista (J.B.Jack ex Gottsche et Rabenh.) Schifn. ..................................................67
Cephaloziella elachista var. spinophylla (C.Gao) C.Gao ..................................................................67
Cephaloziella elegans (Heeg) Schifn. .............................................................................................67
Cephaloziella exigua R.M.Schust. ..................................................................................................67
Cephaloziella exililora (Taylor) Douin ..........................................................................................67
Cephaloziella ilum (Trevis.) Steph. ...............................................................................................71
World checklist of hornworts and liverworts
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531
Cephaloziella lexuosa C.Gao et K.C.Chang...................................................................................71
Cephaloziella fragillima (Spruce) Fulford .......................................................................................67
Cephaloziella garsidei S.W.Arnell...................................................................................................67
Cephaloziella granatensis (J.B.Jack ex Steph.) Fulford ....................................................................71
Cephaloziella grandiretis (R.M.Schust.) R.M.Schust. .....................................................................67
Cephaloziella grimsulana (J.B.Jack ex Gottsche et Rabenh.) Lacout. ..............................................68
Cephaloziella grisea R.M.Schust. ...................................................................................................71
Cephaloziella hampeana (Nees) Schifn. ex Loeske .........................................................................68
Cephaloziella hebridensis Steph. .....................................................................................................68
Cephaloziella herzogiana (Pandé et K.P.Srivast.) Udar et D.Kumar ...............................................68
Cephaloziella heteroica (C.M.Cooke) Douin..................................................................................68
Cephaloziella hirta (Steph.) R.M.Schust. .......................................................................................70
Cephaloziella hyalina Douin ..........................................................................................................68
Cephaloziella hyalina var. rappii (Douin) R.M.Schust. ..................................................................68
Cephaloziella inaequalis R.M.Schust..............................................................................................68
Cephaloziella integerrima (Lindb.) Warnst.....................................................................................70
Cephaloziella intricata Schifn. ex Douin .......................................................................................71
Cephaloziella invisa R.M.Schust. ...................................................................................................68
Cephaloziella kilohanensis (C.M.Cooke) Douin .............................................................................68
Cephaloziella levieri Schifn. ex Douin...........................................................................................72
Cephaloziella longii Váňa ...............................................................................................................68
Cephaloziella lycopodioides (Sim) S.W.Arnell .................................................................................68
Cephaloziella mammillifera R.M.Schust. et Damsh. ......................................................................68
Cephaloziella massalongi (Spruce) Müll.Frib. .................................................................................68
Cephaloziella meghalayensis Udar et Ad.Kumar..............................................................................71
Cephaloziella microphylla (Steph.) Douin ......................................................................................71
Cephaloziella muelleriana R.M.Schust. ..........................................................................................68
Cephaloziella natalensis (Sim) S.W.Arnell ......................................................................................68
Cephaloziella nicholsonii Douin .....................................................................................................68
Cephaloziella nothogena R.M.Schust. .............................................................................................68
Cephaloziella obtusilobula R.M.Schust. ..........................................................................................68
Cephaloziella patulifolia (Steph.) Douin ........................................................................................69
Cephaloziella pellucida R.M.Schust. ..............................................................................................72
Cephaloziella phyllacantha (C.Massal. et Carestia) Müll.Frib. ........................................................69
Cephaloziella polystratosa (R.M.Schust. et Damsh.) Konstant. .......................................................69
Cephaloziella pseudocrassigyna R.M.Schust. ex J.J.Engel.................................................................69
Cephaloziella pulcherrima R.M.Schust. ..........................................................................................69
Cephaloziella pulcherrima subsp. sphagnicola R.M.Schust. .............................................................69
Cephaloziella pungens Steph. ex Fulford.........................................................................................69
Cephaloziella pygmaea (Spruce) Váňa ............................................................................................72
Cephaloziella rubella (Nees) Warnst. .............................................................................................69
Cephaloziella schelpei S.W.Arnell ...................................................................................................69
Cephaloziella secundifolia Pearson ..................................................................................................72
Cephaloziella sinensis Douin ..........................................................................................................72
Cephaloziella spegazziniana (C.Massal.) Douin .............................................................................69
Cephaloziella spinicaulis Douin......................................................................................................69
Cephaloziella spinigera (Lindb.) Jørg..............................................................................................69
Cephaloziella squarrosula (Trevis.) Steph. ......................................................................................71
Cephaloziella stellulifera (Taylor ex Carrington et Pearson) Croz. ..................................................69
Cephaloziella stephanii Schifn. ex Douin ......................................................................................69
Cephaloziella stolonifera R.M.Schust..............................................................................................69
Cephaloziella subspinosa R.M.Schust. ............................................................................................71
532
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Cephaloziella subtilis (Lindenb. et Gottsche) Steph........................................................................69
Cephaloziella sumatrana Schifn. ex Douin ....................................................................................69
Cephaloziella tabularis S.W.Arnell .................................................................................................69
Cephaloziella tenuissima (Lehm.) Steph. ........................................................................................71
Cephaloziella transvaalensis S.W.Arnell..........................................................................................69
Cephaloziella triplicata S.W.Arnell ................................................................................................69
Cephaloziella turneri (Hook.) Müll.Frib. .......................................................................................71
Cephaloziella umtaliensis S.W.Arnell .............................................................................................70
Cephaloziella uncinata R.M.Schust................................................................................................70
Cephaloziella uncinata var. brevigyna R.M.Schust. et Damsh.........................................................70
Cephaloziella uncinata var. sphagnicola R.M.Schust. ......................................................................70
Cephaloziella vaginans Steph. ........................................................................................................70
Cephaloziella varians (Gottsche) Steph. .........................................................................................70
Cephaloziella verrucosa Steph.........................................................................................................70
Cephaloziella villaumei (Steph.) Váňa ............................................................................................70
Cephaloziella violacea Schljakov.....................................................................................................70
Cephaloziella welwitschii (Steph.) Douin .......................................................................................70
Cephaloziopsis intertexta (Gottsche) R.M.Schust............................................................................72
Ceramanus centipes (Lindenb. et Gottsche) E.D.Cooper..............................................................169
Ceramanus clatritexta (Steph.) E.D.Cooper .................................................................................169
Ceramanus elegans (Colenso) E.D.Cooper ...................................................................................169
Ceramanus grossiseta (Steph.) E.D.Cooper...................................................................................169
Ceramanus perfragilis (J.J.Engel et G.L.Merr.) E.D.Cooper.........................................................169
Ceramanus pruinosa (Herzog) E.D.Cooper .................................................................................169
Ceramanus tuberifera (J.J.Engel et R.M.Schust.) E.D.Cooper .....................................................169
Ceratolejeunea aliena Herzog .......................................................................................................303
Ceratolejeunea andringitrae Pócs ..................................................................................................301
Ceratolejeunea atlantica Alvarenga et Ilk.-Borg. ...........................................................................301
Ceratolejeunea belangeriana (Gottsche) Steph. .............................................................................301
Ceratolejeunea beninensis E.W.Jones et Vanden Berghen .............................................................301
Ceratolejeunea brevinervis (Spruce) A.Evans ................................................................................301
Ceratolejeunea ceratantha (Nees et Mont.) Schifn. ......................................................................301
Ceratolejeunea coalita (Ångstr.) Steph. .........................................................................................301
Ceratolejeunea coarina (Gottsche) Schifn. ...................................................................................301
Ceratolejeunea confusa R.M.Schust. .............................................................................................302
Ceratolejeunea cornuta (Lindenb.) Steph. ....................................................................................302
Ceratolejeunea cubensis (Mont.) Schifn. ......................................................................................302
Ceratolejeunea dentistipula Steph. ................................................................................................302
Ceratolejeunea desciscens (Sande Lac.) Schifn. .............................................................................303
Ceratolejeunea fallax (Lehm. et Lindenb.) Bonner .......................................................................302
Ceratolejeunea ilaria (Taylor) Steph. ..........................................................................................302
Ceratolejeunea loribunda Steph. ..................................................................................................302
Ceratolejeunea globulifera Herzog ................................................................................................303
Ceratolejeunea grandiloba J.B.Jack et Steph. ................................................................................303
Ceratolejeunea grandiloba subsp. inlata (Mizut.) Gradst. ............................................................303
Ceratolejeunea guianensis (Nees et Mont.) Steph. ........................................................................302
Ceratolejeunea karstenii Steph......................................................................................................302
Ceratolejeunea kuerschneri Eb.Fisch. et Vanderp. .........................................................................302
Ceratolejeunea laetefusca (Austin) R.M.Schust. ............................................................................302
Ceratolejeunea ledermannii Steph. ...............................................................................................302
Ceratolejeunea malleigera (Spruce) Steph. ....................................................................................302
Ceratolejeunea maranhensis Silva Brito et Ilk.-Borg. .....................................................................302
World checklist of hornworts and liverworts
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533
Ceratolejeunea minor Mizut. ........................................................................................................302
Ceratolejeunea minuta G.Dauphin ..............................................................................................302
Ceratolejeunea moniliata Herzog .................................................................................................302
Ceratolejeunea oculata (Gottsche) Steph. .....................................................................................302
Ceratolejeunea oxygonia Steph......................................................................................................302
Ceratolejeunea papulilora Steph. .................................................................................................303
Ceratolejeunea patentissima (Hampe et Gottsche) A.Evans ..........................................................303
Ceratolejeunea pungens Steph.......................................................................................................303
Ceratolejeunea rubiginosa Steph. ..................................................................................................303
Ceratolejeunea saroltae Pócs .........................................................................................................303
Ceratolejeunea sinensis P.C.Chen et P.C.Wu ...............................................................................303
Ceratolejeunea singapurensis (Lindenb.) Schifn. ..........................................................................303
Ceratolejeunea spinosa (Gottsche) Steph. .....................................................................................303
Ceratolejeunea szyszylowiczii (Loitl.) Steph...................................................................................303
Ceratolejeunea temnantha (Spruce) M.E.Reiner ...........................................................................303
Ceratolejeunea umbonata Steph. ..................................................................................................303
Ceratolejeunea vitiensis Steph.......................................................................................................303
Ceratolejeunea zenkeri Steph........................................................................................................303
Chaetocolea palmata Spruce .........................................................................................................254
Chaetophyllopsis whiteleggei (Carrington et Pearson) R.M.Schust. ex Hamlin ................................72
Chandonanthus squarrosus (Menzies) Mitt. ....................................................................................52
Cheilolejeunea acanthina (Spruce) Gradst. et Ilk.-Borg. ...............................................................305
Cheilolejeunea aciculifera R.M.Schust. .........................................................................................311
Cheilolejeunea acutangula (Nees) Grolle ......................................................................................311
Cheilolejeunea adnata (Kunze ex Lehm.) Grolle ..........................................................................305
Cheilolejeunea adnata var. autoica Gradst. et Ilk.-Borg. ...............................................................305
Cheilolejeunea albovirens (Hook.f. et Taylor) E.A.Hodgs.............................................................305
Cheilolejeunea aneogyna (Spruce) A.Evans ...................................................................................305
Cheilolejeunea ascensionis (Hook.f. et Taylor) Grolle ...................................................................305
Cheilolejeunea asperilora (Spruce) Gradst. et Ilk.-Borg. ...............................................................305
Cheilolejeunea asperrima (Steph.) Grolle ......................................................................................305
Cheilolejeunea australis Solari ......................................................................................................311
Cheilolejeunea baumannii Hürl. ..................................................................................................305
Cheilolejeunea beyrichii (Lindenb.) M.E.Reiner ...........................................................................305
Cheilolejeunea birmensis (Steph.) Mizut. ......................................................................................311
Cheilolejeunea boninensis Mizut. ..................................................................................................305
Cheilolejeunea caducifolia (Gradst. et Schäf.-Verw.) W.Ye et R.L.Zhu .........................................306
Cheilolejeunea camerunensis S.W.Arnell .......................................................................................306
Cheilolejeunea campbelliensis (Steph.) R.M.Schust. ......................................................................311
Cheilolejeunea cedercreutzii (H.Buch et Perss.) Grolle ..................................................................311
Cheilolejeunea celebensis (Steph.) Mizut. ......................................................................................306
Cheilolejeunea ceylanica (Gottsche) R.M.Schust. et Kachroo .......................................................314
Cheilolejeunea chenii R.L.Zhu et M.L.So.....................................................................................306
Cheilolejeunea choachina (Gottsche) Gradst. ................................................................................311
Cheilolejeunea clausa (Nees et Mont.) R.M.Schust. .....................................................................311
Cheilolejeunea clavata Mizut. ......................................................................................................311
Cheilolejeunea clypeata (Schwein.) W.Ye et R.L.Zhu ...................................................................306
Cheilolejeunea comans (Spruce) R.M.Schust. ...............................................................................312
Cheilolejeunea compressa (Herzog) Grolle ....................................................................................306
Cheilolejeunea conchifolia (A.Evans) W.Ye et R.L.Zhu ................................................................306
Cheilolejeunea convexa (S.W.Arnell) S.W.Arnell ..........................................................................312
Cheilolejeunea cookiensis (Steph.) R.M.Schust. et Kachroo...........................................................312
534
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Cheilolejeunea cordigera (Steph.) Grolle .......................................................................................306
Cheilolejeunea cordistipula (Steph.) Grolle ex E.W.Jones .............................................................312
Cheilolejeunea coronalis (Gottsche) R.M.Schust. ..........................................................................306
Cheilolejeunea curvatilobula (Herzog) Grolle ...............................................................................306
Cheilolejeunea decursiva (Sande Lac.) R.M.Schust. ......................................................................306
Cheilolejeunea discoidea (Lehm. et Lindenb.) R.M.Schust. et Kachroo ........................................306
Cheilolejeunea diversifolia Augier .................................................................................................306
Cheilolejeunea ecarinata Vanden Berghen ....................................................................................306
Cheilolejeunea erostrata R.M.Schust. ...........................................................................................306
Cheilolejeunea eximia (Jovet-Ast et Tixier) R.L.Zhu et M.L.So....................................................314
Cheilolejeunea exinnovata E.W.Jones ...........................................................................................306
Cheilolejeunea falsinervis (Sande Lac.) R.M.Schust. et Kachroo ...................................................314
Cheilolejeunea ischeri Malombe ..................................................................................................312
Cheilolejeunea fragrantissima (Spruce) R.M.Schust. .....................................................................306
Cheilolejeunea fukiensis (P.C.Chen et P.C.Wu) Piippo ................................................................307
Cheilolejeunea galliotii Steph. ......................................................................................................307
Cheilolejeunea gaoi R.L.Zhu, M.L.So et Grolle............................................................................307
Cheilolejeunea gardneri (Mitt.) Mizut. .........................................................................................314
Cheilolejeunea germanii (Besch. et Spruce) Grolle........................................................................307
Cheilolejeunea ghatensis G.Asthana, S.C.Srivast. et A.K.Asthana ..................................................307
Cheilolejeunea gigantea (Steph.) R.M.Schust. et Kachroo ............................................................314
Cheilolejeunea gottscheana C.J.Bastos ...........................................................................................315
Cheilolejeunea gradsteinii (Grolle et Piippo) W.Ye et R.L.Zhu.....................................................307
Cheilolejeunea grandibracteata Steph. ..........................................................................................307
Cheilolejeunea hallingii B.M.hiers .............................................................................................312
Cheilolejeunea hamlinii Grolle .....................................................................................................307
Cheilolejeunea hawaica Steph. .....................................................................................................307
Cheilolejeunea herzogiana Steph...................................................................................................307
Cheilolejeunea heteroclada (Spruce) Schifn. .................................................................................315
Cheilolejeunea huerlimannii Tixier ..............................................................................................314
Cheilolejeunea hyalomarginata R.L.Zhu et Frank Müll. ...............................................................314
Cheilolejeunea implexicaulis (Hook.f. et Taylor) R.M.Schust. ......................................................307
Cheilolejeunea incisa (Gottsche) R.M.Schust. et Kachroo ............................................................314
Cheilolejeunea incisa var. teretilora B.M.hiers............................................................................314
Cheilolejeunea inlexa (Hampe) Grolle.........................................................................................312
Cheilolejeunea insecta Grolle et Gradst.........................................................................................307
Cheilolejeunea insignis Jovet-Ast et Tixier ....................................................................................307
Cheilolejeunea intertexta (Lindenb.) Steph. ..................................................................................312
Cheilolejeunea intricata (Steph.) J.J.Engel ....................................................................................307
Cheilolejeunea invaginata R.M.Schust. ........................................................................................307
Cheilolejeunea jamaicensis Steph. .................................................................................................307
Cheilolejeunea japonica (Horik.) W.Ye et R.L.Zhu ......................................................................307
Cheilolejeunea kitagawae W.Ye et R.L.Zhu..................................................................................307
Cheilolejeunea krakakammae (Lindenb.) R.M.Schust. .................................................................312
Cheilolejeunea lacerata C.J.Bastos et Gradst.................................................................................307
Cheilolejeunea laciniata D.F.Peralta et M.E.Reiner......................................................................312
Cheilolejeunea laevicalyx (J.B.Jack et Steph.) Grolle .....................................................................307
Cheilolejeunea laeviuscula (Mitt.) Steph. ......................................................................................307
Cheilolejeunea larsenii Mizut. ......................................................................................................308
Cheilolejeunea laurentii Steph. .....................................................................................................308
Cheilolejeunea leptophylla (Ångstr.) Steph. ...................................................................................308
Cheilolejeunea lindenbergii (Gottsche) Mizut. ..............................................................................312
World checklist of hornworts and liverworts
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535
Cheilolejeunea longidens (Steph.) R.M.Schust. et Kachroo ...........................................................314
Cheilolejeunea longilora (Taylor) R.M.Schust. ............................................................................308
Cheilolejeunea longispina (Herzog) R.M.Schust. ..........................................................................308
Cheilolejeunea loriana (Steph.) W.Ye et R.L.Zhu ........................................................................308
Cheilolejeunea ludoviciae Steph. ...................................................................................................308
Cheilolejeunea lurida (Lindenb.) Steph. .......................................................................................308
Cheilolejeunea macroloba (Herzog) Grolle ...................................................................................308
Cheilolejeunea malaccensis (G.Hofm.) Xiao L.He .......................................................................308
Cheilolejeunea mammifera R.M.Schust. .......................................................................................308
Cheilolejeunea mariana (Gottsche) B.M.hiers et Gradst. ...........................................................308
Cheilolejeunea mexicana Steph.....................................................................................................308
Cheilolejeunea meyeniana (Nees, Lindenb. et Gottsche) R.M.Schust. et Kachroo ........................314
Cheilolejeunea micholitzii (Steph.) R.M.Schust. et Kachroo.........................................................308
Cheilolejeunea microscypha (Hook.f. et Taylor) M.Wigginton .....................................................309
Cheilolejeunea mimosa (Hook.f. et Taylor) R.M.Schust. ..............................................................312
Cheilolejeunea minutilobula Amakawa .........................................................................................315
Cheilolejeunea mizutanii W.Ye et R.L.Zhu ..................................................................................309
Cheilolejeunea montagnei (Gottsche ex Mont.) R.M.Schust. ........................................................314
Cheilolejeunea nana R.M.Schust. ................................................................................................312
Cheilolejeunea neblinensis Ilk.-Borg. et Gradst. ............................................................................309
Cheilolejeunea ngongensis Malombe et Pócs .................................................................................312
Cheilolejeunea nietneri (Steph.) Mizut. ........................................................................................309
Cheilolejeunea nipponica (S.Hatt.) S.Hatt. ...................................................................................312
Cheilolejeunea norisiae G.Dauphin et Gradst. ..............................................................................309
Cheilolejeunea norrisii (Grolle) M.A.M.Renner ...........................................................................315
Cheilolejeunea novaezelandiae R.M.Schust...................................................................................309
Cheilolejeunea obcordata Herzog ..................................................................................................309
Cheilolejeunea obtruncata (Mont.) Solari .....................................................................................309
Cheilolejeunea obtusa (Herzog) Solari ..........................................................................................309
Cheilolejeunea obtusifolia (Steph.) S.Hatt. ...................................................................................309
Cheilolejeunea obtusilobula (S.Hatt.) S.Hatt. ...............................................................................314
Cheilolejeunea occlusa (Herzog) T.Kodama et N.Kitag. ...............................................................309
Cheilolejeunea omphalogastria Pócs ..............................................................................................313
Cheilolejeunea oncophylla (Ångstr.) Grolle et M.E.Reiner ............................................................313
Cheilolejeunea orientalis (Gottsche) Mizut. ..................................................................................309
Cheilolejeunea ornata C.J.Bastos ..................................................................................................313
Cheilolejeunea oscilla M.A.M.Renner...........................................................................................315
Cheilolejeunea osumiensis (S.Hatt.) Mizut. ...................................................................................313
Cheilolejeunea ovistipula Steph. ...................................................................................................309
Cheilolejeunea panurensis (Spruce) Steph. ....................................................................................309
Cheilolejeunea papillata Solari .....................................................................................................309
Cheilolejeunea papulosa Schifn. ...................................................................................................315
Cheilolejeunea paroica Mizut. ......................................................................................................309
Cheilolejeunea parvidens B.M.hiers ............................................................................................314
Cheilolejeunea pirilora Schifn. ...................................................................................................309
Cheilolejeunea pluriplicata (Pearson) R.M.Schust. .......................................................................313
Cheilolejeunea pocsii E.W.Jones ...................................................................................................313
Cheilolejeunea polyantha A.Evans ................................................................................................313
Cheilolejeunea polyantha var. caduciloba R.M.Schust. ..................................................................313
Cheilolejeunea polystachya (Spruce) Gradst. et Ilk.-Borg. ..............................................................310
Cheilolejeunea renneri (G.Hofm.) Xiao L.He..............................................................................310
Cheilolejeunea revoluta (Herzog) Gradst. et Grolle ......................................................................313
536
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Cheilolejeunea rigidula (Nees ex Mont.) R.M.Schust. ..................................................................313
Cheilolejeunea riparia (Steph.) M.E.Reiner..................................................................................315
Cheilolejeunea rotalis (Hook.f. et Taylor) M.Wigginton ..............................................................315
Cheilolejeunea rotundistipula (Lindenb. ex Lehm.) Malombe.......................................................310
Cheilolejeunea rufescens (Lindenb.) Grolle ...................................................................................310
Cheilolejeunea ruwenzorensis (S.W.Arnell) R.M.Schust................................................................313
Cheilolejeunea ryukyuensis Mizut. ................................................................................................313
Cheilolejeunea sandvicensis (Prantl) Steph. ...................................................................................305
Cheilolejeunea stenoschiza (Ångstr.) A.Evans ................................................................................310
Cheilolejeunea streimannii Pócs et Ninh ......................................................................................315
Cheilolejeunea subcrenulata (Spruce) R.M.Schust. .......................................................................310
Cheilolejeunea subopaca (Mitt.) Mizut. ........................................................................................313
Cheilolejeunea suborbicularis (Herzog) H.A.Mill., Bonner et Bischl. ............................................310
Cheilolejeunea suprema (Grolle et Piippo) W.Ye et R.L.Zhu ........................................................310
Cheilolejeunea surrepens (Mitt.) E.W.Jones ..................................................................................310
Cheilolejeunea tenerrima (Steph.) C.J.Bastos................................................................................315
Cheilolejeunea trapezia (Nees) Kachroo et R.M.Schust. ...............................................................315
Cheilolejeunea trifaria (Reinw., Blume et Nees) Mizut. ...............................................................313
Cheilolejeunea turgida (Mitt.) W.Ye et R.L.Zhu ..........................................................................310
Cheilolejeunea udarii G.Asthana, S.C.Srivast. et A.K.Asthana......................................................313
Cheilolejeunea ulugurica Malombe, Eb.Fisch. et Pócs ..................................................................310
Cheilolejeunea unciloba (Lindenb.) Malombe ..............................................................................310
Cheilolejeunea upoluensis S.W.Arnell ...........................................................................................310
Cheilolejeunea urubuensis (Zartman et I.L.Ackerman) R.L.Zhu et Y.M.Wei ................................316
Cheilolejeunea usambarana (Steph.) Grolle ..................................................................................313
Cheilolejeunea valenciae (Steph.) Xiao L.He ................................................................................310
Cheilolejeunea ventricosa (Schifn. ex P.Syd.) Xiao L.He ..............................................................315
Cheilolejeunea verrucosa Steph. ....................................................................................................313
Cheilolejeunea virescens (Lehm. et Lindenb.) Steph. .....................................................................311
Cheilolejeunea viridis Steph. ........................................................................................................311
Cheilolejeunea vittata (Steph. ex G.Hofm.) R.M.Schust. et Kachroo ..........................................315
Cheilolejeunea warnstorii (Steph.) Solari ....................................................................................311
Cheilolejeunea wrightii Steph. ......................................................................................................311
Cheilolejeunea xanthocarpa (Lehm. et Lindenb.) Malombe ..........................................................311
Cheilolejeunea xanthophylla (Lindenb.) Steph. .............................................................................311
Chiastocaulon dendroides (Nees) Carl...........................................................................................219
Chiloscyphus acutus Steph. ...........................................................................................................190
Chiloscyphus alpicola J.J.Engel .....................................................................................................190
Chiloscyphus beesleyanus Pearson ..................................................................................................190
Chiloscyphus biidus Schifn. ........................................................................................................190
Chiloscyphus breviculus B.Y.Yang et W.C.Lee ..............................................................................190
Chiloscyphus brevistipulus Steph. ..................................................................................................190
Chiloscyphus chinnarensis Manju, K.P.Rajesh et Madhus. ............................................................190
Chiloscyphus confertifolius Steph...................................................................................................190
Chiloscyphus confertus Steph. .......................................................................................................190
Chiloscyphus cornutistipulus Steph. ...............................................................................................190
Chiloscyphus durus (Steph.) Hässel...............................................................................................190
Chiloscyphus ernstianus Steph. .....................................................................................................190
Chiloscyphus etesseanus Steph. ......................................................................................................190
Chiloscyphus francanus Steph. ......................................................................................................190
Chiloscyphus graefeanus Steph. ....................................................................................................190
Chiloscyphus greenwelliae (H.A.Mill.) H.A.Mill. ..........................................................................190
World checklist of hornworts and liverworts
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537
Chiloscyphus hookeri J.J.Engel ......................................................................................................190
Chiloscyphus hookeri var. constantifolius J.J.Engel .........................................................................190
Chiloscyphus integerrimus Schifn. ................................................................................................190
Chiloscyphus kashyapii A.Srivast. et S.C.Srivast. ...........................................................................189
Chiloscyphus kehdingianus (Steph.) N.Kitag. ................................................................................190
Chiloscyphus kilauensis Steph. ......................................................................................................190
Chiloscyphus koeppensis (Gottsche) Steph. ....................................................................................190
Chiloscyphus laceratus Steph.........................................................................................................191
Chiloscyphus lambertonii H.A.Mill...............................................................................................191
Chiloscyphus latistipus Steph. .......................................................................................................191
Chiloscyphus lepervanchei (Steph.) J.J.Engel et R.M.Schust. .........................................................191
Chiloscyphus longiissus Steph. ......................................................................................................191
Chiloscyphus pallescens (Ehrh.) Dumort. ......................................................................................189
Chiloscyphus pallescens var. fragilis (Roth) Müll.Frib. ...................................................................189
Chiloscyphus polyanthos (L.) Corda ..............................................................................................189
Chiloscyphus polyanthos var. rivularis (Schrad.) Lindb. et Arnell ...................................................189
Chiloscyphus propagulifer Schifn. ................................................................................................191
Chiloscyphus purpureus Steph.......................................................................................................191
Chiloscyphus quadricilius Steph. ...................................................................................................191
Chiloscyphus quadridentatus (Spruce) J.J.Engel et R.M.Schust. ....................................................191
Chiloscyphus rotundifolius Mitt. ...................................................................................................191
Chiloscyphus rotundiphyllus H.A.Mill...........................................................................................191
Chiloscyphus scaberulus Spruce .....................................................................................................191
Chiloscyphus septatus J.J.Engel .....................................................................................................191
Chiloscyphus similis Steph. ...........................................................................................................191
Chiloscyphus skottsbergianus H.A.Mill. .........................................................................................191
Chiloscyphus subacuminatus Herzog .............................................................................................191
Chiloscyphus subsimilis Steph. ......................................................................................................191
Chiloscyphus tridens Steph. ..........................................................................................................191
Chiloscyphus trigonifolius Steph....................................................................................................191
Chiloscyphus venustulus Colenso ..................................................................................................191
Chonecolea acutiloba (Schifn.) R.M.Schust. ................................................................................138
Chonecolea andina Grolle et Váňa ...............................................................................................138
Chonecolea doellingeri (Nees) Grolle ............................................................................................138
Chonecolea ruwenzorensis E.W.Jones ...........................................................................................138
Chonecolea schusteri Udar et Ad.Kumar .......................................................................................138
Chonecolea verae Potemkin..........................................................................................................138
Clandarium clandestinum (Mont.) R.M.Schust. ..........................................................................103
Clandarium gottscheanum (Grolle) R.M.Schust. ..........................................................................103
Clandarium xiphophyllum (Grolle) R.M.Schust. ..........................................................................103
Clasmatocolea bisexualis Glenny et J.J.Engel ................................................................................191
Clasmatocolea crassiretis (Herzog) Grolle......................................................................................191
Clasmatocolea ctenophylla (Schifn.) Grolle ..................................................................................191
Clasmatocolea cucullistipula (Steph.) Grolle .................................................................................192
Clasmatocolea fasciculata (Nees) Grolle ........................................................................................192
Clasmatocolea fulvella (Hook.f. et Taylor) Grolle.........................................................................192
Clasmatocolea gayana (Mont.) Grolle ..........................................................................................192
Clasmatocolea humilis (Hook.f. et Taylor) Grolle ........................................................................192
Clasmatocolea humilis var. polymorpha J.J.Engel...........................................................................192
Clasmatocolea humilis var. suspecta (C.Massal.) J.J.Engel .............................................................192
Clasmatocolea inlexispina (Hook.f. et Taylor) J.J.Engel...............................................................192
Clasmatocolea marginata (Steph.) Grolle .....................................................................................192
538
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Clasmatocolea minutiretis J.J.Engel et Grolle................................................................................192
Clasmatocolea moniliformis J.J.Engel............................................................................................192
Clasmatocolea navistipula (Steph.) Grolle ....................................................................................192
Clasmatocolea navistipula var. parceramosa J.J.Engel ....................................................................192
Clasmatocolea notophylla (Hook.f. et Taylor) Grolle ....................................................................192
Clasmatocolea obvoluta (Hook.f. et Taylor) Grolle .......................................................................192
Clasmatocolea obvoluta var. cookiana (C.Massal.) J.J.Engel ..........................................................192
Clasmatocolea puccioana (De Not.) Grolle ...................................................................................193
Clasmatocolea rigens (Hook.f. et Taylor) J.J.Engel .......................................................................193
Clasmatocolea strongylophylla (Hook.f. et Taylor) Grolle ..............................................................193
Clasmatocolea trachyopa (Hook.f. et Taylor) Grolle .....................................................................193
Clasmatocolea vermicularis (Lehm.) Grolle...................................................................................193
Clasmatocolea verrucosa J.J.Engel .................................................................................................193
Clevea hyalina (Sommerf.) Lindb. ...............................................................................................484
Clevea hyalina var. californica M.Howe .......................................................................................484
Clevea pedicellata (Grif.) Lindb. .................................................................................................484
Clevea pusilla (Steph.) Rubas. et D.G.Long.................................................................................484
Clevea spathysii (Lindenb.) Müll.Frib. .........................................................................................484
Cololejeunea abnormis Mizut. ......................................................................................................332
Cololejeunea acuminata Mizut. ....................................................................................................325
Cololejeunea adhaesiva (Mitt.) R.M.Schust. .................................................................................332
Cololejeunea adnata Tixier...........................................................................................................332
Cololejeunea aequabilis (Sande Lac.) Schifn. ...............................................................................326
Cololejeunea africana (Steph.) R.M.Schust. .................................................................................332
Cololejeunea albodentata P.C.Chen et P.C.Wu ............................................................................320
Cololejeunea altimontana Pócs .....................................................................................................326
Cololejeunea amaniensis Pócs .......................................................................................................326
Cololejeunea ambeliensis Tixier ....................................................................................................332
Cololejeunea amieuensis Tixier .....................................................................................................332
Cololejeunea amoena Benedix ......................................................................................................337
Cololejeunea amphibola B.M.hiers .............................................................................................324
Cololejeunea andapania Tixier .....................................................................................................332
Cololejeunea angulata (Steph.) Mizut. .........................................................................................332
Cololejeunea angustibracteata Schifn. ex P.Syd. ...........................................................................326
Cololejeunea angustilora (Steph.) Mizut. .....................................................................................323
Cololejeunea ankaiana Tixier .......................................................................................................332
Cololejeunea antillana Pócs..........................................................................................................324
Cololejeunea apiculata (E.W.Jones) R.M.Schust. .........................................................................326
Cololejeunea appressa (A.Evans) Benedix......................................................................................337
Cololejeunea armata Tixier ..........................................................................................................320
Cololejeunea arrectifolia (Mitt.) Steph..........................................................................................326
Cololejeunea attilana Pócs............................................................................................................332
Cololejeunea augieri Tixier...........................................................................................................324
Cololejeunea aurantia (Tixier) houvenot....................................................................................326
Cololejeunea auriculata (E.W.Jones) R.M.Schust.........................................................................332
Cololejeunea australis Tixier .........................................................................................................319
Cololejeunea autoica (Steph.) Grolle ............................................................................................332
Cololejeunea azorica V.Allorge et Jovet-Ast ..................................................................................324
Cololejeunea bachmaensis Tixier ...................................................................................................338
Cololejeunea bandamiae Tixier ....................................................................................................326
Cololejeunea bebourensis Tixier ....................................................................................................326
Cololejeunea bekkeri Tixier ..........................................................................................................332
World checklist of hornworts and liverworts
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539
Cololejeunea bergmansiana Tixier ................................................................................................326
Cololejeunea berneckerae Pócs ......................................................................................................317
Cololejeunea bhutanica Grolle et Mizut. ......................................................................................320
Cololejeunea biddlecomiae (Austin) A.Evans.................................................................................321
Cololejeunea bidentula (Steph.) E.W.Jones ..................................................................................326
Cololejeunea bifalcata Pócs ..........................................................................................................326
Cololejeunea bischleriana Tixier ...................................................................................................332
Cololejeunea blepharophylla Pócs..................................................................................................326
Cololejeunea bolovenensis Tixier ...................................................................................................332
Cololejeunea bontocensis Tixier .....................................................................................................338
Cololejeunea borbonica Tixier ......................................................................................................332
Cololejeunea borhidiana Pócs .......................................................................................................326
Cololejeunea bosseriana Tixier ......................................................................................................326
Cololejeunea brunelii Tixier .........................................................................................................333
Cololejeunea caihuaella But et P.C.Wu ........................................................................................321
Cololejeunea calcarata E.W.Jones ................................................................................................321
Cololejeunea calcarea (Lib.) Steph. ...............................................................................................326
Cololejeunea caledonica Gottsche .................................................................................................319
Cololejeunea camillii (Lehm.) A.Evans .........................................................................................324
Cololejeunea camusii Tixier..........................................................................................................326
Cololejeunea capuronii Tixier .......................................................................................................321
Cololejeunea cardiocarpa (Mont.) A.Evans ...................................................................................333
Cololejeunea ceatocarpa (Ångstr.) Steph. ......................................................................................326
Cololejeunea ceratilobula (P.C.Chen) R.M.Schust. ......................................................................319
Cololejeunea ceylanica Onr. .........................................................................................................326
Cololejeunea chamlongiana Tixier ................................................................................................327
Cololejeunea chenii Tixier ............................................................................................................327
Cololejeunea chinii Tixier ............................................................................................................320
Cololejeunea chittagongensis Tixier ...............................................................................................333
Cololejeunea chuahiana Pócs........................................................................................................337
Cololejeunea ciliata Pócs ..............................................................................................................327
Cololejeunea cingens (Herzog) Bernecker et Pócs..........................................................................317
Cololejeunea cocoscola Tixier ........................................................................................................333
Cololejeunea comptonii (Pearson) H.A.Mill..................................................................................327
Cololejeunea conchifolia (Gottsche) Gradst. .................................................................................340
Cololejeunea contractiloba A.Evans...............................................................................................324
Cololejeunea cookei A.Evans .........................................................................................................327
Cololejeunea cordilora Steph. ......................................................................................................327
Cololejeunea cornuta E.W.Jones...................................................................................................324
Cololejeunea cornutissima (R.M.Schust.) Stotler et Crand.-Stotl. .................................................317
Cololejeunea costaricensis (Bern.-Lück.) Bernecker et Pócs............................................................317
Cololejeunea crassipapillata Tixier ................................................................................................338
Cololejeunea crateris Pócs.............................................................................................................327
Cololejeunea cremersii Tixier ........................................................................................................333
Cololejeunea crenata (A.Evans) Pócs ............................................................................................324
Cololejeunea crenulata (Pearson) H.A.Mill. .................................................................................327
Cololejeunea cristata (Steph.) R.M.Schust. ...................................................................................333
Cololejeunea cubensis Pócs ...........................................................................................................317
Cololejeunea cucullifolia (Herzog) E.A.Hodgs. .............................................................................327
Cololejeunea cuneata (Lehm. et Lindenb.) Herzog .......................................................................333
Cololejeunea cuneifolia Steph. ......................................................................................................327
Cololejeunea dadeuniana Tixier ...................................................................................................327
540
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Cololejeunea dankiaensis Tixier ....................................................................................................340
Cololejeunea dauphinii R.L.Zhu ..................................................................................................337
Cololejeunea decemplicata (Steph.) Tixier.....................................................................................327
Cololejeunea decliviloba Steph......................................................................................................327
Cololejeunea dentata (E.W.Jones) R.M.Schust. ............................................................................327
Cololejeunea denticulata (Horik.) S.Hatt. ....................................................................................327
Cololejeunea dentilobula (Steph.) R.M.Schust..............................................................................327
Cololejeunea deroinii Tixier .........................................................................................................333
Cololejeunea desciscens Steph. .......................................................................................................319
Cololejeunea deslooveri Vanden Berghen ......................................................................................333
Cololejeunea dianae M.Wigginton ...............................................................................................327
Cololejeunea diaphana A.Evans....................................................................................................324
Cololejeunea dilatata (Steph.) Mizut. ...........................................................................................328
Cololejeunea dinghuiana R.L.Zhu et Y.F.Wang ...........................................................................321
Cololejeunea diplasiolejeunoides Tixier ..........................................................................................328
Cololejeunea discilora Tixier........................................................................................................337
Cololejeunea distalopapillata (E.W.Jones) R.M.Schust. ................................................................328
Cololejeunea dolichodonta Tixier ..................................................................................................321
Cololejeunea dozyana (Sande Lac.) Schifn. ..................................................................................321
Cololejeunea drepanolejeuneoides (Horik.) R.M.Schust. ................................................................324
Cololejeunea dzumacensis Tixier ...................................................................................................333
Cololejeunea ecuadoriensis Pócs ....................................................................................................333
Cololejeunea efusa (Mitt.) Steph. ................................................................................................328
Cololejeunea elegans Steph. ..........................................................................................................321
Cololejeunea elephantorum Tixier.................................................................................................328
Cololejeunea ellipsoidea R.M.Schust. ............................................................................................328
Cololejeunea ensifera Tixier ..........................................................................................................340
Cololejeunea epiphylla G.Asthana et A.Shukla ..............................................................................333
Cololejeunea equialbi Tixier .........................................................................................................328
Cololejeunea erostrata (Herzog) Bernecker et Pócs .......................................................................324
Cololejeunea eustacei Pócs ............................................................................................................338
Cololejeunea falcata (Horik.) Benedix ..........................................................................................338
Cololejeunea falcata var. madecassa Tixier ....................................................................................338
Cololejeunea falcidentata R.M.Schust. .........................................................................................321
Cololejeunea ilicis (Herzog) Piippo .............................................................................................328
Cololejeunea ilidens Benedix .......................................................................................................321
Cololejeunea ischeri Tixier ...........................................................................................................328
Cololejeunea issilobula Herzog ....................................................................................................333
Cololejeunea lavicans (Steph.) Mizut...........................................................................................328
Cololejeunea lavida P.C.Wu et J.S.Lou .......................................................................................338
Cololejeunea lavovittata Pócs ......................................................................................................338
Cololejeunea loccosa (Lehm. et Lindenb.) Schifn. .......................................................................338
Cololejeunea loccosa var. amoenoides Tixier ..................................................................................338
Cololejeunea loccosa var. angustibracteata Tixier ..........................................................................338
Cololejeunea loccosa var. aurita Benedix ......................................................................................338
Cololejeunea loccosa var. conivens Benedix ...................................................................................338
Cololejeunea loccosa var. ocellata Tixier .......................................................................................338
Cololejeunea loccosa var. plicata Tixier.........................................................................................338
Cololejeunea loccosa var. trivittata Tixier .....................................................................................338
Cololejeunea lorencei Tixier .........................................................................................................333
Cololejeunea foliicola S.C.Srivast. et G.Srivast. .............................................................................333
Cololejeunea frahmii Tixier ..........................................................................................................321
World checklist of hornworts and liverworts
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541
Cololejeunea fredericii Onr...........................................................................................................328
Cololejeunea fructumarginata Tixier.............................................................................................333
Cololejeunea furcilobulata (Berrie et E.W.Jones) R.M.Schust. ......................................................333
Cololejeunea fusca (Steph.) Mizut. ...............................................................................................328
Cololejeunea geissleriana Tixier ....................................................................................................333
Cololejeunea georgiana Tixier .......................................................................................................334
Cololejeunea gottschei (Steph.) Pandé, K.P.Srivast. et Ahmad .......................................................328
Cololejeunea gracilis (Jovet-Ast) Pócs ...........................................................................................318
Cololejeunea gracilis var. linearifolia (R.M.Schust.) Pócs ..............................................................318
Cololejeunea gradsteinii M.J.Lai et R.L.Zhu.................................................................................325
Cololejeunea gresicola Tixier .........................................................................................................338
Cololejeunea grolleana Pócs ..........................................................................................................321
Cololejeunea grossepapillosa (Horik.) N.Kitag. ..............................................................................318
Cololejeunea grossestyla M.Wigginton ..........................................................................................328
Cololejeunea grushvitzkiana Pócs .................................................................................................328
Cololejeunea guadelupensis Tixier .................................................................................................334
Cololejeunea gynophthalma Benedix .............................................................................................338
Cololejeunea hainanensis R.L.Zhu................................................................................................328
Cololejeunea hamata Steph. .........................................................................................................340
Cololejeunea harrisii Pócs ............................................................................................................328
Cololejeunea haskarliana (Lehm.) Schifn. ...................................................................................321
Cololejeunea hattoriana Mizut. et Pócs ........................................................................................324
Cololejeunea hebridensis Tixier .....................................................................................................334
Cololejeunea herzogii K.I.Goebel .................................................................................................340
Cololejeunea hildebrandii (Austin) Steph. ....................................................................................328
Cololejeunea hinidumae Onr........................................................................................................334
Cololejeunea hirta Steph. .............................................................................................................328
Cololejeunea hoabinhiana Tixier ..................................................................................................334
Cololejeunea hodgsoniae (Herzog) E.A.Hodgs. .............................................................................328
Cololejeunea hoeana Tixier ..........................................................................................................334
Cololejeunea horikawana (S.Hatt.) Mizut. ...................................................................................329
Cololejeunea huerlimannii Tixier .................................................................................................329
Cololejeunea hungii Tixier ...........................................................................................................334
Cololejeunea hyalina G.Asthana et S.C.Srivast. ............................................................................321
Cololejeunea indosinica Tixier ......................................................................................................334
Cololejeunea inlata Steph............................................................................................................338
Cololejeunea inlectens (Mitt.) Benedix .........................................................................................324
Cololejeunea inlexifolia R.M.Schust. ...........................................................................................329
Cololejeunea inoueana Mizut. ......................................................................................................334
Cololejeunea iradieri M.Infante et Heras .....................................................................................329
Cololejeunea irianensis Tixier .......................................................................................................329
Cololejeunea iwatsukiana (Pócs) Pócs ..........................................................................................318
Cololejeunea jamesii (Austin) M.E.Reiner et Pócs ........................................................................340
Cololejeunea japonica (Schifn.) Mizut. ........................................................................................334
Cololejeunea johannis-winkleri (Herzog) R.L.Zhu ........................................................................329
Cololejeunea jonesii Pócs ..............................................................................................................334
Cololejeunea jovetastiana (Pócs) Pócs ...........................................................................................318
Cololejeunea kahuziensis Tixier ....................................................................................................321
Cololejeunea kapingaensis H.A.Mill. ............................................................................................334
Cololejeunea karnatakensis G.Asthana et S.C.Srivast. ...................................................................321
Cololejeunea kegelii Steph. ...........................................................................................................329
Cololejeunea khanii Tixier ...........................................................................................................329
542
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Cololejeunea khiavensis Tixier ......................................................................................................338
Cololejeunea kiriromensis Tixier ...................................................................................................334
Cololejeunea kodamae Kamim. ....................................................................................................321
Cololejeunea kohkongensis Tixier ..................................................................................................329
Cololejeunea kolombangarae Pócs .................................................................................................321
Cololejeunea kolombangarae subsp. sepikensis Pócs .......................................................................321
Cololejeunea konratii Pócs ...........................................................................................................321
Cololejeunea koponenii (Pócs) Pócs ..............................................................................................325
Cololejeunea koratensis Tixier.......................................................................................................339
Cololejeunea kuciana Pócs et Schäf.-Verw. ...................................................................................321
Cololejeunea kulenensis Tixier ......................................................................................................334
Cololejeunea lacinulata Benedix ...................................................................................................320
Cololejeunea laevigata (Mitt.) R.M.Schust. ..................................................................................334
Cololejeunea lanceolata E.W.Jones ...............................................................................................325
Cololejeunea lanciloba Steph. .......................................................................................................334
Cololejeunea latilobula (Herzog) Tixier ........................................................................................334
Cololejeunea latistyla R.L.Zhu .....................................................................................................334
Cololejeunea leloutrei (E.W.Jones) R.M.Schust. ...........................................................................334
Cololejeunea leloutrei var. microlobulata Tixier .............................................................................334
Cololejeunea leloutrei var. ulugurica Pócs ex Tixier .......................................................................334
Cololejeunea lemuriana Tixier......................................................................................................334
Cololejeunea lichenyae R.D.Porley, N.G.Hodgetts et M.Wigginton.............................................329
Cololejeunea linopteroides H.Rob. ................................................................................................319
Cololejeunea lisowskii (Pócs) Pócs ................................................................................................318
Cololejeunea littoralis Tixier.........................................................................................................334
Cololejeunea lobulilineata Tixier ..................................................................................................329
Cololejeunea longiana Grolle et Mizut. ........................................................................................321
Cololejeunea longifolia (Mitt.) Benedix ex Mizut. ........................................................................329
Cololejeunea longistylis A.Evans....................................................................................................335
Cololejeunea macounii (Spruce) A.Evans ......................................................................................322
Cololejeunea madeirensis Schifn. .................................................................................................318
Cololejeunea madothecoides (Steph.) Benedix ...............................................................................320
Cololejeunea magillii Pócs ............................................................................................................322
Cololejeunea magna (Tixier) M.Infante et Heras ..........................................................................329
Cololejeunea magniica Pócs .........................................................................................................329
Cololejeunea magnilobula (Horik.) S.Hatt. ..................................................................................329
Cololejeunea magnistyla (Horik.) Mizut. ......................................................................................335
Cololejeunea malaccensis Tixier ....................................................................................................335
Cololejeunea malanjae Steph. .......................................................................................................322
Cololejeunea malayana Tixier ......................................................................................................335
Cololejeunea mamillata (Ångstr.) E.A.Hodgs. ..............................................................................322
Cololejeunea manlinensis Tixier....................................................................................................339
Cololejeunea maquilingensis Tixier ...............................................................................................339
Cololejeunea marginata (Lehm. et Lindenb.) Pearson ..................................................................335
Cololejeunea maritima Tixier .......................................................................................................335
Cololejeunea metzgeriopsis (K.I.Goebel) Gradst., R.Wilson, Ilk.-Borg. et Heinrichs .....................329
Cololejeunea micrandroecia (Spruce) M.Menzel ...........................................................................317
Cololejeunea micronesica H.A.Mill. et Bonner..............................................................................329
Cololejeunea microscopica (Taylor) Schifn. ..................................................................................318
Cololejeunea microscopica var. africana (Pócs) Pócs et Bernecker ..................................................318
Cololejeunea microscopica var. exigua (A.Evans) Pócs....................................................................318
Cololejeunea minuscula Pócs ........................................................................................................318
World checklist of hornworts and liverworts
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543
Cololejeunea minutilobula Herzog ...............................................................................................335
Cololejeunea mizutaniana Udar et G.Srivast. ...............................................................................322
Cololejeunea mocambiquensis S.W.Arnell .....................................................................................322
Cololejeunea mooreaensis Tixier....................................................................................................329
Cololejeunea moralesiae (Bern.-Lück.) Bernecker et Pócs..............................................................325
Cololejeunea moramangae Tixier ..................................................................................................318
Cololejeunea morobensis (Pócs) Pócs.............................................................................................325
Cololejeunea mouensis (Tixier) H.A.Mill. .....................................................................................324
Cololejeunea mutabilis Benedix ....................................................................................................339
Cololejeunea nakajimae S.Hatt. ...................................................................................................339
Cololejeunea nanhutashanensis J.D.Yang et S.H.Lin .....................................................................322
Cololejeunea nigerica (E.W.Jones) R.M.Schust. ...........................................................................335
Cololejeunea nilgiriensis G.Asthana et S.C.Srivast. .......................................................................322
Cololejeunea ninguana Tixier .......................................................................................................329
Cololejeunea norrisii (Pócs) Pócs ..................................................................................................318
Cololejeunea obcordata (Austin) A.Evans......................................................................................329
Cololejeunea obliqua (Nees et Mont.) Schifn. .............................................................................330
Cololejeunea oblongiperianthia (P.C.Wu et J.S.Lou) Piippo .........................................................330
Cololejeunea obtusifolia (E.W.Jones) Tixier..................................................................................325
Cololejeunea obtusifolia var. madecassa (Tixier) Pócs ....................................................................325
Cololejeunea occidentalis (E.W.Jones) Vanden Berghen ...............................................................335
Cololejeunea ocellata (Horik.) Benedix .........................................................................................339
Cololejeunea ocelloides (Horik.) Mizut. ........................................................................................339
Cololejeunea oleana Sim...............................................................................................................330
Cololejeunea ombrophila Tixier ....................................................................................................320
Cololejeunea onraedtii Tixier .......................................................................................................335
Cololejeunea ornata A.Evans ........................................................................................................322
Cololejeunea ovalifolia A.Evans ....................................................................................................330
Cololejeunea paciica Pócs ............................................................................................................335
Cololejeunea panamensis G.Dauphin et Pócs................................................................................335
Cololejeunea panchoana Tixier .....................................................................................................330
Cololejeunea paniensis (Tixier) Grolle ..........................................................................................330
Cololejeunea papilliloba (Steph.) Steph. .......................................................................................325
Cololejeunea papillosa (K.I.Goebel) Mizut. ..................................................................................318
Cololejeunea papulilora Steph. ....................................................................................................330
Cololejeunea papulosa R.M.Schust. ..............................................................................................325
Cololejeunea parva Vanden Berghen ............................................................................................322
Cololejeunea paucifolia (Spruce) Bernecker et Pócs ......................................................................325
Cololejeunea paucimarginata Tixier .............................................................................................335
Cololejeunea pentagona (Mitt.) Steph...........................................................................................330
Cololejeunea peponiformis Mizut. .................................................................................................325
Cololejeunea perainis (Schifn.) Schifn. .....................................................................................339
Cololejeunea perainis var. ciconiae Tixier ....................................................................................339
Cololejeunea perainis var. elegans Benedix...................................................................................339
Cololejeunea perainis var. serrulata Schifn. ex Benedix...............................................................339
Cololejeunea perakensis Tixier ......................................................................................................335
Cololejeunea plagiochiliana Tixier ................................................................................................330
Cololejeunea plagiophylla Benedix ................................................................................................335
Cololejeunea planilora Benedix ...................................................................................................322
Cololejeunea planissima (Mitt.) Abeyw.........................................................................................335
Cololejeunea planissima var. chagosensis Pócs ................................................................................335
Cololejeunea planiuscula Tixier ....................................................................................................330
544
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Cololejeunea platyneura (Spruce) A.Evans ....................................................................................322
Cololejeunea pluridentata P.C.Wu et J.S.Lou ...............................................................................322
Cololejeunea polisiana Tixier ........................................................................................................339
Cololejeunea praeruptorum Tixier ................................................................................................335
Cololejeunea pretiosa Benedix.......................................................................................................322
Cololejeunea producta (Mitt.) S.Hatt. ..........................................................................................335
Cololejeunea pseudocristallina P.C.Chen et P.C.Wu .....................................................................322
Cololejeunea pseudocuspidata Tixier .............................................................................................325
Cololejeunea pseudoloccosa (Horik.) Benedix ...............................................................................339
Cololejeunea pseudoplagiophylla P.C.Wu et J.S.Lou......................................................................322
Cololejeunea pseudoschmidtii Tixier..............................................................................................322
Cololejeunea pseudoserrata Tixier .................................................................................................330
Cololejeunea pseudostephanii Tixier ..............................................................................................339
Cololejeunea pseudostipulata Schifn. ex P.Syd..............................................................................320
Cololejeunea pterocolea Herzog ....................................................................................................325
Cololejeunea pteroporum Tixier ....................................................................................................330
Cololejeunea pulchella (Mitt.) R.M.Schust. ..................................................................................330
Cololejeunea pulchella var. stylifera R.M.Schust. ...........................................................................330
Cololejeunea punctata (Gottsche) Pearson ....................................................................................336
Cololejeunea pusilla Steph. ...........................................................................................................325
Cololejeunea quadridentata (S.Hatt.) Grolle.................................................................................330
Cololejeunea raduliloba Steph. .....................................................................................................336
Cololejeunea ramromensis Pócs.....................................................................................................322
Cololejeunea reineckeana Steph. ...................................................................................................336
Cololejeunea retusula (Mitt.) H.A.Mill.........................................................................................330
Cololejeunea rosellata Mizut. ........................................................................................................322
Cololejeunea rossettiana (C.Massal.) Schifn. ................................................................................322
Cololejeunea rotundilobula (P.C.Wu et P.J.Lin) Piippo ...............................................................319
Cololejeunea runssorensis (Steph.) Pócs .........................................................................................322
Cololejeunea salgadoi Onr. ...........................................................................................................330
Cololejeunea saltuum Tixier .........................................................................................................336
Cololejeunea sambiroana Tixier....................................................................................................336
Cololejeunea sanctae-helenae M.Wigginton ..................................................................................330
Cololejeunea saroltae Pócs ............................................................................................................336
Cololejeunea schaeferi Grolle ........................................................................................................323
Cololejeunea schmidtii Steph. .......................................................................................................323
Cololejeunea schmidtii var. acutepapillosa Pócs .............................................................................323
Cololejeunea schusteri Pócs ...........................................................................................................336
Cololejeunea schwabei Herzog ......................................................................................................336
Cololejeunea selaginellicola Tixier .................................................................................................323
Cololejeunea selangorensis Tixier...................................................................................................336
Cololejeunea serrata (Steph.) Benedix ..........................................................................................323
Cololejeunea serrulata Steph.........................................................................................................330
Cololejeunea setiloba A.Evans .......................................................................................................331
Cololejeunea setosa Mizut. ............................................................................................................339
Cololejeunea sharpii Mizut. ..........................................................................................................339
Cololejeunea shibiensis Mizut. ......................................................................................................336
Cololejeunea shikokiana (Horik.) S.Hatt. .....................................................................................323
Cololejeunea shimizui N.Kitag. ....................................................................................................320
Cololejeunea shimizui var. phangngana N.Kitag. ..........................................................................320
Cololejeunea siamensis Steph. .......................................................................................................339
Cololejeunea siangensis G.Asthana et S.C.Srivast. .........................................................................331
World checklist of hornworts and liverworts
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545
Cololejeunea sicifolia (Gottsche ex A.Evans) Pócs et Bernecker ....................................................319
Cololejeunea sicifolia subsp. jamaicensis (R.M.Schust.) Bernecker et Pócs.....................................318
Cololejeunea sigmoidea Jovet-Ast et Tixier....................................................................................319
Cololejeunea sigmoidea var. dubia Tixier ......................................................................................319
Cololejeunea sintenisii (Steph.) Pócs .............................................................................................319
Cololejeunea skottsbergii Herzog ...................................................................................................331
Cololejeunea smitinandii Tixier ....................................................................................................336
Cololejeunea societatis Tixier ........................................................................................................331
Cololejeunea sophiana Tixier ........................................................................................................319
Cololejeunea spathulata Jovet-Ast .................................................................................................325
Cololejeunea spathulifolia (Steph.) H.A.Mill. ...............................................................................331
Cololejeunea sphaerodonta Mizut. ................................................................................................339
Cololejeunea spinosa (Horik.) Pandé et R.N.Misra .......................................................................323
Cololejeunea spruceana Tixier ......................................................................................................331
Cololejeunea standleyi Herzog ......................................................................................................319
Cololejeunea stellaris Pócs ............................................................................................................323
Cololejeunea stenophylla Herzog ...................................................................................................331
Cololejeunea stephanii Schifn. ex Benedix ...................................................................................339
Cololejeunea stoniana Tixier ........................................................................................................336
Cololejeunea stotleriana Gradst., Ilk.-Borg. et Vanderp. ...............................................................320
Cololejeunea streimannii Pócs ......................................................................................................331
Cololejeunea streimannii subsp. solomonensis Pócs ........................................................................331
Cololejeunea stylilobula Tixier ......................................................................................................320
Cololejeunea stylosa Steph. ...........................................................................................................336
Cololejeunea subalpina Pócs .........................................................................................................331
Cololejeunea subcardiocarpa Tixier ...............................................................................................336
Cololejeunea subcristata A.Evans ..................................................................................................331
Cololejeunea subloccosa Mizut. ....................................................................................................339
Cololejeunea subinlata Tixier ......................................................................................................336
Cololejeunea subkodamae Mizut. ..................................................................................................323
Cololejeunea sublatistyla Jian Wang bis et R.L.Zhu ......................................................................340
Cololejeunea submarginata Tixier.................................................................................................336
Cololejeunea subminutilobula Mizut. ...........................................................................................336
Cololejeunea subocelloides Mizut. .................................................................................................339
Cololejeunea subscariosa (Spruce) Pócs .........................................................................................336
Cololejeunea subsphaeroidea (R.M.Schust.) Pócs ..........................................................................319
Cololejeunea subtriapiculata Tixier ...............................................................................................336
Cololejeunea succinea Tixier .........................................................................................................336
Cololejeunea surinamensis Tixier ..................................................................................................336
Cololejeunea tahitensis Tixier .......................................................................................................336
Cololejeunea takamakae Tixier .....................................................................................................331
Cololejeunea tamasii Schäf.-Verw. ...............................................................................................331
Cololejeunea tamatavensis Tixier ..................................................................................................336
Cololejeunea tamdaoensis Tixier ...................................................................................................340
Cololejeunea tanneri Pócs ............................................................................................................323
Cololejeunea tanzaniae Pócs.........................................................................................................331
Cololejeunea taprobanea Tixier ....................................................................................................337
Cololejeunea taurifolia Inoue et H.A.Mill. ...................................................................................325
Cololejeunea tenella Benedix ........................................................................................................323
Cololejeunea tenella var. dentiloba Onr.........................................................................................323
Cololejeunea tenuiparietata Tixier ................................................................................................331
Cololejeunea teurnoumensis Tixier ................................................................................................331
546
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Cololejeunea thailandensis Tixier ..................................................................................................337
Cololejeunea thiersiae (Pócs) Pócs ................................................................................................319
Cololejeunea thiersiana Tixier ......................................................................................................323
Cololejeunea timoi Pócs ...............................................................................................................331
Cololejeunea tixieri Onr. ..............................................................................................................340
Cololejeunea touwii Pócs..............................................................................................................331
Cololejeunea tranninhiana Tixier .................................................................................................331
Cololejeunea triapiculata (Herzog) Tixier .....................................................................................337
Cololejeunea tribracteata Tixier ....................................................................................................331
Cololejeunea trichomanis (Gottsche) Besch. .................................................................................331
Cololejeunea tridentata Tixier ......................................................................................................337
Cololejeunea tuiwawana Pócs ......................................................................................................323
Cololejeunea tuksapiana Tixier .....................................................................................................331
Cololejeunea uchimae Amakawa ...................................................................................................337
Cololejeunea variifolia (Mitt.) Steph. ...........................................................................................340
Cololejeunea veillonii Tixier .........................................................................................................319
Cololejeunea verdoornii (S.Hatt.) S.Hatt. .....................................................................................340
Cololejeunea verrucosa Steph. .......................................................................................................323
Cololejeunea verrucosa var. rectispina (Herzog) Benedix ...............................................................323
Cololejeunea verwimpii Tixier ......................................................................................................337
Cololejeunea vesicaria (Sande Lac.) Schifn. .................................................................................324
Cololejeunea vidaliana Tixier .......................................................................................................337
Cololejeunea vietnamensis Tixier ..................................................................................................337
Cololejeunea virotana Tixier ........................................................................................................319
Cololejeunea vitaliana Tixier........................................................................................................337
Cololejeunea vulcania Tixier ........................................................................................................331
Cololejeunea wightii Steph. ..........................................................................................................325
Cololejeunea winkleri (M.I.Morales et A.Lücking) Pócs ...............................................................319
Cololejeunea yakusimensis (S.Hatt.) Mizut. ..................................................................................337
Cololejeunea yelitzae Pócs et Bernecker ........................................................................................325
Cololejeunea yipii R.L.Zhu ..........................................................................................................340
Cololejeunea yoshinagana (S.Hatt.) Mizut. ...................................................................................331
Cololejeunea zangii R.L.Zhu et M.L.So .......................................................................................320
Cololejeunea zantenorum Pócs .....................................................................................................340
Cololejeunea zenkeri (Steph.) E.W.Jones ......................................................................................323
Colura acroloba (Prantl) Jovet-Ast ...............................................................................................344
Colura amboinensis Steph. ...........................................................................................................344
Colura andoi Gradst. et Jovet-Ast ................................................................................................341
Colura ari (Steph.) Steph. ...........................................................................................................342
Colura australiensis Jovet-Ast .......................................................................................................342
Colura berghenii Jovet-Ast ...........................................................................................................340
Colura bicornis Jovet-Ast .............................................................................................................343
Colura bisvoluta Herzog et Jovet-Ast ...........................................................................................343
Colura brevistyla Herzog .............................................................................................................342
Colura calderae Pócs ....................................................................................................................342
Colura calyptrifolia (Hook.) Dumort. ..........................................................................................341
Colura clementis Grolle................................................................................................................342
Colura conica (Sande Lac.) K.I.Goebel ........................................................................................342
Colura corniantha Grolle .............................................................................................................344
Colura corynophora (Nees, Lindenb. et Gottsche) Trevis. ............................................................342
Colura crenulata Grolle ...............................................................................................................342
Colura crispiloba Jovet-Ast ..........................................................................................................342
World checklist of hornworts and liverworts
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547
Colura cristata Jovet-Ast ..............................................................................................................341
Colura cylindrica Herzog .............................................................................................................342
Colura cymbalifera Herzog et Jovet-Ast .......................................................................................342
Colura denticulata Jovet-Ast ........................................................................................................344
Colura digitalis (Mitt.) Steph. .....................................................................................................342
Colura digitalis var. mucronata Pócs ............................................................................................342
Colura dusenii Steph. ..................................................................................................................342
Colura fastigiata Jovet-Ast ...........................................................................................................342
Colura istulosa Jovet-Ast .............................................................................................................342
Colura galeata Jovet-Ast ..............................................................................................................344
Colura greig-smithii Jovet-Ast ......................................................................................................341
Colura hattoriana Pócs ................................................................................................................342
Colura hedbergiana Pócs..............................................................................................................341
Colura heimii Jovet-Ast ...............................................................................................................342
Colura hemisphaerica Jovet-Ast ....................................................................................................342
Colura herzogii Jovet-Ast .............................................................................................................342
Colura hirta Steph.......................................................................................................................344
Colura humbertii Jovet-Ast ..........................................................................................................341
Colura imperfecta Steph...............................................................................................................344
Colura inlata K.I.Goebel ............................................................................................................341
Colura inornata Jovet-Ast ............................................................................................................344
Colura inuii Horik. .....................................................................................................................342
Colura irrorata (Spruce) Heinrichs, Y.Yu, Schäf.-Verw. et Pócs ..................................................341
Colura itatyana Steph..................................................................................................................343
Colura jovet-astiae Grolle ............................................................................................................341
Colura junghuhniana (Prantl) Steph. ...........................................................................................341
Colura karstenii K.I.Goebel .........................................................................................................344
Colura koponenii Pócs .................................................................................................................342
Colura mauritiana Pócs ...............................................................................................................342
Colura maxima Jovet-Ast ............................................................................................................342
Colura medusa J.Eggers et Pócs ...................................................................................................341
Colura meijeri Jovet-Ast ..............................................................................................................341
Colura mizutanii Pócs .................................................................................................................341
Colura mosenii Steph...................................................................................................................342
Colura naumannii (Schifn.) Steph..............................................................................................343
Colura norrisii Pócs .....................................................................................................................342
Colura obesa Jovet-Ast .................................................................................................................343
Colura obvoluta Jovet-Ast ............................................................................................................341
Colura ornata K.I.Goebel ............................................................................................................341
Colura ornithocephala Herzog......................................................................................................343
Colura palawanensis Jovet-Ast .....................................................................................................341
Colura pallida Steph. ..................................................................................................................343
Colura pluridentata Jovet-Ast ......................................................................................................343
Colura pulcherrima Jovet-Ast.......................................................................................................343
Colura pulcherrima var. bartlettii Jovet-Ast ..................................................................................343
Colura queenslandica B.M.hiers.................................................................................................344
Colura rhynchophora Jovet-Ast ....................................................................................................341
Colura saccophylla E.A.Hodgs. et Herzog ....................................................................................343
Colura saroltae Pócs.....................................................................................................................344
Colura schusteri Grolle.................................................................................................................343
Colura siamensis Jovet-Ast ...........................................................................................................344
Colura speciosa Jovet-Ast .............................................................................................................343
548
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Colura streimannii Pócs ...............................................................................................................343
Colura strophiolata Jovet-Ast .......................................................................................................344
Colura superba (Mont.) Steph. ....................................................................................................343
Colura tenuicornis (A.Evans) Steph..............................................................................................341
Colura thomeensis Pócs ................................................................................................................343
Colura tixieri Jovet-Ast................................................................................................................344
Colura tortifolia (Nees et Mont.) Trevis. .....................................................................................343
Colura tutuilana (Pearson) H.A.Mill. ..........................................................................................343
Colura ulei Jovet-Ast ...................................................................................................................343
Colura usambarica E.W.Jones .....................................................................................................344
Colura valida Jovet-Ast ...............................................................................................................341
Colura verdoornii Herzog et Jovet-Ast .........................................................................................341
Colura vietnamensis Jovet-Ast et Tixier ........................................................................................344
Colura vitiensis Pócs et J.Eggers ...................................................................................................343
Conocephalum conicum (L.) Dumort. ..........................................................................................485
Conocephalum japonicum (hunb.) Grolle ...................................................................................486
Conocephalum salebrosum Szweyk., Buczk. et Odrzyk..................................................................485
Conoscyphus koponenii Piippo, Mamontov et Potemkin...............................................................193
Conoscyphus trapezioides (Sande Lac.) Schifn. .............................................................................193
Corsinia coriandrina (Spreng.) Lindb. .........................................................................................486
Cronisia imbriata (Nees) Whittem. et Bischl. .............................................................................486
Cronisia weddellii (Mont.) Grolle ................................................................................................486
Crossocalyx hellerianus (Nees ex Lindenb.) Meyl. ...........................................................................52
Crossocalyx tenuis (Harry Williams) Schljakov ...............................................................................52
Crossotolejeunea curvifolia Steph. .................................................................................................508
Cryptocolea imbricata R.M.Schust. ..............................................................................................123
Cryptocoleopsis imbricata Amakawa..............................................................................................110
Cryptolophocolea aculeata (Mitt.) L.Söderstr. ...............................................................................193
Cryptolophocolea chiloscyphoidea (Lindenb.) L.Söderstr. et Crand.-Stotl. ......................................193
Cryptolophocolea ciliolata (Nees) L.Söderstr., Crand.-Stotl., Stotler et Váňa ................................193
Cryptolophocolea compacta (Mitt.) L.Söderstr...............................................................................193
Cryptolophocolea connata (Sw.) L.Söderstr. et Váňa .....................................................................193
Cryptolophocolea connatifolia (J.J.Engel) L.Söderstr. ....................................................................194
Cryptolophocolea costata (Nees) L.Söderstr. ..................................................................................194
Cryptolophocolea edentata (J.J.Engel) L.Söderstr. .........................................................................194
Cryptolophocolea explanata (Mitt.) Váňa et Crand.-Stotl..............................................................194
Cryptolophocolea leischeri (Steph.) L.Söderstr. .............................................................................194
Cryptolophocolea guadalupensis (Steph.) L.Söderstr. et Váňa ........................................................194
Cryptolophocolea helmsiana (Steph.) L.Söderstr. ...........................................................................194
Cryptolophocolea leucophylla (Hook.f. et Taylor) L.Söderstr. ........................................................194
Cryptolophocolea levieri (Schifn.) L.Söderstr. ..............................................................................194
Cryptolophocolea lilliena (Steph.) L.Söderstr. ...............................................................................194
Cryptolophocolea martiana (Nees) L.Söderstr., Crand.-Stotl. et Stotler.........................................194
Cryptolophocolea martiana subsp. bidentula (Nees) L.Söderstr., Crand.-Stotl. et Stotler ...............194
Cryptolophocolea martiana var. perissodonta (Spruce) Gradst. .......................................................194
Cryptolophocolea massalongoana (Schifn.) L.Söderstr...................................................................195
Cryptolophocolea mitteniana (Colenso) L.Söderstr........................................................................195
Cryptolophocolea mitteniana var. obtusa (J.J.Engel) L.Söderstr. ....................................................195
Cryptolophocolea mitteniana var. symmetrica (J.J.Engel) L.Söderstr. .............................................195
Cryptolophocolea pallida (Mitt.) L.Söderstr. .................................................................................195
Cryptolophocolea pallidovirens (Hook.f. et Taylor) L.Söderstr. .....................................................195
Cryptolophocolea proteus (Herzog) L.Söderstr. ..............................................................................195
World checklist of hornworts and liverworts
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549
Cryptolophocolea pycnophylla (Spruce) L.Söderstr. ........................................................................195
Cryptolophocolea regularis (Steph.) L.Söderstr. .............................................................................195
Cryptolophocolea spinifera (Hook.f. et Taylor) L.Söderstr.............................................................195
Cryptolophocolea stephanii (Schifn.) L.Söderstr. ..........................................................................195
Cryptolophocolea subopposita (J.J.Engel) L.Söderstr. .....................................................................195
Cryptolophocolea thermarum (Schifn.) L.Söderstr. .......................................................................195
Cryptolophocolea trialata (Gottsche) L.Söderstr............................................................................196
Cryptolophocolea tricorata (Hässel) Crand.-Stotl. et Stotler ..........................................................196
Cryptolophocolea tuberculata (J.J.Engel) L.Söderstr. .....................................................................196
Cryptolophocolea whittieriana (Inoue et H.A.Mill.) L.Söderstr. ....................................................196
Cryptomitrium himalayense Kashyap ............................................................................................481
Cryptomitrium oreades Perold ......................................................................................................481
Cryptomitrium tenerum (Hook.) Austin ex Underw.....................................................................481
Cuspidatula contracta (Reinw., Blume et Nees) Steph. ..................................................................45
Cuspidatula laccida (Steph.) K.Feldberg, Váňa, Hentschel et Heinrichs .......................................45
Cuspidatula lexicaulis (Nees) Váňa et L.Söderstr...........................................................................45
Cuspidatula kirkii (Steph.) K.Feldberg, Váňa, Hentschel et Heinrichs ..........................................46
Cuspidatula monodon (Taylor) Steph.............................................................................................46
Cuspidatula orbicularis (Grolle) Váňa et L.Söderstr. ......................................................................46
Cuspidatula robusta (Austin) Váňa et L.Söderstr............................................................................46
Cyathodium aureonitens (Grif.) Schifn.......................................................................................486
Cyathodium bischlerianum N.Salazar ...........................................................................................486
Cyathodium cavernarum Kunze ...................................................................................................486
Cyathodium denticulatum Udar et S.C.Srivast. ............................................................................486
Cyathodium foetidissimum Schifn. ..............................................................................................487
Cyathodium indicum Udar et D.K.Singh .....................................................................................487
Cyathodium mehranum D.K.Singh..............................................................................................487
Cyathodium smaragdinum Schifn. ..............................................................................................487
Cyathodium spruceanum Prosk. ...................................................................................................487
Cyathodium spurium (Dicks.) Lindb. ex Braithw. ........................................................................487
Cyathodium steerei Hässel ............................................................................................................487
Cyathodium tuberculatum Udar et D.K.Singh .............................................................................487
Cyathodium tuberosum Kashyap ..................................................................................................487
Cyclolejeunea accedens (Gottsche) A.Evans ...................................................................................351
Cyclolejeunea chitonia (Taylor) A.Evans.......................................................................................351
Cyclolejeunea convexistipa (Lehm. et Lindenb.) A.Evans ..............................................................351
Cyclolejeunea ecuadorensis Steph. .................................................................................................351
Cyclolejeunea foliorum (Nees) Grolle ...........................................................................................351
Cyclolejeunea integerrima (Steph.) Steph. .....................................................................................351
Cyclolejeunea luteola (Spruce) Grolle ...........................................................................................351
Cyclolejeunea peruviana (Lehm. et Lindenb.) A.Evans .................................................................351
Cyclolejeunea spectabilis Steph......................................................................................................351
Cylindrocolea abyssinica (Gola) Váňa .............................................................................................73
Cylindrocolea andersonii R.M.Schust. ............................................................................................72
Cylindrocolea brasiliensis D.P.Costa, N.D.Santos et Váňa..............................................................72
Cylindrocolea chevalieri (Steph.) R.M.Schust. ................................................................................72
Cylindrocolea gittinsii (E.W.Jones) R.M.Schust. ............................................................................73
Cylindrocolea kiaeri (Austin) Váňa.................................................................................................73
Cylindrocolea madagascariensis (Steph.) R.M.Schust. .....................................................................73
Cylindrocolea nigerica (E.W.Jones) R.M.Schust. ............................................................................73
Cylindrocolea novae-caledoniae (Grolle) R.M.Schust. .....................................................................73
Cylindrocolea obtusifolia Fulford ....................................................................................................73
550
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Cylindrocolea planifolia (Steph.) R.M.Schust. ................................................................................73
Cylindrocolea recurvifolia (Steph.) Inoue........................................................................................73
Cylindrocolea reticulata Udar et Ad.Kumar ....................................................................................73
Cylindrocolea rhizantha (Mont.) R.M.Schust. ...............................................................................73
Cylindrocolea sanctae-helenae M.Wigginton ...................................................................................73
Cylindrocolea sprucei R.M.Schust. .................................................................................................73
Cylindrocolea tagawae (N.Kitag.) R.M.Schust. ..............................................................................73
Cylindrocolea ugandica (E.W.Jones) R.M.Schust. ..........................................................................73
Cyrtolejeunea holostipa (Spruce) A.Evans .....................................................................................316
Cystolejeunea lineata (Lehm. et Lindenb.) A.Evans ......................................................................316
Dactylophorella muricata (Gottsche) R.M.Schust.........................................................................300
Deceptifrons plagiochiloides J.J.Engel et Váňa ...............................................................................196
Delavayella serrata Steph. ............................................................................................................117
Delavayella serrata var. purpurea P.C.Chen .................................................................................117
Dendroceros acutilobus Steph. ........................................................................................................32
Dendroceros adglutinatus (Hook.f. et Taylor) Gottsche, Lindenb. et Nees .....................................33
Dendroceros africanus Steph. ..........................................................................................................33
Dendroceros allionii Steph. ............................................................................................................33
Dendroceros australis Steph. ...........................................................................................................33
Dendroceros borbonicus Steph. .......................................................................................................32
Dendroceros breutelii Nees .............................................................................................................33
Dendroceros breutelii var. surinamensis Lindenb. et Gottsche .........................................................33
Dendroceros cavernosus J.Haseg. .....................................................................................................32
Dendroceros cichoraceus (Mont.) Gottsche .....................................................................................33
Dendroceros crassicostatus Steph. ....................................................................................................33
Dendroceros crassinervis (Nees) Gottsche........................................................................................33
Dendroceros crispatus (Hook.) Nees ...............................................................................................33
Dendroceros crispatus var. simplicior Spruce ....................................................................................33
Dendroceros crispus (Sw.) Nees.......................................................................................................32
Dendroceros cucullatus Steph..........................................................................................................33
Dendroceros diicilis Steph.............................................................................................................32
Dendroceros exalatus Steph. ...........................................................................................................33
Dendroceros foliicola J.Haseg. ........................................................................................................32
Dendroceros gracilis Steph. .............................................................................................................33
Dendroceros granulatus Mitt. .........................................................................................................33
Dendroceros herasii M.Infante........................................................................................................33
Dendroceros humboldtensis Hürl. ...................................................................................................33
Dendroceros japonicus Steph. .........................................................................................................32
Dendroceros javanicus (Nees) Nees ................................................................................................32
Dendroceros muelleri Steph. ...........................................................................................................32
Dendroceros ogeramnangus Piippo ..................................................................................................32
Dendroceros paivae C.A.Garcia, Sérgio et J.C.Villarreal .................................................................33
Dendroceros pedunculatus Steph. ....................................................................................................32
Dendroceros rarus Steph.................................................................................................................34
Dendroceros reticulus Herzog .........................................................................................................34
Dendroceros rigidus Steph. .............................................................................................................34
Dendroceros seramensis J.Haseg. .....................................................................................................34
Dendroceros subdiicilis S.Hatt. .....................................................................................................32
Dendroceros subplanus Steph..........................................................................................................32
Dendroceros subtropicus C.J.Wild...................................................................................................34
Dendroceros tahitensis Ångstr. ........................................................................................................34
Dendroceros tubercularis S.Hatt. ....................................................................................................32
World checklist of hornworts and liverworts
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551
Dendroceros validus Steph..............................................................................................................33
Dendroceros vesconianus Gottsche ..................................................................................................33
Dendroceros wattsianus Steph. .......................................................................................................33
Dendrolembidium dendroides (Carrington et Pearson) Herzog .....................................................163
Dendrolembidium tenax (Grev.) Herzog ......................................................................................164
Dendromastigophora lagellifera (Hook.) R.M.Schust. ..................................................................217
Denotarisia linguifolia (De Not.) Grolle ........................................................................................46
Dicranolejeunea axillaris (Nees et Mont.) Schifn. .......................................................................299
Dicranolejeunea bovonei Gola ......................................................................................................299
Dinckleria fruticella (Hook.f. et Taylor) J.J.Engel et Heinrichs....................................................219
Dinckleria pleurata (Hook.f. et Taylor) Trevis.............................................................................219
Diplasiolejeunea alata Jovet-Ast ...................................................................................................344
Diplasiolejeunea albifolia (Taylor) E.W.Jones ..............................................................................347
Diplasiolejeunea andringitrae Schäf.-Verw. ..................................................................................346
Diplasiolejeunea armatiloba Steph. ..............................................................................................345
Diplasiolejeunea aulae E.W.Jones ................................................................................................347
Diplasiolejeunea auriculata Tixier ................................................................................................347
Diplasiolejeunea borhidiana Reyes Montoya ................................................................................346
Diplasiolejeunea brunnea Steph. ..................................................................................................346
Diplasiolejeunea buckii Grolle......................................................................................................347
Diplasiolejeunea caribea Tixier .....................................................................................................346
Diplasiolejeunea cavifolia Steph. ..................................................................................................346
Diplasiolejeunea cobrensis Steph. ..................................................................................................346
Diplasiolejeunea cobrensis subsp. antsirananae Pócs ......................................................................346
Diplasiolejeunea cogoensis M.Infante, Heras et Pócs .....................................................................346
Diplasiolejeunea columbica Tixier ................................................................................................347
Diplasiolejeunea comorensis Tixier ................................................................................................347
Diplasiolejeunea cornuta Steph.....................................................................................................346
Diplasiolejeunea cubensis Tixier....................................................................................................346
Diplasiolejeunea cyanguguensis Tixier ...........................................................................................347
Diplasiolejeunea deslooveri Vanden Berghen.................................................................................347
Diplasiolejeunea eggersii Pócs .......................................................................................................345
Diplasiolejeunea ensifera Tixier ....................................................................................................345
Diplasiolejeunea erostrata Schäf.-Verw. ........................................................................................347
Diplasiolejeunea evansii Tixier .....................................................................................................347
Diplasiolejeunea glaziovii Tixier ...................................................................................................346
Diplasiolejeunea gradsteinii Tixier ................................................................................................347
Diplasiolejeunea grandirostrata Schäf.-Verw. ................................................................................347
Diplasiolejeunea grolleana Reyes Montoya ...................................................................................347
Diplasiolejeunea guadalupensis Steph. ..........................................................................................345
Diplasiolejeunea hamata Tixier ....................................................................................................347
Diplasiolejeunea heimii Jovet-Ast .................................................................................................345
Diplasiolejeunea ingekarolae Schäf.-Verw. ....................................................................................346
Diplasiolejeunea insignis Tixier ....................................................................................................347
Diplasiolejeunea integerrima Tixier ..............................................................................................347
Diplasiolejeunea involuta S.Winkl. ..............................................................................................345
Diplasiolejeunea involuta subsp. andicola Pócs .............................................................................345
Diplasiolejeunea johnsonii A.Evans...............................................................................................345
Diplasiolejeunea johnsonii var. mexicana Jovet-Ast .......................................................................345
Diplasiolejeunea jonesii Tixier ......................................................................................................346
Diplasiolejeunea jovet-astiae Grolle ..............................................................................................346
Diplasiolejeunea kraussiana (Lindenb.) Steph. .............................................................................346
552
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Diplasiolejeunea lanceolata Grolle ................................................................................................347
Diplasiolejeunea latipuensis Tixier ................................................................................................348
Diplasiolejeunea leiocarpa Jovet-Ast .............................................................................................346
Diplasiolejeunea lemuriana Tixier ................................................................................................348
Diplasiolejeunea longilobula Herzog .............................................................................................348
Diplasiolejeunea magnistipula Tixier ............................................................................................348
Diplasiolejeunea malleiformis (A.Evans) Tixier .............................................................................346
Diplasiolejeunea mayaykuensis Schäf.-Verw. et Heinrichs .............................................................348
Diplasiolejeunea montecristensis S.Winkl. .....................................................................................345
Diplasiolejeunea onraedtii Grolle .................................................................................................348
Diplasiolejeunea ornata Pócs et Schäf.-Verw. ...............................................................................346
Diplasiolejeunea palustrium Tixier ...............................................................................................346
Diplasiolejeunea papilionacea R.M.Schust. ...................................................................................345
Diplasiolejeunea patelligera Herzog ..............................................................................................347
Diplasiolejeunea pauckertii (Nees) Steph. .....................................................................................345
Diplasiolejeunea pellucida (C.F.W.Meissn. ex Spreng.) Schifn. ...................................................346
Diplasiolejeunea phyllarthronii Tixier ...........................................................................................346
Diplasiolejeunea plicatiloba (Hook.f. et Taylor) Grolle ................................................................347
Diplasiolejeunea pluridentata Schäf.-Verw. ..................................................................................345
Diplasiolejeunea pocsii Reyes Montoya.........................................................................................346
Diplasiolejeunea pusilla Grolle .....................................................................................................347
Diplasiolejeunea ramicola Tixier ..................................................................................................348
Diplasiolejeunea ranomafanae Pócs ..............................................................................................347
Diplasiolejeunea replicata (Spruce) Steph. ....................................................................................345
Diplasiolejeunea riclefgrollei Schäf.-Verw. ....................................................................................348
Diplasiolejeunea rudolphiana Steph..............................................................................................345
Diplasiolejeunea runssorensis Steph. ..............................................................................................345
Diplasiolejeunea subcornuta Tixier ...............................................................................................348
Diplasiolejeunea symoensii Vanden Berghen .................................................................................348
Diplasiolejeunea unidentata (Lehm. et Lindenb.) Schifn. ............................................................345
Diplasiolejeunea utriculata Steph. ................................................................................................344
Diplasiolejeunea villaumei Steph. .................................................................................................347
Diplasiolejeunea zakiae Tixier ......................................................................................................347
Diplocolea sikkimensis Amakawa ..................................................................................................123
Diplophyllum africanum S.W.Arnell ..............................................................................................83
Diplophyllum albicans (L.) Dumort. ..............................................................................................83
Diplophyllum andicola R.M.Schust................................................................................................83
Diplophyllum andrewsii A.Evans....................................................................................................83
Diplophyllum androgynum J.J.Engel et G.L.Merr. .........................................................................83
Diplophyllum angustifolium J.J.Engel et G.L.Merr. ........................................................................83
Diplophyllum apiculatum (A.Evans) Steph. ....................................................................................83
Diplophyllum dioicum R.M.Schust. ...............................................................................................83
Diplophyllum exiguum Steph. ........................................................................................................83
Diplophyllum gemmiparum J.J.Engel et G.L.Merr. ........................................................................83
Diplophyllum incrassatum J.J.Engel et G.L.Merr............................................................................84
Diplophyllum nanum Herzog ........................................................................................................84
Diplophyllum novum J.J.Engel et G.L.Merr. ..................................................................................84
Diplophyllum obtusatum (R.M.Schust.) R.M.Schust......................................................................84
Diplophyllum obtusifolium (Hook.) Dumort..................................................................................84
Diplophyllum obtusifolium subsp. domesticum (Gottsche) Váňa......................................................84
Diplophyllum recurvifolium C.Massal. ...........................................................................................83
Diplophyllum serrulatum (Müll.Frib.) Steph. .................................................................................84
World checklist of hornworts and liverworts
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553
Diplophyllum squarrosum Steph. ....................................................................................................83
Diplophyllum taxifolium (Wahlenb.) Dumort................................................................................84
Diplophyllum taxifolium var. mucronatum R.M.Schust. .................................................................84
Diplophyllum trollii Grolle ............................................................................................................84
Diplophyllum verrucosum R.M.Schust. ..........................................................................................83
Douinia imbricata (M.Howe) Konstant. et Vilnet .........................................................................84
Douinia ovata (Dicks.) H.Buch.....................................................................................................84
Douinia plicata (Lindb.) Konstant. et Vilnet .................................................................................84
Drepanolejeunea actinogyna Inuthai, R.L.Zhu et Chantanaorr. ....................................................359
Drepanolejeunea aculeata Bischl...................................................................................................353
Drepanolejeunea anderssonii (Ångstr.) A.Evans ............................................................................353
Drepanolejeunea andina Herzog ..................................................................................................353
Drepanolejeunea angustifolia (Mitt.) Grolle..................................................................................353
Drepanolejeunea ankasica E.W.Jones ...........................................................................................353
Drepanolejeunea anoplantha (Spruce) Steph. ...............................................................................354
Drepanolejeunea appalachiana R.M.Schust. .................................................................................354
Drepanolejeunea araucariae Steph................................................................................................354
Drepanolejeunea araucariae var. chilensis Herzog .........................................................................354
Drepanolejeunea aucklandica Steph. ............................................................................................354
Drepanolejeunea aurita Bischl......................................................................................................354
Drepanolejeunea bakeri Herzog ...................................................................................................354
Drepanolejeunea bidens (Prantl) A.Evans .....................................................................................354
Drepanolejeunea bidoupensis Pócs ................................................................................................360
Drepanolejeunea biocellata A.Evans..............................................................................................354
Drepanolejeunea bischlerae (Grolle) Grolle et R.L.Zhu.................................................................360
Drepanolejeunea blumei Steph. ....................................................................................................354
Drepanolejeunea brunnea Mizut. .................................................................................................354
Drepanolejeunea caledonica Steph. ...............................................................................................354
Drepanolejeunea campanulata (Spruce) Steph. .............................................................................354
Drepanolejeunea canceroides H.A.Mill. et Bonner ........................................................................354
Drepanolejeunea capulata (Taylor) J.B.Jack et Steph. ...................................................................354
Drepanolejeunea capulata var. lagellifera S.W.Arnell ...................................................................354
Drepanolejeunea ciliata Mizut. ....................................................................................................354
Drepanolejeunea commutata Grolle et R.L.Zhu ...........................................................................360
Drepanolejeunea crassiretis A.Evans ..............................................................................................354
Drepanolejeunea crucianella (Taylor) A.Evans ..............................................................................354
Drepanolejeunea cultrella (Mitt.) Steph........................................................................................354
Drepanolejeunea cutervoensis (Loitl.) Grolle .................................................................................354
Drepanolejeunea cyclops (Sande Lac.) Grolle et R.L.Zhu ..............................................................360
Drepanolejeunea dactylophora (Nees, Lindenb. et Gottsche) J.B.Jack et Steph..............................360
Drepanolejeunea dactylophora var. submuricata Herzog ................................................................360
Drepanolejeunea decurviloba Steph. .............................................................................................355
Drepanolejeunea dentistipula Steph. .............................................................................................353
Drepanolejeunea deslooveri Vanden Berghen ................................................................................355
Drepanolejeunea devendrae Sushil K.Singh et M.Dey ..................................................................360
Drepanolejeunea dissitifolia A.Evans.............................................................................................355
Drepanolejeunea elegans Herzog ...................................................................................................355
Drepanolejeunea erecta (Steph.) Mizut. ........................................................................................355
Drepanolejeunea evansii Bischl. ex L.Söderstr., A.Hagborg et von Konrat ....................................355
Drepanolejeunea issicornua Steph. ...............................................................................................359
Drepanolejeunea leischeri (Steph.) Grolle et R.L.Zhu ..................................................................360
Drepanolejeunea foliicola Horik. ..................................................................................................360
554
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Drepanolejeunea fragilis Bischl. ex L.Söderstr., A.Hagborg et von Konrat ....................................355
Drepanolejeunea fulfordiae L.Söderstr. .........................................................................................355
Drepanolejeunea granatensis (Prantl) Bischl..................................................................................355
Drepanolejeunea grandis Herzog ..................................................................................................355
Drepanolejeunea grollei M.E.Reiner et Schäf.-Verw. ....................................................................355
Drepanolejeunea grossidens Steph. ................................................................................................355
Drepanolejeunea hamatifolia (Hook.) Schifn. .............................................................................355
Drepanolejeunea hampeana Steph. ...............................................................................................359
Drepanolejeunea hamulata Steph. ................................................................................................355
Drepanolejeunea helenae Pócs ......................................................................................................355
Drepanolejeunea herzogii R.L.Zhu et M.L.So ..............................................................................355
Drepanolejeunea inchoata (C.F.W.Meissn.) Steph. ......................................................................356
Drepanolejeunea inchoata var. palmicola Pócs ..............................................................................356
Drepanolejeunea inchoata var. roraimae (Steph. ex Zwickel) Bischl. .............................................356
Drepanolejeunea infundibulata (Spruce) A.Evans .........................................................................356
Drepanolejeunea integerrima Herzog ............................................................................................360
Drepanolejeunea integribracteata Bischl........................................................................................356
Drepanolejeunea intermedia Zwickel ............................................................................................359
Drepanolejeunea laciniata Qiong He et R.L.Zhu .........................................................................359
Drepanolejeunea laevis (Mitt.) Steph............................................................................................356
Drepanolejeunea lancifolia (Gottsche) J.B.Jack et Steph. ..............................................................356
Drepanolejeunea levicornua Steph. ...............................................................................................356
Drepanolejeunea lichenicola (Spruce) Steph..................................................................................356
Drepanolejeunea longicornua (Herzog) Mizut. .............................................................................359
Drepanolejeunea longicruris (Steph.) Grolle et R.L.Zhu ...............................................................360
Drepanolejeunea longii Grolle et R.L.Zhu ....................................................................................356
Drepanolejeunea lyrata Grolle ......................................................................................................359
Drepanolejeunea macrodonta (Mitt.) Steph. .................................................................................356
Drepanolejeunea madagascariensis (Steph.) Grolle ........................................................................359
Drepanolejeunea mawtmiana Ajit P.Singh et V.Nath ...................................................................356
Drepanolejeunea microcarpa Pearson............................................................................................356
Drepanolejeunea moluccensis Herzog ............................................................................................356
Drepanolejeunea mosenii (Steph.) Bischl. .....................................................................................356
Drepanolejeunea nymanii Steph. ..................................................................................................359
Drepanolejeunea obliqua Steph. ...................................................................................................356
Drepanolejeunea obtriangulata T.Kodama ...................................................................................356
Drepanolejeunea obtusifolia T.Yamag. ..........................................................................................357
Drepanolejeunea orthophylla (Nees et Mont.) Bischl. ...................................................................357
Drepanolejeunea palmifolia (Nees) Schifn. ..................................................................................357
Drepanolejeunea pentadactyla (Mont.) Steph. ..............................................................................357
Drepanolejeunea perissodonta (Spruce) Bischl. ..............................................................................357
Drepanolejeunea physifolia (Gottsche) Pearson .............................................................................357
Drepanolejeunea pinnatiloba Schifn. ...........................................................................................357
Drepanolejeunea pleiodictya Herzog .............................................................................................357
Drepanolejeunea pocsii Grolle ......................................................................................................359
Drepanolejeunea polyrhiza (Nees) Grolle et R.L.Zhu ...................................................................360
Drepanolejeunea propagulifera Herzog .........................................................................................357
Drepanolejeunea pseudoneura (A.Evans) Grolle ............................................................................357
Drepanolejeunea pterocalyx (Herzog) Bischl. ................................................................................357
Drepanolejeunea pulla (Mitt.) Grolle ...........................................................................................360
Drepanolejeunea pungens Bischl. ..................................................................................................357
Drepanolejeunea ramentilora Steph. ............................................................................................357
World checklist of hornworts and liverworts
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555
Drepanolejeunea ruandensis Vanden Berghen...............................................................................357
Drepanolejeunea senticosa Bischl. .................................................................................................357
Drepanolejeunea serricalyx Herzog ...............................................................................................360
Drepanolejeunea siamensis (Bischl.) Grolle et R.L.Zhu.................................................................360
Drepanolejeunea sikkimensis (Udar et U.S.Awasthi) Grolle ..........................................................357
Drepanolejeunea spicata (Steph.) Grolle et R.L.Zhu.....................................................................360
Drepanolejeunea spinistipula Herzog ............................................................................................353
Drepanolejeunea spinosa Herzog ..................................................................................................358
Drepanolejeunea spinosocornuta Steph. .........................................................................................358
Drepanolejeunea subdissitifolia Herzog .........................................................................................358
Drepanolejeunea submuricata R.M.Schust. ..................................................................................358
Drepanolejeunea subquadrata (Mitt.) Steph. ................................................................................358
Drepanolejeunea subvittata (Herzog) Grolle .................................................................................358
Drepanolejeunea symoensii Vanden Berghen et Grolle ..................................................................359
Drepanolejeunea symoensii var. minor Tixier ................................................................................359
Drepanolejeunea tenax K.I.Goebel ...............................................................................................358
Drepanolejeunea tenera K.I.Goebel ..............................................................................................358
Drepanolejeunea tenera var. litoceras Herzog ................................................................................358
Drepanolejeunea ternatensis (Gottsche) Schifn. ...........................................................................358
Drepanolejeunea teysmannii (Gottsche) Steph. .............................................................................358
Drepanolejeunea thwaitesiana (Mitt.) Steph. ................................................................................360
Drepanolejeunea thwaitesiana var. zhengii R.L.Zhu......................................................................360
Drepanolejeunea tibetana (P.C.Wu et J.S.Lou) Grolle et R.L.Zhu ...............................................360
Drepanolejeunea trematodes (Nees) Bischl. ...................................................................................359
Drepanolejeunea tricornua Herzog ...............................................................................................360
Drepanolejeunea tridactyla (Gottsche) Steph. ...............................................................................358
Drepanolejeunea trigonophylla Steph. ...........................................................................................358
Drepanolejeunea tristaniana S.W.Arnell.......................................................................................358
Drepanolejeunea tuyamae S.Hatt. ................................................................................................358
Drepanolejeunea ualanensis Inoue et H.A.Mill. ............................................................................358
Drepanolejeunea ungulata (Steph.) Grolle ....................................................................................358
Drepanolejeunea urceolata R.M.Schust. .......................................................................................359
Drepanolejeunea valiae Jovet-Ast .................................................................................................359
Drepanolejeunea vandenberghenii Buchb. et Eb.Fisch. .................................................................359
Drepanolejeunea vesiculosa (Mitt.) Steph......................................................................................359
Drepanolejeunea yulensis Steph. ...................................................................................................359
Drepanolejeunea yunnanensis (P.C.Chen) Grolle et R.L.Zhu .......................................................360
Drucella integristipula (Steph.) E.A.Hodgs. .................................................................................163
Dumortiera hirsuta (Sw.) Nees ....................................................................................................487
Dumortiera hirsuta subsp. nepalensis (Taylor) R.M.Schust...........................................................487
Dumortiera hirsuta subsp. tatunoi Horik. ....................................................................................487
Echinolejeunea papillata (Mitt.) R.M.Schust. ex Hamlin .............................................................362
Endogemma caespiticia (Lindenb.) Konstant., Vilnet et A.V.Troitsky ..........................................109
Enigmella thallina G.A.M.Scott et K.G.Beckm. ............................................................................93
Eocalypogeia quelpaertensis (S.Hatt. et Inoue) R.M.Schust. ..........................................................107
Eocalypogeia schusterana (S.Hatt. et Mizut.) R.M.Schust. ............................................................107
Eotrichocolea furukii T.Katag.......................................................................................................256
Eotrichocolea polyacantha (Hook.f. et Taylor) R.M.Schust...........................................................256
Eremonotus myriocarpus (Carrington) Lindb. et Kaal. ex Pearson.................................................118
Eulejeunea setulosa Steph. ............................................................................................................508
Eulejeunea subpililoba Steph. .......................................................................................................508
Euosmolejeunea parvistipula (Lindenb. et Gottsche) Steph. ..........................................................508
556
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Euosmolejeunea tenerrima (Nees) Steph. ......................................................................................508
Evansianthus georgiensis (Gottsche) R.M.Schust. et J.J.Engel .......................................................196
Exormotheca bischlerae Furuki et Higuchi ....................................................................................488
Exormotheca bulbigena Bornefeld, O.H.Volk et R.Wolf ..............................................................488
Exormotheca ceylonensis Meijer ....................................................................................................488
Exormotheca gollanii Steph. .........................................................................................................488
Exormotheca holstii Steph. ...........................................................................................................488
Exormotheca pustulosa Mitt. ........................................................................................................488
Exormotheca tuberifera Kashyap ..................................................................................................488
Exormotheca welwitschii Steph. ....................................................................................................488
Fimbraria gigantea Steph. ...........................................................................................................508
Fimbraria incrassata Steph. .........................................................................................................508
Fimbraria kamerunensis Steph. ....................................................................................................508
Fimbraria pirottae Gola ...............................................................................................................508
Folioceros amboinensis (Schifn.) Piippo .........................................................................................31
Folioceros apiahynus (Steph.) Hässel ..............................................................................................31
Folioceros argillaceus (Steph.) J.C.Villarreal et Cargill ....................................................................31
Folioceros assamicus D.C.Bharadwaj ..............................................................................................31
Folioceros dilatatus (Steph.) J.C.Villarreal et Cargill .......................................................................31
Folioceros dixitianus (Mahab.) D.C.Bharadwaj ..............................................................................31
Folioceros fuciformis (Mont.) D.C.Bharadwaj ................................................................................31
Folioceros glandulosus (Lehm. et Lindenb.) D.C.Bharadwaj ...........................................................31
Folioceros incurvus (Steph.) D.C.Bharadwaj...................................................................................31
Folioceros indicus D.C.Bharadwaj ..................................................................................................31
Folioceros kashyapii S.C.Srivast. et A.K.Asthana.............................................................................31
Folioceros mangaloreus (Steph.) D.C.Bharadwaj .............................................................................31
Folioceros paliformis D.K.Singh .....................................................................................................31
Folioceros physocladus D.C.Bharadwaj............................................................................................31
Folioceros pinnilobus (Steph.) D.C.Bharadwaj................................................................................31
Folioceros satpurensis D.C.Bharadwaj et K.P.Srivast. ......................................................................31
Folioceros udarii A.K.Asthana et S.C.Srivast. .................................................................................32
Folioceros verruculosus (J.Haseg.) R.L.Zhu et M.J.Lai ....................................................................32
Fossombronia alaskana Steere et Inoue .........................................................................................463
Fossombronia alata G.A.M.Scott et D.C.Pike ..............................................................................463
Fossombronia altilamellosa G.A.M.Scott et D.C.Pike ...................................................................463
Fossombronia angulifolia Perold ...................................................................................................463
Fossombronia angulosa (Dicks.) Raddi .........................................................................................463
Fossombronia areolata G.A.M.Scott et D.C.Pike .........................................................................464
Fossombronia auricolor G.A.M.Scott et D.C.Pike ........................................................................464
Fossombronia australis Mitt. ........................................................................................................464
Fossombronia caespitiformis (Raddi) De Not. ex Rabenh. .............................................................464
Fossombronia caespitiformis subsp. multispira (Schifn.) J.R.Bray et Cargill ..................................464
Fossombronia caledonica Steph. ....................................................................................................464
Fossombronia cederbergensis Perold...............................................................................................464
Fossombronia cerebriformis G.A.M.Scott et D.C.Pike ..................................................................464
Fossombronia crassifolia Spruce ....................................................................................................464
Fossombronia crispa Nees .............................................................................................................464
Fossombronia crispula (Brot.) R.M.Schust....................................................................................464
Fossombronia cristula Austin ........................................................................................................464
Fossombronia cultriformis G.A.M.Scott et D.C.Pike ....................................................................464
Fossombronia densa G.A.M.Scott et D.C.Pike .............................................................................464
Fossombronia densilamellata S.W.Arnell ......................................................................................464
World checklist of hornworts and liverworts
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557
Fossombronia echinata Macvicar ..................................................................................................464
Fossombronia elsieae Perold ..........................................................................................................464
Fossombronia fernandeziensis Steph. .............................................................................................464
Fossombronia imbriata Paton ......................................................................................................464
Fossombronia leischeri Osterwald ex Loeske ................................................................................464
Fossombronia foveolata Lindb. .....................................................................................................464
Fossombronia fuhreri G.A.M.Scott et D.C.Pike ...........................................................................464
Fossombronia gemmifera Perold ...................................................................................................465
Fossombronia glenii Perold...........................................................................................................465
Fossombronia grandis Steph. ........................................................................................................465
Fossombronia gregaria Colenso ....................................................................................................465
Fossombronia grossepapillata Steph. ..............................................................................................465
Fossombronia hamatohirta Steph..................................................................................................465
Fossombronia hewsoniae G.A.M.Scott et D.C.Pike ......................................................................465
Fossombronia himalayensis Kashyap .............................................................................................465
Fossombronia hyalorhiza Perold ...................................................................................................465
Fossombronia incurva Lindb. .......................................................................................................465
Fossombronia indica Steph. ..........................................................................................................465
Fossombronia integerrima Steph. ..................................................................................................465
Fossombronia integrifolia Steph. ...................................................................................................465
Fossombronia intestinalis Taylor ...................................................................................................465
Fossombronia japonica Schifn. ....................................................................................................465
Fossombronia laciniata G.A.M.Scott et D.C.Pike ........................................................................465
Fossombronia lamellata Steph. .....................................................................................................465
Fossombronia leucoxantha (Lehm.) Lehm. et Lindenb. .................................................................465
Fossombronia longiseta (Austin) Austin ........................................................................................465
Fossombronia lophoclada Spruce...................................................................................................465
Fossombronia lophoscypha Hässel..................................................................................................465
Fossombronia luetzelburgiana K.I.Goebel .....................................................................................465
Fossombronia macrocalyx Steph. ...................................................................................................465
Fossombronia macrophylla Colenso ..............................................................................................466
Fossombronia magnaspora G.A.M.Scott et D.C.Pike ....................................................................466
Fossombronia marindae Perold.....................................................................................................466
Fossombronia maritima (Paton) Paton .........................................................................................466
Fossombronia marshii J.R.Bray et Stotler .....................................................................................466
Fossombronia microlamellata G.A.M.Scott et D.C.Pike ...............................................................466
Fossombronia mittenii Tind. ........................................................................................................466
Fossombronia montaguensis S.W.Arnell ........................................................................................466
Fossombronia monticola Perold ....................................................................................................466
Fossombronia mylioides Inoue ......................................................................................................466
Fossombronia nigricaulis Colenso .................................................................................................466
Fossombronia nyikaensis Perold ....................................................................................................466
Fossombronia papillata Steph. ......................................................................................................466
Fossombronia paranapanemae Schifn. .........................................................................................466
Fossombronia peruviana Gottsche et Hampe................................................................................466
Fossombronia porphyrorhiza (Nees) Prosk. ...................................................................................466
Fossombronia pulvinata Steph. .....................................................................................................466
Fossombronia punctata G.A.M.Scott et D.C.Pike ........................................................................466
Fossombronia purpureospora G.A.M.Scott et D.C.Pike ................................................................466
Fossombronia pusilla (L.) Nees .....................................................................................................466
Fossombronia renateae Perold.......................................................................................................466
Fossombronia reticulata Steph. .....................................................................................................466
558
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Fossombronia rosulata Colenso.....................................................................................................467
Fossombronia rudis G.A.M.Scott et D.C.Pike ..............................................................................467
Fossombronia ruminata Cargill ....................................................................................................467
Fossombronia rupestris G.A.M.Scott et D.C.Pike .........................................................................467
Fossombronia rwandaensis Perold .................................................................................................467
Fossombronia scrobiculata G.A.M.Scott et D.C.Pike ....................................................................467
Fossombronia spinifolia Steph. .....................................................................................................467
Fossombronia spinosa Perold ........................................................................................................467
Fossombronia stephanii Schifn. ex Steph. ....................................................................................467
Fossombronia straussiana Perold ...................................................................................................467
Fossombronia subsaccata Steph. ....................................................................................................467
Fossombronia swaziensis Perold ....................................................................................................467
Fossombronia tesselata G.A.M.Scott et D.C.Pike..........................................................................467
Fossombronia texana Lindb. .........................................................................................................467
Fossombronia truncata G.A.M.Scott et D.C.Pike .........................................................................467
Fossombronia tumida Mitt. ..........................................................................................................467
Fossombronia valparaisiana Hässel ...............................................................................................467
Fossombronia vermiculata G.A.M.Scott et D.C.Pike ....................................................................467
Fossombronia wattsii Steph. .........................................................................................................467
Fossombronia wondraczekii (Corda) Dumort. ex Lindb. ..............................................................467
Fossombronia wrightii Austin .......................................................................................................467
Fossombronia zuurbergensis Perold ...............................................................................................467
Frullania acicularis Hentschel et von Konrat ...............................................................................292
Frullania aculeata Taylor ............................................................................................................287
Frullania ainis Nees et Mont.....................................................................................................293
Frullania akiyamae S.Hatt...........................................................................................................267
Frullania albertii Steph. ..............................................................................................................266
Frullania allanii E.A.Hodgs. .......................................................................................................281
Frullania allionii Steph. ..............................................................................................................293
Frullania alpina Steph.................................................................................................................293
Frullania alstonii Verd. ...............................................................................................................292
Frullania alternans Nees ..............................................................................................................293
Frullania amamiensis Kamim. .....................................................................................................272
Frullania ambronnii Steph. .........................................................................................................287
Frullania amplicrania Steph. .......................................................................................................272
Frullania ampullifera J.B.Jack et Steph. .......................................................................................272
Frullania anderssonii Ångstr. .......................................................................................................272
Frullania angulata Mitt. ..............................................................................................................289
Frullania angulata var. laciniata Vanden Berghen .......................................................................289
Frullania angustistipa Steph.........................................................................................................272
Frullania anomala E.A.Hodgs. ....................................................................................................282
Frullania antaresensis S.Hatt........................................................................................................266
Frullania aoshimensis Horik. .......................................................................................................292
Frullania apertilobula Gerola.......................................................................................................293
Frullania apicalis Mitt. ................................................................................................................289
Frullania apicalis var. camerunensis Vanden Berghen ...................................................................289
Frullania apiculata (Reinw., Blume et Nees) Nees.......................................................................267
Frullania apiculata var. goebelii Schifn........................................................................................267
Frullania aposinensis S.Hatt. et P.J.Lin ........................................................................................273
Frullania appalachiana R.M.Schust.............................................................................................284
Frullania appendiculata Steph. ....................................................................................................292
Frullania appendistipula S.Hatt. ..................................................................................................273
World checklist of hornworts and liverworts
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559
Frullania appendistipula var. spinifera S.Hatt. ..............................................................................273
Frullania armata Herzog et L.Clark ............................................................................................293
Frullania armatifolia Verd...........................................................................................................267
Frullania armitiana Steph. ..........................................................................................................267
Frullania armitiana var. inlexula S.Hatt. ....................................................................................268
Frullania arsenii Steph. ...............................................................................................................266
Frullania asagrayana Mont..........................................................................................................292
Frullania astrolabea Steph. ..........................................................................................................285
Frullania aterrima (Hook.f. et Taylor) Gottsche, Lindenb. et Nees .............................................268
Frullania atrata (Sw.) Nees ex Mont. ..........................................................................................287
Frullania attenuata Steph. ...........................................................................................................268
Frullania auriculata S.Hatt. ........................................................................................................273
Frullania azorica Sim-Sim, Sérgio, Mues et Kraut .......................................................................284
Frullania baileyana Steph. ...........................................................................................................282
Frullania baladina Gottsche ........................................................................................................282
Frullania baumannii S.Hatt. .......................................................................................................271
Frullania beauverdii Steph. .........................................................................................................265
Frullania bella Steph. ..................................................................................................................268
Frullania belmorensis Steph. ........................................................................................................282
Frullania benjaminiana Inoue .....................................................................................................273
Frullania bergmanii S.Hatt..........................................................................................................273
Frullania berthoumieui Steph. .....................................................................................................273
Frullania beyrichiana (Lehm. et Lindenb.) Lehm. et Lindenb. ....................................................287
Frullania bhutanensis S.Hatt. ......................................................................................................273
Frullania bicornistipula Spruce ....................................................................................................287
Frullania blastopetala S.Hatt. ......................................................................................................273
Frullania blepharozia Spruce .......................................................................................................265
Frullania bogotensis Steph............................................................................................................265
Frullania bolanderi Austin ...........................................................................................................294
Frullania bonariensis M.E.Reiner ................................................................................................266
Frullania bonincola S.Hatt. .........................................................................................................273
Frullania borbonica Lindenb. ......................................................................................................289
Frullania boveana C.Massal.........................................................................................................294
Frullania brachycarpa Spruce.......................................................................................................265
Frullania brasiliensis Raddi ..........................................................................................................287
Frullania brasiliensis var. elegantula Spruce ..................................................................................287
Frullania breuteliana Gottsche ....................................................................................................287
Frullania brevicalycina Steph. ......................................................................................................273
Frullania brittoniae A.Evans ........................................................................................................273
Frullania brotheri Steph. .............................................................................................................286
Frullania brunea (Spreng.) Drège ................................................................................................272
Frullania bullata Steph................................................................................................................273
Frullania caespitans Beauverd ......................................................................................................294
Frullania cafraria Steph. ............................................................................................................273
Frullania calcarata Ångstr. ..........................................................................................................273
Frullania calcarifera Steph. ..........................................................................................................292
Frullania caldensis Ångstr. ...........................................................................................................287
Frullania californica (M.Howe) A.Evans .....................................................................................292
Frullania campanulata Sande Lac. ...............................................................................................282
Frullania campanulata var. caduca Verd. .....................................................................................282
Frullania campanulata var. malesiaca (Verd.) S.Hatt. ..................................................................282
Frullania canaliculata Gottsche ...................................................................................................294
560
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Frullania capensis Gottsche .........................................................................................................289
Frullania capillaris Steph. ............................................................................................................294
Frullania carrii Kamim. et S.Hatt. ..............................................................................................273
Frullania catalinae A.Evans .........................................................................................................284
Frullania cataractarum Steph. .....................................................................................................282
Frullania caulisequa (Nees) Mont................................................................................................272
Frullania cavallii Gola.................................................................................................................294
Frullania changii S.Hatt. et C.Gao ..............................................................................................268
Frullania chenii S.Hatt. et P.J.Lin ...............................................................................................273
Frullania chevalieri (R.M.Schust.) R.M.Schust. ..........................................................................291
Frullania chiapasana Steph. .........................................................................................................294
Frullania chilcootiensis Steph. ......................................................................................................294
Frullania chilensis Steph. .............................................................................................................273
Frullania chiovendae Gola ...........................................................................................................294
Frullania chodatii Beauverd.........................................................................................................273
Frullania ciliata Lindenb. et Gottsche .........................................................................................294
Frullania cinchonae Gottsche ......................................................................................................294
Frullania clavata (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ...............................................281
Frullania claviloba Steph. ............................................................................................................268
Frullania clemensiana Verd. ........................................................................................................292
Frullania cobrensis Gottsche ........................................................................................................273
Frullania colliculosa von Konrat, Braggins, Hentschel et Heinrichs .............................................268
Frullania compacta Gottsche .......................................................................................................287
Frullania complicata Steph. .........................................................................................................294
Frullania confertiloba Steph. ........................................................................................................265
Frullania congesta Gottsche, Lindenb. et Nees.............................................................................271
Frullania consociata Steph. ..........................................................................................................273
Frullania contracta Steph. ...........................................................................................................273
Frullania convoluta Lindenb. et Hampe ......................................................................................289
Frullania convoluta var. ampliata Herzog ....................................................................................289
Frullania cordaeana Lindenb. ......................................................................................................294
Frullania cordistipula (Reinw., Blume et Nees) Nees ...................................................................268
Frullania cordistipula var. dentistipula S.Hatt. .............................................................................268
Frullania cordistipula var. mutica (Gottsche, Lindenb. et Nees) Schifn. ......................................268
Frullania cornuta Steph. ..............................................................................................................273
Frullania crassitexta Steph. ..........................................................................................................273
Frullania crawfordii Steph. ..........................................................................................................282
Frullania crenatiloba Steph..........................................................................................................268
Frullania crenulifolia J.B.Jack et Steph. .......................................................................................287
Frullania crispiloba Steph. ...........................................................................................................288
Frullania crispiplicata Yuzawa et S.Hatt. .....................................................................................273
Frullania cristata S.Hatt. .............................................................................................................274
Frullania cuencensis Taylor ..........................................................................................................266
Frullania cuneatistipula Steph. ....................................................................................................294
Frullania cuneiloba Nees .............................................................................................................274
Frullania curvilobula Schäf.-Verw., D.F.Peralta et S.M.Siqueira .................................................267
Frullania curviramea Steph. ........................................................................................................288
Frullania curvistipula Steph.........................................................................................................274
Frullania curvistipula var. falcatidentata S.Hatt. ..........................................................................274
Frullania curvistipula var. lamii Verd. .........................................................................................274
Frullania cuspidifolia Steph. ........................................................................................................274
Frullania cuspiloba Steph. ...........................................................................................................294
World checklist of hornworts and liverworts
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561
Frullania cyparioides (Schwägr.) Nees ..........................................................................................274
Frullania darwinii Gradst. et Uribe.............................................................................................289
Frullania davurica Hampe ex Gottsche, Lindenb. et Nees...........................................................274
Frullania debilis Steph. ex S.Hatt. ...............................................................................................274
Frullania decidua Spruce .............................................................................................................265
Frullania delexa Mitt..................................................................................................................285
Frullania degelii S.W.Arnell ........................................................................................................268
Frullania densiloba Steph. ex A.Evans .........................................................................................292
Frullania dentata S.Hatt..............................................................................................................282
Frullania dentata var. secernens S.Hatt. ........................................................................................282
Frullania dentifera S.Hatt. et Streimann......................................................................................268
Frullania dentiloba S.Hatt. ..........................................................................................................268
Frullania deplanata Mitt. ............................................................................................................285
Frullania deppii Lehm. ................................................................................................................274
Frullania depressa Mitt. ...............................................................................................................266
Frullania dilatata (L.) Dumort. ...................................................................................................284
Frullania dilatata subsp. asiatica S.Hatt. .....................................................................................284
Frullania diptera (Lehm.) Drège .................................................................................................274
Frullania dispar Nees ..................................................................................................................294
Frullania diversitexta Steph. ........................................................................................................272
Frullania donnellii Austin............................................................................................................289
Frullania dulimensis Uribe ..........................................................................................................289
Frullania durifolia Steph. ............................................................................................................268
Frullania dusenii Steph. ..............................................................................................................266
Frullania duthiana Steph. ...........................................................................................................274
Frullania duthiana var. appendiculata S.Hatt...............................................................................274
Frullania duthiana var. laevis S.Hatt. ..........................................................................................274
Frullania duthiana var. szechuanensis S.Hatt. et C.Gao................................................................274
Frullania eboracensis Lehm. .........................................................................................................284
Frullania echinantha S.Hatt. .......................................................................................................274
Frullania echinatella S.Hatt. ........................................................................................................274
Frullania ecklonii (Spreng.) Gottsche, Lindenb. et Nees ..............................................................265
Frullania ecklonii var. robustior (Gottsche, Lindenb. et Nees) Sim...............................................265
Frullania ecklonii var. rufescens Gottsche, Lindenb. et Nees .........................................................265
Frullania ecklonii var. tenerior (Gottsche, Lindenb. et Nees) Sim ................................................265
Frullania ecuadorensis Steph. .......................................................................................................288
Frullania elegans Lehm. ...............................................................................................................274
Frullania elephantum S.Hatt. ......................................................................................................274
Frullania epiphylla S.Hatt. ..........................................................................................................274
Frullania epiphylla subsp. ijiensis S.Hatt. ....................................................................................274
Frullania eplicata Steph. ..............................................................................................................289
Frullania ericoides (Nees) Mont...................................................................................................284
Frullania ericoides var. laxa (Gottsche, Lindenb. et Nees) Schifn. ...............................................284
Frullania ericoides var. verrucosa (Kamim.) Hentschel et von Konrat ...........................................284
Frullania ericoides var. minor Kamim. .........................................................................................274
Frullania erostrata S.Hatt. ...........................................................................................................274
Frullania errans Verd. .................................................................................................................282
Frullania errans var. angulistipula S.Hatt. ....................................................................................283
Frullania esenbeckiana Beauverd..................................................................................................274
Frullania evelynae S.Hatt. et haithong .......................................................................................274
Frullania evelynae var. devendrae Sushil K.Singh et Barbhuiya.....................................................274
Frullania evelynae var. srivastavae Sushil K.Singh et Barbhuiya....................................................274
562
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Frullania evoluta Mitt. ................................................................................................................287
Frullania exilis Taylor .................................................................................................................268
Frullania eymae S.Hatt. ...............................................................................................................274
Frullania eymae var. crispidentata S.Hatt. et Streimann ...............................................................274
Frullania falciloba Lehm. ............................................................................................................274
Frullania fallax Gottsche .............................................................................................................285
Frullania falsicornuta S.Hatt. ......................................................................................................274
Frullania fauriana Steph. ............................................................................................................274
Frullania fengyangshanensis R.L.Zhu et M.L.So ...........................................................................275
Frullania ferdinandi-muelleri Steph. ............................................................................................275
Frullania fertilis De Not. .............................................................................................................290
Frullania lammea Taylor ............................................................................................................264
Frullania lexuosa S.Hatt. ............................................................................................................275
Frullania formosa Spruce .............................................................................................................288
Frullania fragilifolia (Taylor) Gottsche, Lindenb. et Nees ...........................................................284
Frullania franciscana M.Howe ....................................................................................................292
Frullania fuegiana Steph. ............................................................................................................284
Frullania fugax (Hook.f. et Taylor) Gottsche, Lindenb. et Nees..................................................283
Frullania fulfordiae S.Hatt. .........................................................................................................283
Frullania fuscovirens Steph. .........................................................................................................275
Frullania fuscovirens var. gemmipara (R.M.Schust. et S.Hatt.) S.Hatt. et P.J.Lin .........................275
Frullania gabonensis Vanden Berghen .........................................................................................268
Frullania gaoligongensis X.L.Bai et C.Gao....................................................................................275
Frullania gaudichaudii (Nees et Mont.) Nees et Mont. ...............................................................291
Frullania gaudichaudii var. ceylanica (Nees) S.Hatt. ....................................................................292
Frullania gemmulosa S.Hatt. et haithong...................................................................................275
Frullania gibbosa Nees ................................................................................................................265
Frullania gigantea Steph. .............................................................................................................275
Frullania giraldiana C.Massal. ....................................................................................................275
Frullania giraldiana var. handelii (Verd.) S.Hatt. ........................................................................275
Frullania globosa S.Hatt. et Streimann ........................................................................................275
Frullania glomerata (Lehm. et Lindenb.) Nees et Mont...............................................................283
Frullania gracilicaulis S.Hatt. ......................................................................................................267
Frullania gracilis (Reinw., Blume et Nees) Nees ..........................................................................268
Frullania gracilis var. brevior (Gottsche, Lindenb. et Nees) Schifn..............................................269
Frullania gracilis var. vittata S.Hatt. ............................................................................................269
Frullania gracilis subsp. zennoskei S.Hatt. et haithong ...............................................................269
Frullania gradsteinii Yuzawa, Mues et S.Hatt. .............................................................................266
Frullania granatensis Gottsche .....................................................................................................288
Frullania grandifolia Steph. .........................................................................................................289
Frullania grandilobula S.Hatt. et Piippo......................................................................................275
Frullania grandistipula Lindenb. .................................................................................................275
Frullania griithsiana Gottsche ...................................................................................................288
Frullania grossiclava Steph. ..........................................................................................................271
Frullania grossifolia Steph. ...........................................................................................................272
Frullania guadalupensis Gottsche.................................................................................................288
Frullania gualaquizana Steph. .....................................................................................................288
Frullania guatemalensis Steph. .....................................................................................................294
Frullania haematocysta Spruce .....................................................................................................266
Frullania hainanensis S.Hatt. et P.J.Lin .......................................................................................275
Frullania hamata Steph. ..............................................................................................................286
Frullania hamaticoma Steph. .......................................................................................................281
World checklist of hornworts and liverworts
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563
Frullania hamatiloba Steph. ........................................................................................................275
Frullania hamilora Herzog et L.Clark ........................................................................................288
Frullania handelii Verd. ..............................................................................................................275
Frullania handel-mazzettii S.Hatt................................................................................................275
Frullania hasskarliana Lindenb. ..................................................................................................269
Frullania hasskarliana var. gracilis S.Hatt. ...................................................................................269
Frullania hasskarliana var. parvidentata S.Hatt. ...........................................................................269
Frullania hattoriana J.D.Godfrey et G.Godfrey ..........................................................................284
Frullania hattoriantha Udar et V.Nath ........................................................................................275
Frullania hattorii von Konrat et Braggins ....................................................................................269
Frullania hebridensis Steph. .........................................................................................................275
Frullania hedrantha S.Hatt. et Kamim. .......................................................................................292
Frullania heteromorpha Schifn. ...................................................................................................286
Frullania hicksiae S.Hatt. ............................................................................................................275
Frullania higuchii Yuzawa, Koike et S.Hatt. ................................................................................275
Frullania hiroshii S.Hatt. ............................................................................................................275
Frullania hirtilora Spruce ...........................................................................................................275
Frullania hodgsoniae von Konrat, Braggins, Hentschel et Heinrichs ............................................269
Frullania holostipula S.Hatt. et D.G.Griin ................................................................................265
Frullania hottana S.Hatt. ............................................................................................................269
Frullania howeana Steph. ............................................................................................................276
Frullania huerlimannii S.Hatt. ....................................................................................................286
Frullania huerlimannii var. dioica S.Hatt. ...................................................................................286
Frullania humbertii Vanden Berghen ..........................................................................................267
Frullania humilis Spruce .............................................................................................................276
Frullania hypoleuca Nees .............................................................................................................272
Frullania hypoleucula S.Hatt. ......................................................................................................267
Frullania ignatovii Sofronova, Mamontov et Potemkin ...............................................................276
Frullania imerinensis Steph..........................................................................................................289
Frullania immersa Steph. ............................................................................................................292
Frullania incisoduthiana S.Hatt. ..................................................................................................276
Frullania incisoduthiana var. parva S.Hatt. ..................................................................................276
Frullania incisostipula Steph. .......................................................................................................276
Frullania inconstans Verd. ...........................................................................................................269
Frullania inconstans var. grossedentata Kamim. et S.Hatt. ............................................................269
Frullania incumbens Mitt. ...........................................................................................................283
Frullania incurva S.Hatt. ............................................................................................................267
Frullania inlata Gottsche ...........................................................................................................294
Frullania inlata var. communis R.M.Schust. ...............................................................................294
Frullania inlata var. dioica S.Hatt. et haithong ........................................................................294
Frullania inlata var. mayebarae (S.Hatt.) K.Yamada ...................................................................294
Frullania inlexa Mitt. .................................................................................................................283
Frullania inlexiloba S.Hatt. ........................................................................................................276
Frullania inouei S.Hatt. ..............................................................................................................276
Frullania integristipula (Nees) Nees.............................................................................................285
Frullania integristipula var. emarginata Verd. ..............................................................................285
Frullania intermedia (Reinw., Blume et Nees) Nees ....................................................................286
Frullania intermedia subsp. morokensis (Steph.) S.Hatt................................................................286
Frullania intermedia var. non-apiculata S.Hatt. ...........................................................................286
Frullania intumescens (Lehm. et Lindenb.) Lehm. et Lindenb. ....................................................288
Frullania involvens S.Hatt. et Kamim..........................................................................................286
Frullania iriomotensis S.Hatt. ......................................................................................................286
564
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Frullania irregularis S.Hatt. et Piippo..........................................................................................276
Frullania iwatsukii S.Hatt. ..........................................................................................................293
Frullania jackii Gottsche .............................................................................................................276
Frullania jacobsii S.Hatt. .............................................................................................................276
Frullania jacquinotii Gottsche .....................................................................................................276
Frullania jelskii Loitl. ..................................................................................................................266
Frullania johnsonii Steph.............................................................................................................269
Frullania jovetiana von Konrat et Hentschel ...............................................................................276
Frullania junghuhniana Gottsche ................................................................................................291
Frullania junghuhniana var. bisexualis S.Hatt. .............................................................................291
Frullania junghuhniana var. tenella (Sande Lac.) Grolle et S.Hatt. ..............................................291
Frullania kagoshimensis Steph. .....................................................................................................276
Frullania kagoshimensis subsp. hunanensis (S.Hatt.) S.Hatt. et P.J.Lin .........................................276
Frullania kalimantanensis S.Hatt. ................................................................................................276
Frullania kashyapii Verd. ............................................................................................................276
Frullania kitagawana S.Hatt. ......................................................................................................276
Frullania klotzschii Nees..............................................................................................................267
Frullania knightbridgei von Konrat et de Lange ...........................................................................290
Frullania koponenii S.Hatt. .........................................................................................................284
Frullania kunzei (Lehm. et Lindenb.) Lehm. et Lindenb.............................................................289
Frullania kunzei var. maritima R.M.Schust. ................................................................................290
Frullania laetevirens Hampe ex Gottsche, Lindenb. et Nees ........................................................294
Frullania laeviperiantha X.L.Bai et C.Gao ...................................................................................276
Frullania lancistyla Steph. ...........................................................................................................276
Frullania larjiana Sushil K.Singh et D.K.Singh ..........................................................................294
Frullania larjiana var. didyhatii S.N.Srivast. et M.Rai .................................................................294
Frullania laticaulis Spruce ...........................................................................................................288
Frullania latilora Spruce.............................................................................................................276
Frullania latogaleata Herzog........................................................................................................276
Frullania laxilora Spruce ............................................................................................................266
Frullania laxilora var. crossii Spruce ............................................................................................276
Frullania leana Austin .................................................................................................................294
Frullania leeuwenii Verd. ............................................................................................................286
Frullania lepida S.Hatt. et Piippo................................................................................................276
Frullania letestui Vanden Berghen ...............................................................................................267
Frullania levieri Steph. ................................................................................................................276
Frullania lindenbergii Lehm. .......................................................................................................272
Frullania lindenbergii var. fusca Gottsche, Lindenb. et Nees ........................................................272
Frullania lindmanii Steph. ..........................................................................................................264
Frullania linii S.Hatt. .................................................................................................................276
Frullania lobatohastata Steph. .....................................................................................................288
Frullania lobulata (Hook.) Hook. et Nees ...................................................................................290
Frullania longistipula Steph. ........................................................................................................290
Frullania longistipula var. apiculata Demaret et Vanden Berghen ................................................288
Frullania longistyla Yuzawa et S.Hatt. .........................................................................................276
Frullania loricata Pearson ............................................................................................................272
Frullania loricata var. laxa Pearson ..............................................................................................272
Frullania ludoviciae Steph. ..........................................................................................................277
Frullania lushanensis S.Hatt. et P.J.Lin........................................................................................277
Frullania macgregorii Steph. ........................................................................................................269
Frullania macgregorii var. rostellula (S.Hatt.) S.Hatt. ...................................................................269
Frullania macrocephala (Lehm. et Lindenb.) Lehm. et Lindenb...................................................288
World checklist of hornworts and liverworts
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565
Frullania macrophylla S.Hatt. ......................................................................................................285
Frullania macularis Taylor ..........................................................................................................277
Frullania madagascariensis Gottsche ............................................................................................294
Frullania madens Steph. ..............................................................................................................269
Frullania madothecoides Spruce ...................................................................................................288
Frullania magellanica F.Weber et Nees .......................................................................................290
Frullania magellanica subsp. tristaniana (S.W.Arnell) Váňa et J.J.Engel ......................................290
Frullania mammillosa S.Hatt.......................................................................................................286
Frullania matafaoica H.A.Mill. ...................................................................................................290
Frullania mauritiana Austin ........................................................................................................294
Frullania maymyoensis Svihla .......................................................................................................277
Frullania mcveanii S.Hatt. ..........................................................................................................291
Frullania media (E.A.Hodgs.) S.Hatt. .........................................................................................283
Frullania megalostipa Spruce .......................................................................................................265
Frullania mehrana S.Hatt. ..........................................................................................................269
Frullania meijeri S.Hatt. .............................................................................................................287
Frullania meridana Steph. ...........................................................................................................288
Frullania meyeniana Lindenb. .....................................................................................................290
Frullania meyeniana var. dioica S.Hatt. .......................................................................................290
Frullania microauriculata Verd....................................................................................................277
Frullania microauriculata var. rotundior Verd. .............................................................................277
Frullania microcaulis Gola ...........................................................................................................291
Frullania microcephala Gottsche..................................................................................................288
Frullania microphylla (Gottsche) Pearson ....................................................................................293
Frullania microrhyncha L.Clark et Svihla .....................................................................................277
Frullania microscopica Pearson ....................................................................................................291
Frullania mirabilis J.B.Jack et Steph............................................................................................294
Frullania miradorensis Lindenb. et Gottsche ...............................................................................267
Frullania mizoramensis Sushil K.Singh et Barbhuiya ...................................................................277
Frullania mizutanii Kamim. et S.Hatt. .......................................................................................283
Frullania moniliata (Reinw., Blume et Nees) Mont. ...................................................................293
Frullania monocera (Hook.f. et Taylor) Gottsche, Lindenb. et Nees............................................281
Frullania monocera var. acutiloba (Mitt.) Hentschel et von Konrat ..............................................282
Frullania monocera var. depauperata S.Hatt. ................................................................................282
Frullania monocera var. schifneri (Verd.) S.Hatt. ........................................................................282
Frullania monocera var. subhampeana (E.A.Hodgs.) Hentschel et von Konrat .............................282
Frullania monocera var. undulata (Kamim.) Hentschel et von Konrat .........................................282
Frullania monoica Steph..............................................................................................................295
Frullania montagnei Gottsche .....................................................................................................288
Frullania montana Steph. ............................................................................................................277
Frullania moritziana Lindenb. et Gottsche..................................................................................288
Frullania morobensis S.Hatt. et Streimann ...................................................................................285
Frullania motoyana Steph. ...........................................................................................................269
Frullania multilacera Steph. ........................................................................................................269
Frullania multilacera subsp. gracilior S.Hatt. ...............................................................................269
Frullania multilacera var. lacerissima S.Hatt. ...............................................................................269
Frullania multilaceroides S.Hatt...................................................................................................269
Frullania multituberculata Hentschel et von Konrat ....................................................................277
Frullania muscicola Steph. ...........................................................................................................284
Frullania mutilata Steph. ............................................................................................................277
Frullania nadeaudii Steph. ..........................................................................................................277
Frullania neocaledonica J.J.Engel .................................................................................................291
566
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Frullania neosheana S.Hatt. .........................................................................................................269
Frullania nepalensis (Spreng.) Lehm. et Lindenb. ........................................................................277
Frullania neurota Taylor .............................................................................................................266
Frullania nicholsonii E.A.Hodgs. .................................................................................................277
Frullania nigricaulis (Reinw., Blume et Nees) Nees .....................................................................277
Frullania nigricaulis var. elongata Verd. .......................................................................................277
Frullania nisquallensis Sull...........................................................................................................293
Frullania nivimontana S.Hatt. ....................................................................................................277
Frullania nobilis Steph. ...............................................................................................................277
Frullania nobilis var. cochleata (Steph.) S.Hatt. ...........................................................................277
Frullania nodulosa (Reinw., Blume et Nees) Nees .......................................................................286
Frullania notarisii Steph. .............................................................................................................287
Frullania novocurvirostris S.Hatt. ................................................................................................277
Frullania novoguineensis Schifn. .................................................................................................286
Frullania oahuensis Hampe ex Gottsche, Lindenb. et Nees ..........................................................278
Frullania oakesiana Austin ..........................................................................................................284
Frullania oakesiana subsp. takayuensis (Steph.) R.M.Schust.........................................................284
Frullania obovata S.Hatt. ............................................................................................................278
Frullania obscura (Sw.) Mont. .....................................................................................................265
Frullania obscura var. spiniloba (Steph.) Hentschel et von Konrat ...............................................265
Frullania obscurifolia Mitt. ..........................................................................................................283
Frullania obtusangula Hentschel et von Konrat ...........................................................................278
Frullania ocanniensis Steph..........................................................................................................295
Frullania ocellata S.Hatt. et Kamim. ...........................................................................................271
Frullania odontostipa Spruce........................................................................................................267
Frullania okinawensis Kamim. .....................................................................................................278
Frullania onraedtii Vanden Berghen ...........................................................................................290
Frullania orbicularis Austin .........................................................................................................278
Frullania orientalis Sande Lac......................................................................................................278
Frullania orinocensis Spruce.........................................................................................................278
Frullania ornithocephala (Reinw., Blume et Nees) Nees ..............................................................278
Frullania ornithocephala var. major (Nees) Schifn.......................................................................278
Frullania ornithocephala var. pilosa Verd. ....................................................................................278
Frullania ornithocephala var. tuberculosa S.Hatt. ..........................................................................278
Frullania osculatiana De Not. .....................................................................................................288
Frullania osumiensis (S.Hatt.) S.Hatt...........................................................................................282
Frullania pachyderma S.Hatt. ......................................................................................................278
Frullania pallidevirens Steph. ......................................................................................................278
Frullania pallidula S.Hatt. ..........................................................................................................282
Frullania pancheri Gottsche ........................................................................................................292
Frullania papillata Steph. ............................................................................................................269
Frullania papillilobula S.Hatt. .....................................................................................................292
Frullania papuana Verd. .............................................................................................................290
Frullania papulosa Steph. ............................................................................................................287
Frullania paradoxa Lehm. et Lindenb. ........................................................................................288
Frullania paranensis Steph. ..........................................................................................................264
Frullania parhamii (R.M.Schust.) R.M.Schust. ex von Konrat, L.Söderstr. et A.Hagborg ...........291
Frullania pariharii S.Hatt. et haithong .....................................................................................278
Frullania parvifolia Steph. ...........................................................................................................278
Frullania parvistipula Steph. .......................................................................................................285
Frullania patagonica Steph. .........................................................................................................283
Frullania patula Mitt. .................................................................................................................285
World checklist of hornworts and liverworts
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567
Frullania pauciramea Steph. ........................................................................................................278
Frullania pauciramea var. pauciramella S.Hatt. et Piippo ............................................................278
Frullania paucirameoides S.Hatt. et Piippo ..................................................................................278
Frullania pearceana Steph. ..........................................................................................................288
Frullania pedicellata Steph. .........................................................................................................278
Frullania pentapleura Taylor .......................................................................................................278
Frullania peruviana Gottsche ......................................................................................................289
Frullania phalangilora Steph. .....................................................................................................289
Frullania physantha Mitt. ............................................................................................................278
Frullania pilibracteola S.Hatt. .....................................................................................................267
Frullania pilistipula Steph. ..........................................................................................................278
Frullania piptophylla S.Hatt. .......................................................................................................278
Frullania piptophylla var. minor S.Hatt........................................................................................278
Frullania piptophylloides S.Hatt. ..................................................................................................278
Frullania pittieri Steph. ...............................................................................................................288
Frullania plana Sull.....................................................................................................................285
Frullania planifolia Steph. ...........................................................................................................266
Frullania platycalyx Herzog .........................................................................................................295
Frullania plicata Hentschel et von Konrat ...................................................................................278
Frullania pluricarinata Gottsche .................................................................................................266
Frullania pocsantha haithong et S.Hatt. ....................................................................................279
Frullania polyptera Taylor ...........................................................................................................283
Frullania polyptera var. angustata (Mitt.) S.Hatt. .........................................................................283
Frullania polysticta Lindenb. .......................................................................................................293
Frullania ponapensis S.Hatt. et Koike ..........................................................................................272
Frullania pran-nathii M.Dey et D.K.Singh .................................................................................279
Frullania pringlei Fulford et Sharp ..............................................................................................279
Frullania probosciphora Taylor ....................................................................................................283
Frullania prominula S.Hatt. et Streimann ...................................................................................279
Frullania propaginea S.Hatt. et Streimann...................................................................................279
Frullania pseudericoides S.Hatt. ...................................................................................................279
Frullania pseudoalstonii Tsudo et J.Haseg. ...................................................................................293
Frullania pseudomeyeniana S.Hatt. ..............................................................................................291
Frullania pseudomonocera S.Hatt. ................................................................................................282
Frullania pseudoschensiana S.Hatt. ..............................................................................................279
Frullania pseudoschensiana var. darjeelingensis S.Hatt. ..................................................................279
Frullania ptychantha Mont. .........................................................................................................271
Frullania pulchella Herzog ..........................................................................................................283
Frullania pullei Verd. ..................................................................................................................279
Frullania pulogensis Steph............................................................................................................269
Frullania punctata Reimers .........................................................................................................293
Frullania purpurea Steph.............................................................................................................269
Frullania pusilla Mitt. .................................................................................................................279
Frullania pycnantha (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ..........................................279
Frullania pyricalycina Steph. .......................................................................................................279
Frullania queenslandica Steph. ....................................................................................................279
Frullania quillotensis (Nees et Mont.) Nees et Mont. ..................................................................295
Frullania ramuligera (Nees) Mont...............................................................................................267
Frullania recurvistipula S.Hatt.....................................................................................................279
Frullania relexistipula Sande Lac. ...............................................................................................279
Frullania relexistipula var. squarrosa S.Hatt. et Piippo ................................................................279
Frullania regularis Schifn. ..........................................................................................................286
568
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Frullania reicheana Steph. ...........................................................................................................295
Frullania reimersii Verd. .............................................................................................................270
Frullania remotidens S.Hatt. ........................................................................................................279
Frullania remotiloba Steph. .........................................................................................................286
Frullania repandistipula Sande Lac. .............................................................................................271
Frullania repandistipula subsp. queenslandica S.Hatt. ..................................................................271
Frullania repandistipula subsp. spinibractea S.Hatt. .....................................................................271
Frullania reptans Mitt. ................................................................................................................279
Frullania retusa Mitt. ..................................................................................................................279
Frullania retusa var. gymnantha S.Hatt. et haithong ..................................................................279
Frullania retusa var. hirsuta S.Hatt. et haithong ........................................................................279
Frullania rhystocolea Herzog ........................................................................................................279
Frullania rhytidantha S.Hatt. ......................................................................................................279
Frullania rigescens Spruce ............................................................................................................288
Frullania rigida Steph. ................................................................................................................279
Frullania ringens Spruce ..............................................................................................................265
Frullania rio-janeirensis (Raddi) Ångstr. ......................................................................................265
Frullania riparia Hampe .............................................................................................................279
Frullania rizalii Piippo et S.Hatt. ...............................................................................................280
Frullania rostellata Mitt...............................................................................................................280
Frullania rostrata (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ..............................................291
Frullania rubella Gottsche...........................................................................................................280
Frullania rubella var. elongata (Steph.) S.Hatt. ............................................................................280
Frullania rudoliana S.Hatt. ........................................................................................................287
Frullania rupicola Steph. .............................................................................................................280
Frullania sabahana S.Hatt. ..........................................................................................................285
Frullania sabaliana R.M.Schust. .................................................................................................285
Frullania sabanetica Gottsche ......................................................................................................295
Frullania sackawana Steph. .........................................................................................................285
Frullania saepisdentata S.Hatt. et Streimann ...............................................................................270
Frullania saipanensis S.Hatt. et Koike..........................................................................................280
Frullania sandvicensis Ångstr. ......................................................................................................266
Frullania sarawakensis S.Hatt. .....................................................................................................285
Frullania scalaris S.Hatt. .............................................................................................................291
Frullania scandens Mont. ............................................................................................................285
Frullania schaefer-verwimpii Yuzawa et S.Hatt. ...........................................................................293
Frullania schensiana C.Massal. ....................................................................................................280
Frullania schifneri Verd. .............................................................................................................270
Frullania schimperi Nees .............................................................................................................290
Frullania schimperi var. laciniata Vanden Berghen ......................................................................290
Frullania schusterana S.Hatt. .......................................................................................................270
Frullania schusteri S.Hatt. ...........................................................................................................280
Frullania scottiana S.Hatt............................................................................................................280
Frullania selwyniana Pearson.......................................................................................................293
Frullania semienana Gola ............................................................................................................295
Frullania semivillosa Lindenb. et Gottsche ..................................................................................285
Frullania sergiae Sim-Sim, Fontinha, Mues et Lion .....................................................................293
Frullania seriata Gottsche ...........................................................................................................282
Frullania seriatifolia Steph...........................................................................................................270
Frullania serrata Gottsche ...........................................................................................................270
Frullania serrata var. ceramensis S.Hatt. .......................................................................................270
Frullania serrata subsp. grolleana (S.Hatt.) S.Hatt. ......................................................................270
World checklist of hornworts and liverworts
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569
Frullania serrata var. hamatispina (S.Hatt.) S.Hatt. .....................................................................270
Frullania serrata var. pertenuis (Nees) Schifn. .............................................................................270
Frullania serrata subsp. spinistipula S.Hatt. .................................................................................270
Frullania serrifolia Steph. ............................................................................................................295
Frullania setacea S.Hatt. ..............................................................................................................270
Frullania setchellii Pearson ..........................................................................................................280
Frullania setigera Steph. ..............................................................................................................288
Frullania shanensis Svihla ............................................................................................................280
Frullania sharpantha Udar et Ad.Kumar .....................................................................................280
Frullania sharpii S.Hatt...............................................................................................................280
Frullania sheana S.Hatt. ..............................................................................................................270
Frullania simmondsii Steph. ........................................................................................................270
Frullania sinensis Steph. ..............................................................................................................280
Frullania sinosphaerantha S.Hatt. et P.J.Lin ................................................................................280
Frullania sinskeana J.J.Engel et B.C.Tan .....................................................................................283
Frullania sinuata Sande Lac. .......................................................................................................270
Frullania socotrana Mitt. .............................................................................................................283
Frullania solanderiana Colenso ...................................................................................................283
Frullania speciosa Herzog ............................................................................................................289
Frullania spegazzinii M.E.Reiner.................................................................................................264
Frullania sphaerantha S.Hatt. ......................................................................................................280
Frullania sphaerocephala Spruce ..................................................................................................265
Frullania sphaerolobulata S.H.Lin ...............................................................................................280
Frullania spinifera Taylor ............................................................................................................280
Frullania spinigastria S.Hatt. .......................................................................................................280
Frullania spiniplica S.Hatt...........................................................................................................280
Frullania spinistipula Steph. ........................................................................................................282
Frullania spongiosa Steph. ...........................................................................................................280
Frullania squamuligera Spruce.....................................................................................................280
Frullania squarrosula (Hook.f. et Taylor) Gottsche, Lindenb. et Nees .........................................280
Frullania standaertii Steph. .........................................................................................................266
Frullania steereana S.Hatt. ..........................................................................................................270
Frullania stenostipa Spruce ..........................................................................................................280
Frullania stipatiloba Steph...........................................................................................................270
Frullania streimannii S.Hatt........................................................................................................282
Frullania stylifera (R.M.Schust.) R.M.Schust. .............................................................................285
Frullania subarctica Vilnet, Borovich. et Bakalin .........................................................................293
Frullania subcaduca S.Hatt..........................................................................................................280
Frullania subclavata Steph...........................................................................................................280
Frullania subdentata Steph. .........................................................................................................270
Frullania subdentata var. latistipula (S.Hatt.) S.Hatt. ..................................................................270
Frullania subdilatata C.Massal. ...................................................................................................285
Frullania subincumbens S.Hatt. ...................................................................................................283
Frullania sublignosa Steph. ..........................................................................................................292
Frullania submultilacera S.Hatt. et Koike ....................................................................................270
Frullania subnigricaulis S.Hatt. ...................................................................................................280
Frullania subnigricaulis var. subtruncata S.Hatt. ..........................................................................280
Frullania subocellata S.Hatt. ........................................................................................................270
Frullania subpedicellata S.Hatt. ...................................................................................................280
Frullania subpilibracteola S.Hatt. ................................................................................................267
Frullania subpyricalycina Herzog .................................................................................................295
Frullania subsquarrosa S.Hatt. .....................................................................................................280
570
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Frullania subtilissima (Nees ex Mont.) Lindenb. .........................................................................267
Frullania subtropica Steph. ..........................................................................................................283
Frullania subtruncata Steph.........................................................................................................295
Frullania subvalida S.Hatt. et haithong.....................................................................................281
Frullania supradecomposita (Lehm. et Lindenb.) Lehm. et Lindenb. ............................................289
Frullania svihlana S.Hatt. ...........................................................................................................281
Frullania tagawana (S.Hatt. et haithong) S.Hatt.......................................................................287
Frullania taiheizana Horik. .........................................................................................................281
Frullania tamarisci (L.) Dumort. .................................................................................................293
Frullania tamsuina Steph. ...........................................................................................................281
Frullania taradakensis Steph. .......................................................................................................281
Frullania taxodiocola R.M.Schust. ...............................................................................................270
Frullania tenerifae (F.Weber) Nees .............................................................................................293
Frullania tenuirostris Steph. .........................................................................................................281
Frullania ternatensis Gottsche......................................................................................................270
Frullania ternatensis var. non-appendiculata S.Hatt. .....................................................................270
Frullania tetraptera Nees et Mont. ..............................................................................................266
Frullania thiersiae S.Hatt. ...........................................................................................................287
Frullania tixieri S.Hatt. ...............................................................................................................287
Frullania tjibodensis S.Hatt. et haithong....................................................................................283
Frullania togashiana S.Hatt. ........................................................................................................281
Frullania toropuku von Konrat, de Lange et Larraín ....................................................................290
Frullania trianae Gottsche ..........................................................................................................289
Frullania tricarinata Sande Lac. ..................................................................................................272
Frullania trichodes Mitt. ..............................................................................................................271
Frullania trigona L.Clark, Jovet-Ast et Frye .................................................................................293
Frullania trinervis (Lehm.) Drège ................................................................................................266
Frullania triquetra Lindenb. et Gottsche .....................................................................................281
Frullania trollii Herzog ...............................................................................................................289
Frullania truncatistyla von Konrat, Hentschel, Heinrichs et Braggins ..........................................291
Frullania tubercularis S.Hatt. et P.J.Lin.......................................................................................281
Frullania tunguraguana L.Clark et Frye.......................................................................................265
Frullania turfosa Lindenb. et Gottsche ........................................................................................295
Frullania tuyamae S.Hatt. et haithong ......................................................................................284
Frullania udarii V.Nath et Ajit P.Singh ......................................................................................295
Frullania uleana Steph. ...............................................................................................................289
Frullania umbonata Mitt.............................................................................................................285
Frullania usambarana Schifn......................................................................................................272
Frullania usambarana var. reducta Vanden Berghen ....................................................................272
Frullania usamiensis Steph...........................................................................................................281
Frullania utriculata Steph. ..........................................................................................................285
Frullania vaga Mitt. ....................................................................................................................271
Frullania vaginata (Sw.) Nees .....................................................................................................271
Frullania vaginata var. nigricans (Gottsche, Lindenb. et Nees) Schifn. .......................................271
Frullania valdiviensis J.B.Jack et Steph. .......................................................................................281
Frullania valida Steph. ................................................................................................................281
Frullania valparaisiana Lehm. .....................................................................................................295
Frullania vandenberghenii Pócs ...................................................................................................267
Frullania van-zantenii Kamim. et S.Hatt. ...................................................................................271
Frullania variegata Steph. ...........................................................................................................281
Frullania venusta S.Hatt..............................................................................................................271
Frullania verdoorniana S.Hatt. ....................................................................................................271
World checklist of hornworts and liverworts
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571
Frullania victoriensis Steph. .........................................................................................................281
Frullania virginica Lehm. ............................................................................................................285
Frullania vitalii Yuzawa et S.Hatt. ..............................................................................................271
Frullania vittata S.Hatt. ..............................................................................................................271
Frullania vittiana S.Hatt. ............................................................................................................281
Frullania vivipara Pócs ...............................................................................................................271
Frullania wagneri Steph. .............................................................................................................295
Frullania wairua von Konrat et Braggins ....................................................................................271
Frullania wangii S.Hatt. et P.J.Lin ..............................................................................................281
Frullania warnckeana S.Hatt. ......................................................................................................271
Frullania warnckeana var. dentosa S.Hatt. ...................................................................................271
Frullania weberbaueri Steph. .......................................................................................................289
Frullania winteri Steph. ..............................................................................................................266
Frullania winteri var. vanderhammenii (Haarbrink) Yuzawa ........................................................266
Frullania yuennanensis Steph. ......................................................................................................281
Frullania yuennanensis var. siamensis (N.Kitag., haithong et S.Hatt.) S.Hatt. et P.J.Lin ............281
Frullania yuzawana S.Hatt..........................................................................................................281
Frullania zangii S.Hatt. et P.J.Lin ...............................................................................................281
Frullania zennoskeana S.Hatt. .....................................................................................................281
Frullanoides bahamensis (A.Evans) van Slageren ..........................................................................405
Frullanoides corticalis (Lehm. et Lindenb.) van Slageren ..............................................................405
Frullanoides densifolia Raddi .......................................................................................................405
Frullanoides densifolia subsp. grandidentata (L.Clark) van Slageren .............................................405
Frullanoides laciniatilora (Loitl.) van Slageren ............................................................................405
Frullanoides liebmaniana (Lindenb. et Gottsche) van Slageren ....................................................405
Frullanoides mexicana van Slageren .............................................................................................405
Frullanoides tristis (Steph.) van Slageren ......................................................................................405
Fulfordianthus evansii (Fulford) Gradst. ......................................................................................405
Fulfordianthus pterobryoides (Spruce) Gradst. ..............................................................................405
Fuscocephaloziopsis ainis (Lindb. ex Steph.) Váňa et L.Söderstr. ...................................................59
Fuscocephaloziopsis africana (Váňa) Váňa et L.Söderstr. .................................................................59
Fuscocephaloziopsis albescens (Hook.) Váňa et L.Söderstr. ..............................................................59
Fuscocephaloziopsis albescens var. islandica (Nees) Váňa et L.Söderstr. ............................................59
Fuscocephaloziopsis baldwinii (C.M.Cooke) Váňa et L.Söderstr. ....................................................59
Fuscocephaloziopsis biloba (Herzog) Fulford ...................................................................................59
Fuscocephaloziopsis catenulata (Huebener) Váňa et L.Söderstr........................................................59
Fuscocephaloziopsis catenulata subsp. nipponica (S.Hatt.) Váňa et L.Söderstr..................................59
Fuscocephaloziopsis connivens (Dicks.) Váňa et L.Söderstr. .............................................................60
Fuscocephaloziopsis connivens subsp. issa (Steph.) Váňa et L.Söderstr. ............................................60
Fuscocephaloziopsis connivens subsp. sandwicensis (Mont.) Váňa et L.Söderstr. ...............................60
Fuscocephaloziopsis crassifolia (Lindenb. et Gottsche) Váňa et L.Söderstr. ......................................60
Fuscocephaloziopsis gollanii (Steph.) Váňa et L.Söderstr. ................................................................60
Fuscocephaloziopsis leucantha (Spruce) Váňa et L.Söderstr..............................................................60
Fuscocephaloziopsis loitlesbergeri (Schifn.) Váňa et L.Söderstr. .......................................................60
Fuscocephaloziopsis lunulifolia (Dumort.) Váňa et L.Söderstr. ........................................................60
Fuscocephaloziopsis macrostachya (Kaal.) Váňa et L.Söderstr. ..........................................................60
Fuscocephaloziopsis macrostachya subsp. australis (R.M.Schust.) Váňa et L.Söderstr........................60
Fuscocephaloziopsis macrostachya subsp. macrostachya var. spinilora (Schifn.) Váňa et L.Söderstr. .60
Fuscocephaloziopsis monticola (J.D.Godfrey) Váňa et L.Söderstr.....................................................60
Fuscocephaloziopsis pachycaulis (R.M.Schust.) Váňa et L.Söderstr. .................................................60
Fuscocephaloziopsis pleniceps (Austin) Váňa et L.Söderstr. ..............................................................61
Fuscocephaloziopsis pleniceps var. caroliniana (R.M.Schust.) Váňa et L.Söderstr..............................61
572
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Fuscocephaloziopsis pulvinata (Steph.) Fulford ...............................................................................61
Fuscocephaloziopsis schusteri (Sushil K.Singh et D.K.Singh) Váňa ..................................................61
Fuscocephaloziopsis subintegra Gradst. et Váňa ...............................................................................61
Fuscocephaloziopsis zoopsioides (Horik.) Váňa et L.Söderstr. ...........................................................61
Gackstroemia alpina R.M.Schust. ................................................................................................416
Gackstroemia hariotiana (Besch. et C.Massal.) Grolle ..................................................................415
Gackstroemia ljungneri (Herzog) Grolle .......................................................................................415
Gackstroemia magellanica (Lam.) Trevis. .....................................................................................415
Gackstroemia novae-zelandiae R.M.Schust. et J.J.Engel ...............................................................415
Gackstroemia patagonica (Steph.) Grolle ......................................................................................416
Gackstroemia schwabei (Herzog) Grolle .......................................................................................416
Gackstroemia weindorferi (Herzog) Grolle ...................................................................................415
Geocalyx caledonicus Steph...........................................................................................................109
Geocalyx graveolens (Schrad.) Nees ..............................................................................................109
Geocalyx lancistipulus (Steph.) S.Hatt. .........................................................................................109
Geocalyx orientalis Besch. et Spruce .............................................................................................109
Geothallus tuberosus Campb. .......................................................................................................506
Gerhildiella rossneriana Grolle .......................................................................................................78
Goebeliella cornigera (Mitt.) Steph...............................................................................................415
Goebelobryum grossitextum (Steph.) Grolle.....................................................................................97
Goebelobryum unguiculatum (Hook.f. et Taylor) Grolle ................................................................97
Goebelobryum vermiculare J.J.Engel et Glenny...............................................................................97
Gongylanthus dusenii Steph. ........................................................................................................135
Gongylanthus ericetorum (Raddi) Nees.........................................................................................135
Gongylanthus granatensis (Gottsche) Steph. .................................................................................135
Gongylanthus himalayensis Grolle ................................................................................................135
Gongylanthus javanicus Grolle .....................................................................................................135
Gongylanthus liebmanianus (Lindenb. et Gottsche) Steph............................................................135
Gongylanthus limbatus (Herzog) Grolle et Váňa ..........................................................................135
Gongylanthus muelleri (Gottsche) Steph. .....................................................................................135
Gongylanthus oniscoides (Spruce) Steph........................................................................................135
Gongylanthus richardsii E.W.Jones ..............................................................................................135
Gottschelia maxima (Steph.) Grolle ...............................................................................................74
Gottschelia schizopleura (Spruce) Grolle .........................................................................................74
Greeneothallus gemmiparus Hässel ...............................................................................................471
Grollea antheliopsis R.M.Schust. ..................................................................................................138
Gymnocolea borealis (Frisvoll et Moen) R.M.Schust. .....................................................................52
Gymnocolea fascinifera Potemkin ...................................................................................................52
Gymnocolea inlata (Huds.) Dumort. ............................................................................................52
Gymnocolea inlata subsp. acutiloba (Schifn.) R.M.Schust. et Damsh. ex L.Söderstr. et Váňa .......52
Gymnocoleopsis capensis (S.W.Arnell) R.M.Schust. ........................................................................74
Gymnocoleopsis cylindriformis (Mitt.) R.M.Schust. .........................................................................74
Gymnomitrion adustum Nees.......................................................................................................110
Gymnomitrion africanum (Steph.) Horik. ....................................................................................110
Gymnomitrion alpinum (Gottsche ex Husn.) Schifn. ..................................................................110
Gymnomitrion asperulatum R.M.Schust. .....................................................................................110
Gymnomitrion atroilum Váňa .....................................................................................................110
Gymnomitrion bolivianum (Steph.) Váňa ....................................................................................110
Gymnomitrion brevissimum (Dumort.) Warnst. ...........................................................................110
Gymnomitrion commutatum (Limpr.) Schifn. .............................................................................111
Gymnomitrion concinnatum (Lightf.) Corda ................................................................................111
Gymnomitrion corallioides Nees ...................................................................................................111
World checklist of hornworts and liverworts
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573
Gymnomitrion crenatilobum Grolle..............................................................................................111
Gymnomitrion crenulatum Gottsche ex Carrington......................................................................111
Gymnomitrion crystallocaulon (Grolle) Váňa, Crand.-Stotl. et Stotler ..........................................111
Gymnomitrion incompletum (Gottsche) R.M.Schust. ex Váňa .....................................................111
Gymnomitrion laceratum (Steph.) Horik. ....................................................................................111
Gymnomitrion miniatum Lindenb. et Gottsche ...........................................................................111
Gymnomitrion minutulum (Hässel) Váňa ....................................................................................111
Gymnomitrion moralesae Váňa.....................................................................................................111
Gymnomitrion mucronulatum (N.Kitag.) N.Kitag. ......................................................................111
Gymnomitrion mucrophorum R.M.Schust....................................................................................111
Gymnomitrion nigrum (Grolle et Váňa) Váňa ..............................................................................111
Gymnomitrion noguchianum S.Hatt. ...........................................................................................111
Gymnomitrion obtusilobum N.Kitag. ...........................................................................................111
Gymnomitrion obtusum Lindb. ....................................................................................................111
Gymnomitrion paciicum Grolle ...................................................................................................111
Gymnomitrion revolutum (Nees) H.Philib. ..................................................................................111
Gymnomitrion revolutum subsp. novoguineanensis (R.M.Schust.) Váňa, Crand.-Stotl. et Stotler ..112
Gymnomitrion rubidum (Mitt.) Váňa, Crand.-Stotl. et Stotler ....................................................112
Gymnomitrion setaceum Grolle et Váňa .......................................................................................112
Gymnomitrion sinense Müll.Frib. .................................................................................................112
Gymnomitrion strictum (Berggr.) R.M.Schust..............................................................................112
Gymnomitrion strictum var. inaequale R.M.Schust.......................................................................112
Gymnomitrion subintegrum (S.W.Arnell) Váňa ............................................................................112
Gymnomitrion truncatoapiculatum Herzog ..................................................................................112
Gymnomitrion verrucosum W.E.Nicholson ..................................................................................112
Gyrothyra underwoodiana M.Howe .............................................................................................115
Haesselia acuminata Gradst. ..........................................................................................................64
Haesselia roraimensis Grolle et Gradst............................................................................................64
Hamatostrepta concinna Váňa et D.G.Long ...................................................................................52
Haplolejeunea cucullata (Steph.) Grolle .......................................................................................348
Haplolejeunea sticta Grolle ..........................................................................................................348
Haplomitrium blumei (Nees) R.M.Schust. ....................................................................................41
Haplomitrium gibbsiae (Steph.) R.M.Schust. .................................................................................42
Haplomitrium hookeri (Lyell ex Sm.) Nees ....................................................................................42
Haplomitrium hookeri var. minutum (E.O.Campb.) Barthol.-Began ..............................................42
Haplomitrium intermedium Berrie .................................................................................................42
Haplomitrium mnioides (Lindb.) R.M.Schust. ...............................................................................41
Haplomitrium monoicum J.J.Engel ................................................................................................42
Haplomitrium ovalifolium R.M.Schust. .........................................................................................42
Harpalejeunea acuta S.Winkl. .....................................................................................................362
Harpalejeunea ancistrodes (Spruce) Schifn. .................................................................................362
Harpalejeunea buenaventurae Herzog ..........................................................................................362
Harpalejeunea cinchonae (Nees) Schifn.......................................................................................362
Harpalejeunea cinchonae var. strigulosa Herzog ............................................................................363
Harpalejeunea decurvicuspis (Besch. et C.Massal.) P.Syd..............................................................363
Harpalejeunea emarginata Jovet-Ast ............................................................................................363
Harpalejeunea exocellata Herzog ..................................................................................................363
Harpalejeunea grandis Grolle et M.E.Reiner ................................................................................362
Harpalejeunea grandistipula R.M.Schust. ....................................................................................363
Harpalejeunea grossearmata Steph. ...............................................................................................364
Harpalejeunea harpaphylla (Herzog) Bischl. ................................................................................363
Harpalejeunea herzogii Jovet-Ast .................................................................................................363
574
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Harpalejeunea longibracteata (Spruce) Steph................................................................................363
Harpalejeunea marginalis (Hook.f. et Taylor) Steph. ...................................................................363
Harpalejeunea molleri (Steph.) Grolle ..........................................................................................363
Harpalejeunea molleri subsp. integra (R.M.Schust.) Damsh. ........................................................363
Harpalejeunea oxyphylla (Nees et Mont.) Steph. ..........................................................................363
Harpalejeunea parasitica (Hook.f. et Taylor) Steph. ....................................................................363
Harpalejeunea pinaundensis Grolle ..............................................................................................363
Harpalejeunea relexula A.Evans ..................................................................................................363
Harpalejeunea renneri Herzog .....................................................................................................364
Harpalejeunea scabra Gradst. et Schäf.-Verw. ..............................................................................363
Harpalejeunea schifneri S.W.Arnell .............................................................................................363
Harpalejeunea solitaria (Gottsche) Steph. ....................................................................................363
Harpalejeunea spruceana Steph. ...................................................................................................364
Harpalejeunea stricta (Lindenb. et Gottsche) Steph. ....................................................................364
Harpalejeunea subacuta A.Evans ..................................................................................................364
Harpalejeunea tenuicuspis (Spruce) Schifn. .................................................................................364
Harpalejeunea tridens (Besch. et Spruce) Steph............................................................................364
Harpalejeunea uncinata Steph. ....................................................................................................364
Harpalejeunea uncinata var. setulosa Herzog ................................................................................364
Harpalejeunea vitrea Herzog .......................................................................................................364
Harpanthus drummondii (Taylor) Grolle .....................................................................................116
Harpanthus lotovianus (Nees) Nees ............................................................................................116
Harpanthus scutatus (F.Weber et D.Mohr) Spruce ......................................................................116
Hattoria yakushimensis (Horik.) R.M.Schust. ................................................................................53
Hattorianthus erimonus (Steph.) R.M.Schust. et Inoue ................................................................469
Hattoriolejeunea akiyamae Mizut. ................................................................................................364
Hepatostolonophora abnormis (Besch. et C.Massal.) J.J.Engel et R.M.Schust. ...............................196
Hepatostolonophora conica (Steph.) Hässel ...................................................................................196
Hepatostolonophora paucistipula (Rodway) J.J.Engel ....................................................................196
Hepatostolonophora rotata (Hook.f. et Taylor) J.J.Engel...............................................................196
Hepatostolonophora rotata var. perssonii (R.M.Schust.) J.J.Engel ..................................................197
Herbertus aduncus (Dicks.) Gray .................................................................................................138
Herbertus arcticus (Inoue et Steere) Schljakov ..............................................................................139
Herbertus armitanus (Steph.) H.A.Mill........................................................................................139
Herbertus asparus Tixier ..............................................................................................................139
Herbertus bivittatus Spruce ..........................................................................................................139
Herbertus borealis Crundw. .........................................................................................................139
Herbertus buchii Juslén ................................................................................................................139
Herbertus ceylanicus (Steph.) Abeyw. ...........................................................................................139
Herbertus circinatus (Steph.) H.A.Mill. .......................................................................................139
Herbertus delavayi (Steph.) Steph. ...............................................................................................139
Herbertus dicranus (Gottsche, Lindenb. et Nees) Trevis. .............................................................139
Herbertus durandii (Steph.) Herzog ............................................................................................139
Herbertus evittatus (Steph.) H.A.Mill. .........................................................................................139
Herbertus gaochienii X.Fu ............................................................................................................139
Herbertus gracilis (Mont.) Steph. .................................................................................................139
Herbertus guangdongii P.J.Lin ex Piippo ......................................................................................139
Herbertus hawaiiensis H.A.Mill. ..................................................................................................139
Herbertus helleri (Steph.) W.E.Nicholson....................................................................................139
Herbertus herpocladioides E.B.Scott et H.A.Mill. .........................................................................139
Herbertus hutchinsiae (Gottsche et Rabenh.) A.Evans ..................................................................140
Herbertus juniperoideus (Sw.) Grolle ............................................................................................140
World checklist of hornworts and liverworts
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575
Herbertus juniperoideus subsp. acanthelius (Spruce) K.Feldberg et Heinrichs ...............................140
Herbertus juniperoideus subsp. pensilis (Taylor) K.Feldberg et Heinrichs......................................140
Herbertus kurzii (Steph.) R.S.Chopra ..........................................................................................140
Herbertus leratii (Steph.) H.A.Mill. .............................................................................................140
Herbertus lonchobasis H.A.Mill. ...................................................................................................140
Herbertus longiissus Steph. ..........................................................................................................140
Herbertus longispinus J.B.Jack et Steph. .......................................................................................140
Herbertus mauritianus N.G.Hodgetts ..........................................................................................140
Herbertus norenus D.G.Long, D.Bell et H.H.Blom .....................................................................140
Herbertus oldieldianus (Steph.) Rodway ......................................................................................140
Herbertus pilifer Schifn...............................................................................................................140
Herbertus pocsii N.G.Hodgetts ....................................................................................................140
Herbertus pumilus Steph. .............................................................................................................140
Herbertus ramosus (Steph.) H.A.Mill. ..........................................................................................140
Herbertus runcinatus (Taylor) Trevis. ..........................................................................................140
Herbertus sendtneri (Nees) Lindb.................................................................................................140
Herbertus spicatus N.G.Hodgetts .................................................................................................140
Herbertus stramineus (Dumort.) Trevis. .......................................................................................141
Herbertus streimannii M.L.So ......................................................................................................141
Herbertus subrotundatus X.Fu et Y.J.Yi ........................................................................................141
Herbertus tenuis A.Evans .............................................................................................................141
Herbertus udarii D.Kumar et N.Manocha ...................................................................................141
Herzogianthus sanguineus R.M.Schust. ........................................................................................434
Herzogianthus vaginatus (Herzog) R.M.Schust. ...........................................................................434
Herzogiaria teres (Steph.) Fulford ex Hässel.................................................................................254
Herzogobryum atrocapillum (Hook.f. et Taylor) Grolle ..................................................................74
Herzogobryum iliforme R.M.Schust. .............................................................................................74
Herzogobryum molle Grolle ...........................................................................................................74
Herzogobryum vermiculare (Schifn.) Grolle...................................................................................74
Heterogemma capitata (Hook.) Konstant. et Vilnet .......................................................................79
Heterogemma laxa (Lindb.) Konstant. et Vilnet .............................................................................80
Heterogemma patagonica (Herzog et Grolle) L.Söderstr. et Váňa ...................................................80
Heteroscyphus acutangulus (Schifn.) Schifn. ...............................................................................197
Heteroscyphus allodontus (Hook.f. et Taylor) J.J.Engel et R.M.Schust. .........................................197
Heteroscyphus amboinensis (Schifn.) Schifn. ...............................................................................197
Heteroscyphus ammophilus (Colenso) R.M.Schust. .......................................................................197
Heteroscyphus ammophilus var. obtusifolius J.J.Engel et G.L.Merr. ................................................197
Heteroscyphus argutus (Reinw., Blume et Nees) Schifn................................................................197
Heteroscyphus argutus var. brevidens (Schifn.) Herzog et Nog......................................................197
Heteroscyphus aselliformis (Reinw., Blume et Nees) Schifn. .........................................................197
Heteroscyphus assurgentifolius J.J.Engel .........................................................................................197
Heteroscyphus baduinus (Nees) Schifn.........................................................................................197
Heteroscyphus balnetii (Herzog) Grolle ........................................................................................197
Heteroscyphus billardierei (Schwägr.) Schifn................................................................................197
Heteroscyphus billardierei var. clasmatocoleoides (J.J.Engel et G.L.Merr.) J.J.Engel ........................197
Heteroscyphus brassii (Grolle) Grolle ............................................................................................198
Heteroscyphus caesius (Schifn.) Schifn. .......................................................................................198
Heteroscyphus caledonicus (Steph.) Schifn....................................................................................198
Heteroscyphus chlorophyllus (Hook.f. et Taylor) Schifn................................................................198
Heteroscyphus ciliatus (Steph.) Schifn..........................................................................................198
Heteroscyphus coalitus (Hook.) Schifn. ........................................................................................198
Heteroscyphus coalitus var. simplicifolius J.J.Engel .........................................................................198
576
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Heteroscyphus combinatus (Nees) Schifn......................................................................................198
Heteroscyphus conjugatus (Mitt.) J.J.Engel et R.M.Schust.............................................................198
Heteroscyphus contortuplicatus (Nees et Mont.) Grolle .................................................................198
Heteroscyphus cuneistipulus (Steph.) Schifn. ................................................................................198
Heteroscyphus darjeelingensis A.Srivast. et S.C.Srivast. ..................................................................198
Heteroscyphus deceptifrons J.J.Engel ..............................................................................................198
Heteroscyphus dentammophilus J.J.Engel et G.L.Merr...................................................................198
Heteroscyphus denticulatus (Mitt.) Schifn. ...................................................................................199
Heteroscyphus deplanchei (Steph.) Schifn. ...................................................................................199
Heteroscyphus diestianus (Sande Lac.) Piippo ...............................................................................199
Heteroscyphus divergenticiliatus (Steph.) Fulford ..........................................................................199
Heteroscyphus dubius (Gottsche) Schifn. .....................................................................................199
Heteroscyphus echinellus (Lindenb. et Gottsche) J.J.Engel et Xiao L.He .......................................199
Heteroscyphus echinellus var. hyalinus J.J.Engel .............................................................................199
Heteroscyphus elliottii (Steph.) Pagán ...........................................................................................199
Heteroscyphus elliottii var. portoricensis Fulford.............................................................................199
Heteroscyphus falcifolius (Steph.) Schifn. .....................................................................................199
Heteroscyphus issistipus (Hook.f. et Taylor) Schifn. ....................................................................199
Heteroscyphus issistipus var. multispinus (E.A.Hodgs. et Allison) J.J.Engel ...................................199
Heteroscyphus issistipus var. repandus J.J.Engel.............................................................................199
Heteroscyphus laccidus (Mitt.) A.Srivast. et S.C.Srivast. ...............................................................199
Heteroscyphus leischeri (Steph.) D.G.Long et Rubas. ...................................................................199
Heteroscyphus fragilicilius (Schifn.) Schifn. ................................................................................200
Heteroscyphus furcistipulus (E.A.Hodgs.) J.J.Engel et R.M.Schust. ...............................................200
Heteroscyphus giganteus (Steph.) Hürl. .........................................................................................200
Heteroscyphus graefei (J.B.Jack et Steph.) Grolle..........................................................................200
Heteroscyphus grandilorus (Steph.) Hürl. .....................................................................................200
Heteroscyphus grandistipus (Steph.) Schifn. .................................................................................200
Heteroscyphus gunnianus (Mitt.) J.J.Engel et R.M.Schust.............................................................200
Heteroscyphus hastatus (E.A.Hodgs.) J.J.Engel et R.M.Schust. .....................................................200
Heteroscyphus hebridensis (Steph.) Schifn. ...................................................................................200
Heteroscyphus heterophyllus (Steph.) J.J.Engel et R.M.Schust. ......................................................200
Heteroscyphus hyalinus (Steph.) A.Srivast. et S.C.Srivast. .............................................................200
Heteroscyphus integrifolius (Lehm. et Lindenb.) Fulford ...............................................................200
Heteroscyphus iwatsukii (S.Hatt.) Piippo ......................................................................................200
Heteroscyphus jackii (Steph.) Schifn. ...........................................................................................200
Heteroscyphus knightii (Steph.) Grolle..........................................................................................201
Heteroscyphus levieri (Steph.) Schifn. ..........................................................................................201
Heteroscyphus limosus (Carrington et Pearson) Schifn. ................................................................201
Heteroscyphus lingulatus (Colenso) J.J.Engel et R.M.Schust. ........................................................201
Heteroscyphus lophocoleoides S.Hatt. .............................................................................................201
Heteroscyphus lyallii (Mitt.) R.M.Schust. .....................................................................................201
Heteroscyphus magellanicus (Steph.) J.J.Engel et R.M.Schust. ......................................................201
Heteroscyphus mamillatus Piippo..................................................................................................201
Heteroscyphus marginatus (Steph.) Fulford ...................................................................................201
Heteroscyphus menziesii (Mitt.) J.J.Engel......................................................................................201
Heteroscyphus merapiensis (Steph.) Piippo ....................................................................................201
Heteroscyphus miradorensis (Steph.) Schifn. ................................................................................201
Heteroscyphus mononuculus J.J.Engel ...........................................................................................201
Heteroscyphus mononuculus var. ammophilopsis J.J.Engel ..............................................................201
Heteroscyphus mononuculus var. bilobus J.J.Engel .........................................................................201
Heteroscyphus montagnei (Steph.) Fulford ....................................................................................201
World checklist of hornworts and liverworts
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577
Heteroscyphus multiidus (Steph.) J.J.Engel et R.M.Schust. ..........................................................202
Heteroscyphus multiidus var. subintegerrimus J.J.Engel .................................................................202
Heteroscyphus nadeaudii (Steph.) Schifn. ....................................................................................202
Heteroscyphus orbiculatus A.Srivast. et S.C.Srivast. .......................................................................202
Heteroscyphus palniensis A.Srivast. et S.C.Srivast. .........................................................................202
Heteroscyphus pandei S.C.Srivast. et A.Srivast. .............................................................................202
Heteroscyphus parallelifolius J.J.Engel ...........................................................................................202
Heteroscyphus parvulus (Schifn.) Schifn. ....................................................................................202
Heteroscyphus parvus A.Srivast. et S.C.Srivast. .............................................................................202
Heteroscyphus perfoliatus (Mont.) Schifn. ....................................................................................202
Heteroscyphus pertusus (Lehm.) Fulford........................................................................................202
Heteroscyphus planiusculus (Hook.f. et Taylor) J.J.Engel ..............................................................202
Heteroscyphus planus (Mitt.) Schifn. ...........................................................................................202
Heteroscyphus polyblepharis (Spruce) Schifn. ...............................................................................202
Heteroscyphus polychaetus (Spruce) Hentschel et Heinrichs ..........................................................202
Heteroscyphus polycladus (Hook.f. et Lév.) R.M.Schust. ...............................................................202
Heteroscyphus rectangulatus (Herzog) Piippo ................................................................................203
Heteroscyphus saccogynoides Herzog ..............................................................................................203
Heteroscyphus sarawaketanus Piippo .............................................................................................203
Heteroscyphus sinuosus (Hook.) Schifn. .......................................................................................203
Heteroscyphus spectabilis (Steph.) Schifn. ....................................................................................203
Heteroscyphus spinifer C.Gao, T.Cao et Y.H.Wu .........................................................................203
Heteroscyphus splendens (Lehm. et Lindenb.) Grolle.....................................................................203
Heteroscyphus splendidus (E.A.Hodgs.) J.J.Engel et R.M.Schust. ..................................................203
Heteroscyphus stolonifer J.J.Engel..................................................................................................203
Heteroscyphus succulentus (Gottsche) Schifn. ...............................................................................203
Heteroscyphus supinus (Hook.f. et Taylor) R.M.Schust. ...............................................................203
Heteroscyphus tener (Steph.) Schifn. ............................................................................................203
Heteroscyphus thraustus (Spruce) Fulford .....................................................................................203
Heteroscyphus timppae Piippo ......................................................................................................203
Heteroscyphus triacanthus (Hook.f. et Lév.) Schifn. .....................................................................203
Heteroscyphus triacanthus var. magnistipulatus J.J.Engel ...............................................................203
Heteroscyphus tridentatus (Sande Lac.) Grolle ..............................................................................203
Heteroscyphus turgidus (Schifn.) Schifn. .....................................................................................204
Heteroscyphus valdiviensis (Mont.) Schifn. ..................................................................................204
Heteroscyphus varians (Steph.) J.J.Engel .......................................................................................204
Heteroscyphus wettsteinii (Schifn.) Schifn. ..................................................................................204
Heteroscyphus zollingeri (Gottsche) Schifn. .................................................................................204
Hyalolepidozia bicuspidata (C.Massal.) S.W.Arnell ex Grolle .......................................................183
Hygrobiella intermedia Bakalin et Vilnet......................................................................................116
Hygrobiella laxifolia (Hook.) Spruce............................................................................................116
Hygrobiella squamosa Bakalin et Vilnet ........................................................................................116
Hygrolejeunea cubensis Steph. ......................................................................................................509
Hygrolejeunea harpaphylla Steph..................................................................................................509
Hygrolejeunea paciica Steph. .......................................................................................................509
Hygrolejeunea parvicalycina Steph................................................................................................509
Hygrolejeunea parvistipula Steph..................................................................................................509
Hygrolejeunea patellirostris Steph. ................................................................................................509
Hygrolejeunea staudtiana Steph....................................................................................................509
Hygrolembidium acrocladum (Berggr.) R.M.Schust......................................................................164
Hygrolembidium andinum (Herzog) R.M.Schust. ........................................................................164
Hygrolembidium australe (Steph.) Grolle .....................................................................................164
578
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Hygrolembidium boschianum (Sande Lac.) R.M.Schust................................................................164
Hygrolembidium isophyllum R.M.Schust......................................................................................164
Hygrolembidium rigidum R.M.Schust. et J.J.Engel ......................................................................164
Hygrolembidium triquetrum J.J.Engel et R.M.Schust. ..................................................................164
Hygrolembidium ventrosum (Mitt.) Grolle ...................................................................................164
Hymenophyton labellatum (Labill.) Dumort. ex Trevis. ...............................................................468
Hymenophyton leptopodum (Hook.f. et Taylor) A.Evans ..............................................................468
Hymenophyton pedicellatum Steph. ..............................................................................................469
Hypenantron brachypus Steph. ex Lamothe ..................................................................................509
Hypenantron brasiliense Steph. ex Lamothe .................................................................................509
Isolembidium anomalum (Rodway) Grolle ...................................................................................164
Isolembidium anomalum var. cucullatum (E.A.Hodgs.) J.J.Engel et R.M.Schust. .........................164
Isopaches alboviridis (R.M.Schust.) Schljakov ................................................................................53
Isopaches bicrenatus (Schmidel ex Hofm.) H.Buch........................................................................53
Isopaches decolorans (Limpr.) H.Buch ............................................................................................53
Isopaches pumicicola (Berggr.) Bakalin ...........................................................................................53
Isophyllaria attenuata (Rodway) E.A.Hodgs. ...............................................................................254
Isophyllaria fuegiana (Hässel) R.M.Schust. ..................................................................................254
Isotachis armata (Nees) Gottsche .................................................................................................101
Isotachis aubertii (Schwägr.) Mitt. ...............................................................................................101
Isotachis boliviensis Gottsche ........................................................................................................101
Isotachis chinensis C.Gao, T.Cao et J.Sun ....................................................................................101
Isotachis erythrorhiza (Lehm. et Lindenb.) Spruce........................................................................101
Isotachis fragilis Steph. .................................................................................................................101
Isotachis grandis Carrington et Pearson ........................................................................................101
Isotachis grossidens Steph. .............................................................................................................101
Isotachis hastatistipula (Steph.) J.J.Engel ......................................................................................101
Isotachis hians Steph. ...................................................................................................................101
Isotachis humectata (Hook.f. et Taylor) Steph..............................................................................101
Isotachis inlata Steph. .................................................................................................................101
Isotachis intortifolia (Hook.f. et Taylor) Gottsche ........................................................................101
Isotachis japonica Steph. ..............................................................................................................101
Isotachis lacustris Herzog .............................................................................................................101
Isotachis lopezii (R.M.Schust.) Gradst..........................................................................................101
Isotachis lyallii Mitt. ....................................................................................................................101
Isotachis minima Pearson .............................................................................................................101
Isotachis montana Colenso ...........................................................................................................101
Isotachis multiceps (Lindenb. et Gottsche) Gottsche.....................................................................102
Isotachis multiceps var. fendleri Gottsche ......................................................................................102
Isotachis nigella Herzog ...............................................................................................................102
Isotachis obtusa Steph. .................................................................................................................102
Isotachis olivacea R.M.Schust.......................................................................................................102
Isotachis plicata J.J.Engel .............................................................................................................102
Isotachis pusilla Steph. .................................................................................................................102
Isotachis riparia Rodway ..............................................................................................................102
Isotachis serrulata (Sw.) Gottsche .................................................................................................102
Isotachis spegazziniana C.Massal. .................................................................................................102
Isotachis sprucei Beauverd ............................................................................................................102
Isotachis stephanii E.S.Salmon .....................................................................................................102
Isotachis vexans Steph. .................................................................................................................102
Isotachis westlandica (E.A.Hodgs.) R.M.Schust. ..........................................................................102
Jackiella angustifolia Herzog ........................................................................................................116
World checklist of hornworts and liverworts
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579
Jackiella ceylanica Schifn. ex Steph. ............................................................................................116
Jackiella curvata E.A.Hodgs. et Allison........................................................................................117
Jackiella javanica Schifn. ............................................................................................................117
Jackiella javanica var. cavifolia Schifn. ........................................................................................117
Jackiella javanica var. cordifolia Schifn. ......................................................................................117
Jackiella renifolia Schifn. ............................................................................................................117
Jackiella sinensis (W.E.Nicholson) Grolle ....................................................................................117
Jackiella singapurensis Schifn. .....................................................................................................117
Jackiella singapurensis var. philippinensis N.Kitag. ........................................................................117
Jackiella unica Steph. ..................................................................................................................117
Jamesoniella convoluta Steph........................................................................................................509
Jensenia connivens (Colenso) Grolle.............................................................................................469
Jensenia crassifrons (Steph.) S.Schuette et Stotler .........................................................................470
Jensenia decipiens (Mitt.) Grolle ..................................................................................................470
Jensenia diformis (Nees) Grolle ...................................................................................................470
Jensenia lorschuetzii Gronde .......................................................................................................470
Jensenia spinosa (Lindenb. et Gottsche) Grolle ............................................................................470
Jensenia wallisii (J.B.Jack et Steph.) Grolle ..................................................................................470
Jubula blepharophylla Grolle........................................................................................................295
Jubula hattorii Udar et V.Nath ...................................................................................................295
Jubula hattorii var. muthukuzhiana A.E.D.Daniels et P.Daniel ...................................................295
Jubula himalayensis S.C.Srivast. et D.Sharma ..............................................................................296
Jubula hutchinsiae (Hook.) Dumort. ...........................................................................................296
Jubula hutchinsiae subsp. australiae Pócs et A.Cairns ...................................................................296
Jubula hutchinsiae subsp. bogotensis (Steph.) Verd. ......................................................................296
Jubula hutchinsiae subsp. caucasica Konstant. et Vilnet ...............................................................296
Jubula hutchinsiae subsp. japonica (Steph.) Horik. et Ando .........................................................296
Jubula hutchinsiae subsp. javanica (Steph.) Verd. ........................................................................296
Jubula hutchinsiae subsp. pennsylvanica (Steph.) Verd. ................................................................296
Jubula kwangsiensis C.Gao et K.C.Chang....................................................................................296
Jungermannia amentacea Bertol. ..................................................................................................119
Jungermannia atrovirens Dumort. ...............................................................................................118
Jungermannia borealis Damsh. et Váňa........................................................................................118
Jungermannia brasiliensis Raddi...................................................................................................119
Jungermannia chinensis Osbeck ...................................................................................................119
Jungermannia cordata Vill. ..........................................................................................................119
Jungermannia crenulata Schmidel................................................................................................119
Jungermannia creutzeri Kremer ...................................................................................................119
Jungermannia digitata C.F.W.Meissn. ex Spreng. .......................................................................119
Jungermannia dubioides H.A.Mill. ..............................................................................................119
Jungermannia erectii Ajit P.Singh et V.Nath ...............................................................................118
Jungermannia exsertifolia Steph. ..................................................................................................118
Jungermannia exsertifolia subsp. cordifolia (Dumort.) Váňa .........................................................118
Jungermannia fernandeziana Mitt. ..............................................................................................119
Jungermannia gollanii Steph. .......................................................................................................118
Jungermannia hexagona Schwägr. ................................................................................................119
Jungermannia holandriana Kremer ..............................................................................................119
Jungermannia incerta Gottsche ....................................................................................................119
Jungermannia konstantinovae Bakalin et Vilnet............................................................................118
Jungermannia laterilora Hampe ex Gottsche ..............................................................................119
Jungermannia lescuriana Austin ...................................................................................................119
Jungermannia longiretis Besch. et Spruce .....................................................................................119
580
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Jungermannia mastigophora Spreng. ............................................................................................119
Jungermannia michelii Mérat ......................................................................................................119
Jungermannia minima Scop. .......................................................................................................119
Jungermannia odorata With. .......................................................................................................119
Jungermannia ovatotrigona (Steph.) Grolle ..................................................................................118
Jungermannia peltata Schmidel ...................................................................................................119
Jungermannia polaris Lindb. ........................................................................................................118
Jungermannia pumila With. ........................................................................................................118
Jungermannia quadridigitata Grif. ..............................................................................................120
Jungermannia sauteri De Not. ex Rabenh. ...................................................................................120
Jungermannia secunda Hampe ex Gottsche .................................................................................120
Jungermannia stereocaulis Bory ....................................................................................................120
Jungermannia submersa Kremer ...................................................................................................120
Jungermannia sullivantiana Austin ..............................................................................................120
Jungermannia supina Hofm........................................................................................................120
Jungermannia tenuis Ehrh. ..........................................................................................................120
Jungermannia uncifolia Steph. .....................................................................................................120
Jungermannia vernicosa Cass. ex Mérat ........................................................................................120
Kingiolejeunea guayanensis H.Rob. ..............................................................................................509
Kurzia abbreviata Mizut. ............................................................................................................166
Kurzia abietinella (Herzog) Grolle ..............................................................................................166
Kurzia bisetula (Steph.) Grolle ....................................................................................................166
Kurzia borneensis Mizut. .............................................................................................................166
Kurzia brasiliensis (Steph.) Grolle................................................................................................166
Kurzia brevicalycina (Steph.) Grolle ............................................................................................166
Kurzia caduciloba R.M.Schust.....................................................................................................166
Kurzia calcarata (Steph.) Grolle ..................................................................................................165
Kurzia capillaris (Sw.) Grolle ......................................................................................................164
Kurzia capillaris subsp. capillaris var. verrucosa (Steph.) Pócs ......................................................165
Kurzia capillaris subsp. paramicola Pócs ......................................................................................165
Kurzia capillaris subsp. stephanii (Renauld ex Steph.) Pócs ..........................................................165
Kurzia compacta (Steph.) Grolle ..................................................................................................165
Kurzia cucullifolia (Steph.) R.M.Schust.......................................................................................166
Kurzia lagellifera (Steph.) Grolle ................................................................................................166
Kurzia fragilifolia R.M.Schust. ....................................................................................................166
Kurzia fragillima (Herzog) Grolle ...............................................................................................166
Kurzia geniculata Mizut. .............................................................................................................166
Kurzia gonyotricha (Sande Lac.) Grolle........................................................................................165
Kurzia hawaica (C.M.Cooke) Grolle ..........................................................................................166
Kurzia helophila R.M.Schust. ......................................................................................................165
Kurzia helophila var. laccida R.M.Schust. ex J.J.Engel ................................................................165
Kurzia hippurioides (Hook.f. et Taylor) Grolle ............................................................................165
Kurzia hippurioides var. ornata J.J.Engel et G.L.Merr. .................................................................165
Kurzia hispida (Steph.) Grolle .....................................................................................................166
Kurzia irregularis (Steph.) Grolle ................................................................................................165
Kurzia lateconica (Steph.) Grolle .................................................................................................167
Kurzia lineariloba Mizut. ............................................................................................................167
Kurzia longicaulis Piippo .............................................................................................................167
Kurzia makinoana (Steph.) Grolle...............................................................................................167
Kurzia mauiensis (H.A.Mill.) H.A.Mill. ......................................................................................167
Kurzia mollis (Steph.) J.J.Engel et R.M.Schust. ...........................................................................167
Kurzia moniliformis J.J.Engel ......................................................................................................165
World checklist of hornworts and liverworts
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581
Kurzia nemoides (Hook.f. et Taylor) Grolle.................................................................................165
Kurzia nivicola (R.M.Schust.) E.D.Cooper .................................................................................167
Kurzia pallescens Grolle ...............................................................................................................167
Kurzia pallida Piippo ..................................................................................................................167
Kurzia paucilora (Dicks.) Grolle.................................................................................................166
Kurzia quinquespina J.J.Engel et G.L.Merr. ................................................................................167
Kurzia reversa (Carrington et Pearson) Grolle .............................................................................167
Kurzia saddlensis (Besch. et C.Massal.) Grolle .............................................................................165
Kurzia setiformis (De Not.) J.J.Engel et R.M.Schust....................................................................167
Kurzia sexida (Steph.) Grolle......................................................................................................167
Kurzia sinensis K.C.Chang ..........................................................................................................167
Kurzia sylvatica (A.Evans) Grolle ................................................................................................167
Kurzia tasmanica (Steph.) E.D.Cooper .......................................................................................167
Kurzia tayloriana (H.A.Mill.) H.A.Mill. .....................................................................................167
Kurzia tenerrima (Mitt.) Grolle ...................................................................................................168
Kurzia touwii N.Kitag.................................................................................................................168
Kurzia trichoclados (Müll.Frib.) Grolle ........................................................................................168
Kurzia trilobata (R.M.Schust.) R.M.Schust. ...............................................................................168
Kurzia verticellata (Carrington) Grolle ........................................................................................168
Kymatocalyx africanus Váňa et M.Wigginton.................................................................................74
Kymatocalyx dominicensis (Spruce) Váňa........................................................................................74
Kymatocalyx madagascariensis (Steph.) Gradst. et Váňa..................................................................74
Kymatocalyx rhizomaticus (Herzog) Gradst. et Váňa ......................................................................74
Kymatolejeunea bartlettii Grolle ...................................................................................................362
Lamellocolea granditexta (Steph.) J.J.Engel ..................................................................................204
Lamellocolea integrostia J.J.Engel et Glenny .................................................................................204
Leiolejeunea grandilora A.Evans ..................................................................................................362
Leiomitra breviseta (Steph.) R.M.Schust. .....................................................................................256
Leiomitra capillata Lindb. ...........................................................................................................256
Leiomitra elegans (Lehm.) Hässel .................................................................................................256
Leiomitra elliottii (Steph.) R.M.Schust. .......................................................................................256
Leiomitra laccida Spruce.............................................................................................................257
Leiomitra hirticaulis R.M.Schust. ................................................................................................257
Leiomitra lanata (Hook.) R.M.Schust. ........................................................................................257
Leiomitra mastigophoroides R.M.Schust. ......................................................................................257
Leiomitra merrillana (Steph.) T.Katag. ........................................................................................257
Leiomitra paraphyllina Spruce .....................................................................................................257
Leiomitra robusta (Steph.) R.M.Schust. .......................................................................................257
Leiomitra smaragdina Hässel .......................................................................................................257
Leiomitra tomentosa (Sw.) Lindb. ................................................................................................257
Leiosporoceros dussii (Steph.) Hässel ...............................................................................................41
Lejeunea abyssinica (Gola) Cufod. ...............................................................................................365
Lejeunea acanthogona Spruce .......................................................................................................365
Lejeunea acuminata (Lehm. et Lindenb.) Lehm...........................................................................365
Lejeunea acuta Mitt. ....................................................................................................................365
Lejeunea acutata (Steph.) Solari...................................................................................................365
Lejeunea adpressa Nees ................................................................................................................365
Lejeunea aethiopica E.W.Jones ....................................................................................................365
Lejeunea alaskana (R.M.Schust. et Steere) Inoue et Steere ...........................................................365
Lejeunea alata Gottsche ..............................................................................................................365
Lejeunea alata var. patriciae Pócs .................................................................................................365
Lejeunea albescens (Steph.) Mizut. ...............................................................................................365
582
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Lejeunea albilora Colenso ...........................................................................................................365
Lejeunea aloba Sande Lac. ...........................................................................................................365
Lejeunea alobifolia H.A.Mill........................................................................................................365
Lejeunea amaniensis E.W.Jones ...................................................................................................365
Lejeunea ambigua Lindenb. et Gottsche ......................................................................................365
Lejeunea amentulifera Steph. .......................................................................................................365
Lejeunea androgyna R.M.Schust. .................................................................................................365
Lejeunea angulifolia Mitt. ............................................................................................................365
Lejeunea angusta (Lehm. et Lindenb.) Mont. ..............................................................................366
Lejeunea anisophylla Mont. .........................................................................................................366
Lejeunea anomala Lindenb. et Gottsche ......................................................................................366
Lejeunea antillana Steph. ............................................................................................................366
Lejeunea aphanes Spruce..............................................................................................................366
Lejeunea apiahyna (Steph.) Sushil K.Singh ..................................................................................381
Lejeunea apiculata Sande Lac. .....................................................................................................366
Lejeunea aptycta Gottsche ...........................................................................................................381
Lejeunea aquatica Horik..............................................................................................................366
Lejeunea aquatica var. apiculata S.Hatt. ......................................................................................366
Lejeunea armitii (Steph.) Steph. ..................................................................................................366
Lejeunea asperifolia Steph. ...........................................................................................................366
Lejeunea asperrima Spruce ...........................................................................................................366
Lejeunea asperula (Steph.) Mizut. ................................................................................................366
Lejeunea asprella Spruce ..............................................................................................................366
Lejeunea asthenica Spruce ............................................................................................................381
Lejeunea atheatostipa Spruce ........................................................................................................366
Lejeunea balazsii (Pócs) R.M.Schust............................................................................................380
Lejeunea barbata (Herzog) R.L.Zhu et M.J.Lai ...........................................................................366
Lejeunea bermudiana (A.Evans) R.M.Schust. ..............................................................................366
Lejeunea bethanica Gottsche........................................................................................................381
Lejeunea bidentula Herzog ..........................................................................................................366
Lejeunea biformis Gottsche ..........................................................................................................366
Lejeunea blepharogona Spruce ......................................................................................................366
Lejeunea blomquistii R.M.Schust. ................................................................................................366
Lejeunea boliviensis (Steph.) R.L.Zhu et M.E.Reiner ...................................................................366
Lejeunea bombonasensis Spruce ....................................................................................................381
Lejeunea bornmuelleri (Steph.) M.E.Reiner .................................................................................381
Lejeunea boryana Mont. ..............................................................................................................366
Lejeunea brenanii E.W.Jones .......................................................................................................366
Lejeunea calcicola R.M.Schust. ....................................................................................................366
Lejeunea calcicola var. mexicana R.M.Schust. ..............................................................................367
Lejeunea canariensis (Steph.) Steph..............................................................................................367
Lejeunea cancellata Nees et Mont. ...............................................................................................367
Lejeunea candida (Pócs) R.M.Schust. ..........................................................................................381
Lejeunea cantabrigiensis E.W.Jones ..............................................................................................367
Lejeunea capensis Gottsche ..........................................................................................................367
Lejeunea caracensis Lindenb.........................................................................................................381
Lejeunea caripensis Lindenb. et Gottsche .....................................................................................381
Lejeunea caroliniana Austin .........................................................................................................367
Lejeunea catinulifera Spruce ........................................................................................................367
Lejeunea caulicalyx (Steph.) M.E.Reiner et Goda ........................................................................367
Lejeunea cavifolia (Ehrh.) Lindb..................................................................................................367
Lejeunea caviloba (Steph.) Besch. ................................................................................................367
World checklist of hornworts and liverworts
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583
Lejeunea cerina (Lehm. et Lindenb.) Lehm. et Lindenb...............................................................367
Lejeunea chaishanensis S.H.Lin ....................................................................................................367
Lejeunea chamissonis Lindenb. .....................................................................................................381
Lejeunea chimborazensis Spruce ...................................................................................................381
Lejeunea cladogyna A.Evans .........................................................................................................367
Lejeunea clavata Lindenb. ...........................................................................................................367
Lejeunea claviformis Lindenb. ex Steph........................................................................................367
Lejeunea cochleata Spruce ............................................................................................................367
Lejeunea cocoes Mitt. ...................................................................................................................367
Lejeunea colensoana (Steph.) M.A.M.Renner ...............................................................................367
Lejeunea combuensis O.S.Moura, Ilk.-Borg. et M.E.Reiner ..........................................................381
Lejeunea compacta (Steph.) Steph. ...............................................................................................367
Lejeunea concava Lindenb. et Gottsche .......................................................................................381
Lejeunea concinnula Spruce et Steph. ..........................................................................................367
Lejeunea conformis Nees et Mont. ...............................................................................................381
Lejeunea connatistipula (Steph.) Steph. ........................................................................................367
Lejeunea contracta Mizut. ............................................................................................................368
Lejeunea controversa Gottsche......................................................................................................368
Lejeunea convexiloba M.L.So et R.L.Zhu .....................................................................................368
Lejeunea corcovadae (Steph.) Bischl. ............................................................................................368
Lejeunea cordilora Spruce ...........................................................................................................368
Lejeunea cordistipula Lindenb. et Gottsche ..................................................................................381
Lejeunea corralensis A.Evans ........................................................................................................368
Lejeunea corynantha Spruce .........................................................................................................368
Lejeunea crassiretis Mitt. ..............................................................................................................368
Lejeunea cristulata (Steph.) M.E.Reiner et Goda .........................................................................368
Lejeunea cristulilora (Steph.) M.E.Reiner et Goda ......................................................................368
Lejeunea curviloba Steph. ............................................................................................................380
Lejeunea cuspidistipula (Steph.) Steph. ex Watts ..........................................................................368
Lejeunea cyanomontana R.M.Schust. ...........................................................................................368
Lejeunea cyanophora R.M.Schust. ................................................................................................368
Lejeunea cyathearum E.W.Jones ..................................................................................................368
Lejeunea cyathophora Mitt. ..........................................................................................................368
Lejeunea cyrtotis Spruce ...............................................................................................................381
Lejeunea debilis (Lehm. et Lindenb.) Nees et Mont. ....................................................................381
Lejeunea denticalyx (Steph.) Steph. ..............................................................................................368
Lejeunea denticuspis (Steph.) Mizut. ............................................................................................368
Lejeunea denudata (Pearson) J.J.Engel .........................................................................................368
Lejeunea deplanata Nees ..............................................................................................................368
Lejeunea deplanata var. cuspidata (Steph.) M.E.Reiner ................................................................368
Lejeunea devendrae (Sushil K.Singh) P.K.Verma et K.K.Rawat ...................................................382
Lejeunea diaphana Spruce ...........................................................................................................368
Lejeunea dictyocalyx Spruce .........................................................................................................382
Lejeunea dimorpha T.Kodama .....................................................................................................369
Lejeunea dipterocarpa E.W.Jones .................................................................................................369
Lejeunea dipterota (Eifrig) G.E.Lee ..............................................................................................382
Lejeunea discreta Lindenb. ...........................................................................................................369
Lejeunea disjecta Spruce...............................................................................................................369
Lejeunea diversicuspis Spruce .......................................................................................................369
Lejeunea drehwaldii Heinrichs et Schäf.-Verw. ............................................................................382
Lejeunea drummondii Taylor .......................................................................................................382
Lejeunea duncaniae (Sim) M.E.Reiner .........................................................................................382
584
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Lejeunea ecarinata (Steph.) J.M.Coult., Barnes et Arthur ............................................................369
Lejeunea eckloniana Lindenb. ......................................................................................................369
Lejeunea edentata L.Söderstr. ......................................................................................................382
Lejeunea eifrigii Mizut.................................................................................................................369
Lejeunea elegans Gottsche ............................................................................................................385
Lejeunea elongella Gottsche .........................................................................................................369
Lejeunea emarginulilora Gottsche ex Steph. ................................................................................382
Lejeunea epibrya Taylor ...............................................................................................................382
Lejeunea erostrata M.E.Reiner et Goda........................................................................................369
Lejeunea estrellamontana M.A.M.Renner et Pócs.........................................................................382
Lejeunea exilis (Reinw., Blume et Nees) Grolle............................................................................369
Lejeunea exilis var. abnormis (Herzog) G.E.Lee............................................................................369
Lejeunea falcata (Pócs et J.Eggers) Pócs .......................................................................................381
Lejeunea fawcettiae D.J.Carr........................................................................................................382
Lejeunea fernandeziana S.W.Arnell .............................................................................................369
Lejeunea irma Mitt.....................................................................................................................369
Lejeunea issistipula (Steph.) Steph...............................................................................................369
Lejeunea laccida Lindenb. et Gottsche ........................................................................................382
Lejeunea lagellaris Spruce ...........................................................................................................369
Lejeunea lava (Sw.) Nees ............................................................................................................369
Lejeunea lava subsp. moorei (Lindb.) R.M.Schust.......................................................................369
Lejeunea lava subsp. orientalis R.M.Schust. ................................................................................369
Lejeunea lava var. pellucida Lindenb. et Gottsche .......................................................................370
Lejeunea lava subsp. tabularis (Spreng.) S.W.Arnell ...................................................................370
Lejeunea lavida Mitt. .................................................................................................................382
Lejeunea lavovirens Ångstr. .........................................................................................................370
Lejeunea leischeri (Steph.) Mizut. ...............................................................................................370
Lejeunea lorida Spruce................................................................................................................382
Lejeunea loridana A.Evans ..........................................................................................................370
Lejeunea fulfordiae (Jovet-Ast) R.L.Zhu ......................................................................................370
Lejeunea fulva Spruce ..................................................................................................................382
Lejeunea fusagasugana Gottsche...................................................................................................370
Lejeunea galeata Spruce ...............................................................................................................370
Lejeunea gayana Gottsche............................................................................................................370
Lejeunea gibbiloba (Steph.) H.A.Mill. .........................................................................................370
Lejeunea glaucescens Gottsche ......................................................................................................370
Lejeunea glaucescens var. acrogyna R.M.Schust. ............................................................................370
Lejeunea glaucescens var. obsoleta R.M.Schust. .............................................................................370
Lejeunea globosilora (Steph.) Steph. ............................................................................................370
Lejeunea gomphocalyx Spruce.......................................................................................................370
Lejeunea gottscheana Lindenb. .....................................................................................................382
Lejeunea gracilicaulis Spruce ........................................................................................................370
Lejeunea gracilipes (Taylor) Spruce ..............................................................................................370
Lejeunea gracilis Steph. ................................................................................................................370
Lejeunea gradsteiniana Pócs .........................................................................................................370
Lejeunea gradsteinii G.E.Lee, Damanhuri et Latif ......................................................................370
Lejeunea graminicolor Spruce.......................................................................................................382
Lejeunea grolleana (Bernecker) R.L.Zhu et W.Ye ........................................................................382
Lejeunea grossecristata (Steph.) E.W.Jones ...................................................................................370
Lejeunea grossiretis (Steph.) M.E.Reiner et Goda .........................................................................371
Lejeunea grossistipula Steph. ........................................................................................................371
Lejeunea grossitexta (Steph.) M.E.Reiner et Goda ........................................................................371
World checklist of hornworts and liverworts
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585
Lejeunea grossiuscula Gottsche ex Steph. ......................................................................................371
Lejeunea hahnii Solari .................................................................................................................371
Lejeunea haitica Nees et Mont. ...................................................................................................371
Lejeunea hawaikiana M.A.M.Renner et de Lange .......................................................................371
Lejeunea helenae Pearson .............................................................................................................371
Lejeunea helmsiana (Steph.) Steph. ..............................................................................................371
Lejeunea hepaticola (Steph.) Steph. ..............................................................................................371
Lejeunea herminieri (Steph.) R.L.Zhu .........................................................................................382
Lejeunea heterocheila Taylor ........................................................................................................383
Lejeunea hibernica Bischl., H.A.Mill. et Bonner ex Grolle ...........................................................371
Lejeunea hieronymii Spruce .........................................................................................................385
Lejeunea hodgsoniana Grolle ex R.J.Lewington, Bever. et M.A.M.Renner ...................................371
Lejeunea howeana Grolle .............................................................................................................371
Lejeunea hui R.L.Zhu .................................................................................................................371
Lejeunea humefacta Spruce ..........................................................................................................371
Lejeunea hyalina (Steph.) L.Söderstr. et A.Hagborg.....................................................................371
Lejeunea hygrophila Gottsche ......................................................................................................383
Lejeunea ibadana A.J.Harr. et E.W.Jones ....................................................................................371
Lejeunea immersa Spruce .............................................................................................................371
Lejeunea increscens Spruce ...........................................................................................................371
Lejeunea inlatiloba (Steph.) H.A.Mill., Bonner et Bischl. ...........................................................371
Lejeunea inlexiloba Prantl ...........................................................................................................372
Lejeunea intricata Prantl..............................................................................................................372
Lejeunea isophylla E.W.Jones .......................................................................................................372
Lejeunea japonica Mitt. ...............................................................................................................372
Lejeunea jardinii Spruce ..............................................................................................................372
Lejeunea julacea Steph. ................................................................................................................372
Lejeunea jungneri (Steph.) Steph. ................................................................................................372
Lejeunea juruana Gradst. et M.E.Reiner .....................................................................................380
Lejeunea kashyapii M.Dey, D.K.Singh et D.Singh ......................................................................372
Lejeunea kilimandjarensis Steph...................................................................................................372
Lejeunea kinabalensis Mizut. .......................................................................................................372
Lejeunea kodamae Ikegami et Inoue ............................................................................................372
Lejeunea konosensis Mizut. ...........................................................................................................372
Lejeunea koponenii (Pócs et J.Eggers) Pócs ..................................................................................381
Lejeunea kuerschneriana Pócs.......................................................................................................372
Lejeunea laeta (Lehm. et Lindenb.) Lehm. et Lindenb. ...............................................................372
Lejeunea laetevirens Nees et Mont. ..............................................................................................380
Lejeunea laevicalyx Gottsche ........................................................................................................383
Lejeunea laii R.L.Zhu .................................................................................................................372
Lejeunea lamacerina (Steph.) Schifn. ..........................................................................................372
Lejeunea lamacerina subsp. gemminata R.M.Schust. ....................................................................372
Lejeunea latilobula (Herzog) R.L.Zhu et M.L.So.........................................................................372
Lejeunea laxa (Nees) Lindenb. ....................................................................................................383
Lejeunea leiantha Spruce .............................................................................................................383
Lejeunea leptalea Spruce ..............................................................................................................372
Lejeunea leptoscypha Spruce .........................................................................................................383
Lejeunea leratii (Steph.) Mizut. ...................................................................................................372
Lejeunea leucosis Besch. et Spruce ................................................................................................373
Lejeunea litoralis Steph. ...............................................................................................................373
Lejeunea lomana E.W.Jones ........................................................................................................373
Lejeunea longicollis S.W.Arnell ....................................................................................................373
586
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Lejeunea longilobula Pócs ............................................................................................................373
Lejeunea lowriana Steph. .............................................................................................................373
Lejeunea lumbricoides (Nees) Nees ..............................................................................................373
Lejeunea lunatigastria Tixier ........................................................................................................373
Lejeunea lusoria (Lindenb. et Gottsche) Steph. ............................................................................383
Lejeunea luzonensis (Steph.) R.L.Zhu et M.J.Lai .........................................................................383
Lejeunea lyratilora Prantl ............................................................................................................373
Lejeunea macrorhyncha Spruce.....................................................................................................383
Lejeunea magohukui Mizut. .........................................................................................................373
Lejeunea malangensis (Herzog) R.L.Zhu et Y.M.Wei ...................................................................373
Lejeunea mandonii (Steph.) Müll.Frib. ........................................................................................373
Lejeunea marasmodes Spruce........................................................................................................383
Lejeunea masoalae Pócs................................................................................................................373
Lejeunea massalongoana (Schifn. ex P.Syd) Solari ........................................................................373
Lejeunea megalantha Spruce ........................................................................................................373
Lejeunea mehrana M.Dey, D.K.Singh et D.Singh .......................................................................373
Lejeunea meridensis Ilk.-Borg.......................................................................................................373
Lejeunea micholitzii Mizut. .........................................................................................................373
Lejeunea microloba Taylor ...........................................................................................................373
Lejeunea mimula Hürl.................................................................................................................373
Lejeunea minutiloba A.Evans .......................................................................................................373
Lejeunea minutiloba var. heterogyna R.M.Schust. .........................................................................373
Lejeunea mizutanii Grolle ...........................................................................................................373
Lejeunea molkenboeriana Sande Lac.............................................................................................373
Lejeunea monimiae (Steph.) Steph. ..............................................................................................374
Lejeunea morobensis (Grolle) M.A.M.Renner et Pócs...................................................................383
Lejeunea multidentata M.E.Reiner et Mustelier ...........................................................................374
Lejeunea musae (Spreng.) Gottsche, Lindenb. et Nees .................................................................374
Lejeunea muscicola Spruce ...........................................................................................................374
Lejeunea muscicola var. palmicola Spruce .....................................................................................374
Lejeunea neelgherriana Gottsche ..................................................................................................374
Lejeunea nemoralis Gottsche ........................................................................................................374
Lejeunea nepalensis Steph. ...........................................................................................................374
Lejeunea nesiotica Besch. et Spruce ..............................................................................................374
Lejeunea neumanniana Nees........................................................................................................374
Lejeunea nietneri (Steph.) Steph. ex Watts...................................................................................374
Lejeunea notata Gottsche ............................................................................................................374
Lejeunea novoguineensis Schifn. ..................................................................................................383
Lejeunea nymannii Steph.............................................................................................................374
Lejeunea obfusca Mitt. .................................................................................................................374
Lejeunea obidensis Spruce ............................................................................................................374
Lejeunea obscura Mitt. .................................................................................................................374
Lejeunea obtusangula Spruce........................................................................................................383
Lejeunea obtusata Gottsche..........................................................................................................374
Lejeunea oerstediana Lindenb. et Hampe .....................................................................................383
Lejeunea okomuensis E.W.Jones ...................................................................................................374
Lejeunea oligoclada Spruce...........................................................................................................374
Lejeunea oracola M.A.M.Renner .................................................................................................374
Lejeunea osculatiana De Not. ......................................................................................................374
Lejeunea otiana S.Hatt. ...............................................................................................................374
Lejeunea ovalifolia Steph. ............................................................................................................375
Lejeunea paciica Mont. ...............................................................................................................375
World checklist of hornworts and liverworts
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587
Lejeunea pallescens Mitt. ..............................................................................................................375
Lejeunea pallida Lindenb. et Gottsche .........................................................................................375
Lejeunea pallidevirens S.Hatt. ......................................................................................................380
Lejeunea pallidissima Gola ...........................................................................................................375
Lejeunea papilionacea Prantl ........................................................................................................375
Lejeunea papuana (Pócs) R.M.Schust. .........................................................................................381
Lejeunea paraensis Spruce ............................................................................................................383
Lejeunea paratropa Spruce ...........................................................................................................375
Lejeunea parva (S.Hatt.) Mizut. ..................................................................................................380
Lejeunea parviloba Ångstr............................................................................................................383
Lejeunea patagonica (Steph.) Steph. .............................................................................................375
Lejeunea patens Lindb. ................................................................................................................375
Lejeunea patersonii (Steph.) Steph. ..............................................................................................375
Lejeunea patriciae Schäf.-Verw. ...................................................................................................375
Lejeunea paucidentata (Steph.) Grolle .........................................................................................375
Lejeunea pectinella Mizut. ...........................................................................................................375
Lejeunea perigonialis Gottsche .....................................................................................................375
Lejeunea perpapillosa M.E.Reiner et K.C.Pôrto ...........................................................................375
Lejeunea pertusa (Corda ex Nees et Mont.) Gottsche, Lindenb. et Nees ......................................375
Lejeunea pleidereri Sushil K.Singh ..............................................................................................383
Lejeunea phyllobola Nees et Mont................................................................................................375
Lejeunea phyllobola var. turgidula Spruce .....................................................................................375
Lejeunea planiloba A.Evans .........................................................................................................375
Lejeunea polilloensis Steph. ..........................................................................................................375
Lejeunea polyantha Mont. ...........................................................................................................380
Lejeunea praetervisa Steph. ..........................................................................................................375
Lejeunea primordialis (Hook.f. et Taylor) Gottsche, Lindenb. et Nees.........................................375
Lejeunea princeps (Steph.) Mizut. ................................................................................................376
Lejeunea prionoides Spruce ..........................................................................................................380
Lejeunea proboscidea Gottsche .....................................................................................................385
Lejeunea procumbens Mitt. ..........................................................................................................376
Lejeunea prominula Gottsche ......................................................................................................383
Lejeunea propagulifera Gradst. .....................................................................................................376
Lejeunea pteridis Besch. et Spruce ................................................................................................376
Lejeunea pterigonia (Lehm. et Lindenb.) Mont............................................................................384
Lejeunea ptosimophylla C.Massal. ................................................................................................376
Lejeunea puiggariana Steph. ........................................................................................................376
Lejeunea pulverulenta (Steph.) M.E.Reiner ..................................................................................376
Lejeunea pulvinata Nees et Mont. ...............................................................................................376
Lejeunea quinqueumbonata Spruce ..............................................................................................384
Lejeunea quinqueumbonata var. rotundata (Herzog) Sushil K.Singh ............................................384
Lejeunea raddiana Lindenb. ........................................................................................................376
Lejeunea radicans Lindenb. et Gottsche .......................................................................................376
Lejeunea ramosissima Steph. ........................................................................................................376
Lejeunea ramulosa Spruce ............................................................................................................380
Lejeunea rara Steph. ....................................................................................................................376
Lejeunea ravenelii Austin .............................................................................................................376
Lejeunea recurva M.E.Reiner .......................................................................................................376
Lejeunea relexistipula (Lehm. et Lindenb.) Lehm. et Lindenb. ....................................................376
Lejeunea relexistipula var. costaricensis (Steph.) M.E.Reiner.........................................................376
Lejeunea reinerae Ilk.-Borg. .........................................................................................................376
Lejeunea remotifolia Hampe ex Steph. .........................................................................................384
588
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Lejeunea resupinata (Steph.) Steph. .............................................................................................377
Lejeunea reticulata Herzog ..........................................................................................................377
Lejeunea rhigophila M.A.M.Renner .............................................................................................377
Lejeunea rhodesiae (Sim) R.M.Schust. .........................................................................................377
Lejeunea rionegrensis Spruce ........................................................................................................377
Lejeunea riparia Mitt. .................................................................................................................377
Lejeunea rothii (Schwägr.) Gottsche, Lindenb. et Nees ................................................................377
Lejeunea rotundifolia Mitt. ..........................................................................................................377
Lejeunea ruthii (A.Evans) R.M.Schust. ........................................................................................384
Lejeunea ruthii var. alata R.M.Schust. .........................................................................................384
Lejeunea sanctae-helenae M.Wigginton ........................................................................................377
Lejeunea scabrilora Loitl. ............................................................................................................385
Lejeunea schusteri Grolle ..............................................................................................................377
Lejeunea semiscabrida Gottsche ...................................................................................................377
Lejeunea semperi Steph. ...............................................................................................................377
Lejeunea seriata Lindenb. et Gottsche .........................................................................................377
Lejeunea sessililora (Steph.) Grolle ..............................................................................................377
Lejeunea setacea (Steph.) Steph. ...................................................................................................377
Lejeunea setiloba Spruce ..............................................................................................................377
Lejeunea sharpii (R.M.Schust.) R.M.Schust. ...............................................................................377
Lejeunea siccata Spruce ................................................................................................................377
Lejeunea sikorae (Steph.) Steph. ..................................................................................................384
Lejeunea silvatica Gottsche ..........................................................................................................378
Lejeunea smaragdina Besch. et Spruce .........................................................................................378
Lejeunea soae R.L.Zhu, Y.M.Wei, L.Söderstr., A.Hagborg et von Konrat....................................378
Lejeunea solanicola Spruce ...........................................................................................................378
Lejeunea sordida (Nees) Nees ......................................................................................................378
Lejeunea spiniloba Lindenb. et Gottsche ......................................................................................378
Lejeunea spinulilora Spruce.........................................................................................................378
Lejeunea sporadica Besch. et Spruce .............................................................................................378
Lejeunea squarrosa (Steph.) Steph. ...............................................................................................378
Lejeunea squarrosula (Herzog) Solari ...........................................................................................378
Lejeunea srivastavae P.K.Verma et K.K.Rawat .............................................................................384
Lejeunea stenodentata M.A.M.Renner et Pócs..............................................................................384
Lejeunea stephaniana Mizut. ........................................................................................................378
Lejeunea stevensiana (Steph.) Mizut.............................................................................................378
Lejeunea subacuta Mitt. ...............................................................................................................378
Lejeunea subaquatica Schifn. ......................................................................................................378
Lejeunea subbiida Steph. ex Duss ...............................................................................................378
Lejeunea subelobata Carrington et Pearson ..................................................................................384
Lejeunea subigiensis (Steph.) Steph. .............................................................................................378
Lejeunea subolivacea Mizut. .........................................................................................................378
Lejeunea subplana (Steph.) C.J.Bastos .........................................................................................378
Lejeunea subrufula Spruce ...........................................................................................................378
Lejeunea subsessilis Spruce............................................................................................................378
Lejeunea subspathulata Spruce .....................................................................................................379
Lejeunea succulenta Herzog .........................................................................................................379
Lejeunea sufruticola Spruce .........................................................................................................379
Lejeunea sulphurea (Lehm. et Lindenb.) Spruce...........................................................................384
Lejeunea syoshii Inoue..................................................................................................................379
Lejeunea talamancensis M.E.Reiner et Schäf.-Verw......................................................................379
Lejeunea tamasii M.E.Reiner, N.Salazar et C.Chung...................................................................379
World checklist of hornworts and liverworts
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589
Lejeunea tamaspocsii G.E.Lee ......................................................................................................379
Lejeunea tapajosensis Spruce.........................................................................................................379
Lejeunea tarapotensis Spruce ........................................................................................................379
Lejeunea tenella Taylor ................................................................................................................379
Lejeunea tenera (Sw.) Gottsche, Lindenb. et Nees .......................................................................380
Lejeunea terricola Spruce .............................................................................................................384
Lejeunea thallophora (Eifrig) Gradst. ...........................................................................................379
Lejeunea tonduzana (Steph.) M.E.Reiner ....................................................................................379
Lejeunea topoensis Gradst. et M.E.Reiner.....................................................................................380
Lejeunea touwii (Pócs) R.M.Schust. ............................................................................................381
Lejeunea trachygona Spruce .........................................................................................................379
Lejeunea trinitensis Lindenb. .......................................................................................................379
Lejeunea trochantha Spruce .........................................................................................................385
Lejeunea trukensis H.A.Mill. et Bonner .......................................................................................379
Lejeunea tuberculosa Steph. .........................................................................................................379
Lejeunea tumida Mitt. .................................................................................................................379
Lejeunea tunquiniensis M.E.Reiner et Drehwald ..........................................................................365
Lejeunea uleana (Steph.) Steph. ...................................................................................................379
Lejeunea umbilicata (Nees) Nees .................................................................................................379
Lejeunea urbanii (Steph.) Steph. .................................................................................................384
Lejeunea utriculata (Steph.) Mizut. .............................................................................................379
Lejeunea venezuelana (R.M.Schust.) R.L.Zhu et W.Ye ................................................................384
Lejeunea vesicata Mitt. ................................................................................................................379
Lejeunea villaumei (Steph.) Grolle...............................................................................................379
Lejeunea viridis R.M.Schust. ex L.Söderstr. et A.Hagborg ...........................................................384
Lejeunea vojtkoi Pócs ...................................................................................................................379
Lejeunea vulgariformis Gottsche ..................................................................................................379
Lejeunea wattsiana (Steph.) H.A.Mill., Bonner et Bischl. ............................................................380
Lejeunea wichurae (Steph.) Steph. ...............................................................................................380
Lejeunea wightii Lindenb. ...........................................................................................................380
Lejeunea xiphophylla (Herzog) M.E.Reiner ..................................................................................380
Lejeunea zacuapana (Steph.) Prantl .............................................................................................385
Lembidium berggrenii Herzog......................................................................................................168
Lembidium longifolium R.M.Schust. ...........................................................................................168
Lembidium nutans (Hook.f. et Taylor) Mitt. ...............................................................................168
Lembidium nutans var. lagelliferum E.A.Hodgs. .........................................................................168
Lepicolea attenuata (Mitt.) Steph. ................................................................................................142
Lepicolea magellanica (Gola) Solari ..............................................................................................142
Lepicolea norrisii Piippo ..............................................................................................................142
Lepicolea ochroleuca (Spreng.) Spruce ..........................................................................................142
Lepicolea pruinosa (Taylor) Spruce ..............................................................................................142
Lepicolea ramentiissa Herzog ......................................................................................................142
Lepicolea rara (Steph.) Grolle ......................................................................................................142
Lepicolea rigida (De Not.) E.B.Scott ...........................................................................................142
Lepicolea scolopendra (Hook.) Dumort. ex Trevis. .......................................................................142
Lepicolea yakusimensis (S.Hatt.) S.Hatt........................................................................................142
Lepidogyna hodgsoniae (Grolle) R.M.Schust.................................................................................416
Lepidogyna menziesii (Hook.) R.M.Schust...................................................................................416
Lepidolaena berggrenii E.A.Hodgs. ..............................................................................................416
Lepidolaena brachyclada (Lehm.) Trevis. .....................................................................................416
Lepidolaena clavigera (Hook.) Dumort. ex Trevis. .......................................................................416
Lepidolaena novae-zelandiae (E.A.Hodgs. et S.W.Arnell) von Konrat, L.Söderstr. et A.Hagborg .416
590
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Lepidolaena palpebrifolia (Hook.) Dumort. ex Trevis. .................................................................416
Lepidolaena reticulata (Hook.f. et Taylor) Trevis.........................................................................416
Lepidolaena taylorii (Gottsche) Trevis. ........................................................................................417
Lepidolejeunea auriculata Schäf.-Verw. et Heinrichs ....................................................................391
Lepidolejeunea bidentula (Steph.) R.M.Schust. ............................................................................391
Lepidolejeunea bidentula var. novae-caledoniae Piippo ..................................................................391
Lepidolejeunea borneensis (Steph.) R.M.Schust. ...........................................................................392
Lepidolejeunea cordiissa (Taylor) M.E.Reiner ..............................................................................391
Lepidolejeunea cuspidata (Gottsche) Heinrichs et Schäf.-Verw. ....................................................392
Lepidolejeunea delessertii (Nees et Mont.) Grolle..........................................................................392
Lepidolejeunea eluta (Nees) R.M.Schust. .....................................................................................392
Lepidolejeunea falcata (Herzog) R.M.Schust. ...............................................................................391
Lepidolejeunea graefei (J.B.Jack et Steph.) R.M.Schust................................................................392
Lepidolejeunea grossepapulosa (Steph.) Piippo...............................................................................391
Lepidolejeunea integristipula (J.B.Jack et Steph.) R.M.Schust.......................................................392
Lepidolejeunea involuta (Gottsche) Grolle ...................................................................................391
Lepidolejeunea longilobula (Amakawa) Xiao L.He ........................................................................392
Lepidolejeunea serrulata (Steph.) Grolle .......................................................................................392
Lepidolejeunea sullivantii (Gottsche) M.E.Reiner ........................................................................392
Lepidozia acantha J.J.Engel .........................................................................................................169
Lepidozia aequiloba Steph. ..........................................................................................................170
Lepidozia africana Steph. ............................................................................................................170
Lepidozia alstonii Fulford ............................................................................................................170
Lepidozia ambigua De Not..........................................................................................................170
Lepidozia andicola Beauverd........................................................................................................170
Lepidozia appressifolia Steph. .......................................................................................................170
Lepidozia armata Steph. ..............................................................................................................170
Lepidozia asymmetrica Steph. ......................................................................................................170
Lepidozia auriculata Mitt. ...........................................................................................................170
Lepidozia australis (Lehm. et Lindenb.) Mitt. ..............................................................................170
Lepidozia bidens J.J.Engel............................................................................................................170
Lepidozia biloba Herzog ..............................................................................................................170
Lepidozia bisbiida Steph. ............................................................................................................170
Lepidozia borneensis Steph. ..........................................................................................................170
Lepidozia bragginsiana E.D.Cooper et M.A.M.Renner................................................................170
Lepidozia brasiliensis Steph. .........................................................................................................170
Lepidozia brevidentata Mitt. ........................................................................................................170
Lepidozia brevifolia Mitt. ............................................................................................................170
Lepidozia brevifolia var. planifolia Schifn....................................................................................170
Lepidozia brotheri Steph. .............................................................................................................170
Lepidozia bufalona Steph............................................................................................................170
Lepidozia bursifera S.Hatt. et Grolle............................................................................................170
Lepidozia caespitosa Spruce ..........................................................................................................170
Lepidozia caledonica Steph. .........................................................................................................170
Lepidozia caledonica var. tenuisecta Hürl......................................................................................170
Lepidozia ceramensis Herzog ........................................................................................................170
Lepidozia cherydrion Hürl. ..........................................................................................................170
Lepidozia chiloensis Steph. ...........................................................................................................170
Lepidozia chordulifera Taylor ......................................................................................................171
Lepidozia cladorhiza (Reinw., Blume et Nees) Nees ....................................................................171
Lepidozia coilophylla Taylor.........................................................................................................171
Lepidozia coilophylla var. apiculiloba (Steph.) Fulford ..................................................................171
World checklist of hornworts and liverworts
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591
Lepidozia communis Steph. ..........................................................................................................171
Lepidozia concinna Colenso.........................................................................................................171
Lepidozia cordata Lindenb. .........................................................................................................171
Lepidozia cordistipula Steph. .......................................................................................................171
Lepidozia crassitexta Steph. ..........................................................................................................171
Lepidozia cupressina (Sw.) Lindenb. ............................................................................................171
Lepidozia cupressina subsp. natalensis (Steph.) Pócs .....................................................................171
Lepidozia cupressina subsp. pinnata (Hook.) Pócs ........................................................................171
Lepidozia cupressina subsp. quinqueida (Steph.) Pócs..................................................................171
Lepidozia decaisnei Steph.............................................................................................................171
Lepidozia dendritica Spruce .........................................................................................................171
Lepidozia densa Herzog ...............................................................................................................171
Lepidozia digitata Herzog............................................................................................................171
Lepidozia eenii S.W.Arnell ..........................................................................................................171
Lepidozia elobata R.M.Schust......................................................................................................171
Lepidozia erosa Steph. .................................................................................................................171
Lepidozia erronea Herzog ............................................................................................................171
Lepidozia everettii Steph. .............................................................................................................172
Lepidozia everettii var. javensis Herzog.........................................................................................172
Lepidozia fauriana Steph. ............................................................................................................172
Lepidozia ferdinandi-muelleri Steph.............................................................................................172
Lepidozia ilamentosa (Lehm. et Lindenb.) Lehm. et Lindenb......................................................172
Lepidozia istulosa Mitt. ...............................................................................................................172
Lepidozia lexuosa Mitt. ...............................................................................................................172
Lepidozia fuegiensis Steph. ...........................................................................................................172
Lepidozia fugax J.J.Engel .............................................................................................................172
Lepidozia gedena Steph................................................................................................................172
Lepidozia glaucescens J.J.Engel .....................................................................................................172
Lepidozia glaucophylla (Hook.f. et Taylor) Gottsche, Lindenb. et Nees .......................................172
Lepidozia grandifolia Steph. ........................................................................................................172
Lepidozia griseola Herzog ............................................................................................................172
Lepidozia groenlandica Lehm. ......................................................................................................172
Lepidozia gwamii Piippo .............................................................................................................172
Lepidozia hampeana Lindenb. .....................................................................................................172
Lepidozia haskarliana (Gottsche, Lindenb. et Nees) Steph. .........................................................172
Lepidozia hastatistipula Steph. .....................................................................................................172
Lepidozia hexiloba Pearson ..........................................................................................................176
Lepidozia hirta Steph. .................................................................................................................172
Lepidozia holorhiza (Reinw., Blume et Nees) Nees ......................................................................172
Lepidozia holorhiza var. laxa (Nees) Schifn.................................................................................172
Lepidozia inaequalis (Lehm. et Lindenb.) Lehm. et Lindenb. ......................................................172
Lepidozia incurvata Lindenb. ......................................................................................................172
Lepidozia infuscata Mitt. .............................................................................................................173
Lepidozia integrifolia Doei...........................................................................................................173
Lepidozia jamaicensis Steph. ........................................................................................................173
Lepidozia kashyapii D.Singh et D.K.Singh ..................................................................................173
Lepidozia kinabaluensis Mizut. ....................................................................................................173
Lepidozia kirkii Steph. ................................................................................................................173
Lepidozia lacerifolia Steph. ..........................................................................................................173
Lepidozia laevifolia (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ............................................173
Lepidozia laevifolia var. acutiloba J.J.Engel ..................................................................................173
Lepidozia laevifolia var. alpina R.M.Schust. et J.J.Engel ..............................................................173
592
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Lepidozia lindigiana Steph. .........................................................................................................173
Lepidozia loheri Steph. ................................................................................................................173
Lepidozia longifolia Steph. ...........................................................................................................173
Lepidozia loriana Steph. ..............................................................................................................173
Lepidozia macrocolea Spruce ........................................................................................................173
Lepidozia massartiana Schifn. ex Steph.......................................................................................173
Lepidozia microphylla (Hook.) Lindenb.......................................................................................173
Lepidozia microstipula Steph. ......................................................................................................173
Lepidozia minima Steph. .............................................................................................................173
Lepidozia minor (Gottsche, Lindenb. et Nees) Solari...................................................................173
Lepidozia miqueliana Sande Lac. .................................................................................................173
Lepidozia montana Steph. ...........................................................................................................173
Lepidozia newtonii Steph.............................................................................................................173
Lepidozia nova Steph...................................................................................................................174
Lepidozia novae-zelandiae Steph. .................................................................................................174
Lepidozia novae-zelandiae var. heterostipa R.M.Schust. ................................................................174
Lepidozia novae-zelandiae var. minima R.M.Schust. ....................................................................174
Lepidozia obtusiloba Steph...........................................................................................................174
Lepidozia obtusiloba var. parvula J.J.Engel ...................................................................................174
Lepidozia omeiensis P.C.Chen ex Mizut. et K.C.Chang ...............................................................174
Lepidozia ornata J.J.Engel ...........................................................................................................174
Lepidozia pallida Steph. ..............................................................................................................174
Lepidozia palmicola Steph. ..........................................................................................................174
Lepidozia parvistipa Taylor..........................................................................................................176
Lepidozia paschalis Steph. ............................................................................................................174
Lepidozia patens Lindenb. ...........................................................................................................174
Lepidozia paucifolia Steph. ..........................................................................................................174
Lepidozia paupercula Steph. ........................................................................................................174
Lepidozia pearsonii Spruce ...........................................................................................................174
Lepidozia pendulina (Hook.) Lindenb. ........................................................................................174
Lepidozia peruviensis Steph. .........................................................................................................174
Lepidozia pinnaticruris Spruce ex Steph.......................................................................................174
Lepidozia plumula Herzog ...........................................................................................................174
Lepidozia portoricensis Fulford .....................................................................................................174
Lepidozia procera Mitt. ................................................................................................................174
Lepidozia pseudocupressina Schifn. ..............................................................................................174
Lepidozia pumila J.J.Engel ..........................................................................................................174
Lepidozia quadridens (Nees) Nees ...............................................................................................175
Lepidozia quadriida Lindenb. .....................................................................................................175
Lepidozia reptans (L.) Dumort. ...................................................................................................175
Lepidozia richardsii Herzog .........................................................................................................175
Lepidozia rigida Steph. ................................................................................................................175
Lepidozia robusta Steph. ..............................................................................................................175
Lepidozia rufescens Steph. ............................................................................................................175
Lepidozia sandvicensis Lindenb. ...................................................................................................175
Lepidozia schwabei Herzog ..........................................................................................................175
Lepidozia sellingiana H.A.Mill. ...................................................................................................175
Lepidozia septemida Steph. .........................................................................................................175
Lepidozia serpens Spruce ..............................................................................................................175
Lepidozia serrulata J.J.Engel ........................................................................................................175
Lepidozia setigera Steph. ..............................................................................................................175
Lepidozia sikkimensis Steph. ........................................................................................................175
World checklist of hornworts and liverworts
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593
Lepidozia spinosissima (Hook.f. et Taylor) Mitt. ..........................................................................175
Lepidozia squamifolia Steph. .......................................................................................................175
Lepidozia squarrosa Steph. ...........................................................................................................175
Lepidozia stahlii Steph.................................................................................................................175
Lepidozia stuhlmannii Steph........................................................................................................175
Lepidozia stuhlmannii var. carnosa (Steph.) Pócs .........................................................................175
Lepidozia stuhlmannii subsp. pulvinata (Steph.) Pócs ..................................................................175
Lepidozia subdichotoma Spruce ....................................................................................................175
Lepidozia subintegra Lindenb. .....................................................................................................175
Lepidozia subtransversa Steph. .....................................................................................................175
Lepidozia subtrichodes Steph. .......................................................................................................175
Lepidozia succida Mitt. ................................................................................................................175
Lepidozia supradecomposita Lindenb. ...........................................................................................175
Lepidozia supradecomposita var. falcifolia Herzog .........................................................................175
Lepidozia suyungii C.Gao et X.L.Bai ...........................................................................................176
Lepidozia terricola Steph..............................................................................................................176
Lepidozia triangulifolia Steph. .....................................................................................................176
Lepidozia trichodes (Reinw., Blume et Nees) Nees .......................................................................176
Lepidozia tricuspidata Steph. .......................................................................................................176
Lepidozia tunguraguae Steph. ......................................................................................................176
Lepidozia ubangiensis Steph. ........................................................................................................176
Lepidozia udarii S.C.Srivast., D.Kumar et D.Sharma ..................................................................176
Lepidozia ulothrix (Schwägr.) Lindenb. .......................................................................................176
Lepidozia vitrea Steph. ................................................................................................................176
Lepidozia wattsiana Steph. ..........................................................................................................176
Lepidozia weymouthiana Steph. ...................................................................................................176
Leptocolea sumatrana Herzog.......................................................................................................510
Leptolejeunea amphiophthalma Zwickel .......................................................................................394
Leptolejeunea apiculata (Horik.) S.Hatt. ......................................................................................394
Leptolejeunea arunachalensis Sudipa Das et D.K.Singh ................................................................394
Leptolejeunea astroidea (Mitt.) Steph. ..........................................................................................394
Leptolejeunea australis Steph. .......................................................................................................394
Leptolejeunea balansae Steph. ......................................................................................................394
Leptolejeunea borneensis Herzog ...................................................................................................394
Leptolejeunea brasiliensis Bischl. ...................................................................................................394
Leptolejeunea convexistipa Bischl. .................................................................................................394
Leptolejeunea curvatifolia Steph. ..................................................................................................394
Leptolejeunea denticulata Steph....................................................................................................394
Leptolejeunea dentistipula Steph. ..................................................................................................394
Leptolejeunea diversilobulata Bischl. .............................................................................................394
Leptolejeunea dolabriformis Pearson .............................................................................................394
Leptolejeunea elliptica (Lehm. et Lindenb.) Besch. .......................................................................395
Leptolejeunea emarginata (Horik.) S.Hatt. ...................................................................................395
Leptolejeunea epiphylla (Mitt.) Steph. ..........................................................................................395
Leptolejeunea exocellata (Spruce) A.Evans ....................................................................................395
Leptolejeunea foliicola Steph. .......................................................................................................395
Leptolejeunea integristipula Steph.................................................................................................395
Leptolejeunea jamaicensis R.M.Schust. .........................................................................................395
Leptolejeunea lancifolia Steph. .....................................................................................................395
Leptolejeunea latifolia Herzog ......................................................................................................395
Leptolejeunea lepinii Steph...........................................................................................................395
Leptolejeunea ligulata Herzog ......................................................................................................395
594
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Leptolejeunea maculata (Mitt.) Schifn.........................................................................................395
Leptolejeunea massartiana Schifn. ex Herzog...............................................................................395
Leptolejeunea micronesica Inoue et H.A.Mill. ...............................................................................395
Leptolejeunea minima Herzog ......................................................................................................395
Leptolejeunea mirikana M.Dey et D.K.Singh ..............................................................................395
Leptolejeunea moniliata Steph......................................................................................................395
Leptolejeunea obfuscata (Spruce) Steph. .......................................................................................395
Leptolejeunea punctata Herzog.....................................................................................................395
Leptolejeunea radicosa (Nees ex Mont.) Grolle .............................................................................396
Leptolejeunea renneri Herzog .......................................................................................................396
Leptolejeunea revoluta P.C.Chen .................................................................................................396
Leptolejeunea rosulans Steph. .......................................................................................................396
Leptolejeunea serratifolia Schifn. .................................................................................................396
Leptolejeunea serrulata Herzog .....................................................................................................396
Leptolejeunea spinistipula (Mizut.) Xiao L.He ..............................................................................396
Leptolejeunea subdentata Schifn. ex Herzog ................................................................................396
Leptolejeunea subrotundifolia Herzog ...........................................................................................396
Leptolejeunea tridentata Bischl. ....................................................................................................396
Leptolejeunea trigonostipa (Spruce) Steph. ....................................................................................396
Leptolejeunea tripuncta (Mitt.) Steph. ..........................................................................................396
Leptolejeunea truncatifolia Steph. .................................................................................................396
Leptolejeunea udarii M.Dey et D.K.Singh ...................................................................................396
Leptolejeunea vitrea (Nees) Schifn. .............................................................................................396
Leptophyllopsis laxa (Mitt.) R.M.Schust. ex Hamlin ....................................................................204
Leptoscyphopsis paradoxa R.M.Schust...........................................................................................204
Leptoscyphus aequatus (Hook.f. et Taylor) Mitt. ..........................................................................206
Leptoscyphus amphibolius (Nees) Grolle .......................................................................................207
Leptoscyphus antarcticus (C.Massal.) Solari...................................................................................205
Leptoscyphus australis (Gottsche, Lindenb. et Nees) R.M.Schust. .................................................205
Leptoscyphus autoicus (J.J.Engel et Gradst.) Vanderp. et Gradst. ..................................................207
Leptoscyphus beckettianus (Steph.) R.M.Schust. ex J.J.Engel.........................................................207
Leptoscyphus belmoranus (Steph.) J.J.Engel ..................................................................................205
Leptoscyphus chilensis (De Not.) Hässel ........................................................................................206
Leptoscyphus cleeii Fulford ..........................................................................................................207
Leptoscyphus compactus (Colenso) J.J.Engel..................................................................................207
Leptoscyphus cuneifolius (Hook.) Mitt. .........................................................................................204
Leptoscyphus cuneifolius subsp. fragilis (J.B.Jack et Steph.) Grolle.................................................205
Leptoscyphus diicilis (Steph.) Fulford..........................................................................................206
Leptoscyphus diversifolius (Gottsche) Grolle..................................................................................206
Leptoscyphus erraticus (W.Martin et E.A.Hodgs.) J.J.Engel ..........................................................208
Leptoscyphus excipulatus (Steph.) J.J.Engel ...................................................................................205
Leptoscyphus expansus (Lehm.) Grolle ..........................................................................................206
Leptoscyphus gibbosus (Taylor) Mitt. ............................................................................................205
Leptoscyphus gradsteinii Vanderp., Schäf.-Verw. et D.G.Long......................................................205
Leptoscyphus hedbergii (S.W.Arnell) R.M.Schust. ........................................................................207
Leptoscyphus hexagonus (Nees) Grolle ..........................................................................................205
Leptoscyphus horizontalis (Hook.) Kühnem. .................................................................................208
Leptoscyphus huidobroanus (Mont.) Gottsche ...............................................................................207
Leptoscyphus huonicus Piippo .......................................................................................................207
Leptoscyphus infuscatus (Mitt.) E.W.Jones ex Grolle ....................................................................207
Leptoscyphus innovatus (E.A.Hodgs.) J.J.Engel.............................................................................205
Leptoscyphus intermedius Grolle ...................................................................................................206
World checklist of hornworts and liverworts
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595
Leptoscyphus jackii (Steph.) Grolle ...............................................................................................205
Leptoscyphus lambinonii Vanderp., Schäf.-Verw. et D.G.Long ....................................................206
Leptoscyphus longistipulus (Steph.) J.J.Engel .................................................................................205
Leptoscyphus magellanicus (Gola) Hässel ......................................................................................207
Leptoscyphus normalis (Steph.) J.J.Engel ......................................................................................208
Leptoscyphus obcordatus (Spruce) Grolle .......................................................................................206
Leptoscyphus ovatus (Spruce) Grolle .............................................................................................206
Leptoscyphus physanthus (Hook.f. et Taylor) J.J.Engel ..................................................................208
Leptoscyphus physocalyx (Hampe et Gottsche) Gottsche ...............................................................205
Leptoscyphus porphyrius (Nees) Grolle ..........................................................................................206
Leptoscyphus porphyrius subsp. azoricus (H.Buch et Perss.) Vanderp. et Heinrichs .......................206
Leptoscyphus sotiauxii Vanderp., Schäf.-Verw. et D.G.Long ........................................................206
Leptoscyphus spectabilis (Steph.) Grolle ........................................................................................207
Leptoscyphus subemarginatus (Hook.f. et Taylor) J.J.Engel ...........................................................208
Leptoscyphus trapezoïdes (Mont.) L.Söderstr. ................................................................................208
Lethocolea congesta (Lehm.) S.W.Arnell .........................................................................................97
Lethocolea glossophylla (Spruce) Grolle ...........................................................................................97
Lethocolea indica G.Asthana et Maurya .........................................................................................97
Lethocolea javanica (Schifn.) Grolle ..............................................................................................97
Lethocolea naruto-toganensis Furuki ...............................................................................................97
Lethocolea pansa (Taylor) G.A.M.Scott et K.G.Beckm. .................................................................97
Lethocolea radicosa (Lehm. et Lindenb.) Grolle .............................................................................97
Lethocolea repens S.Winkl. .............................................................................................................97
Lindigianthus cipaconeus (Gottsche) Kruijt et Gradst...................................................................299
Liochlaena lanceolata Nees...........................................................................................................117
Liochlaena subulata (A.Evans) Schljakov .....................................................................................118
Lobatiriccardia alterniloba (Hook.f. et Taylor) Furuki .................................................................437
Lobatiriccardia alterniloba var. gigantea (Steph.) Nebel................................................................437
Lobatiriccardia alterniloba var. robusta (Rodway) Nebel ..............................................................437
Lobatiriccardia athertonensis (Hewson) Furuki ............................................................................438
Lobatiriccardia coronopus (De Not.) Furuki .................................................................................438
Lobatiriccardia coronopus subsp. australis (R.M.Schust.) Nebel, Preussing, Schäf.-Verw. et
D.Quandt...................................................................................................................................438
Lobatiriccardia oberwinkleri Nebel, Preussing, Schäf.-Verw. et D.Quandt...................................438
Lobatiriccardia subaquatica (R.M.Schust.) Nebel ........................................................................438
Lobatiriccardia verdoornioides Nebel, Preussing, Schäf.-Verw. et D.Quandt ................................438
Lobatiriccardia yakusimensis (S.Hatt.) Furuki ..............................................................................438
Lobatiriccardia yunnanensis Furuki et D.G.Long .........................................................................438
Lophocolea aberrans Lindenb. et Gottsche ...................................................................................208
Lophocolea aequifolia Nees et Mont. ............................................................................................208
Lophocolea angustistipula Steph. ..................................................................................................208
Lophocolea anisoloba (J.J.Engel et Glenny) L.Söderstr..................................................................208
Lophocolea anomala Steph. ..........................................................................................................208
Lophocolea anomoda (Mont.) Steph. ............................................................................................208
Lophocolea apalachicola R.M.Schust. ...........................................................................................208
Lophocolea aperticaulis (J.J.Engel) L.Söderstr. ..............................................................................208
Lophocolea aphelophylla (Hässel) Váňa .........................................................................................208
Lophocolea apophylla (Hässel) Váňa .............................................................................................209
Lophocolea appalachiana R.M.Schust...........................................................................................209
Lophocolea ascensionis Steph. .......................................................................................................209
Lophocolea asperrima Steph. ........................................................................................................209
Lophocolea atra Gola ...................................................................................................................209
596
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Lophocolea attenuata Steph. .........................................................................................................209
Lophocolea australis Gottsche.......................................................................................................209
Lophocolea autoica Steph. ............................................................................................................209
Lophocolea baldwinii Steph. ........................................................................................................209
Lophocolea bartlettii H.A.Mill. ....................................................................................................209
Lophocolea bewsii (Sim) Grolle ....................................................................................................209
Lophocolea bicuspidata Steph. ......................................................................................................209
Lophocolea bidentata (L.) Dumort. ..............................................................................................209
Lophocolea biidistipula Steph. .....................................................................................................209
Lophocolea bispinosa (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ..........................................209
Lophocolea bootanensis Steph. ......................................................................................................209
Lophocolea boulyana Steph. .........................................................................................................209
Lophocolea bowiena Steph. ..........................................................................................................209
Lophocolea brookwoodiana Paton et Sheahan ...............................................................................209
Lophocolea caespitans Steph. ........................................................................................................209
Lophocolea calcarea Steph. ...........................................................................................................209
Lophocolea caledonica Steph.........................................................................................................209
Lophocolea canaliculata (Gottsche, Lindenb. et Nees) Steph. .......................................................210
Lophocolea canaliculata var. concava (J.J.Engel) L.Söderstr. .........................................................210
Lophocolea cervicornis Steph. .......................................................................................................210
Lophocolea ciliifera Steph.............................................................................................................210
Lophocolea coadunata (Sw.) Mont. ..............................................................................................210
Lophocolea concreta Mont. ...........................................................................................................210
Lophocolea convexula Mitt. ..........................................................................................................210
Lophocolea corticola Steph............................................................................................................210
Lophocolea decurrens Herzog........................................................................................................210
Lophocolea deningeri Herzog........................................................................................................210
Lophocolea dentilora Steph..........................................................................................................210
Lophocolea diformis Nees ............................................................................................................210
Lophocolea discedens (Lehm. et Lindenb.) Lehm. et Lindenb. ......................................................210
Lophocolea dusenii Steph. ............................................................................................................210
Lophocolea erosa (J.J.Engel) L.Söderstr.........................................................................................210
Lophocolea excisifolia Steph..........................................................................................................210
Lophocolea fertilis (J.J.Engel) L.Söderstr.......................................................................................210
Lophocolea lavicans Steph. ..........................................................................................................210
Lophocolea loribunda Steph. .......................................................................................................210
Lophocolea foliicola Spruce...........................................................................................................210
Lophocolea fragillima Steph. ........................................................................................................210
Lophocolea fragmentissima R.M.Schust. .......................................................................................211
Lophocolea fragrans (Moris et De Not.) Gottsche, Lindenb. et Nees ............................................211
Lophocolea glaziovii Steph. ..........................................................................................................211
Lophocolea gollanii (Steph.) Váňa ................................................................................................211
Lophocolea granatensis Gottsche...................................................................................................211
Lophocolea griithiana Steph. ......................................................................................................211
Lophocolea hahnii Steph. .............................................................................................................211
Lophocolea haskarliana Gottsche..................................................................................................211
Lophocolea hattorii (J.J.Engel) L.Söderstr.....................................................................................211
Lophocolea hawaica Steph............................................................................................................211
Lophocolea heterodonta Steph.......................................................................................................211
Lophocolea heteromorpha Steph. ...................................................................................................211
Lophocolea heterophylla (Schrad.) Dumort. ..................................................................................211
Lophocolea heterophylla subsp. cladogyna R.M.Schust...................................................................211
World checklist of hornworts and liverworts
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597
Lophocolea horikawana S.Hatt. ....................................................................................................211
Lophocolea howeana Steph. ..........................................................................................................211
Lophocolea humifusa (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ..........................................211
Lophocolea humistrata (Hook.f. et Taylor) Gottsche, Lindenb. et Nees .......................................211
Lophocolea itoana Inoue ..............................................................................................................211
Lophocolea javanica Schifn. ........................................................................................................211
Lophocolea koponenii (Piippo) Váňa ............................................................................................211
Lophocolea kurzii Sande Lac. .......................................................................................................212
Lophocolea kurzii var. siamensis N.Kitag. .....................................................................................212
Lophocolea laceristipula Steph. .....................................................................................................212
Lophocolea latistipula Steph. ........................................................................................................212
Lophocolea lauterbachii Steph. .....................................................................................................212
Lophocolea laxissima Herzog ........................................................................................................212
Lophocolea ledermannii Steph. .....................................................................................................212
Lophocolea lenta (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ................................................212
Lophocolea leptantha (Hook.f. et Taylor) Gottsche, Lindenb. et Nees..........................................212
Lophocolea liebmanniana Gottsche ..............................................................................................212
Lophocolea lindmannii Steph. ......................................................................................................212
Lophocolea longiciliata Herzog .....................................................................................................212
Lophocolea lucida (Spreng.) Mont. ..............................................................................................212
Lophocolea madagascariensis Gottsche ..........................................................................................212
Lophocolea magna (Udar et V.Nath) Váňa...................................................................................212
Lophocolea mediinfrons (J.J.Engel et Braggins) L.Söderstr. ...........................................................212
Lophocolea micronesica Inoue et H.A.Mill. ..................................................................................212
Lophocolea microstipula Steph. .....................................................................................................212
Lophocolea minor Nees ................................................................................................................212
Lophocolea minutistipula Steph. ...................................................................................................212
Lophocolea mollis (Nees) Nees .....................................................................................................212
Lophocolea morobeana Piippo ......................................................................................................213
Lophocolea muhavurensis (S.W.Arnell) S.W.Arnell ex Pócs ..........................................................213
Lophocolea muricata (Lehm.) Nees ..............................................................................................213
Lophocolea muricata var. major Pearson .......................................................................................213
Lophocolea nakajimae S.Hatt. et Inoue ........................................................................................213
Lophocolea novae-zeelandiae (Lehm. et Lindenb.) Nees................................................................213
Lophocolea novae-zeelandiae var. meridionalis (Steph.) L.Söderstr. ...............................................213
Lophocolea orbigniana Nees et Mont. ..........................................................................................213
Lophocolea papulimarginata H.A.Mill..........................................................................................213
Lophocolea parca (Gottsche) Fulford et Sharp ..............................................................................213
Lophocolea parva Steph. ..............................................................................................................213
Lophocolea parvispinea (J.J.Engel) L.Söderstr. ..............................................................................213
Lophocolea parvistipula Steph. .....................................................................................................213
Lophocolea patulistipa Steph. .......................................................................................................213
Lophocolea perpusilla (Hook.f. et Taylor) Gottsche, Lindenb. et Nees .........................................213
Lophocolea piacenzai (Gola) Váňa ................................................................................................213
Lophocolea pilistipula Steph. ........................................................................................................213
Lophocolea pinnatistipula Steph. ..................................................................................................213
Lophocolea platensis C.Massal. .....................................................................................................214
Lophocolea purpurea Steph. .........................................................................................................214
Lophocolea pusilla Steph. .............................................................................................................214
Lophocolea randii S.W.Arnell ......................................................................................................214
Lophocolea rara Steph..................................................................................................................214
Lophocolea rectangularis Herzog...................................................................................................214
598
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Lophocolea rectangulata Mitt. ......................................................................................................214
Lophocolea rupicola Steph. ...........................................................................................................214
Lophocolea sabuletorum (Hook.f. et Taylor) Gottsche, Lindenb. et Nees .....................................214
Lophocolea salacensis Steph. .........................................................................................................214
Lophocolea savesiana Steph. .........................................................................................................214
Lophocolea semiteres (Lehm.) Mitt. ..............................................................................................214
Lophocolea semiteres var. retusa (J.J.Engel) L.Söderstr...................................................................214
Lophocolea serrata Mitt. ...............................................................................................................214
Lophocolea siamensis Steph. .........................................................................................................214
Lophocolea sikkimensis (Steph.) Herzog et Grolle .........................................................................214
Lophocolea silvestris Gottsche .......................................................................................................214
Lophocolea steetziae De Not. ........................................................................................................214
Lophocolea striatella (C.Massal.) Schifn. .....................................................................................214
Lophocolea subbidentata Herzog ..................................................................................................214
Lophocolea subcostata Steph. ........................................................................................................214
Lophocolea subporosa Mitt. ..........................................................................................................214
Lophocolea subporosa var. inlexifolia (Steph.) L.Söderstr. .............................................................215
Lophocolea subviridis (Hook.f. et Taylor) Gottsche, Lindenb. et Nees .........................................215
Lophocolea sumatrana Schifn. .....................................................................................................215
Lophocolea sylvatica Mitt. ............................................................................................................215
Lophocolea tenera Ångstr. ............................................................................................................215
Lophocolea tenerrima Spruce ........................................................................................................215
Lophocolea teptepensis Piippo .......................................................................................................215
Lophocolea textilis (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ..............................................215
Lophocolea textiloidea J.J.Engel ....................................................................................................215
Lophocolea trichocoleoides (Glenny, J.J.Engel et He-Nygrén) L.Söderstr. ......................................215
Lophocolea tricuspidata Herzog ....................................................................................................215
Lophocolea tristaniana S.W.Arnell ...............................................................................................215
Lophocolea undulata Mont. .........................................................................................................215
Lophocolea villosa Mitt. ...............................................................................................................215
Lophocolea wacei (S.W.Arnell ex J.J.Engel et Váňa) Váňa et L.Söderstr. ......................................215
Lophocolea wambana Piippo ........................................................................................................215
Lophocolea werthii (J.J.Engel et R.M.Schust.) Váňa et L.Söderstr. ...............................................215
Lophocolea widgrenii Steph. .........................................................................................................215
Lopholejeunea aberrantia Horik. ..................................................................................................408
Lopholejeunea applanata (Reinw., Blume et Nees) Schifn. ..........................................................406
Lopholejeunea borbonica Steph. ...................................................................................................406
Lopholejeunea borneensis (Steph.) Verd. .......................................................................................406
Lopholejeunea ceylanica Steph. .....................................................................................................407
Lopholejeunea colensoi Steph. .......................................................................................................407
Lopholejeunea erugata B.M.hiers ...............................................................................................406
Lopholejeunea eulopha (Taylor) Schifn. .......................................................................................406
Lopholejeunea evansiana Verd. .....................................................................................................406
Lopholejeunea grandicrista Steph..................................................................................................408
Lopholejeunea grollei R.L.Zhu et Gradst. .....................................................................................406
Lopholejeunea herzogiana Verd. ...................................................................................................406
Lopholejeunea hispidissima Steph. ................................................................................................407
Lopholejeunea horticola Schifn. ...................................................................................................407
Lopholejeunea jonesii Vanden Berghen.........................................................................................406
Lopholejeunea laciniata E.W.Jones ..............................................................................................406
Lopholejeunea latilobula Verd. .....................................................................................................407
Lopholejeunea leioptera Gyarmati .................................................................................................408
World checklist of hornworts and liverworts
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599
Lopholejeunea lepidoscypha Kiaer et Pearson .................................................................................408
Lopholejeunea loheri Steph. ..........................................................................................................407
Lopholejeunea magna Mizut. .......................................................................................................407
Lopholejeunea minima Vanden Berghen ......................................................................................406
Lopholejeunea minuta R.L.Zhu et Gradst. ...................................................................................407
Lopholejeunea multilacera Steph. .................................................................................................408
Lopholejeunea nigricans (Lindenb.) Schifn. .................................................................................407
Lopholejeunea obtusilacera Herzog ...............................................................................................406
Lopholejeunea onraedtii Vanden Berghen.....................................................................................408
Lopholejeunea paramultilacera Vanden Berghen ..........................................................................406
Lopholejeunea plicatiscypha (Hook.f. et Taylor) Steph..................................................................407
Lopholejeunea pocsii Gyarmati .....................................................................................................408
Lopholejeunea proxima Steph. ......................................................................................................407
Lopholejeunea quinquecarinata Vanden Berghen .........................................................................406
Lopholejeunea recurvata Mizut.....................................................................................................407
Lopholejeunea renistipula (Mitt.) Steph. .......................................................................................406
Lopholejeunea revoluta E.W.Jones ...............................................................................................406
Lopholejeunea soae R.L.Zhu et Gradst. ........................................................................................407
Lopholejeunea sphaerophora (Lehm. et Lindenb.) Steph. ..............................................................408
Lopholejeunea streimannii B.M.hiers et Gradst. .........................................................................407
Lopholejeunea subfusca (Nees) Schifn. .........................................................................................407
Lopholejeunea subfusca var. elongata Vanden Berghen ..................................................................407
Lopholejeunea tixieriana Vanden Berghen ...................................................................................408
Lopholejeunea udarii M.Dey et D.K.Singh ..................................................................................408
Lopholejeunea utriculata Steph. ...................................................................................................408
Lopholejeunea vietnamica Tixier ..................................................................................................408
Lopholejeunea vojtkoana Gyarmati ...............................................................................................407
Lopholejeunea wiltensii Steph.......................................................................................................407
Lopholejeunea yapensis Steph. ......................................................................................................407
Lopholejeunea zollingeri (Steph.) Schifn. .....................................................................................407
Lophonardia jamesonii (Mont.) L.Söderstr. et Váňa .......................................................................75
Lophonardia laxifolia (Mont.) L.Söderstr. et Váňa.........................................................................75
Lophonardia tristaniana (S.W.Arnell) L.Söderstr. et Váňa .............................................................75
Lophozia ascendens (Warnst.) R.M.Schust. ....................................................................................79
Lophozia austrosibirica Bakalin ......................................................................................................79
Lophozia ciliata Damsh., L.Söderstr. et H.Weibull .......................................................................79
Lophozia guttulata (Lindb. et Arnell) A.Evans ...............................................................................79
Lophozia jamaicensis (Nees) Steph. ................................................................................................79
Lophozia lacerata N.Kitag. ............................................................................................................79
Lophozia lantratoviae Bakalin ........................................................................................................79
Lophozia murmanica Kaal. ............................................................................................................79
Lophozia paciica Bakalin ..............................................................................................................79
Lophozia pallida (Steph.) Grolle ....................................................................................................79
Lophozia savicziae Schljakov..........................................................................................................79
Lophozia schusterana Schljakov ......................................................................................................79
Lophozia serpens (Dumort.) Dumort. ............................................................................................80
Lophozia silvicola H.Buch .............................................................................................................79
Lophozia silvicoloides N.Kitag. .......................................................................................................79
Lophozia subapiculata R.M.Schust. et Damsh. ..............................................................................80
Lophozia udarii S.Srivast., S.C.Srivast. et K.K.Rawat ....................................................................80
Lophozia ventricosa (Dicks.) Dumort.............................................................................................80
Lophozia wenzelii (Nees) Steph. ....................................................................................................80
600
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Lophoziopsis excisa (Dicks.) Konstant. et Vilnet .............................................................................80
Lophoziopsis excisa var. elegans (R.M.Schust.) Konstant. et Vilnet ..................................................80
Lophoziopsis excisa var. infuscata (R.M.Schust. et Damsh.) Konstant. et Vilnet..............................80
Lophoziopsis excisa var. succulenta (R.M.Schust. et Damsh.) Konstant. et Vilnet ............................80
Lophoziopsis longidens (Lindb.) Konstant. et Vilnet .......................................................................81
Lophoziopsis longidens subsp. arctica (R.M.Schust.) Váňa et L.Söderstr..........................................81
Lophoziopsis pellucida (R.M.Schust.) Konstant. et Vilnet...............................................................81
Lophoziopsis pellucida var. minor (R.M.Schust.) L.Söderstr. et Váňa ..............................................81
Lophoziopsis polaris (R.M.Schust.) Konstant. et Vilnet ..................................................................81
Lophoziopsis polaris var. sphagnorum (R.M.Schust.) Konstant. et Vilnet.........................................81
Lophoziopsis propagulifera (Gottsche) Konstant. et Vilnet ..............................................................81
Lophoziopsis rubrigemma (R.M.Schust.) Konstant. et Vilnet ..........................................................81
Lunularia cruciata (L.) Dumort. ex Lindb. ..................................................................................476
Lunularia cruciata subsp. thaxteri (A.Evans et Herzog) R.M.Schust. ...........................................476
Luteolejeunea herzogii (Buchloh) Piippo ......................................................................................304
Macrocolura sagittistipula (Spruce) R.M.Schust. ..........................................................................348
Makinoa crispata (Steph.) Miyake ...............................................................................................468
Mannia androgyna (L.) A.Evans ..................................................................................................481
Mannia californica (Gottsche) L.C.Wheeler ................................................................................481
Mannia controversa (Meyl.) D.B.Schill ........................................................................................481
Mannia controversa subsp. asiatica D.B.Schill et D.G.Long.........................................................481
Mannia fragrans (Balb.) Frye et L.Clark ......................................................................................481
Mannia fragrans subsp. orientalis R.M.Schust. ............................................................................481
Mannia gracilis (F.Weber) D.B.Schill et D.G.Long ....................................................................482
Mannia hegewaldii Bischl. ...........................................................................................................482
Mannia paradoxa R.M.Schust. ....................................................................................................482
Mannia perssonii Udar et V.Chandra ..........................................................................................481
Mannia pilosa (Hornem.) Frye et L.Clark ...................................................................................482
Mannia sibirica (Müll.Frib.) Frye et L.Clark ...............................................................................482
Mannia triandra (Scop.) Grolle ..................................................................................................482
Marchantia acaulis Steph. ...........................................................................................................490
Marchantia antiqua Steph. ..........................................................................................................490
Marchantia assamica Grif. ..........................................................................................................491
Marchantia balboi Gola...............................................................................................................491
Marchantia balboi var. acutisquamata Gerola ..............................................................................491
Marchantia berteroana Lehm. et Lindenb. ...................................................................................490
Marchantia breviloba A.Evans .....................................................................................................489
Marchantia cagnii Gola ...............................................................................................................491
Marchantia carrii Bischl. .............................................................................................................490
Marchantia cengiana Gerola ........................................................................................................491
Marchantia chenopoda L. .............................................................................................................489
Marchantia crenata Austin ..........................................................................................................489
Marchantia debilis K.I.Goebel .....................................................................................................490
Marchantia emarginata Reinw., Blume et Nees ...........................................................................490
Marchantia emarginata subsp. lecordiana (Steph.) Bischl. ............................................................490
Marchantia emarginata subsp. tosana (Steph.) Bischl. ..................................................................490
Marchantia foliacea Mitt. ............................................................................................................489
Marchantia formosana Horik.......................................................................................................489
Marchantia friedrichsthaliana Trevis............................................................................................491
Marchantia geminata Reinw., Blume et Nees ..............................................................................490
Marchantia globosa Brid. .............................................................................................................489
Marchantia hartlessiana Steph. ....................................................................................................491
World checklist of hornworts and liverworts
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601
Marchantia hexaptera Reichardt ..................................................................................................489
Marchantia inlexa Nees et Mont. ...............................................................................................489
Marchantia keniae Gola ..............................................................................................................491
Marchantia linearis Lehm. et Lindenb.........................................................................................489
Marchantia macropora Mitt.........................................................................................................491
Marchantia miqueliana Lehm. ....................................................................................................489
Marchantia novoguineensis Bischl. ...............................................................................................489
Marchantia paleacea Bertol..........................................................................................................489
Marchantia paleacea subsp. diptera (Nees et Mont.) Inoue ..........................................................489
Marchantia papillata Raddi .........................................................................................................490
Marchantia papillata subsp. grossibarba (Steph.) Bischl. ..............................................................490
Marchantia pappeana Lehm. .......................................................................................................489
Marchantia pappeana subsp. robusta (Steph.) Bischl. ...................................................................489
Marchantia papyracea Gola .........................................................................................................491
Marchantia philippinensis Bischl. .................................................................................................491
Marchantia pileata Mitt. .............................................................................................................489
Marchantia pinnata Steph. ..........................................................................................................489
Marchantia plicata Nees et Mont. ...............................................................................................490
Marchantia polymorpha L. ...........................................................................................................490
Marchantia polymorpha subsp. montivagans Bischl. et Boissel.-Dub.............................................490
Marchantia polymorpha subsp. ruderalis Bischl. et Boissel.-Dub. .................................................490
Marchantia quadriloba Steph. .....................................................................................................492
Marchantia rubribarba Steph. .....................................................................................................489
Marchantia sellae Gola ................................................................................................................492
Marchantia solomonensis Bischl. ..................................................................................................491
Marchantia stoloniscyphulus (C.Gao et K.C.Chang) Piippo .........................................................492
Marchantia streimannii Bischl. ....................................................................................................491
Marchantia subgeminata Steph. ...................................................................................................491
Marchantia subintegra Mitt. ........................................................................................................491
Marchantia treubii Schifn. .........................................................................................................491
Marchantia trilocularis Roth........................................................................................................492
Marchantia tusui Gola ................................................................................................................492
Marchantia vitiensis Steph. ..........................................................................................................490
Marchantia wallisii J.B.Jack et Steph...........................................................................................491
Marchesinia bongardiana (Lehm. et Lindenb.) Trevis. .................................................................409
Marchesinia brachiata (Sw.) Schifn. ...........................................................................................409
Marchesinia deslooveri Vanden Berghen.......................................................................................409
Marchesinia excavata (Mitt.) Schifn. ..........................................................................................409
Marchesinia languida (Nees et Mont.) Steph. ..............................................................................409
Marchesinia mackaii (Hook.) Gray..............................................................................................408
Marchesinia nobilis (Gottsche) X.Q.Shi, R.L.Zhu et Gradst. .......................................................409
Marchesinia robusta (Mitt.) Schifn. ............................................................................................409
Marsupella alata S.Hatt. et N.Kitag. ...........................................................................................112
Marsupella andreaeoides (Lindb.) Müll.Frib.................................................................................112
Marsupella apiculata Schifn. .......................................................................................................112
Marsupella aquatica (Lindenb.) Schifn. ......................................................................................112
Marsupella arctica (Berggr.) Bryhn et Kaal. .................................................................................112
Marsupella boeckii (Austin) Lindb. ex Kaal. .................................................................................112
Marsupella bolanderi (Austin) Underw. .......................................................................................112
Marsupella condensata (Ångstr. ex C.Hartm.) Lindb. ex Kaal. .....................................................113
Marsupella disticha Steph. ...........................................................................................................113
Marsupella emarginata (Ehrh.) Dumort. .....................................................................................113
602
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Marsupella emarginata subsp. tubulosa (Steph.) N.Kitag..............................................................113
Marsupella emarginata subsp. tubulosa var. apertifolia (Steph.) N.Kitag. ......................................113
Marsupella emarginata subsp. tubulosa var. patens N.Kitag. .........................................................113
Marsupella emarginata subsp. tubulosa var. tubulosa (Steph.) N.Kitag. ex Váňa et L.Söderstr. .....113
Marsupella funckii (F.Weber et D.Mohr) Dumort. .....................................................................113
Marsupella microphylla R.M.Schust. ............................................................................................113
Marsupella minutissima N.Kitag. .................................................................................................113
Marsupella neesii Sande Lac. ex Schifn. ......................................................................................113
Marsupella paroica R.M.Schust. ..................................................................................................113
Marsupella profunda Lindb..........................................................................................................113
Marsupella pseudofunckii S.Hatt. .................................................................................................113
Marsupella sparsifolia (Lindb.) Dumort. ......................................................................................113
Marsupella sparsifolia subsp. childii R.M.Schust. .........................................................................113
Marsupella sphacelata (Giesecke ex Lindenb.) Dumort. ...............................................................113
Marsupella spiniloba R.M.Schust. et Damsh................................................................................113
Marsupella sprucei (Limpr.) Bernet ..............................................................................................114
Marsupella stableri Spruce ...........................................................................................................114
Marsupella stoloniformis N.Kitag. ................................................................................................114
Marsupella stoloniformis subsp. vermiformis R.M.Schust. .............................................................114
Marsupella yakushimensis (Horik.) S.Hatt....................................................................................114
Mastigobryum aberrans Steph. .....................................................................................................510
Mastigobryum asperum Steph.......................................................................................................510
Mastigobryum deningeri Herzog...................................................................................................510
Mastigobryum karstenii Steph. .....................................................................................................510
Mastigobryum ledermannii Steph. ................................................................................................510
Mastigobryum londbergii Steph. ...................................................................................................510
Mastigobryum longifolium Steph. .................................................................................................510
Mastigobryum minutitextum Steph. .............................................................................................510
Mastigobryum multidens Steph. ...................................................................................................510
Mastigobryum muscicola Steph. ....................................................................................................510
Mastigobryum nigricans Herzog ...................................................................................................510
Mastigobryum nipuranum Steph. .................................................................................................510
Mastigobryum palmicola Steph.....................................................................................................510
Mastigobryum rajanum Herzog ...................................................................................................510
Mastigobryum ribehanum Steph...................................................................................................510
Mastigobryum ruicaule Beauverd ................................................................................................510
Mastigobryum schraderbergii Steph. .............................................................................................510
Mastigobryum squamulistipum Steph. ..........................................................................................510
Mastigobryum subhyalinum Steph. ...............................................................................................510
Mastigobryum venezuelanum Molk. .............................................................................................510
Mastigobryum vermiculare Herzog ...............................................................................................510
Mastigolejeunea auriculata (Wilson et Hook.) Steph....................................................................409
Mastigolejeunea auriculata var. rhodesica (Vanden Berghen) Sukkharak et Gradst. .......................409
Mastigolejeunea calcarata (Steph.) Verd. ......................................................................................409
Mastigolejeunea lorea (Mitt.) Paris ..............................................................................................409
Mastigolejeunea frauenfeldii (Reichardt) Verd. .............................................................................409
Mastigolejeunea gradsteinii Sukkharak .........................................................................................409
Mastigolejeunea humilis (Gottsche) Schifn. .................................................................................410
Mastigolejeunea indica Steph. ......................................................................................................410
Mastigolejeunea innovans (Spruce) Steph. ....................................................................................410
Mastigolejeunea ligulata (Lehm. et Lindenb.) Schifn...................................................................410
Mastigolejeunea nigra Steph. ........................................................................................................410
World checklist of hornworts and liverworts
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603
Mastigolejeunea plicatilora (Spruce) Steph. .................................................................................410
Mastigolejeunea recondita (Steph.) Mizut. ....................................................................................410
Mastigolejeunea repleta (Taylor) A.Evans .....................................................................................410
Mastigolejeunea truncata Mizut. ..................................................................................................410
Mastigolejeunea turgida Steph. .....................................................................................................410
Mastigolejeunea virens (Ångstr.) Steph. nom. conserv. ...................................................................410
Mastigopelma fragile (Steph.) N.Kitag. ........................................................................................163
Mastigopelma pulvinulatum (De Not.) Grolle ..............................................................................163
Mastigopelma simplex Mitt. .........................................................................................................163
Mastigopelma subissum Grolle .....................................................................................................163
Mastigophora appendiculata Steph. ..............................................................................................218
Mastigophora attenuata (Taylor) Trevis. ......................................................................................218
Mastigophora caledonica Steph. ....................................................................................................218
Mastigophora diclados (Brid. ex F.Weber) Nees ...........................................................................218
Mastigophora diclados var. borneensis (De Not.) Schifn. ..............................................................218
Mastigophora diclados var. ramentiissa Herzog.............................................................................218
Mastigophora diclados var. villosa Herzog .....................................................................................218
Mastigophora guineensis Steph. ....................................................................................................218
Mastigophora humillima (Taylor) Trevis. .....................................................................................218
Mastigophora pyramidana Steph. .................................................................................................218
Mastigophora sepikiana Piippo .....................................................................................................218
Mastigophora tuberculata D.H.Mill. et H.A.Mill. ........................................................................218
Mastigophora valida Steph. ..........................................................................................................218
Mastigophora viridula (Nees) Trevis. ...........................................................................................218
Mastigophora woodsii (Hook.) Nees.............................................................................................218
Megaceros aneuriformis Steph. .......................................................................................................34
Megaceros austronesophilus Cargill et Seppelt .................................................................................34
Megaceros ciliatus K.I.Goebel ........................................................................................................34
Megaceros denticulatus (Lehm.) Steph. ...........................................................................................34
Megaceros lagellaris (Mitt.) Steph..................................................................................................34
Megaceros lavens (Spruce) Campb. ...............................................................................................34
Megaceros gracilis (Reichardt) Steph. .............................................................................................34
Megaceros jamesonii (Taylor) Steph. ..............................................................................................34
Megaceros leptohymenius (Hook.f. et Taylor) Steph........................................................................34
Megaceros pellucidus (Colenso) E.A.Hodgs. ...................................................................................34
Megaceros tjibodensis Campb. ........................................................................................................34
Megalembidium insulanum (W.Martin et E.A.Hodgs.) R.M.Schust. ...........................................168
Meinungeria mouensis Frank Müll. ..............................................................................................143
Mesoceros mesophoros Piippo ..........................................................................................................38
Mesoceros porcatus Piippo ..............................................................................................................38
Mesoptychia badensis (Gottsche ex Rabenh.) L.Söderstr. et Váňa .................................................120
Mesoptychia bantriensis (Hook.) L.Söderstr. et Váňa ....................................................................120
Mesoptychia bantriensis subsp. wallischii (Ştefănuţ) L.Söderstr. et Váňa ......................................120
Mesoptychia chichibuensis (Inoue) L.Söderstr. et Váňa .................................................................120
Mesoptychia collaris (Nees) L.Söderstr. et Váňa ............................................................................120
Mesoptychia itzgeraldiae (Paton et A.R.Perry) L.Söderstr. et Váňa ..............................................121
Mesoptychia gillmanii (Austin) L.Söderstr. et Váňa ......................................................................121
Mesoptychia heterocolpos (hed. ex Hartm.) L.Söderstr. et Váňa...................................................121
Mesoptychia heterocolpos var. arctica (S.W.Arnell) L.Söderstr. et Váňa..........................................121
Mesoptychia heterocolpos var. harpanthoides (Bryhn et Kaal.) L.Söderstr. et Váňa..........................121
Mesoptychia igiana (S.Hatt.) L.Söderstr. et Váňa .........................................................................121
Mesoptychia mamatkulovii (Duda) L.Söderstr. et Váňa ................................................................121
604
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Mesoptychia mayebarae (S.Hatt.) L.Söderstr. et Váňa...................................................................121
Mesoptychia morrisoncola (Horik.) L.Söderstr. et Váňa ................................................................121
Mesoptychia polymorpha Stotler, Crand.-Stotl. et Bakalin ............................................................121
Mesoptychia rutheana (Limpr.) L.Söderstr. et Váňa......................................................................121
Mesoptychia rutheana var. laxa (Schifn. ex Burrell) L.Söderstr. et Váňa ......................................121
Mesoptychia sahlbergii (Lindb. et Arnell) A.Evans ........................................................................121
Mesoptychia subcrispa (Herzog) L.Söderstr. et Váňa .....................................................................121
Mesoptychia turbinata (Raddi) L.Söderstr. et Váňa ......................................................................122
Mesoptychia ussuriensis (Bakalin) L.Söderstr. et Váňa ...................................................................122
Metacalypogeia alternifolia (Nees) Grolle .....................................................................................107
Metacalypogeia cordifolia (Steph.) Inoue ......................................................................................107
Metalejeunea crassitexta (J.B.Jack et Steph.) Pócs .........................................................................300
Metalejeunea cucullata (Reinw., Blume et Nees) Grolle ...............................................................300
Metalejeunea winkleri R.L.Zhu et Grolle .....................................................................................300
Metzgeria acuminata Steph. ........................................................................................................457
Metzgeria adscendens Steph. ex K.I.Goebel ..................................................................................457
Metzgeria agnewiae Kuwah..........................................................................................................457
Metzgeria albinea Spruce .............................................................................................................457
Metzgeria albinea var. aberrans Schifn. .......................................................................................457
Metzgeria albinea var. angusta (Steph.) D.P.Costa et Gradst. ......................................................457
Metzgeria allionii Steph...............................................................................................................457
Metzgeria alpina R.M.Schust. et J.J.Engel ...................................................................................457
Metzgeria americana Masuzaki ....................................................................................................457
Metzgeria attenuata Steph. ..........................................................................................................457
Metzgeria aurantiaca Steph. ........................................................................................................457
Metzgeria auriculata Grolle et Kuwah. ........................................................................................457
Metzgeria bahiensis Schifn. .........................................................................................................457
Metzgeria bartlettii Kuwah. .........................................................................................................457
Metzgeria bischlerae Kuwah. ........................................................................................................457
Metzgeria bracteata Spruce ..........................................................................................................457
Metzgeria brasiliensis Schifn. ......................................................................................................457
Metzgeria chilensis Steph. ............................................................................................................457
Metzgeria ciliata Raddi................................................................................................................457
Metzgeria clavilora Spruce ..........................................................................................................457
Metzgeria cleeii Kuwah. ..............................................................................................................458
Metzgeria comata Steph. ..............................................................................................................458
Metzgeria conjugata Lindb...........................................................................................................458
Metzgeria consanguinea Schifn....................................................................................................458
Metzgeria convoluta Steph. ..........................................................................................................458
Metzgeria coorgensis S.C.Srivast. et S.Srivast. ...............................................................................458
Metzgeria corralensis Steph. .........................................................................................................458
Metzgeria crassipilis (Lindb.) A.Evans ..........................................................................................458
Metzgeria cratoneura Schifn. ......................................................................................................458
Metzgeria cylindra Kuwah. ..........................................................................................................458
Metzgeria decrescens Steph. ..........................................................................................................458
Metzgeria dichotoma (Sw.) Nees ..................................................................................................458
Metzgeria divaricata A.Evans.......................................................................................................458
Metzgeria dorsipara (Herzog) Kuwah. .........................................................................................458
Metzgeria duricosta Steph. ...........................................................................................................458
Metzgeria engelii Kuwah..............................................................................................................458
Metzgeria epiphylla A.Evans ........................................................................................................458
Metzgeria ilicina Mitt. ................................................................................................................458
World checklist of hornworts and liverworts
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605
Metzgeria lavovirens Colenso ......................................................................................................458
Metzgeria foliicola Schifn............................................................................................................458
Metzgeria francana Steph. ...........................................................................................................458
Metzgeria frontipilis Lindb. .........................................................................................................459
Metzgeria fruticola Spruce ...........................................................................................................459
Metzgeria furcata (L.) Corda .......................................................................................................459
Metzgeria furcata var. expansa Douin ..........................................................................................459
Metzgeria furcata var. paciica Brinkm. ........................................................................................459
Metzgeria grandilora A.Evans .....................................................................................................459
Metzgeria hasselii Kuwah. ............................................................................................................459
Metzgeria hebridensis Steph. ........................................................................................................459
Metzgeria hegewaldii Kuwah. ......................................................................................................459
Metzgeria herminieri Schifn........................................................................................................459
Metzgeria heteroramea Steph. ......................................................................................................459
Metzgeria imberbis J.B.Jack et Steph............................................................................................459
Metzgeria inlata Steph................................................................................................................459
Metzgeria jamesonii Kuwah. ........................................................................................................459
Metzgeria kanaii Kuwah..............................................................................................................459
Metzgeria kinabaluensis Masuzaki................................................................................................459
Metzgeria kuwaharae Piippo........................................................................................................459
Metzgeria laciniata Kuwah. .........................................................................................................459
Metzgeria lechleri Steph. ..............................................................................................................459
Metzgeria leptoneura Spruce ........................................................................................................459
Metzgeria leptoneura var. breviseta (Schifn.) O.Yano...................................................................459
Metzgeria leptoneura var. polychaeta R.M.Schust. ........................................................................459
Metzgeria liebmaniana Lindenb. et Gottsche ...............................................................................459
Metzgeria lindbergii Schifn. ........................................................................................................459
Metzgeria litoralis J.J.Engel et Kuwah..........................................................................................459
Metzgeria longitexta Steph. ..........................................................................................................460
Metzgeria macrospora Kuwah. .....................................................................................................460
Metzgeria macveanii Kuwah. .......................................................................................................460
Metzgeria maegdefraui Kuwah. ....................................................................................................460
Metzgeria magellanica Schifn......................................................................................................460
Metzgeria metaensis Kuwah. ........................................................................................................460
Metzgeria mexicana Steph. ..........................................................................................................460
Metzgeria monoica Kuwah. et J.J.Engel .......................................................................................460
Metzgeria myriopoda Lindb. ........................................................................................................460
Metzgeria neotropica Kuwah. .......................................................................................................460
Metzgeria nudifrons Steph. ..........................................................................................................460
Metzgeria parviinvolucrata Kuwah...............................................................................................460
Metzgeria patagonica Steph. ........................................................................................................460
Metzgeria polytricha Spruce .........................................................................................................460
Metzgeria procera Mitt. ...............................................................................................................460
Metzgeria psilocraspeda Schifn. ...................................................................................................460
Metzgeria pubescens (Schrank) Raddi ...........................................................................................460
Metzgeria pulvinata Steph. ..........................................................................................................460
Metzgeria quadrifaria Steph. .......................................................................................................460
Metzgeria raoi S.C.Srivast. et S.Srivast. .......................................................................................460
Metzgeria rigida Lindb. ...............................................................................................................460
Metzgeria robinsonii Steph. .........................................................................................................460
Metzgeria roivainenii Kuwah. ......................................................................................................461
Metzgeria rufula Spruce...............................................................................................................461
606
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Metzgeria saccata Mitt. ................................................................................................................461
Metzgeria saxbyi Pearson .............................................................................................................461
Metzgeria scobina Mitt. ...............................................................................................................461
Metzgeria scyphigera A.Evans .......................................................................................................461
Metzgeria senjoana Masuzaki .......................................................................................................461
Metzgeria setigera R.M.Schust. ex Crand.-Stotl. et L.Söderstr......................................................461
Metzgeria sikkimensis S.C.Srivast. et K.K.Rawat ..........................................................................461
Metzgeria simplex Lorb. ex Müll.Frib. .........................................................................................461
Metzgeria sinuata Loitl. ...............................................................................................................461
Metzgeria sparrei Kuwah. ............................................................................................................461
Metzgeria spindleri Steph.............................................................................................................461
Metzgeria subaneura Schifn. .......................................................................................................461
Metzgeria submarginata M.L.So ..................................................................................................461
Metzgeria subundulata (Lindb.) Kuwah. ......................................................................................461
Metzgeria temperata Kuwah. .......................................................................................................461
Metzgeria uncigera A.Evans .........................................................................................................461
Metzgeria undulata Kuwah..........................................................................................................461
Metzgeria violacea (Ach.) Dumort. ..............................................................................................461
Microlejeunea acutifolia Steph. ....................................................................................................385
Microlejeunea africana Steph. ......................................................................................................385
Microlejeunea aligera (Mitt.) Steph..............................................................................................385
Microlejeunea ankasica E.W.Jones ...............................................................................................385
Microlejeunea aphanella (Spruce) Steph. ......................................................................................385
Microlejeunea atsuana Steph. .......................................................................................................385
Microlejeunea bischlerae (B.M.hiers) B.M.hiers .......................................................................385
Microlejeunea bullata (Taylor) Steph. ..........................................................................................385
Microlejeunea byssoides (Gottsche) Pearson ..................................................................................385
Microlejeunea capillaris (Gottsche) Steph. ...................................................................................385
Microlejeunea cochlearifolia Steph. ...............................................................................................386
Microlejeunea colombiana Bischl. .................................................................................................386
Microlejeunea constricta (Grolle) Grolle .......................................................................................386
Microlejeunea crenulifolia (Gottsche) Steph. ................................................................................386
Microlejeunea cystifera Herzog .....................................................................................................386
Microlejeunea dispar Jovet-Ast .....................................................................................................386
Microlejeunea epiphylla Bischl......................................................................................................386
Microlejeunea ilicuspis (Steph.) Heinrichs, Schäf.-Verw., Pócs et S.Dong ...................................386
Microlejeunea ischeri (Tixier) Heinrichs, Schäf.-Verw., Pócs et S.Dong ......................................386
Microlejeunea issistipula Steph. ...................................................................................................386
Microlejeunea globosa (Spruce) Steph. ..........................................................................................386
Microlejeunea herzogiana Steph. ..................................................................................................386
Microlejeunea indica (Udar et U.S.Awasthi) Y.M.Wei et R.L.Zhu...............................................386
Microlejeunea inlata Steph. ........................................................................................................386
Microlejeunea kinabaluensis (Mizut.) Grolle ................................................................................386
Microlejeunea latitans (Hook.f. et Taylor) Heinrichs, Schäf.-Verw., Pócs et S.Dong ...................386
Microlejeunea lunulatiloba Horik. ...............................................................................................386
Microlejeunea magnilobula Gola ..................................................................................................386
Microlejeunea mammillosa (Mizut.) Grolle ..................................................................................387
Microlejeunea minutissima (Mizut.) Grolle ..................................................................................387
Microlejeunea minutistipula Steph. ..............................................................................................387
Microlejeunea moniliata (Mizut.) R.L.Zhu et Y.M.Wei ...............................................................387
Microlejeunea nepalensis Steph. ....................................................................................................387
Microlejeunea nyandaruensis Pócs ................................................................................................387
World checklist of hornworts and liverworts
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607
Microlejeunea oblongistipula (Gottsche) Pearson ..........................................................................387
Microlejeunea ocellata (Herzog) Grolle ........................................................................................387
Microlejeunea ovistipula Steph. ....................................................................................................387
Microlejeunea papulosa (Gottsche) Pearson ..................................................................................387
Microlejeunea perpusilla (Spruce) Steph. ......................................................................................387
Microlejeunea punctiformis (Taylor) Steph. ..................................................................................387
Microlejeunea pyriformis (Lindenb. et Gottsche) Steph. ...............................................................387
Microlejeunea spinosa (Mizut.) Grolle ..........................................................................................387
Microlejeunea squarrosa (Steph.) Heinrichs, Schäf.-Verw., Pócs et S.Dong ..................................387
Microlejeunea strasbergii Bardat et Ah-Peng .................................................................................387
Microlejeunea subulistipa Steph. ..................................................................................................388
Microlejeunea szechuanensis P.C.Chen .........................................................................................388
Microlejeunea udarii P.K.Verma et S.C.Srivast. ...........................................................................388
Microlejeunea ulicina (Taylor) Steph. ..........................................................................................388
Microlejeunea usambarensis Steph. ...............................................................................................388
Microlejeunea valenciana Steph. ..................................................................................................388
Microlejeunea victoriensis D.J.Carr...............................................................................................388
Microlejeunea wallichiana (Lehm.) R.L.Zhu et Y.M.Wei ............................................................388
Micropterygium angustistipulum Spruce .......................................................................................181
Micropterygium bialatum Fulford ................................................................................................181
Micropterygium bolivarense Fulford .............................................................................................181
Micropterygium campanense Spruce .............................................................................................181
Micropterygium carinatum (Grev.) Reimers .................................................................................181
Micropterygium conchifolium Reimers ..........................................................................................181
Micropterygium duidae Reimers ...................................................................................................181
Micropterygium exalatum Steph. ..................................................................................................181
Micropterygium grandistipulum Steph. .........................................................................................181
Micropterygium laeve H.Rob. ......................................................................................................181
Micropterygium lechleri Reimers ..................................................................................................181
Micropterygium leiophyllum Spruce ..............................................................................................181
Micropterygium longicellulatum Uribe et E.L.Linares ...................................................................182
Micropterygium parvistipulum Spruce ..........................................................................................182
Micropterygium pterygophyllum (Nees) Trevis. .............................................................................182
Micropterygium pterygophyllum var. lancifolium Reimers ..............................................................182
Micropterygium reimersianum Herzog..........................................................................................182
Micropterygium steyermarkii Fulford ............................................................................................182
Micropterygium tatei Reimers ......................................................................................................182
Micropterygium tenax (Steph.) Grolle ..........................................................................................182
Micropterygium trachyphyllum Reimers ........................................................................................182
Micropterygium trachyphyllum var. brasiliense Reimers .................................................................182
Micropterygium trachyphyllum var. cubense Reimers .....................................................................182
Micropterygium trachyphyllum var. guadeloupense Reimers ...........................................................182
Micropterygium trachyphyllum var. jamaicense Reimers ................................................................182
Micropterygium tumidulum Fulford .............................................................................................182
Mizutania riccardioides Furuki et Z.Iwats....................................................................................107
Mnioloma bolivianum (Fulford) R.M.Schust. ..............................................................................108
Mnioloma caespitosum (Spruce) R.M.Schust. ...............................................................................108
Mnioloma cellulosum (Spreng.) R.M.Schust. ...............................................................................108
Mnioloma crenulatum (Bischl.) R.M.Schust. ...............................................................................108
Mnioloma cyclostipum (Spruce) R.M.Schust. ...............................................................................108
Mnioloma elliottii (Steph.) R.M.Schust. ......................................................................................108
Mnioloma issistipulum (Bischl.) R.M.Schust. ..............................................................................108
608
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Mnioloma fuscum (Lehm.) R.M.Schust. ......................................................................................108
Mnioloma nephrostipum (Spruce) R.M.Schust. ............................................................................108
Mnioloma novaezelandiae J.J.Engel .............................................................................................108
Mnioloma parallelogramum (Spruce) R.M.Schust. .......................................................................108
Mnioloma retusum (Bischl.) R.M.Schust. ....................................................................................108
Mnioloma rhynchophyllum Herzog ..............................................................................................108
Mnioloma stamatotonum M.A.M.Renner et E.A.Br. ....................................................................108
Mnioloma venezuelanum (Fulford) R.M.Schust. ..........................................................................108
Moerckia blyttii (Mørch) Brockm. ...............................................................................................469
Moerckia lotoviana (Nees) Schifn. .............................................................................................469
Moerckia hibernica (Hook.) Gottsche ..........................................................................................469
Monocarpus sphaerocarpus D.J.Carr .............................................................................................504
Monoclea forsteri Hook................................................................................................................492
Monoclea gottschei Lindb. ............................................................................................................492
Monoclea gottschei subsp. elongata Gradst. et Mues ......................................................................492
Monodactylopsis monodactyla (Spruce) R.M.Schust. .....................................................................183
Monosolenium tenerum Grif. ......................................................................................................492
Mylia anomala (Hook.) Gray ......................................................................................................258
Mylia taylorii (Hook.) Gray ........................................................................................................258
Mylia verrucosa Lindb. ................................................................................................................258
Mylia verrucosa subsp. nuda (Inoue et B.Y.Yang) Potemkin et Kazan. .........................................259
Myriocoleopsis luviatilis (Steph.) M.E.Reiner et Gradst. ..............................................................348
Myriocoleopsis gymnocolea (Spruce) M.E.Reiner et Gradst. ...........................................................348
Myriocoleopsis minutissima (Sm.) R.L.Zhu, Y.Yu et Pócs .............................................................348
Myriocoleopsis minutissima subsp. myriocarpa (Nees et Mont.) R.L.Zhu, Y.Yu et Pócs .................348
Myriocoleopsis vuquangensis (Pócs et Ninh) Pócs ..........................................................................349
Mytilopsis albifrons Spruce ...........................................................................................................182
Nanomarsupella xenophylla (R.M.Schust.) R.M.Schust. ex A.Hagborg, L.Söderstr. et von Konrat......110
Nardia arnelliana Grolle .............................................................................................................114
Nardia assamica (Mitt.) Amakawa...............................................................................................114
Nardia breidleri (Limpr.) Lindb. .................................................................................................114
Nardia compressa (Hook.) Gray...................................................................................................114
Nardia lagelliformis Inoue ..........................................................................................................114
Nardia geoscyphus (De Not.) Lindb. ............................................................................................114
Nardia geoscyphus var. dioica Bakalin ...........................................................................................114
Nardia geoscyphus var. suberecta (Lindb. ex Kaal.) Váňa ...............................................................114
Nardia grollei Váňa et D.G.Long ................................................................................................115
Nardia insecta Lindb. ..................................................................................................................115
Nardia japonica Steph. ................................................................................................................115
Nardia kamtschatica Arnell et C.E.O.Jensen ...............................................................................115
Nardia leptocaulis C.Gao.............................................................................................................115
Nardia lescurii (Austin) Underw. ................................................................................................115
Nardia minutifolia Furuki ...........................................................................................................115
Nardia nuda (Lindenb. et Gottsche) Váňa ..................................................................................115
Nardia poeltii Váňa .....................................................................................................................115
Nardia scalaris Gray ....................................................................................................................115
Nardia scalaris var. botryoidea (R.M.Schust.) Váňa ......................................................................115
Nardia scalaris var. harae (Amakawa) Váňa .................................................................................115
Nardia subclavata (Steph.) Amakawa ..........................................................................................115
Nardia succulenta (A.Rich.) Spruce .............................................................................................115
Nardia unispiralis Amakawa ........................................................................................................115
Neesioscyphus allionii (Steph.) Grolle ...........................................................................................102
World checklist of hornworts and liverworts
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609
Neesioscyphus argillaceus (Nees) Grolle.........................................................................................102
Neesioscyphus bicuspidatus (Steph.) Grolle....................................................................................102
Neesioscyphus carneus (Nees) Grolle .............................................................................................102
Neesioscyphus homophyllus (Nees) Grolle ......................................................................................102
Nemoursia tuberculata Mérat .......................................................................................................511
Neogrollea notabilis E.A.Hodgs....................................................................................................183
Neohattoria herzogii (S.Hatt.) Kamim. ........................................................................................296
Neohodgsonia mirabilis (Perss.) Perss. ..........................................................................................504
Neolepidozia aubertii (Jovet-Ast) E.D.Cooper .............................................................................176
Neolepidozia autoica (J.J.Engel et G.L.Merr.) E.D.Cooper ..........................................................176
Neolepidozia capilligera (Schwägr.) Fulford et J.Taylor ................................................................176
Neolepidozia consobrina (J.J.Engel et G.L.Merr.) E.D.Cooper .....................................................176
Neolepidozia cuneifolia (Steph.) Fulford et J.Taylor .....................................................................177
Neolepidozia disparata (J.J.Engel et G.L.Merr.) E.D.Cooper .......................................................177
Neolepidozia disticha (Steph.) Fulford et J.Taylor ........................................................................177
Neolepidozia gibbsiana (Steph.) E.D.Cooper ...............................................................................177
Neolepidozia heterotexta (Steph.) E.D.Cooper .............................................................................177
Neolepidozia hodgsoniae (J.J.Engel et G.L.Merr.) E.D.Cooper .....................................................177
Neolepidozia longitudinalis (Herzog) E.D.Cooper .......................................................................177
Neolepidozia mamillosa (Schifn.) E.D.Cooper ............................................................................177
Neolepidozia meridiana (E.A.Hodgs.) E.D.Cooper......................................................................177
Neolepidozia oligophylla (Lehm. et Lindenb.) Fulford et J.Taylor ................................................177
Neolepidozia ophiria (Gottsche) E.D.Cooper ..............................................................................177
Neolepidozia palmata (J.J.Engel et G.L.Merr.) E.D.Cooper ........................................................177
Neolepidozia paludicola (E.A.Hodgs.) E.D.Cooper ......................................................................177
Neolepidozia papulosa (Steph.) Fulford et J.Taylor ......................................................................177
Neolepidozia parvifolia (Steph.) Fulford et J.Taylor .....................................................................178
Neolepidozia patentissima (Hook.f. et Taylor) E.D.Cooper..........................................................178
Neolepidozia patentissima var. ampliata (J.J.Engel et G.L.Merr.) E.D.Cooper .............................178
Neolepidozia patentissima var. zebrina (J.J.Engel et G.L.Merr.) E.D.Cooper................................178
Neolepidozia pennata (J.J.Engel et G.L.Merr.) E.D.Cooper .........................................................178
Neolepidozia planifolia (Steph.) E.D.Cooper ...............................................................................178
Neolepidozia praenitens (Lehm. et Lindenb.) E.D.Cooper ...........................................................178
Neolepidozia praenitens var. dentifolia (J.J.Engel et G.L.Merr.) E.D.Cooper ................................178
Neolepidozia quadristipula (Steph.) E.D.Cooper .........................................................................178
Neolepidozia rectangularis (R.M.Schust.) E.D.Cooper .................................................................178
Neolepidozia seriatitexta (Steph.) Fulford.....................................................................................178
Neolepidozia tetrapila (Hook.f. et Taylor) E.D.Cooper ...............................................................178
Neolepidozia tetrapila var. cancellata (Colenso) E.D.Cooper ........................................................178
Neolepidozia tetrapila var. roseana (Steph.) E.D.Cooper ..............................................................179
Neolepidozia tridactylis (Lehm. et Lindenb.) E.D.Cooper ............................................................179
Neolepidozia triida (Steph.) E.D.Cooper ....................................................................................179
Neolepidozia verruculosa (J.J.Engel et G.L.Merr.) E.D.Cooper ....................................................179
Neolepidozia wallichiana (Gottsche) Fulford et J.Taylor ..............................................................179
Neoorthocaulis attenuatus (Mart.) L.Söderstr., De Roo et Hedd.....................................................53
Neoorthocaulis binsteadii (Kaal.) L.Söderstr., De Roo et Hedd. .....................................................53
Neoorthocaulis loerkei (F.Weber et D.Mohr) L.Söderstr., De Roo et Hedd. ..................................53
Neoorthocaulis hyperboreus (R.M.Schust.) L.Söderstr., De Roo et Hedd. .......................................53
Neotrichocolea bissetii (Mitt.) S.Hatt............................................................................................434
Nephelolejeunea bidentata B.M.hiers ex L.Söderstr. et A.Hagborg .............................................349
Nephelolejeunea carcharias M.A.M.Renner ..................................................................................349
Nephelolejeunea conchophylla Grolle ............................................................................................349
610
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Nephelolejeunea fragilis (R.M.Schust.) L.Söderstr. et A.Hagborg .................................................349
Nephelolejeunea hamata Grolle ....................................................................................................349
Nephelolejeunea hispida R.M.Schust. ex L.Söderstr. et A.Hagborg ...............................................349
Nephelolejeunea jarmaniana Grolle ex L.Söderstr. et A.Hagborg .................................................349
Nephelolejeunea nudipes (Hook.f. et Taylor) L.Söderstr. et A.Hagborg ........................................349
Nephelolejeunea occidentalis Pócs ex L.Söderstr. et A.Hagborg .....................................................349
Nephelolejeunea papillosa Glenny.................................................................................................349
Nephelolejeunea radulifolia (C.Massal.) L.Söderstr. et A.Hagborg ................................................349
Nephelolejeunea secunda M.A.M.Renner ex L.Söderstr. et A.Hagborg .........................................349
Nephelolejeunea talinayi (S.W.Arnell) Grolle ...............................................................................349
Neurolejeunea breutelii (Gottsche) A.Evans .................................................................................299
Neurolejeunea breutelii var. africana Pócs.....................................................................................299
Neurolejeunea catenulata (Nees) Schifn. .....................................................................................299
Neurolejeunea sastreana Gradst. ...................................................................................................299
Neurolejeunea seminervis (Spruce) Schifn....................................................................................300
Nipponolejeunea pilifera (Steph.) S.Hatt. .....................................................................................296
Nipponolejeunea subalpina (Horik.) S.Hatt..................................................................................296
Noteroclada conluens Taylor .......................................................................................................474
Noteroclada longiuscula Colenso ..................................................................................................474
Nothoceros aenigmaticus J.C.Villarreal et K.D.McFarland ..............................................................35
Nothoceros canaliculatus (Pagán) J.C.Villarreal, Hässel et N.Salazar ...............................................35
Nothoceros endiviifolius (Mont.) J.Haseg. ex J.C.Villarreal, Hässel et N.Salazar .............................35
Nothoceros fuegiensis (Steph.) J.C.Villarreal....................................................................................35
Nothoceros giganteus (Lehm. et Lindenb.) J.Haseg. ex J.C.Villarreal, Hässel et N.Salazar ...............35
Nothoceros minarum (Nees) J.C.Villarreal .....................................................................................35
Nothoceros renzagliensis J.C.Villarreal, L.V.Campos et Uribe .........................................................35
Nothoceros schizophyllus (Steph.) J.C.Villarreal ..............................................................................35
Nothoceros superbus J.C.Villarreal, Hässel et N.Salazar...................................................................35
Nothoceros vincentianus (Lehm. et Lindenb.) J.C.Villarreal............................................................35
Nothogymnomitrion erosum (Carrington et Pearson) R.M.Schust...................................................75
Nothostrepta biida (Steph.) R.M.Schust........................................................................................46
Nothostrepta longissima (Steph.) R.M.Schust. ................................................................................46
Notoscyphus lutescens (Lehm. et Lindenb.) Mitt. ..........................................................................122
Notothylas anaporata Udar et D.K.Singh .......................................................................................37
Notothylas breutelii (Gottsche) Gottsche........................................................................................37
Notothylas decurva (Mitt.) Steph. ..................................................................................................37
Notothylas depressispora J.Haseg. ....................................................................................................37
Notothylas dissecta Steph. ...............................................................................................................37
Notothylas labellata Steph. ............................................................................................................37
Notothylas galapagensis M.Howe....................................................................................................37
Notothylas himalayensis Udar et D.K.Singh ...................................................................................37
Notothylas indica Kashyap .............................................................................................................37
Notothylas irregularis Chantanaorr.................................................................................................37
Notothylas javanica (Sande Lac.) Gottsche.....................................................................................37
Notothylas kashyapii D.K.Singh .....................................................................................................38
Notothylas khasiana Udar et D.K.Singh.........................................................................................37
Notothylas nepalensis D.K.Singh ....................................................................................................38
Notothylas orbicularis (Schwein.) Sull. ...........................................................................................37
Notothylas pandei Udar et V.Chandra ...........................................................................................37
Notothylas pleidereri Udar et D.K.Singh .......................................................................................37
Notothylas temperata J.Haseg. ........................................................................................................38
Notothylas udarii D.K.Singh et Semwal.........................................................................................38
World checklist of hornworts and liverworts
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611
Notothylas verdoornii Khanna ........................................................................................................38
Notothylas vitalii Udar et D.K.Singh .............................................................................................38
Notothylas yunnanensis T.Peng et R.L.Zhu ....................................................................................38
Nowellia aciliata (P.C.Chen et P.C.Wu) Mizut. ...........................................................................62
Nowellia borneensis (De Not.) Schifn. ..........................................................................................61
Nowellia curvifolia (Dicks.) Mitt. ..................................................................................................62
Nowellia dominicensis Steph. .........................................................................................................61
Nowellia evansii Grolle..................................................................................................................61
Nowellia langii Pearson .................................................................................................................61
Nowellia pusilla Grolle ..................................................................................................................61
Nowellia reedii H.Rob...................................................................................................................61
Nowellia wrightii (Gottsche ex Spruce) Steph. ex Duss .................................................................61
Nowellia yunckeri Fulford .............................................................................................................61
Obtusifolium obtusum (Lindb.) S.W.Arnell ...................................................................................75
Odontolejeunea decemdentata (Spruce) Steph. ..............................................................................229
Odontolejeunea lunulata (F.Weber) Schifn. ................................................................................229
Odontolejeunea rhomalea (Spruce) Steph. ....................................................................................229
Odontoschisma biidum (Fulford) Gradst., S.C.Aranda et Vanderp. ...............................................63
Odontoschisma brasiliense Steph. ....................................................................................................62
Odontoschisma cleeii Gradst., S.C.Aranda et Vanderp. ..................................................................62
Odontoschisma denudatum (Mart.) Dumort. .................................................................................62
Odontoschisma denudatum subsp. naviculare (Steph.) Gradst., S.C.Aranda et Vanderp..................62
Odontoschisma denudatum subsp. sandvicense (Ångstr.) Gradst., S.C.Aranda et Vanderp. ..............62
Odontoschisma elongatum (Lindb.) A.Evans ...................................................................................62
Odontoschisma engelii Gradst. et Burghardt ...................................................................................62
Odontoschisma luitans (Nees) L.Söderstr. et Váňa .........................................................................63
Odontoschisma francisci (Hook.) L.Söderstr. et Váňa .....................................................................62
Odontoschisma grosseverrucosum Steph. ..........................................................................................63
Odontoschisma jishibae (Steph.) L.Söderstr. et Váňa ......................................................................63
Odontoschisma longilorum (Taylor) Trevis. ...................................................................................63
Odontoschisma macounii (Austin) Underw. ...................................................................................63
Odontoschisma obcordatum (Spruce) Steph. ...................................................................................64
Odontoschisma portoricense (Hampe et Gottsche) Steph. ................................................................63
Odontoschisma pseudogrosseverrucosum Gradst., S.C.Aranda et Vanderp.........................................63
Odontoschisma purpuratum Herzog ...............................................................................................63
Odontoschisma soratamum Fulford ................................................................................................63
Odontoschisma sphagni (Dicks.) Dumort. ......................................................................................64
Odontoschisma spinosum (Fulford) Gradst., S.C.Aranda et Vanderp. .............................................63
Odontoschisma steyermarkii Gradst. et Ilk.-Borg. ...........................................................................63
Odontoschisma variabile (Lindenb. et Gottsche) Trevis..................................................................63
Odontoschisma zhui Gradst., S.C.Aranda et Vanderp. ...................................................................63
Odontoseries chimantana Fulford .................................................................................................183
Oleolophozia perssonii (H.Buch et S.W.Arnell) L.Söderstr., De Roo et Hedd. ...............................75
Omphalanthus baracoensis Mustelier, M.E.Reiner et Gradst. .......................................................316
Omphalanthus iliformis (Sw.) Nees .............................................................................................316
Omphalanthus iliformis var. platycoleus (Herzog) Gradst. ............................................................316
Omphalanthus iliformis var. wallisii (Prantl) Gradst. ...................................................................316
Omphalanthus huanucensis (Gottsche) Gradst. ............................................................................316
Omphalanthus jackii (Prantl) Gradst. ..........................................................................................316
Omphalanthus ovalis (Lindenb. et Gottsche) Gradst. ...................................................................316
Omphalanthus roccatii (Gola) R.M.Schust...................................................................................316
Orthocaulis atlanticus (Kaal.) H.Buch............................................................................................53
612
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Orthocaulis cavifolius H.Buch et S.W.Arnell..................................................................................53
Otigoniolejeunea crenulata Steph. ................................................................................................304
Otigoniolejeunea huctumalcensis (Lindenb. et Gottsche) Y.M.Wei, R.L.Zhu et Gradst. ...............304
Otigoniolejeunea ledermannii Steph. ............................................................................................304
Otigoniolejeunea portoricensis (Hampe et Gottsche) Y.M.Wei, R.L.Zhu et Gradst. ......................304
Otolejeunea australiensis B.M.hiers ............................................................................................392
Otolejeunea hoana (Tixier) Grolle................................................................................................392
Otolejeunea moniliata Grolle .......................................................................................................393
Otolejeunea philippinensis R.L.Zhu et M.L.So .............................................................................393
Otolejeunea rabenorii Tixier ........................................................................................................392
Otolejeunea schmidii (Tixier) Grolle ............................................................................................392
Otolejeunea schnellii (Tixier) R.L.Zhu et M.L.So.........................................................................393
Otolejeunea semperiana (Steph.) Grolle........................................................................................393
Otolejeunea streimannii Grolle.....................................................................................................393
Otolejeunea subana Pócs ..............................................................................................................393
Otolejeunea zantenii Grolle .........................................................................................................393
Otoscyphus crassicaulis (Steph.) J.J.Engel, Bardat et houvenot ....................................................216
Oxymitra cristata Garside ............................................................................................................493
Oxymitra incrassata (Brot.) Sérgio et Sim-Sim .............................................................................493
Pachyglossa austrigena (Hook.f. et Taylor) L.Söderstr. .................................................................216
Pachyglossa austrigena subsp. okaritana (Steph.) L.Söderstr. .........................................................216
Pachyglossa boveana (C.Massal.) L.Söderstr. ................................................................................216
Pachyglossa dissitifolia Herzog et Grolle .......................................................................................216
Pachyglossa exilis (Herzog et Grolle) Hässel et Solari ....................................................................216
Pachyglossa issa (Mitt.) Herzog et Grolle .....................................................................................216
Pachyglossa gottscheoides (Besch. et C.Massal.) L.Söderstr.............................................................216
Pachyglossa grolleana Váňa ...........................................................................................................216
Pachyglossa otiphylla (Hook.f. et Taylor) Váňa.............................................................................216
Pachyglossa spegazziniana (C.Massal.) Herzog et Grolle ...............................................................216
Pachyglossa tenacifolia (Hook.f. et Taylor) Herzog et Grolle ........................................................216
Pallavicinia ambigua (Mitt.) Steph..............................................................................................470
Pallavicinia baldwinii (Austin) A.Evans ......................................................................................470
Pallavicinia bipinnata Steph........................................................................................................470
Pallavicinia camisassai Gola.........................................................................................................470
Pallavicinia cylindrica (Austin) A.Evans ......................................................................................470
Pallavicinia himalayensis Schifn..................................................................................................470
Pallavicinia indica Schifn. ..........................................................................................................470
Pallavicinia levieri Schifn. ..........................................................................................................470
Pallavicinia lyellii (Hook.) Gray ..................................................................................................470
Pallavicinia pilifera Steph............................................................................................................470
Pallavicinia purpurea Steph. ........................................................................................................470
Pallavicinia ridleyi Steph. ............................................................................................................471
Pallavicinia rubristipa Schifn......................................................................................................471
Pallavicinia subciliata (Austin) Steph. .........................................................................................471
Pallavicinia xiphoides (Hook.f. et Taylor) Trevis. ........................................................................471
Paracromastigum denticulatum (Steph.) E.D.Cooper ...................................................................183
Paracromastigum drucei (R.M.Schust.) R.M.Schust.....................................................................184
Paracromastigum dusenii (Steph.) R.M.Schust. ............................................................................184
Paracromastigum iordlandiae R.M.Schust. et J.J.Engel................................................................184
Paracromastigum furcifolium (Steph.) R.M.Schust. ......................................................................184
Paracromastigum granatense (Gottsche) R.M.Schust. ...................................................................184
Paracromastigum longiscyphum (Taylor) R.M.Schust. et J.J.Engel ................................................184
World checklist of hornworts and liverworts
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613
Paracromastigum macrostipum (Steph.) R.M.Schust. ...................................................................184
Paracromastigum micromera (Spruce) R.M.Schust. ......................................................................184
Paracromastigum microphyllum (R.M.Schust. ex J.J.Engel) E.D.Cooper ......................................184
Paracromastigum pachyrhizum (Nees) Fulford .............................................................................184
Paracromastigum ryszardii Váňa, Bedn.-Ochyra et Cykowska......................................................184
Paracromastigum stipulatum (Herzog) Fulford.............................................................................184
Paracromastigum subsimplex (Steph.) Fulford et J.Taylor .............................................................184
Paracromastigum succulentum (Sim) J.J.Engel et G.L.Merr. .........................................................184
Paracromastigum tristanianum (R.M.Schust.) J.J.Engel et R.M.Schust. .......................................184
Paracromastigum vastilobum (Steph.) J.J.Engel et G.L.Merr. .......................................................185
Paramomitrion paradoxum R.M.Schust. ......................................................................................110
Paraphymatoceros diadematus Hässel..............................................................................................38
Paraphymatoceros hallii (Austin) Hässel .........................................................................................38
Paraphymatoceros pearsonii (M.Howe) J.C.Villarreal et Cargill ......................................................38
Paraphymatoceros proskaueri (Stotler, Crand.-Stotl. et W.T.Doyle) J.C.Villarreal et Cargill...........38
Pedinophyllopsis abdita (Sull.) R.M.Schust. et Inoue....................................................................219
Pedinophyllum autoicum (Steph.) Inoue ......................................................................................219
Pedinophyllum interruptum (Nees) Kaal. .....................................................................................219
Pedinophyllum monoicum (Steph.) Grolle ....................................................................................219
Pedinophyllum truncatum (Steph.) Inoue.....................................................................................219
Pellia alpicola R.M.Schust. ex L.Söderstr., A.Hagborg et von Konrat ..........................................475
Pellia appalachiana R.M.Schust. .................................................................................................475
Pellia cordaeana Trevis. ...............................................................................................................475
Pellia crispa P.Kumm. .................................................................................................................475
Pellia endiviifolia (Dicks.) Dumort. ............................................................................................475
Pellia epiphylla (L.) Corda ...........................................................................................................475
Pellia epiphylla subsp. borealis (Lorb.) Messe ...............................................................................475
Pellia gottscheana Kreh ................................................................................................................475
Pellia longifolia P.Kumm.............................................................................................................475
Pellia megaspora R.M.Schust. ......................................................................................................475
Pellia neesiana (Gottsche) Limpr. ................................................................................................475
Pellia undulata P.Kumm. ............................................................................................................475
Peltolepis japonica (Shimizu et S.Hatt.) S.Hatt. ...........................................................................484
Peltolepis quadrata (Saut.) Müll.Frib. ..........................................................................................484
Perdusenia rheophila Hässel .........................................................................................................217
Petalophyllum americanum C.H.Ford et Crand.-Stotl. ................................................................468
Petalophyllum hodgsoniae C.H.Ford et Crand.-Stotl. ...................................................................468
Petalophyllum indicum Kashyap ..................................................................................................468
Petalophyllum preissii Gottsche ....................................................................................................468
Petalophyllum ralfsii (Wilson) Nees et Gottsche ..........................................................................468
Phaeoceros austroandinus Hässel.....................................................................................................38
Phaeoceros bolusii (Sim) S.W.Arnell...............................................................................................38
Phaeoceros brevicapsulus (Steph.) Hässel.........................................................................................39
Phaeoceros carolinianus (Michx.) Prosk. .........................................................................................39
Phaeoceros delicatus E.O.Campb. et Outred ..................................................................................39
Phaeoceros dendroceroides (Steph.) Hässel.......................................................................................39
Phaeoceros engelii Cargill et Fuhrer ................................................................................................39
Phaeoceros erectus Udar et D.K.Singh ............................................................................................39
Phaeoceros evanidus (Steph.) Cargill et Fuhrer ...............................................................................39
Phaeoceros exiguus (Steph.) J.Haseg. ..............................................................................................39
Phaeoceros lexivalvis (Gottsche et Nees) Hässel .............................................................................39
Phaeoceros fulvisporus (Steph.) J.Haseg. .........................................................................................39
614
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Phaeoceros gemmifer (Horik.) J.Haseg. ...........................................................................................39
Phaeoceros gualaquizanus (Steph.) Gradst. .....................................................................................39
Phaeoceros himalayensis (Kashyap) Prosk. ex Bapna et G.G.Vyas ...................................................39
Phaeoceros huebschmannii Hässel ...................................................................................................39
Phaeoceros inlatus (Steph.) Cargill et Fuhrer .................................................................................39
Phaeoceros kashyapii A.K.Asthana et S.C.Srivast. ...........................................................................40
Phaeoceros laevis (L.) Prosk. ...........................................................................................................40
Phaeoceros maranguensis (Steph.) Bapna ........................................................................................40
Phaeoceros microsporus (Steph.) Hässel ...........................................................................................40
Phaeoceros minutus (Mitt.) S.W.Arnell ..........................................................................................40
Phaeoceros mohrii (Austin) Hässel..................................................................................................40
Phaeoceros oreganus (Austin) Hässel ...............................................................................................40
Phaeoceros parvulus (Schifn.) J.Haseg. ..........................................................................................40
Phaeoceros perpusillus Chantanaorr. ...............................................................................................40
Phaeoceros pichinchensis (Spruce) Hässel ........................................................................................40
Phaeoceros propagulifer (Steph.) Prosk. ..........................................................................................40
Phaeoceros striatisporus J.Haseg ......................................................................................................40
Phaeoceros tenuis (Spruce) Hässel...................................................................................................40
Phaeoceros tigrinus (Gola) J.C.Villarreal.........................................................................................40
Phaeoceros tuberosus (Taylor) Prosk. ..............................................................................................40
Phaeoceros udarii A.K.Asthana et V.Nath ......................................................................................40
Phaeoceros wrightii (Steph.) Hässel ................................................................................................40
Phaeolejeunea amicorum (Hürl.) Pócs ..........................................................................................410
Phaeolejeunea etesseana (Steph.) Mizut. .......................................................................................410
Phaeolejeunea inermis (Steph.) Mizut. .........................................................................................410
Phaeolejeunea latistipula (Schifn. ex P.Syd.) Mizut. ....................................................................410
Phaeomegaceros coriaceus (Steph.) R.J.Duf, J.C.Villarreal, Cargill et Renzaglia .............................35
Phaeomegaceros imbriatus (Gottsche) R.J.Duf, J.C.Villarreal, Cargill et Renzaglia .......................35
Phaeomegaceros foveatus (J.Haseg.) J.C.Villarreal ...........................................................................35
Phaeomegaceros hirticalyx (Steph.) R.J.Duf, J.C.Villarreal, Cargill et Renzaglia.............................36
Phaeomegaceros plicatus (Mitt.) J.C.Villarreal, J.J.Engel et Váňa ....................................................36
Phaeomegaceros skottsbergii (Steph.) R.J.Duf, J.C.Villarreal, Cargill et Renzaglia ..........................36
Phaeomegaceros squamuliger (Spruce) J.C.Villarreal .......................................................................36
Phaeomegaceros squamuliger subsp. hasselii J.C.Villarreal, Cargill et Goinet .................................36
Phycolepidozia exigua R.M.Schust. ................................................................................................75
Phycolepidozia indica Gradst., J.-P.Frahm et U.Schwarz ................................................................75
Phyllothallia fuegiana R.M.Schust. ..............................................................................................474
Phyllothallia nivicola E.A.Hodgs. ................................................................................................474
Phymatoceros bulbiculosus (Brot.) Stotler, W.T.Doyle et Crand.-Stotl. ..........................................36
Phymatoceros phymatodes (M.Howe) R.J.Duf, J.C.Villarreal, Cargill et Renzaglia .........................36
Physocolea tambillensis (Loitl.) Steph. ..........................................................................................511
Pictolejeunea levis Grolle et M.E.Reiner.......................................................................................300
Pictolejeunea mizutanii Grolle .....................................................................................................301
Pictolejeunea piconii Pócs ............................................................................................................301
Pictolejeunea picta (Steph.) Grolle ...............................................................................................301
Pictolejeunea reginae Ilk.-Borg. ....................................................................................................301
Pictolejeunea sprucei Grolle ..........................................................................................................301
Pigafettoa crenulata C.Massal. .....................................................................................................217
Pisanoa chilensis Hässel .................................................................................................................46
Plagiochasma appendiculatum Lehm. et Lindenb. ........................................................................482
Plagiochasma argentinicum Bischl. ...............................................................................................482
Plagiochasma beccarianum Steph. ................................................................................................483
World checklist of hornworts and liverworts
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615
Plagiochasma cordatum Lehm. et Lindenb. ..................................................................................483
Plagiochasma crenulatum Gottsche ..............................................................................................483
Plagiochasma cuneatum A.Evans ..................................................................................................483
Plagiochasma eximium (Schifn.) Steph. ......................................................................................483
Plagiochasma intermedium Lindenb. et Gottsche .........................................................................483
Plagiochasma jamaicense (Haynes) A.Evans..................................................................................483
Plagiochasma japonicum (Steph.) C.Massal. .................................................................................483
Plagiochasma landii A.Evans........................................................................................................483
Plagiochasma megacarpon (Grif.) Steph. .....................................................................................483
Plagiochasma microcephalum (Steph.) Steph. ...............................................................................483
Plagiochasma microcephalum var. tunesicum Bischl.......................................................................483
Plagiochasma muenchianum Steph. ..............................................................................................483
Plagiochasma pterospermum C.Massal. .........................................................................................483
Plagiochasma rupestre (J.R.Forst. et G.Forst.) Steph. ...................................................................483
Plagiochasma rupestre var. volkii Bischl. .......................................................................................483
Plagiochasma udarii A.Alam et S.C.Srivast. .................................................................................483
Plagiochasma wrightii Sull. ..........................................................................................................483
Plagiochila abietina (Nees) Mont. et Nees ...................................................................................232
Plagiochila abrupta Lehm. et Lindenb. ........................................................................................232
Plagiochila abscendens Gottsche ...................................................................................................238
Plagiochila acanthocaulis Sull. ......................................................................................................224
Plagiochila aculeata (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ...........................................238
Plagiochila acuta Steph. ...............................................................................................................238
Plagiochila adianthoides (Sw.) Lindenb. .......................................................................................220
Plagiochila adianthoides var. aspergillifera Spruce .........................................................................220
Plagiochila aequatorialis Gottsche ................................................................................................233
Plagiochila aerea Taylor ...............................................................................................................225
Plagiochila africana Steph............................................................................................................233
Plagiochila akiyamae Inoue..........................................................................................................233
Plagiochila albertii Steph. ............................................................................................................238
Plagiochila aliena Gottsche ..........................................................................................................238
Plagiochila allionii Steph. ............................................................................................................238
Plagiochila alternans Lindenb. et Gottsche ..................................................................................223
Plagiochila ambigua Lindenb. et Hampe .....................................................................................238
Plagiochila amboynensis Taylor ....................................................................................................226
Plagiochila amicta Steph. .............................................................................................................226
Plagiochila ampliata Steph...........................................................................................................238
Plagiochila andicola Mont. et Gottsche .......................................................................................238
Plagiochila andina Steph. ............................................................................................................238
Plagiochila angolensis Steph. ........................................................................................................238
Plagiochila angulata Steph. ..........................................................................................................224
Plagiochila angusta Lindenb. .......................................................................................................233
Plagiochila angusteoblonga Steph..................................................................................................238
Plagiochila angustisedens Steph. ...................................................................................................238
Plagiochila angustispina Steph. ....................................................................................................238
Plagiochila angustitexta Steph. .....................................................................................................233
Plagiochila annotina Lindenb. .....................................................................................................231
Plagiochila ansata (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ..............................................223
Plagiochila apicalis Gottsche ........................................................................................................238
Plagiochila appalachiana Inoue ....................................................................................................239
Plagiochila arbuscula (Brid. ex Lehm. et Lindenb.) Lindenb. .......................................................233
Plagiochila arbuscula var. rekohuensis J.J.Engel et G.L.Merr. ........................................................233
616
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Plagiochila arcta S.Winkl. ...........................................................................................................239
Plagiochila arctica Bryhn et Kaal. ................................................................................................228
Plagiochila arctica var. intermedia R.M.Schust.............................................................................228
Plagiochila arcuata Lindenb. .......................................................................................................239
Plagiochila arenacea Schifn. ........................................................................................................239
Plagiochila arnelliana Steph.........................................................................................................220
Plagiochila artsii Pócs ..................................................................................................................239
Plagiochila aspera Steph...............................................................................................................233
Plagiochila aspericaulis Grolle et M.L.So......................................................................................227
Plagiochila aspleniformis R.M.Schust. ..........................................................................................233
Plagiochila asplenioides (L.) Dumort. ...........................................................................................228
Plagiochila assamica Steph. ..........................................................................................................225
Plagiochila atrovirens Taylor ........................................................................................................239
Plagiochila austinii A.Evans .........................................................................................................232
Plagiochila badia Mitt. ................................................................................................................220
Plagiochila balansae Gottsche ......................................................................................................239
Plagiochila baldwinii Austin ........................................................................................................239
Plagiochila bamingensis Steph. .....................................................................................................239
Plagiochila bancroftii Steph. ........................................................................................................239
Plagiochila banksiana Gottsche....................................................................................................223
Plagiochila banksiana var. echinophora Inoue et R.M.Schust. .......................................................223
Plagiochila bantamensis (Reinw., Blume et Nees) Mont...............................................................222
Plagiochila barbadensis Steph.......................................................................................................239
Plagiochila barbata Steph. ...........................................................................................................239
Plagiochila barbeyi Steph. ............................................................................................................239
Plagiochila barteri Mitt. ..............................................................................................................220
Plagiochila barteri var. valida (Steph.) Vanden Berghen ..............................................................220
Plagiochila batava Steph. .............................................................................................................239
Plagiochila baylisii Inoue et R.M.Schust. .....................................................................................231
Plagiochila bazzanioides J.J.Engel et G.L.Merr. ...........................................................................231
Plagiochila beddomei Steph. .........................................................................................................233
Plagiochila benitoi Inoue ex Piippo ..............................................................................................225
Plagiochila berggrenii Steph. ........................................................................................................239
Plagiochila beskeana Steph. ..........................................................................................................239
Plagiochila bialata Mitt. ..............................................................................................................239
Plagiochila biapiculata Steph. ......................................................................................................239
Plagiochila bicaudata Steph. ........................................................................................................239
Plagiochila biciliata Steph............................................................................................................239
Plagiochila bicornis Hampe et Gottsche .......................................................................................239
Plagiochila bicornuta Steph..........................................................................................................224
Plagiochila bicuspidata Gottsche ..................................................................................................230
Plagiochila bidens Gottsche .........................................................................................................220
Plagiochila bidentula Steph..........................................................................................................239
Plagiochila bifaria (Sw.) Lindenb. ...............................................................................................220
Plagiochila bifaria var. rosea (R.M.Schust.) Heinrichs .................................................................220
Plagiochila binghamiae A.Evans ...................................................................................................239
Plagiochila binominata Steph. .....................................................................................................239
Plagiochila biondiana C.Massal. ..................................................................................................229
Plagiochila bischleriana Grolle et M.L.So.....................................................................................228
Plagiochila bitexta Dugas .............................................................................................................239
Plagiochila blepharobasis Herzog ..................................................................................................239
Plagiochila blepharophora (Nees) Lindenb. ..................................................................................222
World checklist of hornworts and liverworts
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617
Plagiochila bogotensis Gottsche ....................................................................................................239
Plagiochila boivinii Steph. ...........................................................................................................233
Plagiochila boliviana Spruce ........................................................................................................239
Plagiochila borneensis Steph. ........................................................................................................239
Plagiochila boryana Gottsche .......................................................................................................226
Plagiochila brassii Inoue et Grolle................................................................................................240
Plagiochila breuteliana Lindenb. ..................................................................................................226
Plagiochila brevicalycina Lindenb. et Gottsche.............................................................................240
Plagiochila britannica Paton ........................................................................................................228
Plagiochila brunneola Steph. ........................................................................................................240
Plagiochila bryhnii Steph. ............................................................................................................240
Plagiochila bryopteroides Spruce ...................................................................................................233
Plagiochila buchtiniana Steph. .....................................................................................................225
Plagiochila bunburii Taylor .........................................................................................................240
Plagiochila byssacea Hampe .........................................................................................................240
Plagiochila caducidentata R.M.Schust..........................................................................................240
Plagiochila caducifolia Inoue et R.M.Schust. ...............................................................................240
Plagiochila caduciloba H.L.Blomq. ..............................................................................................230
Plagiochila caldana Steph. ...........................................................................................................240
Plagiochila callaensis Steph. .........................................................................................................240
Plagiochila camusii Steph. ...........................................................................................................240
Plagiochila canelensis Steph. .........................................................................................................226
Plagiochila capillicaulis Steph. .....................................................................................................240
Plagiochila caribbeania R.M.Schust. ............................................................................................240
Plagiochila carringtonii (Balf. ex Carrington) Grolle ....................................................................229
Plagiochila carringtonii subsp. lobuchensis Grolle .........................................................................229
Plagiochila caulimammillosa Grolle et M.L.So .............................................................................227
Plagiochila cava Steph. ................................................................................................................240
Plagiochila ceylanica Mitt. ...........................................................................................................240
Plagiochila chacabucensis Steph. ...................................................................................................220
Plagiochila chacoënsis Herzog.......................................................................................................240
Plagiochila chauviniana Mont. ....................................................................................................222
Plagiochila chenii Grolle et M.L.So .............................................................................................231
Plagiochila chiloënsis Steph. .........................................................................................................240
Plagiochila chimborazensis Spruce ................................................................................................230
Plagiochila chinantlana Gottsche .................................................................................................240
Plagiochila chinensis Steph. ..........................................................................................................228
Plagiochila chiovendae Gola .........................................................................................................240
Plagiochila chonotica Taylor ........................................................................................................223
Plagiochila cinchonae Steph. ........................................................................................................240
Plagiochila circinalis (Lehm. et Lindenb.) Lehm. .........................................................................231
Plagiochila circinalis var. hemicardia (Hook.f. et Taylor) J.J.Engel et G.L.Merr. ..........................231
Plagiochila circumdentata Steph...................................................................................................228
Plagiochila circumdentata var. carinata J.J.Engel et G.L.Merr. .....................................................228
Plagiochila circumserrata Inoue et Grolle .....................................................................................228
Plagiochila circumvoluta Gerola ...................................................................................................240
Plagiochila clavatosaccata Steph. ..................................................................................................222
Plagiochila cleeii Inoue ...............................................................................................................240
Plagiochila cobana Steph. ............................................................................................................222
Plagiochila colensoi Hook.f. et Taylor ..........................................................................................231
Plagiochila colensoi var. quinquespina (Steph.) J.J.Engel et G.L.Merr. ..........................................231
Plagiochila colorans Steph. ...........................................................................................................220
618
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Plagiochila columbiana A.Evans ...................................................................................................228
Plagiochila concava Nees .............................................................................................................240
Plagiochila conduplicata Steph. ....................................................................................................240
Plagiochila conferta Steph. ...........................................................................................................240
Plagiochila confertissima Steph. ....................................................................................................240
Plagiochila connata Lindenb. et Gottsche ....................................................................................240
Plagiochila contigua Gottsche ......................................................................................................233
Plagiochila contorta Lindenb. et Hampe ......................................................................................240
Plagiochila convoluta Steph..........................................................................................................240
Plagiochila convolutifolia Steph. ...................................................................................................240
Plagiochila corrugata (Nees) Nees et Mont. .................................................................................233
Plagiochila corticola Steph. ..........................................................................................................231
Plagiochila corymbulosa Pearson...................................................................................................241
Plagiochila costariensis Horik. ......................................................................................................241
Plagiochila crispabilis Lindenb. ....................................................................................................241
Plagiochila crispabilis var. minima Lindenb. ................................................................................241
Plagiochila cristata (Sw.) Lindenb. ...............................................................................................220
Plagiochila cristophylla Steph. ......................................................................................................241
Plagiochila crozetensis Kaal...........................................................................................................224
Plagiochila cuatrecasii H.Rob. .....................................................................................................241
Plagiochila cubensis Steph. ...........................................................................................................241
Plagiochila cucullata Lindenb. et Gottsche...................................................................................241
Plagiochila cucullifolia J.B.Jack et Steph. .....................................................................................226
Plagiochila cucullifolia var. anomala Heinrichs et Gradst. ............................................................226
Plagiochila cuervina Gottsche ......................................................................................................241
Plagiochila cumingiana Steph. .....................................................................................................221
Plagiochila cuneata Lindenb. et Gottsche ....................................................................................230
Plagiochila cuspidata Steph. .........................................................................................................233
Plagiochila cymata Inoue et Grolle...............................................................................................233
Plagiochila debilis Mitt. ...............................................................................................................230
Plagiochila delexa Mont. et Gottsche ..........................................................................................230
Plagiochila delexirama Taylor .....................................................................................................233
Plagiochila defolians Grolle et M.L.So .........................................................................................230
Plagiochila delapsa Inoue .............................................................................................................241
Plagiochila delavayi Steph. ...........................................................................................................228
Plagiochila delognei Steph. ...........................................................................................................241
Plagiochila deltoidea Lindenb. .....................................................................................................224
Plagiochila deltoidea var. densa J.J.Engel et G.L.Merr. .................................................................224
Plagiochila denigrata Inoue..........................................................................................................241
Plagiochila densa Herzog .............................................................................................................241
Plagiochila densilora Herzog .......................................................................................................241
Plagiochila densiramosa Steph. .....................................................................................................241
Plagiochila denticulata Mitt. ........................................................................................................222
Plagiochila dependula Taylor .......................................................................................................225
Plagiochila deppeana Steph. .........................................................................................................241
Plagiochila desciscens Steph. .........................................................................................................241
Plagiochila detecta M.L.So et Grolle ............................................................................................222
Plagiochila devexa Steph. .............................................................................................................227
Plagiochila dichotoma (P.Beauv.) Nees et Mont. ..........................................................................233
Plagiochila dichotoma var. luitans Spruce ....................................................................................233
Plagiochila dilatata Steph. ...........................................................................................................241
Plagiochila dimorpha Lindenb. et Gottsche .................................................................................226
World checklist of hornworts and liverworts
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619
Plagiochila dissecta Steph. ............................................................................................................233
Plagiochila distans Colenso ..........................................................................................................241
Plagiochila disticha (Lehm. et Lindenb.) Lindenb. .......................................................................234
Plagiochila distinctifolia Lindenb. ................................................................................................234
Plagiochila divaricata Lindenb.....................................................................................................241
Plagiochila divergens Steph. .........................................................................................................234
Plagiochila diversifolia Lindenb. et Gottsche ................................................................................225
Plagiochila doerleri Steph. ...........................................................................................................241
Plagiochila dolichoblasta Herzog ..................................................................................................241
Plagiochila dominicensis Taylor ....................................................................................................226
Plagiochila drepanophylla Sande Lac. ...........................................................................................234
Plagiochila drepanophylla var. minor Gottsche .............................................................................234
Plagiochila dura De Not..............................................................................................................224
Plagiochila durelii Schifn. ...........................................................................................................227
Plagiochila durelii subsp. guizhouensis Grolle et M.L.So ..............................................................227
Plagiochila dusenii Steph. ............................................................................................................224
Plagiochila dussiana Steph. ..........................................................................................................241
Plagiochila duthiana Steph. .........................................................................................................229
Plagiochila echinata R.M.Schust. .................................................................................................241
Plagiochila echinella Gottsche ......................................................................................................241
Plagiochila ecuadorensis Steph. .....................................................................................................241
Plagiochila ecuadorica (Inoue) L.Söderstr.....................................................................................226
Plagiochila efusa Steph................................................................................................................234
Plagiochila efusa var. decurrens Steph. .........................................................................................234
Plagiochila efuseintricata Steph. ..................................................................................................242
Plagiochila eggersii Inoue .............................................................................................................242
Plagiochila ekmanii S.W.Arnell ...................................................................................................242
Plagiochila elata Taylor ...............................................................................................................224
Plagiochila electa Steph. ...............................................................................................................242
Plagiochila elegans Mitt. ..............................................................................................................228
Plagiochila elegantula Herzog ......................................................................................................242
Plagiochila emarginatobidentula Steph. ........................................................................................242
Plagiochila emeiensis Grolle et M.L.So .........................................................................................220
Plagiochila ensiformis Taylor ........................................................................................................226
Plagiochila equitans Gottsche ......................................................................................................223
Plagiochila ericicola Steph. ...........................................................................................................234
Plagiochila erlangensis M.L.So .....................................................................................................229
Plagiochila erythraeae Herzog ......................................................................................................242
Plagiochila estrellensis Herzog ......................................................................................................242
Plagiochila eurydictyon Herzog ....................................................................................................242
Plagiochila excisa Steph. ..............................................................................................................242
Plagiochila exesa Lindenb. et Gottsche.........................................................................................242
Plagiochila exigua (Taylor) Taylor ...............................................................................................230
Plagiochila exilis Colenso .............................................................................................................242
Plagiochila exinnovata Steph........................................................................................................234
Plagiochila expansa Gottsche .......................................................................................................242
Plagiochila facallonia Steph. ........................................................................................................242
Plagiochila falcato-oblonga Steph. ................................................................................................242
Plagiochila fallax Lindenb. et Hampe ..........................................................................................242
Plagiochila fasciculata Lindenb. ...................................................................................................231
Plagiochila fastigiata Lindenb. et Gottsche ..................................................................................234
Plagiochila faxinensis Schifn. ......................................................................................................242
620
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Plagiochila fendleri Mont.............................................................................................................242
Plagiochila ilicicola Steph. ..........................................................................................................242
Plagiochila issidentoides Taylor....................................................................................................242
Plagiochila labellata Steph. .........................................................................................................234
Plagiochila labellilora Steph. ......................................................................................................226
Plagiochila labellifrons Spruce .....................................................................................................242
Plagiochila lava Steph.................................................................................................................242
Plagiochila lavescens (Gottsche, Lindenb. et Nees) Gottsche .......................................................242
Plagiochila lavorufescens Steph. ...................................................................................................242
Plagiochila lexicaulis Mont. ........................................................................................................225
Plagiochila lexuosa Mitt. .............................................................................................................232
Plagiochila loridana A.Evans.......................................................................................................234
Plagiochila footei A.Evans ............................................................................................................242
Plagiochila fordiana Steph. ..........................................................................................................234
Plagiochila formosa Nees..............................................................................................................242
Plagiochila fracta Pócs .................................................................................................................230
Plagiochila fragilis Taylor ............................................................................................................221
Plagiochila fragillima Steph. ........................................................................................................222
Plagiochila fragmentata R.M.Schust. ...........................................................................................243
Plagiochila fragmentissima Inoue et R.M.Schust. .........................................................................223
Plagiochila francana Steph. ..........................................................................................................234
Plagiochila frausa Gottsche ..........................................................................................................243
Plagiochila frausa var. boliviana Spruce .......................................................................................243
Plagiochila frayjorgensis Hässel .....................................................................................................243
Plagiochila frondescens (Nees) Lindenb. .......................................................................................225
Plagiochila fruticosa Mitt. ............................................................................................................225
Plagiochila furcifolia Mitt. ...........................................................................................................234
Plagiochila fusca Sande Lac. .........................................................................................................231
Plagiochila fuscella (Hook.f. et Taylor) Gottsche, Lindenb. et Nees .............................................231
Plagiochila fuscella var. novae-zelandiae (E.A.Hodgs.) J.J.Engel et G.L.Merr................................231
Plagiochila fuscolutea Taylor ........................................................................................................225
Plagiochila fusifera Taylor............................................................................................................234
Plagiochila gafatensis Gottsche ex Schweinf. ...............................................................................243
Plagiochila gaudichaudii Mont. et Gottsche.................................................................................243
Plagiochila geniculata Lindenb.....................................................................................................243
Plagiochila geppii Steph. ..............................................................................................................243
Plagiochila germana Gottsche ......................................................................................................243
Plagiochila germanii Steph............................................................................................................24,
Plagiochila ghatiensis Steph. .........................................................................................................230
Plagiochila gibbilora Steph..........................................................................................................243
Plagiochila gigantea Lindenb. ......................................................................................................223
Plagiochila gigantea var. inermis J.J.Engel et G.L.Merr. ...............................................................223
Plagiochila gittinsii Inoue ............................................................................................................243
Plagiochila glauca Carl.................................................................................................................243
Plagiochila gracilicaulis Spruce.....................................................................................................243
Plagiochila gracilis Lindenb. et Gottsche......................................................................................232
Plagiochila gracillima Austin........................................................................................................243
Plagiochila granatensis Gottsche...................................................................................................243
Plagiochila grandicrista Steph. .....................................................................................................220
Plagiochila granditexta Steph. ......................................................................................................243
Plagiochila grateloupii Mont. .......................................................................................................243
Plagiochila gregaria (Hook.f. et Taylor) Gottsche, Lindenb. et Nees............................................223
World checklist of hornworts and liverworts
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621
Plagiochila grollei Inoue...............................................................................................................227
Plagiochila grossa Grolle et M.L.So ..............................................................................................230
Plagiochila grossispina Steph. .......................................................................................................222
Plagiochila guatemalensis Steph. ...................................................................................................243
Plagiochila guevarii H.Rob. .........................................................................................................226
Plagiochila guttisquama Inoue et Grolle .......................................................................................243
Plagiochila gymnocalycina (Lehm. et Lindenb.) Mont. et Nees.....................................................230
Plagiochila gymnocalycina var. surinamensis (Molk.) Heinrichs et D.S.Rycroft .............................230
Plagiochila gymnocalyx Inoue .......................................................................................................243
Plagiochila gymnoclada Sande Lac................................................................................................232
Plagiochila haeseliae Inoue ...........................................................................................................243
Plagiochila hakkodensis Steph. .....................................................................................................228
Plagiochila hampeana Gottsche ...................................................................................................234
Plagiochila hans-meyeri Steph. .....................................................................................................243
Plagiochila harlingii S.W.Arnell ..................................................................................................243
Plagiochila haumanii Herzog .......................................................................................................244
Plagiochila hawaica Steph. ..........................................................................................................244
Plagiochila herminieri Steph. .......................................................................................................220
Plagiochila heteracantha Steph. ....................................................................................................244
Plagiochila heterodonta (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ......................................224
Plagiochila heterofolia Steph. .......................................................................................................244
Plagiochila heteromalla (Lehm. et Lindenb.) Lindenb. .................................................................220
Plagiochila heterophylla Lindenb. .................................................................................................225
Plagiochila heterophylla var. beauverdii (Steph.) Heinrichs ...........................................................225
Plagiochila heterospina Steph. ......................................................................................................234
Plagiochila heterostipa Steph. .......................................................................................................234
Plagiochila hieronymii Steph. .......................................................................................................244
Plagiochila himalayana Schifn. ...................................................................................................232
Plagiochila hiroshiana Pócs ..........................................................................................................244
Plagiochila hoehnei Herzog ..........................................................................................................244
Plagiochila hoei Inoue..................................................................................................................227
Plagiochila holstii Steph. ..............................................................................................................244
Plagiochila hookeriana Lindenb. ..................................................................................................223
Plagiochila horrida Gottsche ........................................................................................................244
Plagiochila huatuscana Gottsche ..................................................................................................244
Plagiochila huerlimannii Inoue ....................................................................................................244
Plagiochila humboldtiana Gottsche ..............................................................................................244
Plagiochila husnotii Steph. ...........................................................................................................226
Plagiochila hyalina Lindenb.........................................................................................................224
Plagiochila hyalodermica Grolle et M.L.So ...................................................................................227
Plagiochila incerta Gottsche.........................................................................................................234
Plagiochila incisa Dugas ..............................................................................................................244
Plagiochila incurvicolla (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ......................................232
Plagiochila incurvicolla var. lonchoscypha (Herzog) J.J.Engel et G.L.Merr. ...................................232
Plagiochila indica Mitt. ex Steph. ................................................................................................234
Plagiochila inlata Steph. .............................................................................................................244
Plagiochila informifolia Steph. .....................................................................................................244
Plagiochila infuscata Mitt. ...........................................................................................................244
Plagiochila injasutiensis S.W.Arnell..............................................................................................244
Plagiochila inouei Grolle..............................................................................................................244
Plagiochila insularia Mitt. ...........................................................................................................244
Plagiochila integerrima Steph. ......................................................................................................222
622
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Plagiochila integrilobula Schifn. ..................................................................................................222
Plagiochila intertexta Hook.f. et Taylor .......................................................................................244
Plagiochila inuensis Steph. ...........................................................................................................244
Plagiochila invisa (R.M.Schust.) R.M.Schust. .............................................................................244
Plagiochila irregularis Gottsche ....................................................................................................244
Plagiochila itatiajensis Steph. .......................................................................................................244
Plagiochila jacquinotii Mont........................................................................................................244
Plagiochila jamaicensis Lindenb. et Hampe..................................................................................244
Plagiochila jaramillii H.Rob. .......................................................................................................244
Plagiochila javanica (Sw.) Nees et Mont......................................................................................234
Plagiochila johannis-winkleri Herzog ...........................................................................................222
Plagiochila junghuhniana Sande Lac. ...........................................................................................235
Plagiochila karstenii Steph. ..........................................................................................................244
Plagiochila kaulfussiana Steph. ....................................................................................................244
Plagiochila keckiana Steph. ..........................................................................................................244
Plagiochila kerneriana S.W.Arnell ...............................................................................................245
Plagiochila khasiana Mitt. ...........................................................................................................235
Plagiochila kiaeri Gottsche ..........................................................................................................235
Plagiochila kiaeri var. capensis (Steph.) M.Wigginton et Grolle ...................................................235
Plagiochila kiaeri var. myriocarpa (Pearson) Pócs .........................................................................235
Plagiochila koponenii Inoue et Piippo ..........................................................................................245
Plagiochila korthalsiana Molk. .....................................................................................................228
Plagiochila kuhliana Sande Lac....................................................................................................228
Plagiochila kunmingensis Piippo ..................................................................................................235
Plagiochila kurokawae Inoue .......................................................................................................235
Plagiochila kurzii Steph. ..............................................................................................................223
Plagiochila lacerifolia Steph. ........................................................................................................245
Plagiochila lachenaudii Steph. .....................................................................................................245
Plagiochila laciniosa Dugas ..........................................................................................................245
Plagiochila laetevirens Lindenb. ...................................................................................................245
Plagiochila laevifolia Gola............................................................................................................245
Plagiochila lamellistipula Spruce ..................................................................................................235
Plagiochila lansbergii Gottsche.....................................................................................................245
Plagiochila lastii Mitt. .................................................................................................................235
Plagiochila latifolia Steph. ...........................................................................................................235
Plagiochila latifrons Gottsche et Hampe ......................................................................................224
Plagiochila latitrigona Schifn. .....................................................................................................245
Plagiochila laxiramea Steph. ........................................................................................................245
Plagiochila lechleri Gottsche ........................................................................................................225
Plagiochila lecontei Steph. ............................................................................................................245
Plagiochila ledermanniana Beauverd ............................................................................................245
Plagiochila ledieui Steph. .............................................................................................................245
Plagiochila lehmanniana S.W.Arnell ............................................................................................245
Plagiochila liebmanniana Lehm. et Lindenb. ...............................................................................245
Plagiochila lignicola Spruce..........................................................................................................245
Plagiochila lindaui Steph. ............................................................................................................245
Plagiochila lindigiana Gottsche ...................................................................................................245
Plagiochila lingua Steph. .............................................................................................................245
Plagiochila loloënsis Steph. ...........................................................................................................235
Plagiochila longiissa Steph...........................................................................................................245
Plagiochila longilora Mont. .........................................................................................................227
Plagiochila longiramea Steph. ......................................................................................................225
World checklist of hornworts and liverworts
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623
Plagiochila longispina Lindenb. et Gottsche .................................................................................225
Plagiochila lophocoleoides Mont. ..................................................................................................227
Plagiochila loriana Steph. ............................................................................................................228
Plagiochila loriloba Herzog ex Carl ..............................................................................................230
Plagiochila lotsyana Steph. ...........................................................................................................245
Plagiochila lunata S.W.Arnell ......................................................................................................221
Plagiochila luteola Steph. .............................................................................................................245
Plagiochila lutescens Steph. ..........................................................................................................245
Plagiochila luzonensis Grolle et M.L.So .......................................................................................245
Plagiochila macra Taylor .............................................................................................................246
Plagiochila macrifolia Taylor .......................................................................................................246
Plagiochila macrifolia var. angustifolia Spruce ..............................................................................246
Plagiochila macrostachya Lindenb. ...............................................................................................226
Plagiochila maderensis Gottsche ex Steph.....................................................................................230
Plagiochila magna Inoue..............................................................................................................227
Plagiochila manillana Mont. et Gottsche.....................................................................................235
Plagiochila martiana (Nees) Lindenb...........................................................................................246
Plagiochila massalongoana Schifn. ...............................................................................................235
Plagiochila mastigophoroides Inoue ...............................................................................................235
Plagiochila matanga Steph. ..........................................................................................................246
Plagiochila mauiensis Steph. ........................................................................................................225
Plagiochila maunakeana Steph.....................................................................................................246
Plagiochila maximiliana Gottsche................................................................................................246
Plagiochila meghalayensis K.K.Rawat et S.C.Srivast. ....................................................................246
Plagiochila meridana Gottsche ....................................................................................................246
Plagiochila metcalii Steph. ..........................................................................................................233
Plagiochila microdentata M.L.So .................................................................................................228
Plagiochila microdictyon Mitt.......................................................................................................232
Plagiochila microdonta Mitt. ........................................................................................................246
Plagiochila micropteryx Gottsche..................................................................................................235
Plagiochila mildbreadiana Beauverd ............................................................................................247
Plagiochila minarum Herzog .......................................................................................................247
Plagiochila minutiretis Reimers ....................................................................................................247
Plagiochila minutula (Hook.f. et Taylor) Gottsche, Lindenb. et Nees..........................................224
Plagiochila miqueliana Lehm. et Lindenb. ...................................................................................246
Plagiochila miradorensis Gottsche ................................................................................................235
Plagiochila miradorensis var. convoluta R.M.Schust. .....................................................................235
Plagiochila moenkemeyeri Steph. ..................................................................................................235
Plagiochila molliuscula Inoue .......................................................................................................246
Plagiochila mollusca Lehm. ..........................................................................................................246
Plagiochila moniliformis R.M.Schust. ..........................................................................................246
Plagiochila monospiris Inoue et Grolle .........................................................................................232
Plagiochila montagnei Nees..........................................................................................................235
Plagiochila morobensis Inoue et Piippo.........................................................................................236
Plagiochila muelleriana Gottsche .................................................................................................246
Plagiochila multilora Steph. ........................................................................................................246
Plagiochila multipinnula Herzog et S.Hatt. .................................................................................236
Plagiochila mundaliensis Steph.....................................................................................................228
Plagiochila mutila Gottsche .........................................................................................................246
Plagiochila naranjoënsis Steph......................................................................................................246
Plagiochila neckeroidea Mitt. .......................................................................................................236
Plagiochila neesiana Lindenb. ......................................................................................................246
624
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Plagiochila neglecta Steph. ...........................................................................................................246
Plagiochila negrensis Spruce .........................................................................................................246
Plagiochila negrii Gola.................................................................................................................246
Plagiochila nepalensis Lindenb. ....................................................................................................236
Plagiochila nigricaulis Steph. .......................................................................................................247
Plagiochila nitens Inoue ...............................................................................................................232
Plagiochila nobilis Gottsche .........................................................................................................224
Plagiochila norfolkiensis Steph......................................................................................................236
Plagiochila notha Steph. ..............................................................................................................247
Plagiochila nova Steph.................................................................................................................247
Plagiochila nudifolia (Steph.) Grolle ............................................................................................247
Plagiochila nutans Steph. .............................................................................................................247
Plagiochila obliquetruncata Steph. ...............................................................................................247
Plagiochila oblonga Inoue ............................................................................................................247
Plagiochila obovata Steph. ...........................................................................................................224
Plagiochila obtusa Lindenb. .........................................................................................................236
Plagiochila oerstediana Lindenb. et Hampe..................................................................................247
Plagiochila olivacea Steph. ...........................................................................................................247
Plagiochila orbicularis (S.Hatt.) S.Hatt. .......................................................................................228
Plagiochila oresitropha Spruce ......................................................................................................230
Plagiochila ornata Wilson ex Lindenb. ........................................................................................247
Plagiochila ovalifolia Mitt. ...........................................................................................................228
Plagiochila ovata Lindenb. et Gottsche ........................................................................................224
Plagiochila ovato-obconica Steph. .................................................................................................247
Plagiochila ovifolia Steph. ............................................................................................................247
Plagiochila owaihiensis Nees et Lindenb. .....................................................................................234
Plagiochila pachyloma Taylor .......................................................................................................221
Plagiochila pachyloma var. elatior Spruce .....................................................................................221
Plagiochila paciica Mitt. .............................................................................................................234
Plagiochila palangiensis S.C.Srivast., K.K.Rawat et P.K.Verma ....................................................247
Plagiochila panamensis Inoue .......................................................................................................247
Plagiochila papillifolia Steph. .......................................................................................................221
Plagiochila parallela Steph. ..........................................................................................................236
Plagiochila paramicola Herzog .....................................................................................................247
Plagiochila paraphyllina Herzog...................................................................................................225
Plagiochila paraphyllosa Grolle et M.L.So ....................................................................................227
Plagiochila parvifolia Lindenb. ....................................................................................................234
Plagiochila parviramifera Inoue ...................................................................................................221
Plagiochila parvitexta Steph. ........................................................................................................247
Plagiochila parvivittata Inoue ......................................................................................................247
Plagiochila parvula Steph. ...........................................................................................................247
Plagiochila pastasensis Steph.........................................................................................................247
Plagiochila patentispina Steph. .....................................................................................................247
Plagiochila patentissima Lindenb. ................................................................................................247
Plagiochila patriciae Heinrichs et H.Anton ..................................................................................226
Plagiochila patula (Sw.) Lindenb. ................................................................................................236
Plagiochila patula var. brevifolia Gottsche....................................................................................236
Plagiochila patuloides Schifn. ......................................................................................................247
Plagiochila paucidens Steph. ........................................................................................................236
Plagiochila paucidentata Mont. et Gottsche .................................................................................247
Plagiochila paupercula Gottsche...................................................................................................247
Plagiochila pearceana Steph. ........................................................................................................247
World checklist of hornworts and liverworts
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625
Plagiochila pectinata Lindenb. .....................................................................................................230
Plagiochila peculiaris Schifn. .......................................................................................................227
Plagiochila pellucida Lindenb. et Gottsche...................................................................................247
Plagiochila pensilis Spruce............................................................................................................236
Plagiochila peradenyensis Schifn. .................................................................................................236
Plagiochila perdentata M.L.So et Grolle.......................................................................................236
Plagiochila perrottetiana Mont. et Gottsche .................................................................................248
Plagiochila perrottetiana var. minor Gottsche ...............................................................................248
Plagiochila perserrata Herzog .......................................................................................................227
Plagiochila philippinensis Steph....................................................................................................227
Plagiochila pilifera Steph. ............................................................................................................248
Plagiochila pinnatidens Steph.......................................................................................................248
Plagiochila pinnilora Steph. ........................................................................................................236
Plagiochila pittieri Steph..............................................................................................................248
Plagiochila platyphylla Herzog .....................................................................................................248
Plagiochila poeltii Inoue et Grolle ................................................................................................229
Plagiochila polopolensis Herzog ....................................................................................................248
Plagiochila porelloides (Torr. ex Nees) Lindenb. ...........................................................................229
Plagiochila porelloides var. norvegica (H.H.Blom et Holten) Schumacker et Váňa ........................229
Plagiochila porelloides var. subarctica (Jørg.) Lammes ...................................................................229
Plagiochila praemorsa Steph. ........................................................................................................236
Plagiochila propinqua Sande Lac..................................................................................................236
Plagiochila prostrata Steph. ..........................................................................................................244
Plagiochila pseudoattenuata S.W.Arnell........................................................................................221
Plagiochila pseudocapillaris Inoue.................................................................................................232
Plagiochila pseudoirma Herzog ...................................................................................................232
Plagiochila pseudopatula Herzog ..................................................................................................248
Plagiochila pseudopoeltii Inoue .....................................................................................................227
Plagiochila pseudoradicans Herzog ...............................................................................................248
Plagiochila pseudorenitens Schifn. ...............................................................................................222
Plagiochila ptychanthoidea Steph..................................................................................................248
Plagiochila pulchella Steph...........................................................................................................248
Plagiochila pulcherrima Horik. ....................................................................................................225
Plagiochila pulvinata Steph..........................................................................................................248
Plagiochila punctata (Taylor) Taylor............................................................................................221
Plagiochila purpurascens Steph. ....................................................................................................236
Plagiochila raddiana Lindenb. .....................................................................................................236
Plagiochila radiculosa Mitt...........................................................................................................232
Plagiochila ragazzii Gola .............................................................................................................236
Plagiochila ramosissima (Hook.) Lindenb. ...................................................................................224
Plagiochila ratkowskiana Inoue ....................................................................................................248
Plagiochila recurvata (W.E.Nicholson) Grolle .............................................................................229
Plagiochila regeliana Steph. .........................................................................................................248
Plagiochila reischeckiana Steph. ...................................................................................................223
Plagiochila remyana Steph. ..........................................................................................................248
Plagiochila renauldii Steph. .........................................................................................................221
Plagiochila renitens (Nees) Lindenb. ............................................................................................227
Plagiochila repanda (Schwägr.) Lindenb. .....................................................................................236
Plagiochila repanda var. perrotana (Steph.) Vanden Berghen .......................................................237
Plagiochila retrorsa Gottsche........................................................................................................221
Plagiochila retrospectans Lindenb. ................................................................................................224
Plagiochila retusa Mitt. ................................................................................................................229
626
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Plagiochila revolvens Mitt. ...........................................................................................................248
Plagiochila rigidula Lindenb. et Gottsche ....................................................................................248
Plagiochila riparia Steph. ............................................................................................................224
Plagiochila rodriguezii Steph. .......................................................................................................237
Plagiochila rosana Steph. .............................................................................................................248
Plagiochila rubescens (Lehm. et Lindenb.) Lindenb. .....................................................................221
Plagiochila rubricaulis Steph. .......................................................................................................221
Plagiochila rudischusteri H.Rob. ..................................................................................................225
Plagiochila rudolii Pócs...............................................................................................................237
Plagiochila ruifolia Steph. ...........................................................................................................248
Plagiochila rufoviridis Spruce .......................................................................................................248
Plagiochila rusbyi Spruce .............................................................................................................248
Plagiochila rutilans Lindenb. .......................................................................................................230
Plagiochila rutilans var. moritziana (Lindenb. et Gottsche) Heinrichs..........................................230
Plagiochila rutilans var. standleyi (Herzog ex Carl) Heinrichs et D.S.Rycroft ...............................230
Plagiochila rutlandii Steph. .........................................................................................................224
Plagiochila sabensis Steph. ...........................................................................................................248
Plagiochila sachapatensis Steph.....................................................................................................248
Plagiochila salacensis Gottsche .....................................................................................................237
Plagiochila salazariae Inoue .........................................................................................................248
Plagiochila saltuensis Spruce ex Steph. .........................................................................................248
Plagiochila saltuensis var. spinosissima Herzog ..............................................................................249
Plagiochila salvadorica Steph. ......................................................................................................237
Plagiochila samoana Steph. ..........................................................................................................249
Plagiochila sandei Dozy ex Sande Lac. .........................................................................................223
Plagiochila sarmentosa (Lehm. et Lindenb.) Lindenb. ..................................................................249
Plagiochila schifneri Steph. .........................................................................................................249
Plagiochila schinzei Steph. ...........................................................................................................249
Plagiochila schmidtii Steph. .........................................................................................................249
Plagiochila schoieldiana Inoue .....................................................................................................229
Plagiochila schraderbergii Steph. ..................................................................................................249
Plagiochila schubertiana Steph. ....................................................................................................249
Plagiochila schusteri Inoue et Grolle.............................................................................................249
Plagiochila sciophila Nees ............................................................................................................223
Plagiochila sciophila subsp. ciliigera (R.M.Schust.) L.Söderstr. .....................................................223
Plagiochila scissifolia Steph...........................................................................................................249
Plagiochila scotica Macvicar ex Steph. ..........................................................................................249
Plagiochila secretifolia Mitt. .........................................................................................................229
Plagiochila semiamplexicaulis Steph. ............................................................................................249
Plagiochila semidecurrens (Lehm. et Lindenb.) Lindenb. ..............................................................227
Plagiochila semidecurrens var. alaskana (A.Evans) Inoue...............................................................227
Plagiochila semiermis Dugas ........................................................................................................249
Plagiochila serrialata Herzog........................................................................................................237
Plagiochila shangaica Steph..........................................................................................................237
Plagiochila sichuanensis Grolle et M.L.So ....................................................................................221
Plagiochila silvatica Gottsche .......................................................................................................249
Plagiochila similis Steph. .............................................................................................................249
Plagiochila simplex (Sw.) Lindenb. ..............................................................................................237
Plagiochila singularis Schifn. .......................................................................................................222
Plagiochila sisparensis Steph. ........................................................................................................249
Plagiochila slateri Steph. ..............................................................................................................249
Plagiochila solitaria Gottsche .......................................................................................................249
World checklist of hornworts and liverworts
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627
Plagiochila spathulifolia Mitt. ......................................................................................................232
Plagiochila spinifera Ångstr. .........................................................................................................249
Plagiochila spinosa M.L.So ..........................................................................................................229
Plagiochila spinulosa (Dicks.) Dumort. ........................................................................................221
Plagiochila sprucei Steph. .............................................................................................................249
Plagiochila squamulosa Mitt. .......................................................................................................237
Plagiochila squamulosa var. crispulocaudata (Gottsche) Vanden Berghen......................................237
Plagiochila squamulosa var. sinuosa (Mitt.) Vanden Berghen ........................................................237
Plagiochila staudtiana Steph. .......................................................................................................249
Plagiochila stephanii Schifn. .......................................................................................................223
Plagiochila stephensoniana Mitt....................................................................................................232
Plagiochila stevensiana Steph. ......................................................................................................222
Plagiochila steyermarkii H.Rob. ...................................................................................................231
Plagiochila sticticola Mont. et Gottsche .......................................................................................221
Plagiochila stipata Steph. .............................................................................................................249
Plagiochila stolzii Steph. ..............................................................................................................249
Plagiochila streimannii Inoue.......................................................................................................237
Plagiochila stricta Lindenb...........................................................................................................221
Plagiochila strictifolia Steph. ........................................................................................................237
Plagiochila strombifolia Taylor .....................................................................................................231
Plagiochila subbidentata Taylor ...................................................................................................225
Plagiochila subcontigua Herzog ....................................................................................................249
Plagiochila subconvoluta Gottsche................................................................................................249
Plagiochila subdenudata Steph. ....................................................................................................249
Plagiochila subedentata Steph. .....................................................................................................250
Plagiochila sublabellata Colenso .................................................................................................237
Plagiochila subfragilis Inoue .........................................................................................................250
Plagiochila subhyalina Steph. .......................................................................................................250
Plagiochila subjavanica M.L.So ...................................................................................................250
Plagiochila subligulata Steph........................................................................................................250
Plagiochila sublyallii M.L.So........................................................................................................250
Plagiochila subpectinata Besch. et C.Massal. ................................................................................224
Plagiochila subplana Lindenb. .....................................................................................................223
Plagiochila subrara Herzog ..........................................................................................................250
Plagiochila subrotundifolia Steph. ................................................................................................250
Plagiochila subtropica Steph. ........................................................................................................237
Plagiochila subundulata Lindenb. ................................................................................................250
Plagiochila sullivantii Gottsche ....................................................................................................232
Plagiochila sullivantii var. spinigera R.M.Schust. .........................................................................232
Plagiochila sumatrana Schifn. .....................................................................................................229
Plagiochila superba (Nees ex Spreng.) Mont. et Nees ...................................................................226
Plagiochila superba var. macrotricha (Spruce) Heinrichs ...............................................................226
Plagiochila suringarii Steph. ........................................................................................................250
Plagiochila sylvicultrix Spruce ......................................................................................................250
Plagiochila symmetrica Steph. ......................................................................................................250
Plagiochila tabinana Gottsche .....................................................................................................250
Plagiochila tabinensis Steph. ........................................................................................................225
Plagiochila tagawae Inoue............................................................................................................222
Plagiochila taiwanensis Inoue.......................................................................................................229
Plagiochila talinayi S.W.Arnell ....................................................................................................250
Plagiochila tamariscina Steph. .....................................................................................................237
Plagiochila tambillensis Loitl. .......................................................................................................250
628
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Plagiochila tamiensis Steph. .........................................................................................................250
Plagiochila tarapotensis Steph.......................................................................................................250
Plagiochila tecta Inoue et Grolle ..................................................................................................237
Plagiochila tenera Steph. ..............................................................................................................250
Plagiochila tenuis Lindenb. ..........................................................................................................250
Plagiochila tenuispica Steph. ........................................................................................................250
Plagiochila terebrans Nees et Mont. .............................................................................................237
Plagiochila teysmannii Sande Lac. ................................................................................................237
Plagiochila thamniopsis Spruce.....................................................................................................250
Plagiochila thollonii Steph. ..........................................................................................................250
Plagiochila thrausta Inoue et Grolle .............................................................................................250
Plagiochila thyoides Spruce...........................................................................................................237
Plagiochila tixieri Inoue ...............................................................................................................222
Plagiochila tocarema Gottsche .....................................................................................................237
Plagiochila tonduzana Steph. .......................................................................................................250
Plagiochila tortuosa Lindenb. et Gottsche ....................................................................................250
Plagiochila tovarina Gottsche ......................................................................................................250
Plagiochila trabeculata Steph. ......................................................................................................232
Plagiochila trabeculatospinosa Herzog ..........................................................................................250
Plagiochila trabutii Steph. ...........................................................................................................250
Plagiochila trapezoidea Lindenb. ..................................................................................................229
Plagiochila trianae Gottsche ........................................................................................................250
Plagiochila trichomanes Spruce ....................................................................................................250
Plagiochila trichostoma Gottsche ..................................................................................................231
Plagiochila tricuspis Steph. ...........................................................................................................250
Plagiochila tristaniana Váňa et J.J.Engel ......................................................................................251
Plagiochila tristis Steph. ...............................................................................................................251
Plagiochila trollii Herzog .............................................................................................................251
Plagiochila tronadoris Herzog ......................................................................................................221
Plagiochila truncata Gottsche ......................................................................................................251
Plagiochila turgida Herzog ..........................................................................................................226
Plagiochila ulata Inoue et Grolle .................................................................................................237
Plagiochila uleana Steph. .............................................................................................................251
Plagiochila umbrosioides L.Söderstr. .............................................................................................251
Plagiochila uncialis (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ............................................221
Plagiochila undata Sull. ...............................................................................................................237
Plagiochila undata subsp. crispata (Gottsche) R.M.Schust. ..........................................................237
Plagiochila unduavensis Steph. .....................................................................................................251
Plagiochila ungarangana Sande Lac. ............................................................................................238
Plagiochila uniformis Mitt. ..........................................................................................................229
Plagiochila usambarana Steph. ....................................................................................................251
Plagiochila validissima Steph. ......................................................................................................225
Plagiochila variedentata Steph. ....................................................................................................251
Plagiochila vastifolia Steph. .........................................................................................................251
Plagiochila velata Inoue et Piippo ................................................................................................251
Plagiochila venezuelana Steph. .....................................................................................................251
Plagiochila ventricosotrigona Steph. ..............................................................................................251
Plagiochila verrucosa Steph. .........................................................................................................251
Plagiochila vetustisilva Steph........................................................................................................251
Plagiochila vexans Schifn. ex Steph. ............................................................................................227
Plagiochila viminea Spruce ..........................................................................................................251
Plagiochila vincentina Lindenb. ...................................................................................................226
World checklist of hornworts and liverworts
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629
Plagiochila virginica A.Evans .......................................................................................................238
Plagiochila virginica var. caroliniana R.M.Schust.........................................................................238
Plagiochila virginica var. euryphylla R.M.Schust. .........................................................................238
Plagiochila viridis Steph. .............................................................................................................251
Plagiochila viridonigra (E.A.Hodgs.) Inoue .................................................................................251
Plagiochila vitiensis Mitt. .............................................................................................................223
Plagiochila vittiana Inoue ............................................................................................................251
Plagiochila vittifolia Steph. ..........................................................................................................251
Plagiochila vulcanica Steph. .........................................................................................................251
Plagiochila wacei S.W.Arnell ex Váňa et J.J.Engel .......................................................................251
Plagiochila wallichiana Steph. .....................................................................................................229
Plagiochila wallisiana Steph.........................................................................................................251
Plagiochila wangii Inoue .............................................................................................................229
Plagiochila wattsiana J.J.Engel et G.L.Merr. ................................................................................252
Plagiochila wettsteiniana S.W.Arnell ............................................................................................252
Plagiochila weymouthiana Steph. .................................................................................................252
Plagiochila wiemanniana S.W.Arnell ...........................................................................................252
Plagiochila wightii Nees...............................................................................................................238
Plagiochila wilhelmina Inoue .......................................................................................................238
Plagiochila wilmsiana Steph. .......................................................................................................221
Plagiochila winteri Steph. ............................................................................................................252
Plagiochila wrightii Steph. ...........................................................................................................252
Plagiochila xalapensis Gottsche ....................................................................................................252
Plagiochila xanthochroma Spruce .................................................................................................252
Plagiochila yoshinagana Steph. .....................................................................................................252
Plagiochila yulungensis Piippo ......................................................................................................229
Plagiochila zacuapana Gottsche ...................................................................................................252
Plagiochila zangii Grolle et M.L.So .............................................................................................228
Plagiochila zantenii Inoue ...........................................................................................................238
Plagiochila zhuensis Grolle et M.L.So ..........................................................................................222
Plagiochila zonata Steph. .............................................................................................................228
Plagiochilidium bidentulum (Steph.) Grolle .................................................................................252
Plagiochilion braunianum (Nees) S.Hatt. ....................................................................................252
Plagiochilion combinatum (Mitt.) Inoue ......................................................................................252
Plagiochilion conjugatum (Hook.) R.M.Schust. ...........................................................................252
Plagiochilion imbriatum (Mitt.) Inoue ........................................................................................252
Plagiochilion giulianettii (Steph.) Inoue .......................................................................................252
Plagiochilion herzogii Inoue .........................................................................................................252
Plagiochilion intermedium R.M.Schust. .......................................................................................252
Plagiochilion mayebarae S.Hatt. ...................................................................................................253
Plagiochilion oppositum (Reinw., Blume et Nees) S.Hatt. ............................................................253
Plagiochilion pachycephalum (De Not.) Inoue ..............................................................................253
Plagiochilion proliferum (Mitt.) R.M.Schust. ...............................................................................253
Plagiochilion theriotanum (Steph.) Inoue .....................................................................................253
Platycaulis renifolius R.M.Schust. ................................................................................................218
Plectocolea subamoena S.Winkl. ...................................................................................................511
Pleurozia acinosa (Mitt.) Trevis. ..................................................................................................462
Pleurozia articulata (Lindb.) Lindb. et Lackström .......................................................................462
Pleurozia caledonica (Gottsche) Steph. ........................................................................................462
Pleurozia conchifolia (Hook. et Arn.) Austin................................................................................462
Pleurozia conchifolia var. papillosa B.M.hiers .............................................................................462
Pleurozia curiosa B.M.hiers .......................................................................................................462
630
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Pleurozia gigantea (F.Weber) Lindb. ...........................................................................................462
Pleurozia heterophylla Steph. ex Fulford ......................................................................................462
Pleurozia johannis-winkleri Herzog .............................................................................................462
Pleurozia paradoxa (J.B.Jack) Schifn. .........................................................................................462
Pleurozia pocsii Frank Müll. ........................................................................................................463
Pleurozia purpurea Lindb. ...........................................................................................................462
Pleurozia subinlata (Austin) Austin ............................................................................................462
Plicanthus birmensis (Steph.) R.M.Schust. .....................................................................................54
Plicanthus diicilis (Steph.) L.Söderstr. et Váňa .............................................................................54
Plicanthus giganteus (Steph.) R.M.Schust. .....................................................................................54
Plicanthus hirtellus (F.Weber) R.M.Schust. ...................................................................................54
Podomitrium malaccense (Steph.) Campb. ...................................................................................471
Podomitrium marginatum (Steph.) Hürl......................................................................................471
Podomitrium phyllanthus (Hook.) Mitt. ......................................................................................471
Poeltia campylata Grolle ..............................................................................................................114
Polyotus peckianus Austin ............................................................................................................511
Porella abyssinica (Nees) Trevis. ..................................................................................................417
Porella abyssinica var. hoehnelii (Steph.) Pócs...............................................................................417
Porella acutifolia (Lehm. et Lindenb.) Trevis. ..............................................................................417
Porella acutifolia var. hattoriana (Pócs) S.Hatt.............................................................................417
Porella acutifolia var. linguifolia (Steph.) M.L.So.........................................................................417
Porella acutifolia subsp. tosana (Steph.) S.Hatt. ...........................................................................417
Porella andica (Gottsche) Hässel .................................................................................................417
Porella arboris-vitae (With.) Grolle..............................................................................................418
Porella arboris-vitae subsp. nitidula (C.Massal.) S.Hatt. ..............................................................418
Porella baueri (Schifn.) C.E.O.Jensen ........................................................................................418
Porella bolanderi (Austin) Pearson ...............................................................................................418
Porella borellii (Gola) Parihar ......................................................................................................418
Porella brachiata (Taylor) Spruce ................................................................................................418
Porella brasiliensis (Raddi) Schifn. ..............................................................................................418
Porella brasiliensis var. ciliata (Gottsche, Lindenb. et Nees) Schifn. ............................................418
Porella brasiliensis var. laevior (Gottsche, Lindenb. et Nees) Schifn. ...........................................418
Porella caespitans (Steph.) S.Hatt.................................................................................................418
Porella caespitans var. cordifolia (Steph.) S.Hatt. ex T.Katag. et T.Yamag. ...................................418
Porella caespitans subsp. latior (S.Hatt.) S.Hatt............................................................................418
Porella caespitans var. nipponica S.Hatt. .......................................................................................418
Porella caespitans var. relexigastria (Pócs) S.Hatt. ........................................................................419
Porella campylophylla (Lehm. et Lindenb.) Trevis. .......................................................................419
Porella campylophylla subsp. lancistipula (Steph.) S.Hatt. ............................................................419
Porella campylophylla var. ligulifera (Taylor) S.Hatt. ....................................................................419
Porella campylophylla var. tixieri (Pócs) S.Hatt. ...........................................................................419
Porella canariensis (F.Weber) Underw. ........................................................................................419
Porella capehorniensis Swails ........................................................................................................419
Porella capensis (Gottsche) Mitt...................................................................................................419
Porella caucasica Steph. ...............................................................................................................419
Porella chenii S.Hatt....................................................................................................................419
Porella chilensis (Lehm. et Lindenb.) Trevis. ................................................................................419
Porella chilensis var. antucensis (Gottsche) Hässel .........................................................................419
Porella chilensis var. fernandeziensis (Herzog) Swails ....................................................................419
Porella chilensis var. microloba (Herzog) Swails ............................................................................419
Porella chinensis (Steph.) S.Hatt. .................................................................................................419
Porella chinensis var. crispata Udar et Shaheen .............................................................................420
World checklist of hornworts and liverworts
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631
Porella chinensis var. decurrens (Steph.) S.Hatt.............................................................................420
Porella chinensis var. hattorii Udar et Shaheen .............................................................................420
Porella chinensis var. irregularis (Steph.) S.Hatt. ..........................................................................420
Porella circinnata Lindb. .............................................................................................................420
Porella cordaeana (Huebener) Moore ..........................................................................................420
Porella cranfordii Steph. ..............................................................................................................420
Porella crispata (Hook.) Trevis. ...................................................................................................420
Porella cucullistipula Steph. .........................................................................................................420
Porella densifolia (Steph.) S.Hatt. ................................................................................................420
Porella densifolia subsp. andamana S.Hatt. ..................................................................................420
Porella densifolia subsp. appendiculata (Steph.) S.Hatt.................................................................420
Porella densifolia var. oviloba (Steph.) N.Kitag. ...........................................................................420
Porella densifolia var. paraphyllina (P.C.Chen) Pócs ....................................................................420
Porella densifolia var. pilosa S.Hatt. et K.C.Chang .......................................................................420
Porella densifolia var. robusta (Steph.) S.Hatt...............................................................................420
Porella elegantula (Mont.) E.A.Hodgs. ........................................................................................421
Porella faurieri (Steph.) S.Hatt. ...................................................................................................421
Porella fengii P.C.Chen et S.Hatt. ...............................................................................................421
Porella geheebii (Steph.) S.Hatt. ..................................................................................................421
Porella gracillima Mitt.................................................................................................................421
Porella grandifolia (Steph.) S.Hatt. ..............................................................................................421
Porella grandiloba Lindb. ............................................................................................................421
Porella grollei S.Hatt. ..................................................................................................................421
Porella handelii S.Hatt. ...............................................................................................................421
Porella hattorii Udar et Shaheen ..................................................................................................421
Porella hoeana S.Hatt. .................................................................................................................421
Porella imbricata Lour. ................................................................................................................421
Porella inaequalis (Gottsche) Perss. .............................................................................................421
Porella japonica (Sande Lac.) Mitt. ..............................................................................................421
Porella japonica subsp. appalachiana R.M.Schust. .......................................................................421
Porella japonica var. densespinosa S.Hatt. et M.X.Zhang ..............................................................421
Porella javanica (Gottsche) Inoue................................................................................................421
Porella latifolia J.S.Lou et Q.Li ...................................................................................................421
Porella leiboldii (Lehm.) Trevis. ..................................................................................................421
Porella longifolia (Steph.) S.Hatt. ................................................................................................421
Porella madagascariensis (Nees et Mont.) Trevis. .........................................................................421
Porella maxima (Steph.) M.L.So .................................................................................................422
Porella mexicana (Hampe ex Gottsche, Lindenb. et Nees) Trevis. ...............................................422
Porella montantii (Steph.) E.W.Jones ..........................................................................................422
Porella navicularis (Lehm. et Lindenb.) Pfeif. .............................................................................422
Porella nitens (Steph.) S.Hatt. .....................................................................................................422
Porella oblongifolia S.Hatt. ..........................................................................................................422
Porella obtusata (Taylor) Trevis. ..................................................................................................422
Porella obtusata var. macroloba (Steph.) S.Hatt. et M.X.Zhang....................................................422
Porella obtusiloba S.Hatt. ............................................................................................................422
Porella perrottetiana (Mont.) Trevis.............................................................................................422
Porella perrottetiana var. angustifolia Pócs ....................................................................................422
Porella perrottetiana var. ciliatodentata (P.C.Chen et P.C.Wu) S.Hatt. ........................................422
Porella perrottetiana var. triciliata (Steph.) Pócs ...........................................................................422
Porella pinnata L. ........................................................................................................................422
Porella planifolia J.S.Lou .............................................................................................................422
Porella platyphylla (L.) Pfeif........................................................................................................422
632
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Porella platyphylloidea (Schwein.) Lindb. .....................................................................................423
Porella plicata J.S.Lou .................................................................................................................423
Porella plumosa (Mitt.) Parihar ....................................................................................................423
Porella prolixa (Gottsche) E.W.Jones ..........................................................................................423
Porella pulcherrima Herzog et S.Hatt. .........................................................................................423
Porella relexa (Lehm. et Lindenb.) Trevis. ..................................................................................423
Porella revoluta (Lehm. et Lindenb.) Trevis. ................................................................................423
Porella revoluta var. propinqua (C.Massal.) S.Hatt. ......................................................................423
Porella roellii Steph. ....................................................................................................................423
Porella saccata M.L.So .................................................................................................................423
Porella sichuanensis S.Hatt. et K.C.Chang ...................................................................................423
Porella spinulosa (Steph.) S.Hatt..................................................................................................423
Porella squamulifera (Taylor) Trevis. ...........................................................................................423
Porella stephaniana (C.Massal.) S.Hatt. .......................................................................................423
Porella subdentata (Mitt.) Steph. .................................................................................................423
Porella subdentata var. camerunensis E.W.Jones ...........................................................................423
Porella subobtusa (Steph.) S.Hatt.................................................................................................423
Porella subparaphyllina J.S.Lou....................................................................................................423
Porella subsquarrosa (Nees et Mont.) Trevis. ...............................................................................424
Porella swailsii Grolle ..................................................................................................................424
Porella swartziana (F.Weber) Trevis. ...........................................................................................424
Porella triquetra (Steph.) E.W.Jones............................................................................................424
Porella truncata J.S.Lou...............................................................................................................424
Porella ulophylla (Steph.) S.Hatt..................................................................................................424
Porella undatorevoluta J.S.Lou .....................................................................................................424
Porella urceolata S.Hatt. ..............................................................................................................424
Porella urogea (C.Massal.) S.Hatt. ...............................................................................................424
Porella vallis-gratiae (Gottsche) E.W.Jones ..................................................................................424
Porella variabilis (Kashyap et R.S.Chopra) Parihar ......................................................................424
Porella vernicosa Lindb. ...............................................................................................................424
Porella viridissima (Mitt.) Grolle .................................................................................................424
Porella wataugensis (Sull.) Underw. ex M.Howe ..........................................................................424
Prasanthus jamalicus Potemkin....................................................................................................114
Prasanthus suecicus (Gottsche) Lindb...........................................................................................114
Preissia quadrata (Scop.) Nees .....................................................................................................492
Preissia quadrata subsp. hyperborea R.M.Schust. .........................................................................492
Prionolejeunea aemula (Gottsche) A.Evans ..................................................................................351
Prionolejeunea ampliretis Herzog .................................................................................................351
Prionolejeunea arguta (Nees) Steph..............................................................................................351
Prionolejeunea corbisieri Pearson ..................................................................................................351
Prionolejeunea decora (Taylor) Steph. ..........................................................................................351
Prionolejeunea denticulata (F.Weber) Schifn...............................................................................351
Prionolejeunea diversitexta (Hampe et Gottsche) Steph................................................................352
Prionolejeunea exauriculata A.Evans ............................................................................................352
Prionolejeunea galliotii Steph. ......................................................................................................352
Prionolejeunea grata (Gottsche) Schifn. ......................................................................................352
Prionolejeunea grollei Ilk.-Borg. et Schäf.-Verw. ..........................................................................352
Prionolejeunea guadalupensis (Lindenb.) Steph. ...........................................................................352
Prionolejeunea limpida Herzog ....................................................................................................352
Prionolejeunea maculata Herzog ..................................................................................................353
Prionolejeunea magnistipula Herzog.............................................................................................352
Prionolejeunea meissneri (Gottsche) Steph. ..................................................................................353
World checklist of hornworts and liverworts
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633
Prionolejeunea mucronata (Sande Lac.) Steph. .............................................................................352
Prionolejeunea muricatoserrulata (Spruce) Steph. .........................................................................352
Prionolejeunea principensis Vanden Berghen ................................................................................352
Prionolejeunea recurvula (Spruce) Steph. .....................................................................................352
Prionolejeunea scaberula (Spruce) Steph.......................................................................................352
Prionolejeunea schlimiana (Gottsche) Steph. ................................................................................352
Prionolejeunea trachyodes (Spruce) Steph. ....................................................................................352
Protocephalozia ephemeroides (Spruce) K.I.Goebel .......................................................................182
Protolophozia androgyna R.M.Schust. ex Váňa et L.Söderstr. .........................................................76
Protolophozia autoica (R.M.Schust.) Váňa et L.Söderstr. ...............................................................76
Protolophozia crispata (R.M.Schust.) Váňa et L.Söderstr................................................................76
Protolophozia druceae (Grolle et E.A.Hodgs.) Váňa et L.Söderstr. .................................................76
Protolophozia elongata (Steph.) Schljakov ......................................................................................76
Protolophozia herzogiana (E.A.Hodgs. et Grolle) Váňa et L.Söderstr. ............................................76
Protolophozia lancistipa (Grolle) Váňa et L.Söderstr. .....................................................................76
Protolophozia leucorhiza (Mitt.) Váňa et L.Söderstr. ......................................................................76
Protolophozia longilora (Herzog) L.Söderstr. et Váňa ....................................................................76
Protolophozia monoica (E.A.Hodgs.) Váňa et L.Söderstr. ...............................................................76
Protolophozia multicuspidata (Hook.f. et Taylor) Váňa et L.Söderstr. ............................................76
Protolophozia nivicola (R.M.Schust.) Váňa et L.Söderstr. ..............................................................76
Protolophozia perssoniana (H.A.Mill.) Váňa et L.Söderstr. .............................................................76
Protolophozia subalpina (R.M.Schust.) Váňa et L.Söderstr.............................................................76
Protolophozia tasmanica (R.M.Schust.) Váňa et L.Söderstr. ...........................................................76
Protolophozia verruculosa (R.M.Schust.) Váňa et L.Söderstr. .........................................................76
Protosyzygiella pseudoconnexa (Herzog) R.M.Schust. ......................................................................46
Pseudocephalozia cucullata J.J.Engel et R.M.Schust. ....................................................................168
Pseudocephalozia lepidozioides R.M.Schust...................................................................................169
Pseudocephalozia leptodictyon R.M.Schust....................................................................................169
Pseudocephalozia lobulata (Herzog) R.M.Schust. .........................................................................168
Pseudocephalozia paludicola R.M.Schust. .....................................................................................169
Pseudocephalozia quadriloba (Steph.) R.M.Schust. ......................................................................168
Pseudocephaloziella epiphytica R.M.Schust. ....................................................................................81
Pseudoisotachis pocsii Váňa.............................................................................................................99
Pseudolepicolea andoi (R.M.Schust.) Inoue ..................................................................................254
Pseudolepicolea fryei (Perss.) Grolle et Ando.................................................................................254
Pseudolepicolea grolleana (R.M.Schust.) Grolle ............................................................................254
Pseudolepicolea kuehnemannii (R.M.Schust.) Hässel ....................................................................254
Pseudolepicolea quadrilaciniata (Sull.) Fulford et J.Taylor ............................................................254
Pseudolepicolea temnomoides (R.M.Schust.) Váňa et J.J.Engel ......................................................255
Pseudolepicolea trollii (Herzog) Grolle et Ando ............................................................................255
Pseudolepicolea trollii var. darjeelingensis S.Hatt. et Mizut. ...........................................................255
Pseudolophocolea denticulata R.M.Schust. et J.J.Engel .................................................................253
Pseudomarsupidium aureocinctum (R.M.Schust.) J.J.Engel ............................................................45
Pseudomarsupidium borneensis (Grolle) Váňa, L.Söderstr., A.Hagborg et von Konrat ....................45
Pseudomarsupidium decipiens (Hook.) Grolle ................................................................................45
Pseudomarsupidium piliferum (Steph.) Herzog ex Grolle ...............................................................45
Pseudotritomaria heterophylla (R.M.Schust.) Konstant. et Vilnet ...................................................85
Psiloclada clandestina Mitt...........................................................................................................185
Psiloclada clandestina subsp. melanesica R.M.Schust. ...................................................................185
Psiloclada clandestina subsp. spinosa (S.W.Arnell) R.M.Schust. ...................................................185
Pteropsiella frondiformis Spruce ...................................................................................................185
Pteropsiella metzgeriiformis R.M.Schust. ......................................................................................185
634
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Ptilidium californicum (Austin) Pearson ......................................................................................434
Ptilidium ciliare (L.) Hampe .......................................................................................................435
Ptilidium pulcherrimum (Weber) Vain. .......................................................................................435
Ptychanthus africanus Steph. ........................................................................................................411
Ptychanthus stephensonianus (Mitt.) Steph. ..................................................................................411
Ptychanthus striatus (Lehm. et Lindenb.) Nees ............................................................................411
Ptychanthus striatus var. intermedius (Gottsche) Verd. .................................................................411
Pycnolejeunea anotomensis Tixier .................................................................................................396
Pycnolejeunea borneensis Steph. ....................................................................................................396
Pycnolejeunea cavistipula (Steph.) Mizut. .....................................................................................397
Pycnolejeunea connivens Schifn. ex P.Syd. ...................................................................................397
Pycnolejeunea contigua (Nees) Grolle ...........................................................................................397
Pycnolejeunea convexifolia (Mitt.) Steph. .....................................................................................397
Pycnolejeunea decurviloba Steph. .................................................................................................397
Pycnolejeunea densistipula (Lehm. et Lindenb.) Steph. .................................................................397
Pycnolejeunea gradsteinii Ilk.-Borg. ..............................................................................................397
Pycnolejeunea grandiocellata Steph. ..............................................................................................397
Pycnolejeunea grossiloba Steph......................................................................................................397
Pycnolejeunea macroloba (Nees et Mont.) Schifn. .......................................................................397
Pycnolejeunea monophthalma (R.M.Schust.) Xiao L.He ...............................................................397
Pycnolejeunea novae-caledoniae (Steph.) Horik. ...........................................................................397
Pycnolejeunea palmicola Steph. ....................................................................................................397
Pycnolejeunea papillosa Xiao L.He ...............................................................................................397
Pycnolejeunea porrectilobula C.J.Bastos et O.Yano .......................................................................397
Pycnolejeunea retusa R.M.Schust. ................................................................................................397
Pycnolejeunea schlimiana Steph....................................................................................................397
Pycnolejeunea schwaneckei (Steph.) Schifn. ex P.Syd. ..................................................................398
Pycnolejeunea sphaeroides (Sande Lac.) J.B.Jack et Steph. .............................................................398
Radula acuminata Steph..............................................................................................................426
Radula acuta Mitt. ......................................................................................................................427
Radula acutangula Steph. ............................................................................................................430
Radula acutiloba Steph. ...............................................................................................................428
Radula aguirrei R.M.Schust. .......................................................................................................426
Radula allisonii Castle .................................................................................................................427
Radula amentulosa Mitt...............................................................................................................425
Radula amoena Herzog ...............................................................................................................427
Radula anceps Sande Lac. ............................................................................................................427
Radula aneurismalis (Hook.f. et Taylor) Gottsche, Lindenb. et Nees...........................................425
Radula angulata Steph. ...............................................................................................................430
Radula anisotoma M.A.M.Renner ...............................................................................................426
Radula ankeinensis Gottsche .......................................................................................................429
Radula antilleana Castle ..............................................................................................................429
Radula appressa Mitt. ..................................................................................................................428
Radula aquilegia (Hook.f. et Taylor) Gottsche, Lindenb. et Nees................................................428
Radula assamica Steph.................................................................................................................426
Radula auriculata Steph. .............................................................................................................426
Radula australiana K.Yamada......................................................................................................426
Radula australis Austin ................................................................................................................428
Radula bipinnata Mitt.................................................................................................................426
Radula bogotensis Steph. ..............................................................................................................430
Radula bolanderi Gottsche ..........................................................................................................430
Radula boninensis Furuki et K.Yamada ........................................................................................430
World checklist of hornworts and liverworts
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635
Radula borneensis Steph...............................................................................................................428
Radula boryana (F.Weber) Nees ex Mont....................................................................................426
Radula brasilica K.Yamada ..........................................................................................................430
Radula brunnea Steph. ................................................................................................................426
Radula buccinifera (Hook.f. et Taylor) Gottsche, Lindenb. et Nees .............................................426
Radula caduca K.Yamada ............................................................................................................428
Radula caespitosa Steph. ..............................................................................................................430
Radula campanigera Mont...........................................................................................................426
Radula campanigera subsp. obiensis (S.Hatt.) K.Yamada ..............................................................426
Radula campanulata Lindenb. et Gottsche ..................................................................................430
Radula carringtonii J.B.Jack ........................................................................................................428
Radula castlei Grolle....................................................................................................................430
Radula cavifolia Hampe ex Gottsche, Lindenb. et Nees ..............................................................430
Radula ceylanica K.Yamada .........................................................................................................425
Radula chinensis Steph. ...............................................................................................................426
Radula cochabambaensis K.Yamada .............................................................................................430
Radula comorensis Steph. .............................................................................................................429
Radula complanata (L.) Dumort..................................................................................................428
Radula conferta Lindenb. et Gottsche ..........................................................................................430
Radula constricta Steph................................................................................................................428
Radula cordata Mitt. ...................................................................................................................430
Radula costaricensis Gottsche .......................................................................................................431
Radula crenulata Schifn..............................................................................................................427
Radula cubensis K.Yamada ..........................................................................................................429
Radula curvilobula M.L.So ..........................................................................................................431
Radula cuspidata Steph................................................................................................................427
Radula decora Gottsche ...............................................................................................................427
Radula decurrens Mitt. ................................................................................................................431
Radula demissa M.A.M.Renner ...................................................................................................426
Radula densifolia Castle ...............................................................................................................431
Radula diaphana K.I.Goebel .......................................................................................................431
Radula diversifolia Steph. ............................................................................................................429
Radula dolabrata K.Yamada ........................................................................................................431
Radula eggersii K.Yamada ............................................................................................................429
Radula elliottii Castle ..................................................................................................................431
Radula emarginata K.Yamada et Piippo ......................................................................................428
Radula episcia Spruce ..................................................................................................................429
Radula evansii Castle...................................................................................................................426
Radula evelynae K.Yamada ..........................................................................................................428
Radula falcata Steph....................................................................................................................431
Radula fauriana Steph.................................................................................................................431
Radula fendleri Gottsche .............................................................................................................428
Radula fernandezana Steph. ........................................................................................................431
Radula issifolia Steph..................................................................................................................425
Radula laccida Lindenb. et Gottsche ..........................................................................................426
Radula laccida var. brachycalyx Spruce ........................................................................................426
Radula lavifolia (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ................................................431
Radula loridana Castle ...............................................................................................................429
Radula foricata M.A.M.Renner ..................................................................................................426
Radula formosa (C.F.W.Meissn. ex Spreng.) Nees .......................................................................425
Radula fujitae Furuki ..................................................................................................................431
Radula fulvifolia (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ................................................429
636
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Radula galapagona Steph. ............................................................................................................431
Radula gedena Gottsche ..............................................................................................................431
Radula gottscheana Taylor ...........................................................................................................426
Radula gracilis Mitt. ....................................................................................................................431
Radula gradsteinii K.Yamada .......................................................................................................431
Radula grandifolia Steph. ............................................................................................................431
Radula grandis Steph. ..................................................................................................................428
Radula grevilleana Taylor ............................................................................................................426
Radula grollei K.Yamada et Piippo ..............................................................................................431
Radula guyanensis K.Yamada .......................................................................................................431
Radula hastata Steph. ..................................................................................................................429
Radula hattorii K.Yamada ...........................................................................................................431
Radula hawaiica M.L.So .............................................................................................................431
Radula hedingeri K.I.Goebel .......................................................................................................431
Radula helix (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ......................................................425
Radula hicksiae K.Yamada ...........................................................................................................425
Radula holstiana Steph. ...............................................................................................................429
Radula holtii Spruce ....................................................................................................................430
Radula husnotii Castle .................................................................................................................430
Radula imposita M.A.M.Renner ..................................................................................................427
Radula inlexa Gottsche ..............................................................................................................430
Radula inouei K.Yamada .............................................................................................................431
Radula involvens Spruce ..............................................................................................................431
Radula iwatsukiana K.Yamada ....................................................................................................431
Radula iwatsukii K.Yamada.........................................................................................................425
Radula jamaicensis Pearson..........................................................................................................431
Radula jamesonii Taylor ..............................................................................................................431
Radula japonica Gottsche ............................................................................................................428
Radula javanica Gottsche ............................................................................................................428
Radula jonesii Bouman, Dirkse et K.Yamada...............................................................................428
Radula jovetiana K.Yamada .........................................................................................................427
Radula kegelii Gottsche ex Steph. ................................................................................................430
Radula kilgourii M.A.M.Renner..................................................................................................427
Radula kinabaluensis K.Yamada ..................................................................................................431
Radula kitagawae K.Yamada .......................................................................................................431
Radula kojana Steph. ..................................................................................................................428
Radula koponenii K.Yamada et Piippo .........................................................................................431
Radula kurzii Steph. ...................................................................................................................431
Radula lacerata Steph. .................................................................................................................428
Radula laxiramea Steph...............................................................................................................431
Radula leiboldii Steph. ................................................................................................................432
Radula lewisii K.Yamada .............................................................................................................432
Radula ligula Steph. ....................................................................................................................432
Radula lindenbergiana Gottsche ex C.Hartm...............................................................................428
Radula lingulata Gottsche ...........................................................................................................432
Radula longiloba K.Yamada .........................................................................................................432
Radula longispica Steph. ..............................................................................................................432
Radula loriana Castle ..................................................................................................................427
Radula macroloba Steph. .............................................................................................................430
Radula madagascariensis Gottsche ...............................................................................................428
Radula mammosa Spruce .............................................................................................................427
Radula marginata Gottsche, Lindenb. et Nees ............................................................................432
World checklist of hornworts and liverworts
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637
Radula marojezica E.W.Jones ......................................................................................................428
Radula mauiensis M.L.So ............................................................................................................432
Radula mazarunensis K.Yamada ..................................................................................................430
Radula mexicana Lindenb. et Gottsche .......................................................................................430
Radula microloba Gottsche ..........................................................................................................430
Radula microlobula Castle ...........................................................................................................432
Radula minutilobula K.Yamada et Piippo ....................................................................................432
Radula mittenii Steph..................................................................................................................427
Radula mizutanii K.Yamada .......................................................................................................432
Radula morobeana K.Yamada et Piippo .......................................................................................425
Radula multiamentula E.A.Hodgs. ..............................................................................................425
Radula multilora Gottsche ex Schifn. ........................................................................................429
Radula myriopoda M.A.M.Renner...............................................................................................427
Radula neotropica Castle..............................................................................................................430
Radula nigra Pearson ..................................................................................................................432
Radula nilgiriensis Udar et D.Kumar ...........................................................................................432
Radula norrisii K.Yamada et Piippo ............................................................................................432
Radula notabilis M.A.M.Renner..................................................................................................427
Radula novae-hollandiae Hampe .................................................................................................428
Radula novivrieseana K.Yamada ..................................................................................................432
Radula novocaledonica Hürl. et K.Yamada...................................................................................432
Radula novocaledoniensis K.Yamada.............................................................................................432
Radula novoguineensis K.Yamada et Piippo..................................................................................429
Radula nudicaulis Steph. .............................................................................................................430
Radula nudicaulis var. delicatula P.Allorge et V.Allorge ...............................................................430
Radula nymannii Steph. ..............................................................................................................427
Radula obconica Sull....................................................................................................................429
Radula obovata Castle .................................................................................................................432
Radula obscura Mitt. ...................................................................................................................432
Radula obtusiloba Steph. .............................................................................................................429
Radula obtusiloba subsp. polyclada (A.Evans) S.Hatt....................................................................429
Radula oceania Castle ..................................................................................................................429
Radula ocellata K.Yamada ...........................................................................................................428
Radula okamurana Steph. ...........................................................................................................432
Radula onraedtii K.Yamada .........................................................................................................432
Radula opaciuscula (Spruce) Castle ..............................................................................................432
Radula oreopsis M.A.M.Renner ...................................................................................................429
Radula ornata E.A.Br. et Pócs .....................................................................................................425
Radula ovalilobula K.Yamada ......................................................................................................432
Radula oyamensis Steph. ..............................................................................................................432
Radula paganii Castle ..................................................................................................................432
Radula pallens (Sw.) Nees ex Mont. ............................................................................................432
Radula pandei Udar et Dh.Kumar...............................................................................................432
Radula patens K.Yamada .............................................................................................................433
Radula perrottetii Gottsche ..........................................................................................................426
Radula peruviana K.Yamada .......................................................................................................433
Radula philippinensis K.Yamada ..................................................................................................433
Radula physoloba Mont. ..............................................................................................................425
Radula pinnulata Mitt.................................................................................................................433
Radula plicata Mitt. ....................................................................................................................428
Radula pocsii K.Yamada ..............................................................................................................430
Radula portoricensis Steph. ..........................................................................................................429
638
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Radula prolifera Arnell ................................................................................................................429
Radula protensa Lindenb. ............................................................................................................427
Radula protensa var. erectilobula Schifn.......................................................................................427
Radula pseudolaccida E.W.Jones .................................................................................................427
Radula pseudoscripta M.A.M.Renner ...........................................................................................425
Radula pseudostachya Spruce........................................................................................................433
Radula psychosis M.A.M.Renner ..................................................................................................427
Radula pugioniformis M.A.M.Renner ..........................................................................................428
Radula pulchella Mitt. .................................................................................................................428
Radula punctata Steph. ...............................................................................................................433
Radula pusilla Spruce ..................................................................................................................433
Radula quadrata Gottsche ...........................................................................................................429
Radula queenslandica K.Yamada ..................................................................................................425
Radula ratkowskiana K.Yamada ..................................................................................................427
Radula recubans Taylor ...............................................................................................................430
Radula relexa Nees et Mont. ......................................................................................................429
Radula retrolexa Taylor ..............................................................................................................428
Radula rhombiloba Steph. ...........................................................................................................433
Radula robinsonii Steph...............................................................................................................427
Radula rupicola K.Yamada ..........................................................................................................433
Radula saccatiloba Steph..............................................................................................................430
Radula santacruziana K.Yamada et Gradst. .................................................................................433
Radula scariosa Mitt. ...................................................................................................................425
Radula schaefer-verwimpii K.Yamada...........................................................................................430
Radula schoieldiana K.Yamada ...................................................................................................430
Radula sharpii K.Yamada ............................................................................................................429
Radula silvestris Gottsche ............................................................................................................429
Radula sinskeana K.Yamada ........................................................................................................433
Radula sinuata Gottsche ex Steph. ..............................................................................................433
Radula socorana Gerola ...............................................................................................................433
Radula sonsonensis Steph. ............................................................................................................433
Radula splendida M.A.M.Renner et Devos ..................................................................................425
Radula squarrosa K.Yamada ........................................................................................................425
Radula stellatogemmipara C.Gao et Y.H.Wu ...............................................................................433
Radula stenocalyx Mont. ..............................................................................................................427
Radula stipatilora Steph..............................................................................................................430
Radula strangulata Hook.f. et Taylor ..........................................................................................427
Radula striata Mitt. .....................................................................................................................430
Radula subinlata Lindenb. et Gottsche .......................................................................................430
Radula subsimplex Steph..............................................................................................................433
Radula subsquarrosa S.W.Arnell ..................................................................................................433
Radula sullivantii Austin .............................................................................................................430
Radula sumatrana Steph..............................................................................................................429
Radula tabularis Steph. ...............................................................................................................433
Radula tasmanica Steph...............................................................................................................428
Radula taylorii Steph. ..................................................................................................................433
Radula tectiloba Steph. ................................................................................................................433
Radula tenax Lindb. ....................................................................................................................426
Radula tenera Mitt. .....................................................................................................................430
Radula tenuis K.Yamada..............................................................................................................433
Radula thiersiae K.Yamada ..........................................................................................................425
Radula tjibodensis K.I.Goebel ......................................................................................................427
World checklist of hornworts and liverworts
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639
Radula tokiensis Steph. ................................................................................................................429
Radula underwoodii Castle ..........................................................................................................429
Radula uvifera (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ...................................................425
Radula vagans Steph....................................................................................................................425
Radula van-zantenii K.Yamada ...................................................................................................429
Radula varilobula Castle..............................................................................................................429
Radula venezuelensis K.Yamada ...................................................................................................433
Radula ventricosa Steph. ..............................................................................................................427
Radula verrucosa K.Yamada.........................................................................................................425
Radula vieillardii Gottsche ..........................................................................................................433
Radula visianica C.Massal. ..........................................................................................................433
Radula voluta Taylor ...................................................................................................................430
Radula vrieseana Sande Lac. ........................................................................................................433
Radula weymouthiana Steph. .......................................................................................................428
Radula wichurae Steph. ...............................................................................................................429
Radula wrightii Castle .................................................................................................................433
Radula xalapensis Nees et Mont. .................................................................................................434
Radula yanoella R.M.Schust. .......................................................................................................427
Reboulia hemisphaerica (L.) Raddi ...............................................................................................483
Reboulia hemisphaerica subsp. acrogyna (R.M.Schust.) R.M.Schust. ............................................483
Reboulia hemisphaerica subsp. australis R.M.Schust. ....................................................................483
Reboulia hemisphaerica subsp. dioica R.M.Schust. .......................................................................483
Reboulia hemisphaerica var. issisquama Herzog............................................................................483
Reboulia hemisphaerica subsp. orientalis R.M.Schust....................................................................484
Reboulia hemisphaerica var. turkestanica C.E.O.Jensen ex Herzog................................................484
Rectolejeunea colombiana R.M.Schust. .........................................................................................393
Rectolejeunea emarginulilora (Schifn.) A.Evans ..........................................................................393
Rectolejeunea lagelliformis A.Evans ..............................................................................................393
Rectolejeunea lagelliformis subsp. hamata R.M.Schust. ................................................................393
Rectolejeunea lindenbergii Steph...................................................................................................393
Rectolejeunea lindigiana Steph. ....................................................................................................394
Rectolejeunea pachyderma R.M.Schust. ........................................................................................394
Rectolejeunea queenslandica (B.M.hiers) Xiao L.He ...................................................................393
Rectolejeunea truncatilobula C.J.Bastos ........................................................................................393
Rectolejeunea versifolia (Schifn.) L.Söderstr. et A.Hagborg ..........................................................393
Riccardia aberrans (Steph.) Gradst. .............................................................................................445
Riccardia aequicellularis (Steph.) Hewson....................................................................................439
Riccardia aequitexta (Steph.) E.A.Br. ...........................................................................................441
Riccardia aeruginosa Furuki .........................................................................................................439
Riccardia agumana Hewson ........................................................................................................445
Riccardia alba (Colenso) E.A.Br. .................................................................................................441
Riccardia albomarginata (Steph.) Schifn. ....................................................................................439
Riccardia alcicornis (Hook.f. et Taylor) Trevis. ............................................................................440
Riccardia algoides (Taylor) Meenks..............................................................................................445
Riccardia amazonica (Spruce) Schifn. ex Gradst. et Hekking ......................................................445
Riccardia amnicola Hässel ...........................................................................................................441
Riccardia andina (Spruce) Herzog ...............................................................................................441
Riccardia angustata Horik. ..........................................................................................................445
Riccardia angustealata (Steph.) Hewson ......................................................................................445
Riccardia angustissima (Steph.) H.A.Mill. ....................................................................................445
Riccardia arcuata Furuki .............................................................................................................439
Riccardia argentolimbata Hewson et Grolle .................................................................................438
640
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Riccardia aspera (Steph.) Grolle...................................................................................................445
Riccardia asperulata R.M.Schust..................................................................................................439
Riccardia australis (Lehm.) Hewson ............................................................................................439
Riccardia autoica (Steph.) A.Evans ..............................................................................................443
Riccardia baldwinii (Steph.) H.A.Mill. ........................................................................................446
Riccardia barbilora (Steph.) Piippo.............................................................................................446
Riccardia baumannii Hürl. ..........................................................................................................445
Riccardia bipinnatiida (Colenso) Hewson ..................................................................................441
Riccardia bogotensis (Gottsche) Pagán ..........................................................................................446
Riccardia boliviensis (Steph.) Meenks ..........................................................................................446
Riccardia bongeriana Hewson ......................................................................................................446
Riccardia breviala E.A.Br. ...........................................................................................................443
Riccardia breviramosa (Steph.) A.Evans .......................................................................................443
Riccardia brunnea (Steph.) S.Hatt. ..............................................................................................446
Riccardia calcarea (Steph.) Meenks..............................................................................................446
Riccardia calva (Schifn.) A.Evans ...............................................................................................441
Riccardia canaliculata (Nees) Kuntze...........................................................................................439
Riccardia capillacea (Steph.) Meenks et C.De Jong......................................................................446
Riccardia capillacea var. dentata Meenks ......................................................................................446
Riccardia cardotii (Steph.) Pandé et S.C.Srivast. ..........................................................................446
Riccardia cataractarum (Spruce) Schifn. .....................................................................................446
Riccardia cervicornis (Spruce) Herzog ex Gradst. et Hekking .......................................................446
Riccardia chamedryfolia (With.) Grolle ........................................................................................439
Riccardia changbaishanensis C.Gao ..............................................................................................439
Riccardia chinensis C.Gao............................................................................................................446
Riccardia ciliolata (Spruce) Horik................................................................................................446
Riccardia cochleata (Hook.f. et Taylor) Kuntze............................................................................439
Riccardia colensoi (Steph.) W.Martin...........................................................................................441
Riccardia columbica (Steph.) Hässel ex Gradst. et Hekking .........................................................447
Riccardia comata (Steph.) H.A.Mill.............................................................................................447
Riccardia compacta (Steph.) S.W.Arnell.......................................................................................447
Riccardia comptonii (Pearson) H.A.Mill. .....................................................................................447
Riccardia conimitra (Steph.) A.Evans...........................................................................................440
Riccardia corralensis (Steph.) A.Evans ..........................................................................................440
Riccardia costata (Steph.) Hürl. ...................................................................................................447
Riccardia crassa (Schwägr.) C.Massal. ..........................................................................................441
Riccardia crassicaulis (Steph.) Meenks et C.De Jong ....................................................................447
Riccardia crassicrispa (Steph.) A.Evans .........................................................................................441
Riccardia crassiretis Schifn. .........................................................................................................447
Riccardia crenulata Schifn. .........................................................................................................447
Riccardia crenuliformis R.M.Schust. ............................................................................................447
Riccardia decolyana Schifn. .........................................................................................................447
Riccardia deguchii Furuki et K.T.Yong ........................................................................................445
Riccardia densiramea (Steph.) S.Hatt...........................................................................................447
Riccardia devexa Schifn. .............................................................................................................447
Riccardia diablotina (Spruce) Pagán ............................................................................................447
Riccardia diderma Hässel.............................................................................................................441
Riccardia digitiloba (Spruce) Pagán .............................................................................................447
Riccardia dilatata (Spruce) Schäf.-Verw. et Pócs .........................................................................447
Riccardia diminuta Schifn. .........................................................................................................447
Riccardia diminuta var. thermarum Schifn..................................................................................447
Riccardia distans (Spruce) Pagán..................................................................................................448
World checklist of hornworts and liverworts
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641
Riccardia diversilora A.Evans ......................................................................................................441
Riccardia diversilora subsp. paucigyna R.M.Schust. .....................................................................441
Riccardia duriuscula Hässel .........................................................................................................441
Riccardia elata (Steph.) Schifn....................................................................................................448
Riccardia elata var. laccida Schifn. .............................................................................................448
Riccardia elata var. intercedens Schifn. ........................................................................................448
Riccardia elegans (Steph.) Hürl. ...................................................................................................448
Riccardia elisabethae houvenot et Reeb ......................................................................................448
Riccardia emarginata (Steph.) K.G.Hell ......................................................................................448
Riccardia eriocaula (Hook.) C.Massal. .........................................................................................439
Riccardia erosa (Steph.) E.W.Jones ..............................................................................................448
Riccardia falsiloribunda Hässel ...................................................................................................442
Riccardia fastigiata (Lehm.) Trevis. .............................................................................................448
Riccardia fendleri (Steph.) Pagán .................................................................................................448
Riccardia ilicina (Colenso) E.A.Hodgs. ......................................................................................444
Riccardia laccida (Steph.) S.Hatt. ...............................................................................................448
Riccardia laccidissima Schifn......................................................................................................448
Riccardia lagellaris (A.Gepp) H.A.Mill. ......................................................................................448
Riccardia lagellifrons C.Gao ........................................................................................................448
Riccardia lavovirens Furuki .........................................................................................................439
Riccardia leischeri (Steph.) H.A.Mill...........................................................................................448
Riccardia loribunda (Steph.) A.Evans .........................................................................................442
Riccardia luvigena Hässel............................................................................................................442
Riccardia foliacea Meenks et C.De Jong ......................................................................................448
Riccardia formosensis (Steph.) Horik. ...........................................................................................448
Riccardia fruticosa (Steph.) Furuki...............................................................................................439
Riccardia fucoidea (Sw.) C.Massal. ..............................................................................................448
Riccardia fuegiensis C.Massal. ......................................................................................................439
Riccardia furtiva E.A.Br. et Braggins ...........................................................................................440
Riccardia fuscobrunnea (Steph.) A.Evans......................................................................................441
Riccardia geniana Hewson...........................................................................................................449
Riccardia georgiensis (Steph.) Hässel ............................................................................................442
Riccardia georgiensis subsp. sympodea R.M.Schust. .......................................................................442
Riccardia glauca Furuki ...............................................................................................................439
Riccardia glaziovii (Spruce) Meenks ............................................................................................449
Riccardia gogolensis (Steph.) Hewson ...........................................................................................449
Riccardia gracilis (Steph.) R.M.Schust. ........................................................................................449
Riccardia graefei (Steph.) Hewson ..............................................................................................442
Riccardia grandilora (Steph.) H.A.Mill. ......................................................................................449
Riccardia granulata (Steph.) A.Evans...........................................................................................443
Riccardia grollei Furuki ...............................................................................................................449
Riccardia grossidens (Steph.) Pagán ..............................................................................................449
Riccardia grossitexta (Steph.) Furuki ............................................................................................445
Riccardia gunniana (Steph.) H.A.Mill. ........................................................................................449
Riccardia hamatilora (Steph.) H.A.Mill. .....................................................................................449
Riccardia hans-meyeri (Steph.) Meenks et C.De Jong ..................................................................449
Riccardia hans-meyeri var. dentata Meenks ..................................................................................449
Riccardia hattorii Furuki .............................................................................................................439
Riccardia hawaica (Steph.) H.A.Mill. ..........................................................................................449
Riccardia hebridensis (Steph.) H.A.Mill. ......................................................................................449
Riccardia herzogiana (Steph.) Meenks et C.De Jong ....................................................................449
Riccardia heteroclada Schifn. ......................................................................................................449
642
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Riccardia hirtilora (Steph.) Schifn. ............................................................................................449
Riccardia humilis (Gottsche) O.Yano ..........................................................................................450
Riccardia hyalina (Steph.) H.A.Mill. ...........................................................................................450
Riccardia hyalitricha Hässel .........................................................................................................445
Riccardia hydra Hürl. ..................................................................................................................450
Riccardia hymenophylloides Schifn. .............................................................................................450
Riccardia hymenophylloides var. laccida Schifn............................................................................450
Riccardia hymenophytoides (Spruce) Meenks ................................................................................450
Riccardia hypipamensis Hewson ...................................................................................................450
Riccardia ibana Hewson..............................................................................................................450
Riccardia inconspicua (Steph.) Reeb et Bardat..............................................................................445
Riccardia incurvata Lindb. ..........................................................................................................450
Riccardia innovans (Steph.) Pagán ...............................................................................................450
Riccardia insularis Schifn............................................................................................................450
Riccardia intercellula E.A.Br. .......................................................................................................443
Riccardia intricata (Steph.) H.A.Mill. .........................................................................................450
Riccardia jackii Schifn. ...............................................................................................................450
Riccardia jackii var. densa Schifn. ...............................................................................................450
Riccardia judithae Meenks et C.De Jong .....................................................................................450
Riccardia jugata R.M.Schust. ......................................................................................................450
Riccardia karstenii (Steph.) Schifn. .............................................................................................450
Riccardia kodamae Mizut. et S.Hatt. ...........................................................................................439
Riccardia laticostata (Spruce) Schifn. ..........................................................................................450
Riccardia latifrondoides Schifn. ...................................................................................................450
Riccardia latifrons (Lindb.) Lindb................................................................................................440
Riccardia latifrons subsp. arctica R.M.Schust. et Damsh. .............................................................440
Riccardia latifrons var. miyakeana (Schifn.) Furuki .....................................................................440
Riccardia lepidomitra (Spruce) Gradst. ........................................................................................450
Riccardia leptophylla (Spruce) Herzog..........................................................................................451
Riccardia leptostachya A.Evans .....................................................................................................444
Riccardia leptothallus R.M.Schust. ...............................................................................................451
Riccardia levieri Schifn. ..............................................................................................................451
Riccardia lichenoides (Steph.) H.A.Mill. ......................................................................................451
Riccardia ligulata (Steph.) Pócs et Schäf.-Verw. ..........................................................................451
Riccardia lilliena (Steph.) H.A.Mill. ............................................................................................451
Riccardia limbata (Steph.) E.W.Jones .........................................................................................451
Riccardia lobulata (Colenso) E.A.Hodgs......................................................................................444
Riccardia loefgrenii Schifn. .........................................................................................................451
Riccardia longilora (Steph.) Hewson ...........................................................................................451
Riccardia longioleata Hässel .........................................................................................................441
Riccardia longispica (Steph.) Pearson ...........................................................................................451
Riccardia loriana (Steph.) H.A.Mill.............................................................................................451
Riccardia macdonaldiana Hewson ...............................................................................................451
Riccardia macrantha (Pearson) H.A.Mill. ....................................................................................451
Riccardia magnicellularis Furuki ..................................................................................................451
Riccardia marginata (Colenso) Pearson .......................................................................................442
Riccardia marginata var. paciica Furuki ......................................................................................442
Riccardia marionensis R.M.Schust. ..............................................................................................438
Riccardia mejlandii S.W.Arnell ...................................................................................................444
Riccardia metzgeriiformis (Steph.) R.M.Schust. ...........................................................................451
Riccardia microscopica (Nees) Kuntze ..........................................................................................451
Riccardia minuta (Steph.) W.Martin ...........................................................................................451
World checklist of hornworts and liverworts
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643
Riccardia multicorpora E.A.Br. ....................................................................................................452
Riccardia multiida (L.) Gray .......................................................................................................444
Riccardia multiida subsp. decrescens (Steph.) Furuki ...................................................................444
Riccardia multiida subsp. synoica R.M.Schust. ............................................................................444
Riccardia multiidoides Schifn. ....................................................................................................452
Riccardia multioleata Hässel ........................................................................................................444
Riccardia multispica (Steph.) S.Hatt. ...........................................................................................452
Riccardia mycophora A.Evans.......................................................................................................442
Riccardia nadeaudii (Steph.) Hürl. ..............................................................................................452
Riccardia nagasakiensis (Steph.) S.Hatt. .......................................................................................440
Riccardia negeri (Steph.) A.Evans ................................................................................................442
Riccardia newellana (Steph.) H.A.Mill. .......................................................................................452
Riccardia nigra (Steph.) H.A.Mill. ..............................................................................................452
Riccardia nitida (Colenso) E.A.Hodgs.........................................................................................442
Riccardia nobilis (Steph.) Schifn. ................................................................................................452
Riccardia novo-amstelodamensis Schifn. .......................................................................................452
Riccardia nudilora (Steph.) Grolle ..............................................................................................452
Riccardia obtusa S.W.Arnell ........................................................................................................452
Riccardia obtusifrons (Steph.) H.A.Mill. ......................................................................................452
Riccardia omkaliensis Hewson .....................................................................................................452
Riccardia opuntiiformis S.W.Arnell ..............................................................................................443
Riccardia pallida (Spruce) Meenks et C.De Jong .........................................................................452
Riccardia pallidevirens (Steph.) A.Evans ......................................................................................443
Riccardia palmata (Hedw.) Carruth. ...........................................................................................440
Riccardia palmatiida (Steph.) H.A.Mill. .....................................................................................452
Riccardia palmatiformis Schifn. ..................................................................................................452
Riccardia papillata (Gottsche) Hässel ex Gradst. et Hekking .......................................................452
Riccardia papillosa (C.Massal. et Steph.) Hässel...........................................................................442
Riccardia papulosa (Steph.) E.A.Br. .............................................................................................444
Riccardia paramorum Meenks .....................................................................................................452
Riccardia parasitans (Steph.) Meenks et C.De Jong .....................................................................453
Riccardia parvula Schifn.............................................................................................................453
Riccardia patens Hässel ................................................................................................................444
Riccardia pauciramea (Steph.) H.A.Mill. .....................................................................................453
Riccardia paulensis Schifn. ..........................................................................................................453
Riccardia pectinata (Austin) H.A.Mill. ........................................................................................453
Riccardia pectinata var. fasciculata (Steph.) Hürl. ........................................................................438
Riccardia pellucida Piippo ...........................................................................................................453
Riccardia pembaiensis (Steph.) Hürl.............................................................................................453
Riccardia pengagensis Hewson......................................................................................................453
Riccardia pennata E.A.Br.............................................................................................................443
Riccardia perspicua E.A.Br. ..........................................................................................................443
Riccardia perssonii S.C.Srivast. et Udar ........................................................................................453
Riccardia philippinensis Furuki ....................................................................................................453
Riccardia phleganiana Hewson ....................................................................................................453
Riccardia pindensis Hewson .........................................................................................................453
Riccardia plana (Steph.) Hürl......................................................................................................453
Riccardia plana var. minor (Pearson) Hürl. ex H.A.Mill., H.Whittier et B.Whittier ....................453
Riccardia planilora (Steph.) S.Hatt. ............................................................................................440
Riccardia planilora var. aequatorialis Furuki ...............................................................................440
Riccardia planifrons (Spruce) Pagán .............................................................................................453
Riccardia plumiformis (Spruce) Hässel ex Meenks........................................................................453
644
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Riccardia plumosa (Mitt.) E.O.Campb. .......................................................................................453
Riccardia poeppigiana (Lehm. et Lindenb.) Hässel ex Meenks et C.De Jong ................................453
Riccardia polyclada (Mitt.) Hässel................................................................................................444
Riccardia porcina (Hewson) L.Söderstr. .......................................................................................454
Riccardia portoricensis (Steph.) Pagán ..........................................................................................454
Riccardia prehensilis (Hook.f. et Taylor) C.Massal. ......................................................................438
Riccardia pseudodendroceros R.M.Schust. .....................................................................................440
Riccardia pumila Furuki ..............................................................................................................440
Riccardia punahuina (Steph.) H.A.Mill. ......................................................................................454
Riccardia pusilla Grolle ...............................................................................................................440
Riccardia ramosissima (Steph.) Grolle ..........................................................................................454
Riccardia regina Meenks et C.De Jong ........................................................................................454
Riccardia regnellii (Ångstr.) K.G.Hell ..........................................................................................454
Riccardia regularis (Steph.) Kühnem. ..........................................................................................442
Riccardia reyesiana Meenks..........................................................................................................454
Riccardia riccioides Pearson..........................................................................................................454
Riccardia rigida Schifn. ..............................................................................................................454
Riccardia rivularis Hässel.............................................................................................................442
Riccardia robbinsii Hewson et Grolle ..........................................................................................454
Riccardia robusta (Steph.) H.A.Mill. ...........................................................................................454
Riccardia rockii (Steph.) H.A.Mill. ..............................................................................................454
Riccardia rupicola (Steph.) Hewson .............................................................................................454
Riccardia russellii R.M.Schust......................................................................................................454
Riccardia saccatilora (Steph.) S.W.Arnell ....................................................................................454
Riccardia santapaui Udar et S.C.Srivast. ......................................................................................454
Riccardia saxicola Hässel..............................................................................................................442
Riccardia schwaneckei (Steph.) Pagán ..........................................................................................454
Riccardia singapurensis Schifn. ....................................................................................................454
Riccardia smaragdina Meenks et C.De Jong ................................................................................454
Riccardia spectabilis (Steph.) A.Evans ..........................................................................................442
Riccardia spegazziniana C.Massal. ...............................................................................................443
Riccardia spinulifera C.Massal. ....................................................................................................444
Riccardia spongiosa Furuki ...........................................................................................................439
Riccardia sprucei (Steph.) Meenks et C.De Jong ..........................................................................454
Riccardia squamifera Schifn. .......................................................................................................455
Riccardia statensis Hässel .............................................................................................................444
Riccardia stipatilora (Steph.) Pagán ............................................................................................455
Riccardia stricta R.M.Schust........................................................................................................455
Riccardia subalpina Furuki ..........................................................................................................440
Riccardia subantarctica Grolle et L.Söderstr. ................................................................................455
Riccardia subexalata Schifn.........................................................................................................455
Riccardia subexalata var. procera Schifn. .....................................................................................455
Riccardia submersa (Hook.f. et Taylor) Trevis. ............................................................................455
Riccardia submultiida Horik. ......................................................................................................455
Riccardia subpalmata (Steph.) Hürl. ............................................................................................455
Riccardia subsimplex (Steph.) Pagán ............................................................................................455
Riccardia sumatrana Schifn. .......................................................................................................455
Riccardia tahitensis (Steph.) Hürl. ...............................................................................................455
Riccardia tamariscina (Steph.) Schifn. ........................................................................................440
Riccardia tenax (Steph.) A.Evans .................................................................................................443
Riccardia tenella Hewson.............................................................................................................455
Riccardia tenerrima (Steph.) A.Evans...........................................................................................443
World checklist of hornworts and liverworts
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645
Riccardia tenuicula (Spruce) Meenks ...........................................................................................455
Riccardia tenuis (Steph.) Schifn. .................................................................................................455
Riccardia thaxteri A.Evans ...........................................................................................................444
Riccardia theliophora Hässel ........................................................................................................443
Riccardia tjibodensis Schifn. ........................................................................................................455
Riccardia trichomanoides (Spruce) Hässel ex Meenks ...................................................................455
Riccardia tristaniana S.W.Arnell .................................................................................................444
Riccardia trukensis H.A.Mill. et Bonner ......................................................................................455
Riccardia tumbareriensis Hewson .................................................................................................455
Riccardia umida E.A.Br. ..............................................................................................................441
Riccardia upoluna (Steph.) Grolle................................................................................................456
Riccardia valida (Steph.) J.J.Engel ...............................................................................................456
Riccardia venosa (Steph.) Hürl. ...................................................................................................456
Riccardia villosa (Steph.) Pandé et S.C.Srivast. ............................................................................456
Riccardia virens (Steph.) Hürl. ....................................................................................................456
Riccardia virgata (Gottsche) Pagán ..............................................................................................456
Riccardia vitrea Furuki ................................................................................................................440
Riccardia wallisii (Steph.) Gradst. ...............................................................................................456
Riccardia wettsteinii Schifn. ........................................................................................................456
Riccardia wettsteinii var. angustilimbia Schifn. ............................................................................456
Riccardia wettsteinii var. crassa Schifn. ........................................................................................456
Riccardia wettsteinii var. procera Schifn. .....................................................................................456
Riccardia wettsteinii var. tenuiretis Schifn. ..................................................................................456
Riccardia womersleyana Hewson ..................................................................................................456
Riccardia xylophila Hässel............................................................................................................443
Riccia abuensis Bapna ..................................................................................................................499
Riccia acutisulca Steph.................................................................................................................500
Riccia alatospora O.H.Volk et Perold ..........................................................................................495
Riccia albida Sull. ex Austin ........................................................................................................493
Riccia albolimbata S.W.Arnell .....................................................................................................495
Riccia albomarginata Bisch. ex C.Krauss......................................................................................495
Riccia alboporosa Perold ..............................................................................................................495
Riccia albopunctata Jovet-Ast.......................................................................................................493
Riccia albornata O.H.Volk et Perold ...........................................................................................493
Riccia albovestita O.H.Volk ........................................................................................................495
Riccia amboinensis Schifn. ..........................................................................................................500
Riccia ampullacea Perold .............................................................................................................495
Riccia angolensis Steph.................................................................................................................495
Riccia aravalliensis Pandé et Udar................................................................................................500
Riccia argenteolimbata O.H.Volk et Perold .................................................................................495
Riccia arnellii Sultan Khan ..........................................................................................................500
Riccia asprella Carrington et Pearson ...........................................................................................500
Riccia asservanda De Not. ex Lamothe ........................................................................................500
Riccia atlantica Sérgio et Perold ..................................................................................................495
Riccia atromarginata Levier .........................................................................................................496
Riccia atromarginata var. jovet-astiae Rauh et Buchloh ................................................................496
Riccia atropurpurea Sim...............................................................................................................496
Riccia attenuata Pandé ................................................................................................................500
Riccia australis Steph. ..................................................................................................................493
Riccia bahiensis Steph. .................................................................................................................498
Riccia balansae Steph...................................................................................................................500
Riccia beyrichiana Hampe ...........................................................................................................496
646
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Riccia bialbistrata Hässel .............................................................................................................500
Riccia bicarinata Lindb. ..............................................................................................................496
Riccia bicolorata Perold ...............................................................................................................496
Riccia bifurca Hofm. ..................................................................................................................496
Riccia billardierei Mont. et Nees .................................................................................................496
Riccia biokoensis Perold ...............................................................................................................500
Riccia blackii Na-halang............................................................................................................500
Riccia boliviensis Jovet-Ast ...........................................................................................................493
Riccia brasiliensis Schifn. ............................................................................................................493
Riccia breidleri Jur. ex Steph........................................................................................................496
Riccia breutelii Hampe ................................................................................................................493
Riccia brittonii M.Howe .............................................................................................................493
Riccia bullosa Link ......................................................................................................................499
Riccia californica Austin ..............................................................................................................496
Riccia campbelliana M.Howe ......................................................................................................496
Riccia canaliculata Hofm. ..........................................................................................................498
Riccia cancellata Taylor ...............................................................................................................497
Riccia caroliniana Na-halang .....................................................................................................500
Riccia cartilaginosa Steph.............................................................................................................500
Riccia cavernosa Hofm. ..............................................................................................................499
Riccia chartacea K.I.Goebel .........................................................................................................500
Riccia chiapasensis Jovet-Ast ........................................................................................................498
Riccia chinensis Herzog................................................................................................................500
Riccia chudoana Steph. ................................................................................................................493
Riccia ciliata Hofm. ...................................................................................................................496
Riccia ciliifera Link .....................................................................................................................496
Riccia cincta Jovet-Ast .................................................................................................................497
Riccia collata Na-halang ............................................................................................................500
Riccia compacta Garside ..............................................................................................................500
Riccia concava Bisch. ex C.Krauss................................................................................................495
Riccia congoana Steph. ................................................................................................................496
Riccia convexa Steph....................................................................................................................500
Riccia coracina Jovet-Ast .............................................................................................................493
Riccia coronata Sim .....................................................................................................................500
Riccia corrugata Jovet-Ast ............................................................................................................493
Riccia crassa Steph. ......................................................................................................................500
Riccia crassifrons Spruce...............................................................................................................498
Riccia crassivenia Jovet-Ast ..........................................................................................................493
Riccia crenatodentata O.H.Volk ..................................................................................................500
Riccia crinita Taylor ....................................................................................................................496
Riccia crozalsii Levier...................................................................................................................496
Riccia cruciata Kashyap ...............................................................................................................497
Riccia crustata Trab. ....................................................................................................................496
Riccia crystallina L. ......................................................................................................................499
Riccia cubensis S.W.Arnell ...........................................................................................................493
Riccia cupulifera A.V.Duthie .......................................................................................................499
Riccia curtisii (Austin) Austin ......................................................................................................499
Riccia delavayi Steph. ..................................................................................................................500
Riccia deserticola Steph. ...............................................................................................................500
Riccia dictyospora M.Howe ..........................................................................................................496
Riccia discolor Lehm. et Lindenb. ................................................................................................494
Riccia duplex Lorb. ex Müll.Frib. ................................................................................................498
World checklist of hornworts and liverworts
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647
Riccia duplex var. megaspora Na-halang .....................................................................................498
Riccia dussiana Steph. .................................................................................................................498
Riccia eburnea Jovet-Ast ..............................................................................................................497
Riccia ekmanii S.W.Arnell...........................................................................................................494
Riccia elliottii Steph.....................................................................................................................494
Riccia elongata Perold ..................................................................................................................495
Riccia enyae Jovet-Ast ..................................................................................................................494
Riccia erubescens Perold ...............................................................................................................500
Riccia erythrocarpa Jovet-Ast........................................................................................................494
Riccia esulcata Steph. ...................................................................................................................500
Riccia fertilissima Steph. ..............................................................................................................500
Riccia luitans L. ..........................................................................................................................498
Riccia frostii Austin .....................................................................................................................498
Riccia frostii var. crystallinoides Schifn.........................................................................................498
Riccia fruchartii Steph. ................................................................................................................494
Riccia furfuracea Perold ...............................................................................................................495
Riccia gangetica Ahmad ex L.Söderstr., A.Hagborg et von Konrat ...............................................494
Riccia garsidei Sim.......................................................................................................................499
Riccia geissleriana Jovet-Ast .........................................................................................................498
Riccia gemmifera O.H.Volk.........................................................................................................500
Riccia glauca L. ...........................................................................................................................496
Riccia glauca var. ciliaris Warnst..................................................................................................496
Riccia gothica Damsh. et Hallingb. ..............................................................................................496
Riccia gougetiana Durieu et Mont. ..............................................................................................496
Riccia gougetiana var. armatissima Levier ex Müll.Frib.................................................................496
Riccia grandis Nees ......................................................................................................................494
Riccia grollei Udar .......................................................................................................................500
Riccia handelii Schifn. ................................................................................................................500
Riccia hantamensis Perold ............................................................................................................495
Riccia hasskarliana Steph. ............................................................................................................497
Riccia hawaiiensis Hürl. ..............................................................................................................500
Riccia hegewaldiana Jovet-Ast ......................................................................................................498
Riccia helenae Jovet-Ast ...............................................................................................................494
Riccia hirsuta O.H.Volk et Perold ...............................................................................................495
Riccia hirta (Austin) Underw. .....................................................................................................494
Riccia horrida Jovet-Ast ...............................................................................................................494
Riccia hortorum Bory...................................................................................................................494
Riccia howellii M.Howe ..............................................................................................................494
Riccia huebeneriana Lindenb. ......................................................................................................498
Riccia huebeneriana subsp. sullivantii (Austin) R.M.Schust. ........................................................498
Riccia ianthina Jovet-Ast .............................................................................................................494
Riccia indica Udar et A.Gupta.....................................................................................................501
Riccia indira-gandhiensis Dabhade et A.Hasan ............................................................................501
Riccia inlexa Taylor ....................................................................................................................494
Riccia intermedia Roum. .............................................................................................................501
Riccia iodocheila M.Howe ...........................................................................................................494
Riccia jodhpurensis Bapna ............................................................................................................501
Riccia jovet-astiae E.Vianna .........................................................................................................498
Riccia junghuhniana Nees et Lindenb..........................................................................................497
Riccia junghuhniana var. simplex Schifn. ....................................................................................497
Riccia kirinensis C.Gao et K.C.Chang .........................................................................................501
Riccia lamellosa Raddi .................................................................................................................496
648
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Riccia lanceolata Steph. ...............................................................................................................494
Riccia laxisquamata (Steph.) Steph. .............................................................................................501
Riccia leptothallus R.M.Schust. ....................................................................................................499
Riccia liaoningensis C.Gao et K.C.Chang ....................................................................................501
Riccia ligula Steph. ......................................................................................................................496
Riccia limbata Bisch. ex C.Krauss................................................................................................496
Riccia limicola Jovet-Ast ..............................................................................................................498
Riccia lindmanii Steph. ...............................................................................................................494
Riccia linearis (Schifn.) Steph. ....................................................................................................501
Riccia luticola Na-halang...........................................................................................................501
Riccia macallisteri M.Howe .........................................................................................................494
Riccia macrocarpa Levier .............................................................................................................496
Riccia macrospora Steph. .............................................................................................................494
Riccia mamillata Trab. ex Steph. .................................................................................................494
Riccia mammifera O.H.Volk et Perold ........................................................................................496
Riccia mamrensis Perold ..............................................................................................................501
Riccia mangalorica Ahmad ex Jovet-Ast .......................................................................................497
Riccia marginata Lindb. ..............................................................................................................501
Riccia mauryana Steph. ...............................................................................................................494
Riccia melanospora Kashyap.........................................................................................................501
Riccia melitensis C.Massal............................................................................................................496
Riccia membranacea Gottsche et Lindenb. ...................................................................................493
Riccia michelii Raddi ...................................................................................................................496
Riccia microciliata O.H.Volk et Perold........................................................................................497
Riccia miyakeana Schifn. ............................................................................................................501
Riccia moenkemeyeri Steph. .........................................................................................................499
Riccia montana Perold .................................................................................................................497
Riccia multiida (Steph.) Steph. ...................................................................................................498
Riccia muscicola Steph. ................................................................................................................501
Riccia namaquensis Perold ...........................................................................................................495
Riccia natalensis Sim....................................................................................................................497
Riccia nigerica E.W.Jones ............................................................................................................501
Riccia nigrella DC. ......................................................................................................................497
Riccia nigrescens Mont. ................................................................................................................501
Riccia nipponica S.Hatt. ..............................................................................................................501
Riccia novo-hannoverana Schifn. ................................................................................................501
Riccia numeensis Steph. ...............................................................................................................501
Riccia obtusa Meijer ....................................................................................................................501
Riccia oerstediana Lindenb. et Hampe .........................................................................................501
Riccia okahandjana S.W.Arnell ...................................................................................................497
Riccia olgensis Na-halang ...........................................................................................................494
Riccia ozarkiana McGregor .........................................................................................................497
Riccia pandei Udar ......................................................................................................................501
Riccia papillispora Steph. .............................................................................................................501
Riccia papillosa Moris ..................................................................................................................497
Riccia papulosa (Steph.) Steph. ....................................................................................................501
Riccia papulosa var. variabilis Na-halang ...................................................................................502
Riccia paraguayensis Spruce .........................................................................................................498
Riccia paranaensis Hässel .............................................................................................................499
Riccia parvoareolata O.H.Volk et Perold .....................................................................................495
Riccia pathankotensis Kashyap .....................................................................................................502
Riccia perennis Steph. ..................................................................................................................499
World checklist of hornworts and liverworts
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649
Riccia perssonii Sultan Khan ........................................................................................................499
Riccia perthiana Steph. ex K.I.Goebel .........................................................................................502
Riccia planobiconvexa Steph. .......................................................................................................494
Riccia polycarpa (Trab.) Jelenc .....................................................................................................498
Riccia porosa Taylor ....................................................................................................................498
Riccia pottsiana Sim ....................................................................................................................497
Riccia prominens Meijer...............................................................................................................502
Riccia pseudoluitans C.Gao et K.C.Chang ..................................................................................502
Riccia pubescens S.Hatt. ...............................................................................................................502
Riccia pullulans Jovet-Ast ............................................................................................................498
Riccia pulveracea Perold ..............................................................................................................495
Riccia purpurascens Lehm. ...........................................................................................................499
Riccia radiata Perold ...................................................................................................................502
Riccia radicosa Pearson ................................................................................................................495
Riccia rechingeri Steph.................................................................................................................502
Riccia reticulatula Udar ...............................................................................................................502
Riccia rhenana Lorb. ex Müll.Frib. ..............................................................................................499
Riccia rhenana var. violacea M.F.Boiko .......................................................................................499
Riccia ridleyi A.Gepp...................................................................................................................494
Riccia rorida Na-halang ............................................................................................................502
Riccia rosea O.H.Volk et Perold ..................................................................................................497
Riccia rubricollis Garside et A.V.Duthie ex Perold .......................................................................499
Riccia runssorensis Steph. .............................................................................................................497
Riccia saharensis Steph. ex Jovet-Ast ............................................................................................502
Riccia sanguineisporis Jovet-Ast ....................................................................................................494
Riccia satoi S.Hatt. ......................................................................................................................502
Riccia schelpei O.H.Volk et Perold ..............................................................................................493
Riccia schroederi Steph. ................................................................................................................502
Riccia schweinfurthii Steph. .........................................................................................................502
Riccia sibayenii Perold .................................................................................................................502
Riccia simii Perold.......................................................................................................................495
Riccia singularis Jovet-Ast ............................................................................................................499
Riccia somaliensis Perold ..............................................................................................................502
Riccia sommieri Levier .................................................................................................................497
Riccia sorocarpa Bisch. .................................................................................................................497
Riccia sorocarpa subsp. sorocarpa var. heegii Schifn......................................................................497
Riccia spongiosula Na-halang .....................................................................................................502
Riccia squamata Nees ..................................................................................................................495
Riccia stricta (Lindenb.) Perold ...................................................................................................499
Riccia subbifurca Warnst. ex Croz. ..............................................................................................497
Riccia subdepilata Jovet-Ast .........................................................................................................495
Riccia subplana Steph. .................................................................................................................495
Riccia subtilis (Steph.) Steph. ......................................................................................................502
Riccia sumatrana Meijer ..............................................................................................................502
Riccia symoensii Vanden Berghen ................................................................................................502
Riccia taeniiformis Jovet-Ast ........................................................................................................495
Riccia tasmanica Steph. ex Rodway .............................................................................................502
Riccia tenella D.L.Jacobs .............................................................................................................497
Riccia tomentosa O.H.Volk et Perold ..........................................................................................502
Riccia trabutiana Steph. ..............................................................................................................497
Riccia trachyglossa Perold .............................................................................................................495
Riccia treubiana Steph. ................................................................................................................502
650
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Riccia treubiana var. subrubescens Schifn.....................................................................................502
Riccia triangularis Steph. .............................................................................................................502
Riccia tuberculata Poir. ................................................................................................................502
Riccia udarii Kanwal ...................................................................................................................502
Riccia velenovskyi Kavina .............................................................................................................503
Riccia velimalaiana A.E.D.Daniels et P.Daniel............................................................................503
Riccia viannae Jovet-Ast ..............................................................................................................495
Riccia victoriensis Steph. ..............................................................................................................503
Riccia villosa Steph. .....................................................................................................................495
Riccia violacea M.Howe ..............................................................................................................497
Riccia violacea var. laevis Jovet-Ast ..............................................................................................497
Riccia vitalii Jovet-Ast .................................................................................................................495
Riccia vitrea Perold .....................................................................................................................495
Riccia volkii S.W.Arnell ..............................................................................................................499
Riccia vulcanicola Eb.Fisch. .........................................................................................................499
Riccia warnstorii Limpr. ex Warnst. ...........................................................................................497
Riccia weinionis Steph. ................................................................................................................495
Riccia weymouthiana Steph. ex Rodway.......................................................................................503
Riccia wichurae Steph..................................................................................................................503
Ricciocarpos natans (L.) Corda .....................................................................................................503
Riella ainis M.Howe et Underw. ...............................................................................................506
Riella alatospora Wigglesw...........................................................................................................505
Riella americana M.Howe et Underw. ........................................................................................505
Riella battandieri Trab. ...............................................................................................................505
Riella bialata Trab. .....................................................................................................................505
Riella capensis Cavers...................................................................................................................505
Riella choconensis Hässel ..............................................................................................................505
Riella cossoniana Trab. ................................................................................................................506
Riella cyrenaica Maire..................................................................................................................505
Riella echinata (Müll.Frib.) Segarra, Puche et Sabovlj..................................................................506
Riella echinospora Wigglesw.........................................................................................................505
Riella gallica Balansa ex Trab. .....................................................................................................505
Riella gamundiae Hässel ..............................................................................................................506
Riella halophila Banwell ..............................................................................................................505
Riella helicophylla (Bory et Mont.) Mont.....................................................................................505
Riella helicophylla var. macrocarpa P.Allorge ................................................................................505
Riella heliospora Segarra, Puche et Sabovlj. ..................................................................................506
Riella indica Steph. ex Kashyap ...................................................................................................505
Riella mediterranea Segarra, Puche, Sabovlj., M.Infante et Heras ................................................506
Riella notarisii (Mont.) Mont. .....................................................................................................505
Riella numidica Trab. ..................................................................................................................505
Riella pampae Hässel ...................................................................................................................505
Riella parisii Gottsche .................................................................................................................505
Riella purpureospora Wigglesw.....................................................................................................505
Riella reuteri Mont. .....................................................................................................................505
Riella sersuensis Trab. ..................................................................................................................505
Riella spiculata J.Taylor ...............................................................................................................505
Riella trigonospora Segarra et Puche .............................................................................................506
Riella undulata Hässel .................................................................................................................506
Rivulariella gemmipara (A.Evans) D.H.Wagner ..........................................................................122
Ruizanthus venezuelanus R.M.Schust. .........................................................................................103
Saccobasis polita (Nees) H.Buch ....................................................................................................85
World checklist of hornworts and liverworts
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651
Saccobasis polymorpha (R.M.Schust.) Schljakov .............................................................................85
Saccogyna ligulata Steph. .............................................................................................................122
Saccogyna subacuta Steph. ...........................................................................................................122
Saccogyna tridens Steph. ..............................................................................................................122
Saccogyna viticulosa (L.) Dumort. ..................................................................................................98
Saccogynidium australe (Mitt.) Grolle ............................................................................................98
Saccogynidium caldense (Ångstr.) Grolle.........................................................................................98
Saccogynidium chiloscyphoides R.M.Schust. ....................................................................................98
Saccogynidium decurvum (Mitt.) Grolle .........................................................................................98
Saccogynidium goebelii (Herzog) Grolle .........................................................................................98
Saccogynidium irregularospinum C.Gao, T.Cao et M.J.Lai.............................................................98
Saccogynidium muricellum (De Not.) Grolle..................................................................................98
Saccogynidium rigidulum (Nees) Grolle .........................................................................................98
Saccogynidium vasculosum (Hook.f. et Taylor) Grolle ....................................................................98
Sandeothallus japonicus (Inoue) Crand.-Stotl. et Stotler ...............................................................474
Sandeothallus radiculosus (Schifn.) R.M.Schust. .........................................................................474
Sauteria alpina (Nees) Nees ........................................................................................................485
Sauteria chilensis (Lindenb.) Grolle .............................................................................................485
Sauteria crassipes Austin ..............................................................................................................485
Sauteria inlata C.Gao et K.C.Chang ..........................................................................................485
Sauteria japonica (Shimizu et S.Hatt.) S.Hatt. ............................................................................485
Sauteria nyikaensis Perold ............................................................................................................485
Sauteria spongiosa (Kashyap) S.Hatt. ...........................................................................................485
Scapania aequiloba (Schwägr.) Dumort. ........................................................................................87
Scapania americana Müll.Frib. ......................................................................................................87
Scapania ampliata Steph. ..............................................................................................................85
Scapania ampliata subsp. queenslandica M.L.Hicks .......................................................................85
Scapania apiculata Spruce .............................................................................................................87
Scapania aspera M.Bernet et Bernet ..............................................................................................87
Scapania bhutanensis Amakawa .....................................................................................................88
Scapania bolanderi Austin .............................................................................................................85
Scapania brevicaulis Taylor............................................................................................................90
Scapania calcicola (Arnell et J.Perss.) Ingham ................................................................................90
Scapania carinthiaca J.B.Jack ex Lindb. .........................................................................................87
Scapania carinthiaca var. massalongi Müll.Frib. .............................................................................87
Scapania ciliata Sande Lac.............................................................................................................88
Scapania ciliata subsp. hawaiica (Müll.Frib.) Potemkin ................................................................88
Scapania ciliatospinosa Horik.........................................................................................................86
Scapania compacta (Roth) Dumort................................................................................................88
Scapania contorta Mitt. .................................................................................................................85
Scapania crassiretis Bryhn ..............................................................................................................90
Scapania curta (Mart.) Dumort. ....................................................................................................88
Scapania curta var. grandiretis R.M.Schust. ...................................................................................88
Scapania curta var. isoloba R.M.Schust. .........................................................................................88
Scapania cuspiduligera (Nees) Müll.Frib. .......................................................................................89
Scapania cuspiduligera var. diplophyllopsis R.M.Schust. ..................................................................89
Scapania davidii Potemkin ............................................................................................................86
Scapania degenii Schifn. ex Müll.Frib...........................................................................................90
Scapania diplophylloides Amakawa et S.Hatt. .................................................................................88
Scapania esterhuyseniae S.W.Arnell ................................................................................................88
Scapania ferruginaeoides T.Cao, C.Gao et J.Sun ............................................................................86
Scapania ferruginea (Lehm. et Lindenb.) Lehm. et Lindenb. .........................................................87
652
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Scapania fulfordiae W.S.Hong ......................................................................................................88
Scapania gamundiae R.M.Schust. ..................................................................................................88
Scapania gaochii X.Fu ex T.Cao ....................................................................................................86
Scapania geppii Steph. ...................................................................................................................86
Scapania gigantea Horik. ...............................................................................................................91
Scapania glaucocephala (Taylor) Austin .........................................................................................92
Scapania glaucocephala var. saxicola (R.M.Schust.) Potemkin ........................................................92
Scapania glaucoviridis Horik. ........................................................................................................90
Scapania gracilis Lindb. .................................................................................................................85
Scapania grandiloba Steph. ............................................................................................................91
Scapania griithii Schifn. .............................................................................................................92
Scapania grossidens Steph. ex Müll.Frib. ........................................................................................91
Scapania gymnostomophila Kaal. ....................................................................................................90
Scapania harae Amakawa ..............................................................................................................86
Scapania hedbergii S.W.Arnell .......................................................................................................91
Scapania helvetica Gottsche ...........................................................................................................89
Scapania hians Steph. ex Müll.Frib. ..............................................................................................87
Scapania hians subsp. salishensis J.D.Godfrey et G.Godfrey ...........................................................87
Scapania himalayica Müll.Frib. ex Herzog ....................................................................................90
Scapania hirosakiensis Steph. ex Müll.Frib. ....................................................................................88
Scapania hollandiae W.S.Hong .....................................................................................................88
Scapania hyperborea Jørg. ..............................................................................................................90
Scapania integerrima Steph............................................................................................................91
Scapania irrigua (Nees) Nees.........................................................................................................89
Scapania irrigua subsp. rufescens (Loeske) R.M.Schust. .................................................................89
Scapania javanica Gottsche ...........................................................................................................92
Scapania javanica var. scabra Schifn. ............................................................................................92
Scapania karl-muelleri Grolle ........................................................................................................90
Scapania kaurinii Ryan .................................................................................................................88
Scapania komagadakensis Amakawa ...............................................................................................91
Scapania koponenii Potemkin ........................................................................................................88
Scapania lepida Mitt. ....................................................................................................................88
Scapania ligulata Steph. ................................................................................................................92
Scapania ligulata subsp. stephanii (Müll.Frib.) Potemkin, Piippo et T.J.Kop.................................92
Scapania ligulifolia R.M.Schust. ....................................................................................................90
Scapania lingulata H.Buch ............................................................................................................89
Scapania lingulata var. microphylla (Warnst.) R.M.Schust. ............................................................89
Scapania macroparaphyllia T.Cao, C.Gao et J.Sun ........................................................................85
Scapania magadanica S.S.Choi, Bakalin et B.Y.Sun.......................................................................89
Scapania matveyevae Potemkin ......................................................................................................91
Scapania maxima Horik. ...............................................................................................................85
Scapania microdonta (Mitt.) Müll.Frib. .........................................................................................87
Scapania mucronata H.Buch .........................................................................................................89
Scapania nemorea (L.) Grolle ........................................................................................................91
Scapania nimbosa Taylor ...............................................................................................................86
Scapania nipponica (Amakawa et S.Hatt.) Amakawa .....................................................................85
Scapania obcordata (Berggr.) S.W.Arnell .......................................................................................89
Scapania obscura (Arnell et C.E.O.Jensen) Schifn. .......................................................................91
Scapania orientalis Steph. ex Müll.Frib..........................................................................................87
Scapania ornithopodioides (With.) Waddell ...................................................................................86
Scapania paludicola Loeske et Müll.Frib. .......................................................................................90
Scapania paludicola var. viridigemma R.M.Schust. ........................................................................90
World checklist of hornworts and liverworts
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653
Scapania paludosa (Müll.Frib.) Müll.Frib. .....................................................................................91
Scapania paraphyllia T.Cao, C.Gao, J.Sun et B.R.Zuo..................................................................85
Scapania parvidens Steph...............................................................................................................91
Scapania parvifolia Warnst. ...........................................................................................................89
Scapania parvitexta Steph. .............................................................................................................91
Scapania portoricensis Hampe et Gottsche .....................................................................................86
Scapania portoricensis var. boliviensis (Steph.) Herzog ....................................................................86
Scapania portoricensis var. organensis (Herzog) Herzog ...................................................................86
Scapania portoricensis var. roraimensis Warnst. ...............................................................................86
Scapania praetervisa Meyl. .............................................................................................................89
Scapania pseudocalcicola R.M.Schust. ............................................................................................90
Scapania pseudocontorta Potemkin.................................................................................................87
Scapania rigida Nees .....................................................................................................................91
Scapania rotundifolia W.E.Nicholson............................................................................................86
Scapania ruidula Warnst. .............................................................................................................91
Scapania sandei Schifn. ex Müll.Frib. ...........................................................................................88
Scapania scandica (Arnell et H.Buch) Macvicar .............................................................................89
Scapania scandica var. argutedentata H.Buch .................................................................................89
Scapania scandica var. dimorpha R.M.Schust. ................................................................................89
Scapania scandica var. grandiretis (Schljakov) Schljakov.................................................................89
Scapania scapanioides (C.Massal.) Grolle .......................................................................................92
Scapania schljakovii Potemkin .......................................................................................................90
Scapania secunda Steph. ................................................................................................................86
Scapania serrulata R.M.Schust. .....................................................................................................91
Scapania simmonsii Bryhn et Kaal. ................................................................................................92
Scapania sinikkae Potemkin ..........................................................................................................87
Scapania sphaerifera H.Buch et Tuom...........................................................................................92
Scapania spiniloba Potemkin .........................................................................................................87
Scapania spitsbergensis (Lindb.) Müll.Frib......................................................................................88
Scapania subalpina (Nees ex Lindenb.) Dumort. ...........................................................................91
Scapania subalpina var. haynesiae Frye et L.Clark ..........................................................................91
Scapania subalpina var. muddiae C.D.Bird et W.S.Hong ..............................................................91
Scapania subnimbosa Steph. ..........................................................................................................85
Scapania tundrae (Arnell) H.Buch.................................................................................................90
Scapania udarii S.C.Srivast. et A.Srivast. .......................................................................................92
Scapania uliginosa (Lindenb.) Dumort. .........................................................................................89
Scapania umbrosa (Schrad.) Dumort. ............................................................................................87
Scapania undulata (L.) Dumort. ...................................................................................................91
Scapania valdonii Váňa, Bedn.-Ochyra et Cykowska.....................................................................89
Scapania verrucosa Heeg ................................................................................................................92
Scapania zemliae S.W.Arnell .........................................................................................................89
Scapania zhukovae Potemkin.........................................................................................................86
Schifneria hyalina Steph. ..............................................................................................................64
Schifneriolejeunea altimontana Vanden Berghen .........................................................................411
Schifneriolejeunea amazonica Gradst. ..........................................................................................411
Schifneriolejeunea cumingiana (Mont.) Gradst. ...........................................................................412
Schifneriolejeunea ferruginea (Steph.) Gradst. .............................................................................412
Schifneriolejeunea fragilis Gradst. et E.W.Jones...........................................................................411
Schifneriolejeunea madagascariensis (Steph.) Gradst. ...................................................................411
Schifneriolejeunea occulta (Steph.) Gradst. ..................................................................................412
Schifneriolejeunea omphalanthoides Verd. ....................................................................................412
Schifneriolejeunea parviloba (Steph.) Gradst. ..............................................................................411
654
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Schifneriolejeunea tumida (Nees) Gradst. ....................................................................................412
Schifneriolejeunea tumida var. haskarliana (Gottsche) Gradst. et Terken.....................................412
Schifneriolejeunea nymannii (Steph.) Gradst. et Terken ..............................................................412
Schifneriolejeunea pappeana (Nees) Gradst..................................................................................411
Schifneriolejeunea pappeana var. bidentata Gradst. et Vanden Berghen .......................................411
Schifneriolejeunea pappeana var. integra Gradst. et Vanden Berghen ...........................................411
Schifneriolejeunea polycarpa (Nees) Gradst. .................................................................................412
Schifneriolejeunea pulopenangensis (Gottsche) Gradst. .................................................................412
Schisma orizabense (Gottsche) Steph. ..........................................................................................511
Schisma uleanum Steph. ..............................................................................................................511
Schistochila acuminata Steph. ......................................................................................................259
Schistochila aequiloba Steph.........................................................................................................259
Schistochila alata (Lehm.) Schifn. ...............................................................................................259
Schistochila aligera (Nees et Blume) J.B.Jack et Steph. .................................................................259
Schistochila aligera var. laxa (Nees) Schifn. .................................................................................259
Schistochila altissima E.A.Hodgs. .................................................................................................259
Schistochila altissima subsp. polystratosa R.M.Schust. et J.J.Engel .................................................259
Schistochila antara Grolle ............................................................................................................259
Schistochila appendiculata (Hook.) Dumort. ex Trevis. ................................................................259
Schistochila baileyana Steph. ........................................................................................................259
Schistochila balfouriana (Hook.f. et Taylor) Steph. ......................................................................259
Schistochila beccariana (De Not.) Trevis. .....................................................................................260
Schistochila berggrenii (J.J.Engel et R.M.Schust.) Xiao L.He et Glenny .......................................260
Schistochila berteroana (Hook.) Steph. .........................................................................................260
Schistochila blumei (Nees) Trevis. ................................................................................................260
Schistochila brassii Grolle.............................................................................................................260
Schistochila caledonica Steph. .......................................................................................................260
Schistochila carnosa (Mitt.) Steph. ...............................................................................................260
Schistochila caudata R.M.Schust. et J.J.Engel ..............................................................................260
Schistochila childii (R.M.Schust. et J.J.Engel) Xiao L.He et Glenny ............................................260
Schistochila chlorophylla (Colenso) E.A.Hodgs. ............................................................................260
Schistochila ciliata (Mitt.) Steph. .................................................................................................260
Schistochila compacta (Colenso) E.A.Hodgs. ................................................................................260
Schistochila conchophylla Herzog ex E.A.Hodgs. et Allison...........................................................260
Schistochila conchophylla var. multidentata (J.J.Engel) Xiao L.He et Glenny.................................260
Schistochila congoana Steph. ........................................................................................................260
Schistochila cookei (H.A.Mill.) R.M.Schust..................................................................................260
Schistochila crinita Grolle ............................................................................................................260
Schistochila cristata Steph. ...........................................................................................................261
Schistochila cunninghamii Steph. .................................................................................................261
Schistochila doriae (De Not.) Trevis. ...........................................................................................261
Schistochila engleriana Steph........................................................................................................261
Schistochila exalata Herzog ..........................................................................................................261
Schistochila ijiensis H.Buch et Herzog.........................................................................................261
Schistochila glaucescens (Hook.) A.Evans ......................................................................................261
Schistochila hattorii Grolle ...........................................................................................................261
Schistochila integerrima Steph. .....................................................................................................261
Schistochila isotachyphylla (J.J.Engel et R.M.Schust.) Xiao L.He et Glenny..................................261
Schistochila kirkiana Steph. .........................................................................................................261
Schistochila kunkelii S.W.Arnell ..................................................................................................261
Schistochila lacerata Steph. ..........................................................................................................261
Schistochila lamellata (Hook.) Dumort. ex A.Evans .....................................................................261
World checklist of hornworts and liverworts
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655
Schistochila laminigera (Hook.f. et Taylor) A.Evans ....................................................................261
Schistochila latiloba (R.M.Schust. et J.J.Engel) Xiao L.He et Glenny ...........................................261
Schistochila lehmanniana (Lindenb.) Steph. .................................................................................261
Schistochila leucophylla (Lehm. ex Gottsche, Lindenb. et Nees) Steph. ........................................261
Schistochila macrodonta W.E.Nicholson ......................................................................................261
Schistochila minor C.Gao et Y.H.Wu ..........................................................................................261
Schistochila monticola R.M.Schust. ..............................................................................................262
Schistochila muricata E.A.Hodgs. et Allison.................................................................................262
Schistochila nadeaudiana Steph....................................................................................................262
Schistochila neesii (Mont.) Lindb. ................................................................................................262
Schistochila nitidissima R.M.Schust. ............................................................................................262
Schistochila nivicola (R.M.Schust. et J.J.Engel) Xiao L.He et Glenny ..........................................262
Schistochila nobilis (Hook.) Trevis. ..............................................................................................262
Schistochila nuda Horik. ..............................................................................................................262
Schistochila pachyphylla (Lehm.) Steph. .......................................................................................262
Schistochila papillifera R.M.Schust. .............................................................................................262
Schistochila parvistipula Rodway..................................................................................................262
Schistochila pellucida R.M.Schust. et J.J.Engel .............................................................................262
Schistochila piligera Steph. ...........................................................................................................262
Schistochila pinnatifolia (Hook.) Trevis. ......................................................................................262
Schistochila pluriciliata R.M.Schust. et J.J.Engel..........................................................................262
Schistochila pseudociliata R.M.Schust. .........................................................................................262
Schistochila quadriida A.Evans....................................................................................................262
Schistochila ramentacea Steph. .....................................................................................................262
Schistochila relexa (Mont.) Steph. ...............................................................................................262
Schistochila relexistipula J.J.Engel et R.M.Schust. .......................................................................263
Schistochila reinwardtii (Nees) Schifn. ........................................................................................263
Schistochila repleta (Hook.f. et Taylor) Steph. .............................................................................263
Schistochila rubriseta Steph. .........................................................................................................263
Schistochila schultzei Steph. .........................................................................................................263
Schistochila sciophila R.M.Schust. ................................................................................................263
Schistochila sciurea (Nees) Schifn................................................................................................263
Schistochila simulans (C.Massal.) Xiao L.He et Yu Sun ................................................................263
Schistochila spegazziniana (C.Massal.) Steph................................................................................263
Schistochila sphagnoides (Schwägr.) Lindb. ex Steph.....................................................................263
Schistochila splachnophylla (Hook.f. et Taylor) Steph. ..................................................................263
Schistochila stratosa (Mont.) A.Evans ...........................................................................................263
Schistochila subhyalina R.M.Schust. ............................................................................................263
Schistochila subhyalina var. grandidentata (J.J.Engel et R.M.Schust.) Xiao L.He et Glenny..........263
Schistochila subimmersa J.J.Engel et R.M.Schust..........................................................................263
Schistochila succulenta (J.J.Engel et R.M.Schust.) Xiao L.He et Glenny .......................................263
Schistochila tasmanica Steph. .......................................................................................................264
Schistochila trispiralis R.M.Schust................................................................................................264
Schistochila tuloides (Hook.f. et Taylor) Steph. ............................................................................264
Schistochila undulatifolia Piippo ..................................................................................................264
Schistochila virescens R.M.Schust. ................................................................................................264
Schistochila vitreocincta (Herzog) Xiao L.He et Glenny ...............................................................264
Schistochila volans Grolle .............................................................................................................264
Schistochila yakushimensis N.Ohnishi et Deguchi ........................................................................264
Schistochila zantenii Grolle ..........................................................................................................264
Schistochilopsis cornuta (Steph.) Konstant. .....................................................................................93
Schistochilopsis grandiretis (Lindb. ex Kaal.) Konstant. ...................................................................93
656
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Schistochilopsis hyperarctica Konstant. et L.Söderstr. ......................................................................93
Schistochilopsis incisa (Schrad.) Konstant. ......................................................................................93
Schistochilopsis nakanishii (Inoue) Konstant...................................................................................93
Schistochilopsis opacifolia (Culm. ex Meyl.) Konstant. ....................................................................93
Schistochilopsis setosa (Mitt.) Konstant. ..........................................................................................93
Schizophyllopsis aristata (Herzog ex N.Kitag.) Váňa et L.Söderstr. .................................................54
Schizophyllopsis bidens (Reinw., Blume et Nees) Váňa et L.Söderstr...............................................54
Schizophyllopsis lanciloba (Steph.) Váňa et L.Söderstr. ...................................................................54
Schizophyllopsis papillosa (J.J.Engel et Braggins) Váňa et L.Söderstr. ..............................................54
Schizophyllopsis sphenoloboides (R.M.Schust.) Váňa et L.Söderstr. ..................................................54
Schljakovia kunzeana (Huebener) Konstant. et Vilnet ...................................................................55
Schljakovianthus quadrilobus (Lindb.) Konstant. et Vilnet .............................................................55
Schusterolejeunea inundata (Spruce) Grolle ..................................................................................350
Seppeltia succuba Grolle...............................................................................................................471
Sewardiella tuberifera Kashyap ....................................................................................................468
Siphonolejeunea elegantissima (Steph.) Grolle ...............................................................................350
Siphonolejeunea neesii (Mont.) Bischl. .........................................................................................350
Siphonolejeunea schifneri (Schifn.) Herzog .................................................................................350
Solenostoma amoenum (Lindenb. et Gottsche) R.M.Schust. ex Váňa, Hentschel et Heinrichs......129
Solenostoma amplexifolium (Hampe) Váňa et Schäf.-Verw. ..........................................................129
Solenostoma appalachianum R.M.Schust. ex Bakalin....................................................................129
Solenostoma appressifolium (Mitt.) Váňa et D.G.Long .................................................................129
Solenostoma appressifolium var. minor (Amakawa) Váňa et D.G.Long ..........................................129
Solenostoma appressifolium var. nigricans (Amakawa) Váňa et D.G.Long .....................................129
Solenostoma ariadne (Taylor) R.M.Schust. ex Váňa et D.G.Long ................................................129
Solenostoma atrobrunneum (Amakawa) Váňa et D.G.Long ..........................................................129
Solenostoma atrorevolutum (Grolle ex Amakawa) Váňa et D.G.Long ...........................................129
Solenostoma atrovirens Steph........................................................................................................129
Solenostoma balfourii (Váňa) Váňa, Hentschel et Heinrichs .........................................................125
Solenostoma baueri (Schifn.) Steph. ............................................................................................130
Solenostoma bengalense (Amakawa) Váňa et D.G.Long ................................................................130
Solenostoma bilobum (S.Hatt. ex Amakawa) Potemkin et Nyushko .............................................123
Solenostoma borneense (Amakawa) Váňa, Hentschel et Heinrichs.................................................125
Solenostoma brevilorum Kashyap et R.S.Chopra..........................................................................130
Solenostoma caeleste (Inoue et Váňa) Váňa, Hentschel et Heinrichs .............................................130
Solenostoma callithrix (Lindenb. et Gottsche) Steph. ...................................................................125
Solenostoma caoi (C.Gao et X.L.Bai) Váňa et D.G.Long..............................................................130
Solenostoma caucasicum (Váňa) Konstant.....................................................................................130
Solenostoma champawatense (S.N.Srivast. et Amakawa) Váňa et D.G.Long .................................125
Solenostoma chenianum (C.Gao, Y.H.Wu et Grolle) Váňa et D.G.Long......................................130
Solenostoma clavellatum Mitt. ex Steph. .......................................................................................130
Solenostoma comatum (Nees) C.Gao ............................................................................................125
Solenostoma comatum var. novae-guineae (Váňa) Váňa, Hentschel et Heinrichs ...........................125
Solenostoma confertissimum (Nees) Schljakov ...............................................................................130
Solenostoma conilorum (Schifn.) Steph.......................................................................................130
Solenostoma crassulum (Nees et Mont.) Steph. .............................................................................130
Solenostoma crenuliforme (Austin) Steph. .....................................................................................125
Solenostoma cryptogynum R.M.Schust. ex J.J.Engel ......................................................................130
Solenostoma cyclops (S.Hatt.) R.M.Schust. ...................................................................................130
Solenostoma decolor (Schifn.) R.M.Schust. ex Váňa, Hentschel et Heinrichs ...............................125
Solenostoma diversiclavellatum (Amakawa et Grolle) R.M.Schust. ex Váňa et D.G.Long .............130
Solenostoma dulongense Váňa et D.G.Long ..................................................................................130
World checklist of hornworts and liverworts
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657
Solenostoma dusenii (Steph.) Váňa, Hentschel et Heinrichs .........................................................125
Solenostoma emarginatum (Amakawa) Váňa, Hentschel et Heinrichs ...........................................124
Solenostoma erectum (Amakawa) C.Gao.......................................................................................125
Solenostoma exsertum (A.Evans) Steph. ........................................................................................131
Solenostoma faurieanum (Beauverd) R.M.Schust. ........................................................................131
Solenostoma lagellalioides C.Gao .................................................................................................125
Solenostoma lagellare (Amakawa) Váňa et D.G.Long ..................................................................131
Solenostoma lagellatum (S.Hatt.) Váňa et D.G.Long ...................................................................124
Solenostoma lavialbicans (Amakawa et Grolle) Váňa et D.G.Long ..............................................126
Solenostoma lavorevolutum (Váňa) Váňa et D.G.Long ................................................................131
Solenostoma fossombronioides (Austin) R.M.Schust. .....................................................................126
Solenostoma fusiforme (Steph.) R.M.Schust..................................................................................124
Solenostoma glaucum (Amakawa) Váňa et D.G.Long ...................................................................126
Solenostoma gongshanense (C.Gao et J.Sun) Váňa et D.G.Long....................................................126
Solenostoma gracillimum (Sm.) R.M.Schust. ................................................................................124
Solenostoma grollei (D.G.Long et Váňa) K.Feldberg, Hentschel, Bombosch, D.G.Long, Váňa et
Heinrichs ................................................................................................................................... 131
Solenostoma grosseverrucosum (Amakawa et S.Hatt.) Váňa, Crand.-Stotl. et Stotler ......................131
Solenostoma handelii (Schifn.) Müll.Frib. ...................................................................................124
Solenostoma haskarlianum (Nees) R.M.Schust. ex Váňa et D.G.Long ..........................................126
Solenostoma hattorianum (Amakawa) Potemkin et Nyushko........................................................126
Solenostoma heterolimbatum (Amakawa) Váňa et D.G.Long ........................................................131
Solenostoma hewsoniae (Amakawa et Grolle) R.M.Schust. ex Váňa, Hentschel et Heinrichs ........131
Solenostoma hirticalyx (Steph.) R.M.Schust. ex Váňa, Hentschel et Heinrichs .............................126
Solenostoma hiugaense Amakawa ..................................................................................................131
Solenostoma hokkaidense (Váňa) Váňa, Hentschel et Heinrichs ....................................................124
Solenostoma horikawanum (Amakawa) Váňa, Hentschel et Heinrichs ..........................................126
Solenostoma hyalinum (Lyell) Mitt...............................................................................................126
Solenostoma indrodayanum (Sushil K.Singh et D.K.Singh) Váňa et D.G.Long ............................131
Solenostoma infuscum (Mitt.) Hentschel ......................................................................................126
Solenostoma infuscum var. ovicalyx (Steph.) Potemkin et Sofronova .............................................126
Solenostoma inundatum (Hook.f. et Taylor) Mitt. ex Steph. ........................................................131
Solenostoma javanicum (Schifn.) Steph. ......................................................................................131
Solenostoma kanaii (Amakawa) Váňa et D.G.Long ......................................................................131
Solenostoma kashyapii (S.C.Srivast., S.Srivast. et D.Sharma) Váňa et D.G.Long ..........................131
Solenostoma kurilense (Bakalin) Váňa...........................................................................................126
Solenostoma lanigerum (Mitt.) Váňa et D.G.Long .......................................................................132
Solenostoma lignicola (Schifn.) Váňa, Hentschel et Heinrichs .....................................................124
Solenostoma limbatifolium (Amakawa) Váňa et D.G.Long ...........................................................124
Solenostoma lixingjiangii (C.Gao et X.L.Bai) Váňa et D.G.Long .................................................126
Solenostoma macrocarpum (Schifn. ex Steph.) Váňa et D.G.Long ...............................................132
Solenostoma mamatkulovii (Váňa et Zerov) Váňa, Hentschel et Heinrichs ...................................132
Solenostoma marginatum (S.Hatt.) R.M.Schust. ..........................................................................126
Solenostoma micranthum (Mitt.) Váňa, Hentschel et Heinrichs ...................................................127
Solenostoma microphyllum C.Gao ................................................................................................132
Solenostoma microrevolutum (C.Gao et X.L.Bai) Váňa et D.G.Long ............................................132
Solenostoma mildbraedii (Steph.) R.M.Schust. .............................................................................132
Solenostoma montanum (Steph.) Váňa .........................................................................................127
Solenostoma multicarpum (C.Gao et J.Sun) Váňa et D.G.Long ...................................................132
Solenostoma nilgiriense (A.Alam, Ad.Kumar et S.C.Srivast.) Váňa et D.G.Long...........................127
Solenostoma niveum (Grolle) R.M.Schust. ex Váňa, Hentschel et Heinrichs ................................132
Solenostoma novazelandiae R.M.Schust........................................................................................132
658
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Solenostoma obliquifolium (Schifn.) R.M.Schust. ex Váňa, Hentschel et Heinrichs .....................127
Solenostoma obovatum (Nees) C.Massal. ......................................................................................124
Solenostoma obscurum (A.Evans) R.M.Schust. .............................................................................124
Solenostoma ohbae (Amakawa) C.Gao .........................................................................................132
Solenostoma onraedtii (Váňa) Váňa, Hentschel et Heinrichs ........................................................127
Solenostoma orbicularifolium (Piippo ex C.Gao et Bai) Váňa .......................................................127
Solenostoma orbiculatum (Colenso) R.M.Schust. .........................................................................132
Solenostoma otianum (S.Hatt.) R.M.Schust. ................................................................................127
Solenostoma ovalifolium (Amakawa) Váňa ...................................................................................127
Solenostoma paroicum (Schifn.) R.M.Schust. ..............................................................................127
Solenostoma parvitextum (Amakawa) Váňa et D.G.Long .............................................................132
Solenostoma patoniae (Grolle, D.B.Schill et D.G.Long) K.Feldberg, Hentschel, Bombosch,
D.G.Long, Váňa et Heinrichs .....................................................................................................132
Solenostoma philippinense Váňa....................................................................................................124
Solenostoma plagiochilaceum (Grolle) Váňa et D.G.Long .............................................................127
Solenostoma pocsii (Váňa) Bakalin................................................................................................132
Solenostoma poeltii (Amakawa) Váňa et D.G.Long ......................................................................133
Solenostoma polyrhizoides (Grolle ex Amakawa) Váňa et D.G.Long .............................................127
Solenostoma pseudocyclops (Inoue) Váňa et D.G.Long ..................................................................133
Solenostoma pseudopyrilorum Bakalin et Vilnet............................................................................133
Solenostoma purpuratum (Mitt.) Steph. .......................................................................................133
Solenostoma pyrilorum Steph.......................................................................................................133
Solenostoma pyrilorum var. gracillimum (Amakawa) Váňa et D.G.Long ......................................133
Solenostoma pyrilorum var. major (S.Hatt.) Bakalin.....................................................................133
Solenostoma pyrilorum var. minutissimum (Amakawa) Bakalin ....................................................133
Solenostoma radicellosum Mitt. ....................................................................................................127
Solenostoma raujeanum (Grolle ex Amakawa) Váňa et D.G.Long ................................................133
Solenostoma renauldii (Steph.) Váňa, Hentschel et Heinrichs ......................................................127
Solenostoma ricleii Váňa et D.G.Long.........................................................................................133
Solenostoma rigidulum (S.Hatt.) R.M.Schust. ..............................................................................127
Solenostoma rosulans (Steph.) Váňa et D.G.Long .........................................................................127
Solenostoma rotundatum Amakawa ..............................................................................................127
Solenostoma rubripunctatum (S.Hatt.) R.M.Schust. .....................................................................127
Solenostoma rubrum (Gottsche) R.M.Schust................................................................................125
Solenostoma rupicola (Amakawa) Váňa et D.G.Long ...................................................................133
Solenostoma sanguinolentum (Grif.) Steph...................................................................................133
Solenostoma schaulianum (Steph.) Váňa et D.G.Long ..................................................................133
Solenostoma schusteranum (J.D.Godfrey et G.Godfrey) Váňa, Hentschel et Heinrichs .................124
Solenostoma shimizuanum (S.Hatt. ex Váňa) Váňa, Hentschel et Heinrichs .................................133
Solenostoma sikkimense (Schifn. ex Steph.) Váňa et D.G.Long ....................................................127
Solenostoma speciosum (Horik.) Hentschel, K.Feldberg, Bombosch, D.G.Long, Váňa et Heinrichs....133
Solenostoma speciosum subsp. villosum (R.M.Schust.) Hentschel, K.Feldberg, Bombosch, D.G.Long,
Váňa et Heinrichs .......................................................................................................................134
Solenostoma sphaerocarpum (Hook.) Steph. .................................................................................134
Solenostoma stephanii (Schifn.) Steph. ........................................................................................134
Solenostoma stoloniferum (Steph.) S.W.Arnell ..............................................................................134
Solenostoma strictum (Schifn.) Váňa, Hentschel et Heinrichs......................................................134
Solenostoma subacutum (Herzog) Váňa, Crand.-Stotl. et Stotler ..................................................134
Solenostoma suborbiculatum (Amakawa) Váňa et D.G.Long ........................................................125
Solenostoma subrubrum (Schifn. ex Steph.) Váňa et D.G.Long ...................................................134
Solenostoma subtilissimum (Schifn.) R.M.Schust. ........................................................................124
Solenostoma sunii Bakalin et Vilnet..............................................................................................134
World checklist of hornworts and liverworts
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659
Solenostoma tetragonum (Lindenb.) R.M.Schust. ex Váňa et D.G.Long .......................................127
Solenostoma tetragonum var. kodaikanalense A.Alam, D.Sharma et So.Yadav................................127
Solenostoma torticalyx (Steph.) C.Gao..........................................................................................134
Solenostoma totopapillosum (E.A.Hodgs.) R.M.Schust. ................................................................134
Solenostoma truncatum (Nees) R.M.Schust. ex Váňa et D.G.Long ..............................................134
Solenostoma truncatum var. setulosum (Herzog) Váňa et D.G.Long..............................................134
Solenostoma tuberculiferum (Herzog) Váňa, Hentschel et Heinrichs ............................................127
Solenostoma udarii (S.C.Srivast. et P.Singh) Váňa et D.G.Long ..................................................134
Solenostoma unispire (Amakawa) Váňa, Hentschel et Heinrichs ...................................................127
Solenostoma ventroversum (Grolle) Váňa et D.G.Long .................................................................134
Solenostoma virgatum (Mitt.) Váňa et D.G.Long .........................................................................127
Solenostoma vulcanicola (Schifn.) Nyushko ex Potemkin et Sofronova........................................127
Solenostoma zangmuii (C.Gao et X.L.Bai) Váňa et D.G.Long .....................................................127
Solenostoma zantenii (Amakawa) R.M.Schust. ex Váňa et D.G.Long ..........................................134
Solenostoma zengii (C.Gao et X.L.Bai) Váňa et D.G.Long ...........................................................134
Southbya gollanii Steph. ..............................................................................................................135
Southbya nigrella (De Not.) Henriq. ...........................................................................................135
Southbya organensis Herzog .........................................................................................................136
Southbya tophacea (Spruce) Spruce ..............................................................................................136
Sphaerocarpos cristatus M.Howe ..................................................................................................506
Sphaerocarpos donnellii Austin .....................................................................................................506
Sphaerocarpos drewiae Wigglesw. .................................................................................................506
Sphaerocarpos europaeus Lorb. .....................................................................................................507
Sphaerocarpos hians Haynes .........................................................................................................507
Sphaerocarpos michelii Bellardi ....................................................................................................506
Sphaerocarpos muccilloi E.Vianna ................................................................................................507
Sphaerocarpos stipitatus Bisch. ex Lindenb. ..................................................................................506
Sphaerocarpos texanus Austin .......................................................................................................506
Sphenolobopsis pearsonii (Spruce) R.M.Schust................................................................................55
Sphenolobus austroamericanus (Váňa) Váňa....................................................................................55
Sphenolobus minutus (Schreb. ex D.Crantz) Berggr. ......................................................................55
Sphenolobus saxicola (Schrad.) Steph..............................................................................................55
Spruceanthus macrostipulus (Steph.) Gradst. ................................................................................412
Spruceanthus mamillilobulus (Herzog) Verd.................................................................................412
Spruceanthus pluriplicatus (Steph.) Gradst. ..................................................................................412
Spruceanthus polymorphus (Sande Lac.) Verd. ..............................................................................412
Spruceanthus semirepandus (Nees) Verd. ......................................................................................412
Spruceanthus sulcatus (Nees) Gradst. ...........................................................................................413
Spruceanthus theobromae (Spruce) Gradst. ...................................................................................413
Spruceanthus thozetianus (Gottsche et F.Muell.) B.M.hiers et Gradst. .......................................413
Steereella lilliana (Steph.) Kuwah. ...............................................................................................461
Steereella linearis (Sw.) Kuwah. ...................................................................................................461
Stephaniella boliviensis Steph. ......................................................................................................136
Stephaniella hamata Steph. ..........................................................................................................136
Stephaniella paraphyllina J.B.Jack ................................................................................................136
Stephaniella rostrata U.Schmitt ...................................................................................................136
Stephaniella uncifolia S.Winkl. ....................................................................................................136
Stephaniellidium sleumeri (Müll.Frib.) S.Winkl. ex Grolle ...........................................................136
Stephensoniella brevipedunculata Kashyap ....................................................................................488
Stictolejeunea balfourii (Mitt.) E.W.Jones ....................................................................................300
Stictolejeunea balfourii var. bekkeri Gradst. ..................................................................................300
Stictolejeunea iwatsukii Mizut......................................................................................................300
660
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Stictolejeunea squamata (Willd.) Schifn. .....................................................................................300
Stolonivector clasmatocoleoides J.J.Engel .......................................................................................127
Stolonivector iordlandiae (E.A.Hodgs.) J.J.Engel .........................................................................127
Stolonivector iordlandiae var. nodulosus J.J.Engel .........................................................................127
Stolonivector gremmenii (Váňa) Váňa ...........................................................................................127
Stolonivector obtusilobus J.J.Engel ................................................................................................127
Stolonivector waipouensis J.J.Engel ...............................................................................................127
Strepsilejeunea apollinea (Gottsche) Steph....................................................................................512
Strepsilejeunea durelii Schifn.......................................................................................................512
Strepsilejeunea hamatifolia Steph. ................................................................................................512
Strepsilejeunea lanceolata (Gottsche) Steph. .................................................................................512
Strepsilejeunea muscicola Herzog ..................................................................................................512
Strepsilejeunea novae-guineae Steph..............................................................................................512
Strepsilejeunea obtusistipula Steph. ...............................................................................................512
Strepsilejeunea papulifolia Steph. .................................................................................................512
Strepsilejeunea pectiniformis (Gottsche) Steph. .............................................................................512
Strepsilejeunea renistipula Steph. ..................................................................................................512
Strepsilejeunea vatovae Gerola ......................................................................................................512
Symbiezidium barbilorum (Lindenb. et Gottsche) A.Evans .........................................................398
Symbiezidium dentatum Herzog ..................................................................................................398
Symbiezidium madagascariense Steph. ..........................................................................................398
Symbiezidium transversale (Sw.) Trevis. .......................................................................................398
Symbiezidium transversale var. hookerianum (Gottsche, Lindenb. et Nees) Gradst. et J.Beek .......398
Symphyogyna apiculispina Steph. .................................................................................................471
Symphyogyna aspera Steph. ex F.A.McCormick ...........................................................................471
Symphyogyna atronervia Taylor....................................................................................................471
Symphyogyna bogotensis Steph. .....................................................................................................471
Symphyogyna boliviensis Steph. ....................................................................................................471
Symphyogyna brasiliensis Nees et Mont. .......................................................................................471
Symphyogyna brasiliensis var. angustior (Gottsche, Lindenb. et Nees) Gottsche ............................471
Symphyogyna brasiliensis var. subsinuata Schifn. ..........................................................................472
Symphyogyna brongniartii Mont. .................................................................................................472
Symphyogyna circinata Nees et Mont. ..........................................................................................472
Symphyogyna digitisquama Steph. ................................................................................................472
Symphyogyna fuscovirens A.Evans .................................................................................................472
Symphyogyna hochstetteri Nees et Mont. ......................................................................................472
Symphyogyna hymenophyllum (Hook.) Nees et Mont. ..................................................................472
Symphyogyna hymenophyllum var. heterogenum Spruce .................................................................472
Symphyogyna ignambiensis Hürl. ..................................................................................................472
Symphyogyna interrupta Carrington et Pearson ............................................................................472
Symphyogyna irregularis Steph. ....................................................................................................472
Symphyogyna lacerosquama Steph. ...............................................................................................472
Symphyogyna leptothelia Taylor ....................................................................................................472
Symphyogyna lindmanii A.Evans ..................................................................................................472
Symphyogyna luetzelburgii Herzog ...............................................................................................472
Symphyogyna marginata Steph. ....................................................................................................472
Symphyogyna mexicana Steph. .....................................................................................................472
Symphyogyna multilora Steph. ....................................................................................................472
Symphyogyna paucidens Steph. .....................................................................................................472
Symphyogyna podophylla (hunb.) Nees et Mont. ........................................................................472
Symphyogyna purpureolimbata E.A.Hodgs. ..................................................................................473
Symphyogyna rectidens Grolle .......................................................................................................473
World checklist of hornworts and liverworts
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661
Symphyogyna rhodina (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ........................................473
Symphyogyna rubescens Steph. ......................................................................................................473
Symphyogyna rubritincta A.Evans ................................................................................................473
Symphyogyna semi-involucrata Austin ..........................................................................................473
Symphyogyna similis Grolle ..........................................................................................................473
Symphyogyna sinuata (Sw.) Nees et Mont. ...................................................................................473
Symphyogyna subsimplex Mitt. .....................................................................................................473
Symphyogyna tenuinervis (Hook.f. et Taylor) Grolle ....................................................................473
Symphyogyna trivittata Spruce .....................................................................................................473
Symphyogyna ulvoides (Reinw., Blume et Nees) Nees ...................................................................473
Symphyogyna undulata Colenso ...................................................................................................473
Symphyogyna volkensii Steph........................................................................................................473
Symphyogynopsis gottscheana (Mont. et Nees) Grolle ....................................................................473
Syzygiella acinacifolia (Hook.f. et Taylor) K.Feldberg, Váňa, Hentschel et Heinrichs ....................47
Syzygiella anomala (Lindenb. et Gottsche) Steph. ..........................................................................47
Syzygiella autumnalis (DC.) K.Feldberg, Váňa, Hentschel et Heinrichs .........................................48
Syzygiella bilobata Inoue................................................................................................................47
Syzygiella campanulata Herzog ......................................................................................................48
Syzygiella ciliata Gradst. et A.R.Benitez .........................................................................................47
Syzygiella colorata (Lehm.) K.Feldberg, Váňa, Hentschel et Heinrichs ...........................................48
Syzygiella colorata var. collenchymata J.J.Engel et Váňa ..................................................................48
Syzygiella concreta (Gottsche) Spruce .............................................................................................47
Syzygiella contigua Steph. ..............................................................................................................48
Syzygiella eatonii (Austin) Inoue ....................................................................................................49
Syzygiella elongella (Taylor) K.Feldberg, Váňa, Hentschel et Heinrichs .........................................48
Syzygiella jacquinotii (Mont.) Hentschel, K.Feldberg, Váňa et Heinrichs ......................................48
Syzygiella macrocalyx (Mont.) Spruce ............................................................................................48
Syzygiella manca (Mont.) Steph.....................................................................................................47
Syzygiella nigrescens (Steph.) K.Feldberg, Váňa, Hentschel et Heinrichs ........................................47
Syzygiella nipponica (S.Hatt.) K.Feldberg, Váňa, Hentschel et Heinrichs ......................................48
Syzygiella oenops (Lindenb. et Gottsche) K.Feldberg, Váňa, Hentschel et Heinrichs ......................49
Syzygiella ovalifolia Inoue ..............................................................................................................47
Syzygiella paludosa (Steph.) K.Feldberg, Váňa, Hentschel et Heinrichs..........................................47
Syzygiella pectiniformis Spruce .......................................................................................................47
Syzygiella perfoliata (Sw.) Spruce ...................................................................................................49
Syzygiella pseudocclusa (E.A.Hodgs.) K.Feldberg, Váňa, Hentschel et Heinrichs ............................47
Syzygiella purpurascens (Steph.) K.Feldberg, Váňa, Hentschel et Heinrichs....................................49
Syzygiella rubricaulis (Nees) Steph. ................................................................................................49
Syzygiella securifolia (Nees) Inoue ..................................................................................................48
Syzygiella setulosa Steph. ................................................................................................................49
Syzygiella sonderi (Gottsche) K.Feldberg, Váňa, Hentschel et Heinrichs ........................................47
Syzygiella spegazziniana (Spruce ex C.Massal.) K.Feldberg, Váňa, Hentschel et Heinrichs.............47
Syzygiella subintegerrima (Reinw., Blume et Nees) Spruce .............................................................48
Syzygiella tasmanica (Hook.f. et Taylor) K.Feldberg, Váňa, Hentschel et Heinrichs ......................48
Syzygiella teres (Carrington et Pearson) Váňa .................................................................................49
Syzygiella tonduzana Steph. ...........................................................................................................47
Syzygiella trigonifolia (Steph.) Herzog............................................................................................47
Syzygiella uleana Steph. .................................................................................................................49
Syzygiella undata (Mont.) K.Feldberg, Váňa, Hentschel et Heinrichs ............................................49
Targionia dioica Schifn. .............................................................................................................503
Targionia elongata Bisch..............................................................................................................503
Targionia iorii Gola ...................................................................................................................503
662
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Targionia formosica Horik. ..........................................................................................................503
Targionia hypophylla L. ...............................................................................................................503
Targionia hypophylla subsp. linealis W.Frey et Kürschner ............................................................503
Targionia indica Udar et A.Gupta ...............................................................................................503
Targionia lorbeeriana Müll.Frib. .................................................................................................503
Targionia stellaris (Müll.Frib.) Hässel..........................................................................................503
Taxilejeunea acutifolia (Steph.) Steph. .........................................................................................388
Taxilejeunea antillana Steph........................................................................................................388
Taxilejeunea apiculata (Gottsche) J.B.Jack et Steph. ....................................................................388
Taxilejeunea argentina Steph. ......................................................................................................388
Taxilejeunea arsenii Steph. ..........................................................................................................388
Taxilejeunea auriculata Steph. .....................................................................................................388
Taxilejeunea berteroana Steph. ....................................................................................................388
Taxilejeunea beyrichiana Steph. ...................................................................................................388
Taxilejeunea biapiculata Steph. ...................................................................................................388
Taxilejeunea boliviana Steph. ......................................................................................................388
Taxilejeunea brasiliensis Steph. ....................................................................................................388
Taxilejeunea coilantha Herzog .....................................................................................................388
Taxilejeunea compressiuscula Steph. .............................................................................................389
Taxilejeunea convoluta Herzog ....................................................................................................389
Taxilejeunea cuervi (Gottsche) Steph. ..........................................................................................389
Taxilejeunea cuneistipula Steph. ..................................................................................................389
Taxilejeunea cuspidata Steph. ......................................................................................................389
Taxilejeunea decurrens Steph. ......................................................................................................389
Taxilejeunea delexa Steph. ..........................................................................................................389
Taxilejeunea densilora A.Evans ...................................................................................................389
Taxilejeunea diaphana (Lehm.) Steph. ........................................................................................389
Taxilejeunea dissitifolia Steph. .....................................................................................................389
Taxilejeunea eggersiana Schifn. ...................................................................................................389
Taxilejeunea elobulata Sim ..........................................................................................................389
Taxilejeunea issistipula Steph. .....................................................................................................389
Taxilejeunea foliicola Steph. ........................................................................................................389
Taxilejeunea furcicornuta Grolle ..................................................................................................389
Taxilejeunea fuscorufa Steph. .......................................................................................................389
Taxilejeunea galapagensis Onr......................................................................................................389
Taxilejeunea ghatensis P.K.Verma et S.C.Srivast. .........................................................................389
Taxilejeunea giulianettii Steph. ....................................................................................................389
Taxilejeunea gomphocalyx Herzog ................................................................................................389
Taxilejeunea grandifolia Steph. ....................................................................................................389
Taxilejeunea grandistipula Steph..................................................................................................389
Taxilejeunea hamatifolia Steph. ...................................................................................................389
Taxilejeunea himalayensis Herzog ................................................................................................389
Taxilejeunea immersa Eifrig .........................................................................................................389
Taxilejeunea irregularis Steph. .....................................................................................................389
Taxilejeunea jamaicensis A.Evans .................................................................................................389
Taxilejeunea jeringii Steph. .........................................................................................................389
Taxilejeunea killipii Herzog.........................................................................................................389
Taxilejeunea laevis (Gottsche) Steph............................................................................................340
Taxilejeunea langiana Pearson .....................................................................................................340
Taxilejeunea lindbergiana Steph. .................................................................................................340
Taxilejeunea linguifolia Steph. .....................................................................................................340
Taxilejeunea maxima Steph. ........................................................................................................340
World checklist of hornworts and liverworts
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663
Taxilejeunea mexicana Steph. ......................................................................................................340
Taxilejeunea microstipula Steph. ..................................................................................................340
Taxilejeunea mucronata Steph. ....................................................................................................340
Taxilejeunea multilora Steph. .....................................................................................................340
Taxilejeunea muscicola Steph. ......................................................................................................340
Taxilejeunea nilgiriensis P.K.Verma et S.C.Srivast. ......................................................................340
Taxilejeunea nymannii Steph. ......................................................................................................340
Taxilejeunea obtusifolia Steph. .....................................................................................................340
Taxilejeunea papulilora Steph. ....................................................................................................340
Taxilejeunea parvibracteata Steph. ...............................................................................................340
Taxilejeunea parvistipula Steph....................................................................................................340
Taxilejeunea paucidens Steph. ......................................................................................................340
Taxilejeunea pendula Steph. ........................................................................................................340
Taxilejeunea peruviana Steph. .....................................................................................................340
Taxilejeunea planilobula Herzog ..................................................................................................340
Taxilejeunea pulchrilora Pearson .................................................................................................340
Taxilejeunea pusilla Steph. ..........................................................................................................340
Taxilejeunea renistipula (Lindenb.) Steph. ...................................................................................340
Taxilejeunea rufescens Steph. .......................................................................................................340
Taxilejeunea serpillifolioides (Raddi) D.P.Costa ...........................................................................340
Taxilejeunea setchellii Pearson .....................................................................................................340
Taxilejeunea speciosa Herzog .......................................................................................................340
Taxilejeunea splendida Eifrig .......................................................................................................340
Taxilejeunea stephanii Eifrig ........................................................................................................340
Taxilejeunea steyermarkii H.Rob. ................................................................................................340
Taxilejeunea surinamensis (Lindenb. et Gottsche) Steph. .............................................................340
Taxilejeunea suringarii Steph. ......................................................................................................341
Taxilejeunea tenerrima Steph.......................................................................................................341
Taxilejeunea tenuiplica Steph. .....................................................................................................341
Taxilejeunea tjibodensis (Steph.) Eifrig.........................................................................................341
Taxilejeunea tonduzana Steph. ....................................................................................................341
Taxilejeunea uleana Steph. ..........................................................................................................341
Taxilejeunea umbonata Steph. .....................................................................................................341
Taxilejeunea urbanii Steph. .........................................................................................................341
Taxilejeunea vallis-gratiae Steph. .................................................................................................341
Telaranea anomala R.M.Schust. ex J.J.Engel et G.L.Merr. ..........................................................186
Telaranea apiahyna (Steph.) Fulford ...........................................................................................185
Telaranea azorica (H.Buch et Perss.) Pócs ...................................................................................187
Telaranea bicruris (Steph.) M.Howe ...........................................................................................185
Telaranea bischleriana Pócs .........................................................................................................186
Telaranea blepharostoma (Steph.) Fulford ....................................................................................185
Telaranea breviseta (Herzog) J.J.Engel et G.L.Merr. ....................................................................185
Telaranea chaetophylla (Spruce) Schifn. ......................................................................................185
Telaranea coactilis (Spruce) J.J.Engel et G.L.Merr. ......................................................................186
Telaranea confervoides J.J.Engel et G.L.Merr. ..............................................................................186
Telaranea diacantha (Mont.) J.J.Engel et G.L.Merr. ...................................................................186
Telaranea europaea J.J.Engel et G.L.Merr. ..................................................................................185
Telaranea fragilis Mizut...............................................................................................................186
Telaranea granulata J.J.Engel et G.L.Merr. .................................................................................186
Telaranea herzogii (E.A.Hodgs.) E.A.Hodgs................................................................................186
Telaranea inaequalis R.M.Schust. ex J.J.Engel et G.L.Merr. ........................................................187
Telaranea indica (S.C.Srivast. et P.K.Verma) A.E.D.Daniels et P.Daniel ....................................187
664
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Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
Telaranea longifolia (M.Howe) J.J.Engel et G.L.Merr. ................................................................186
Telaranea major (Herzog) J.J.Engel et G.L.Merr. ........................................................................187
Telaranea maorensis Pócs .............................................................................................................187
Telaranea microstipulata R.M.Schust. .........................................................................................187
Telaranea monocera (Mitt. ex R.M.Schust. et Grolle) J.J.Engel et G.L.Merr. ..............................187
Telaranea nematodes (Gottsche ex Austin) M.Howe ....................................................................186
Telaranea panchoi Del Ros. .........................................................................................................186
Telaranea pecten (Spruce) J.J.Engel et G.L.Merr. ........................................................................187
Telaranea pellucida J.J.Engel et G.L.Merr. ..................................................................................186
Telaranea pseudozoopsis (Herzog) Fulford ....................................................................................186
Telaranea redacta (Steph.) J.J.Engel et G.L.Merr. .......................................................................186
Telaranea rosarioana H.A.Mill. ...................................................................................................186
Telaranea sejuncta (Ångstr.) S.W.Arnell ......................................................................................187
Telaranea setosa J.J.Engel et G.L.Merr.........................................................................................186
Telaranea tenuifolia J.J.Engel et G.L.Merr. .................................................................................187
Telaranea trisetosa (Steph.) Grolle ...............................................................................................186
Temnoma angustifolium R.M.Schust. ..........................................................................................255
Temnoma chaetophyllum R.M.Schust. .........................................................................................255
Temnoma palmatum (Lindb. ex Pearson) R.M.Schust. ................................................................255
Temnoma palmatum var. cuneatum R.M.Schust. .........................................................................255
Temnoma palmatum var. laxifolium R.M.Schust. ........................................................................255
Temnoma palmatum var. pseudospiniferum R.M.Schust. ..............................................................255
Temnoma patagonicum R.M.Schust. ...........................................................................................255
Temnoma paucisetigerum R.M.Schust..........................................................................................255
Temnoma pilosum (A.Evans) R.M.Schust. ...................................................................................255
Temnoma pulchellum (Hook.) Mitt. ............................................................................................255
Temnoma quadriidum (Mitt.) Mitt. ex E.A.Hodgs. et Allison ....................................................255
Temnoma quadripartitum (Hook.) Mitt. .....................................................................................255
Temnoma quadripartitum var. pseudopungens R.M.Schust. ..........................................................255
Temnoma quadripartitum var. randii (S.W.Arnell) R.M.Schust. .................................................256
Temnoma setigerum (Lindenb.) R.M.Schust. ...............................................................................256
Temnoma setigerum var. hawaiicum Inoue ...................................................................................256
Temnoma townrowii R.M.Schust. ...............................................................................................256
Tetracymbaliella comptonii (Pearson) Grolle ................................................................................137
Tetracymbaliella cymbalifera (Hook.f. et Taylor) Grolle ..............................................................137
Tetracymbaliella decipiens (Gottsche) Grolle ................................................................................138
Tetracymbaliella subsimplex (Austin) J.J.Engel .............................................................................138
Tetralophozia cavallii (Gola) Váňa ................................................................................................55
Tetralophozia iliformis (Steph.) Urmi ...........................................................................................55
Tetralophozia pilifera (Steph.) R.M.Schust. ...................................................................................55
Tetralophozia setiformis (Ehrh.) Schljakov .....................................................................................55
hysananthus aculeatus Herzog ....................................................................................................413
hysananthus amazonicus (Spruce) Schifn...................................................................................413
hysananthus anguiformis (Hook.f. et Taylor) Gottsche, Lindenb. et Nees ..................................413
hysananthus appendiculatus Steph. .............................................................................................413
hysananthus ciliaris (Sande Lac.) Sukkharak ..............................................................................413
hysananthus comosus Lindenb. ...................................................................................................413
hysananthus convolutus Lindenb. ...............................................................................................413
hysananthus convolutus var. laceratus (Steph.) Sukkharak ...........................................................413
hysananthus discretus Sukkharak et Gradst. ................................................................................413
hysananthus fruticosus (Lindenb. et Gottsche) Schifn. ...............................................................414
hysananthus gottschei (J.B.Jack et Steph.) Steph. ........................................................................413
World checklist of hornworts and liverworts
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665
hysananthus gottschei var. continuus Sukkharak ..........................................................................413
hysananthus mollis Steph. ..........................................................................................................414
hysananthus montanus Gradst., Xiao L.He et Piippo..................................................................414
hysananthus pancheri (Steph.) Hürl. ..........................................................................................413
hysananthus retusus (Reinw., Blume et Nees) B.M.hiers et Gradst. ..........................................414
hysananthus retusus subsp. sellingii (Herzog) Sukkharak.............................................................414
hysananthus spathulistipus (Reinw., Blume et Nees) Lindenb. ....................................................414
hysanolejeunea africana Sim .......................................................................................................512
Trabacellula tumidula Fulford .......................................................................................................64
Trachylejeunea conifera Steph. .....................................................................................................512
Trachylejeunea cristulilora Steph. ................................................................................................512
Trachylejeunea englishii Steph......................................................................................................512
Trachylejeunea jamaicensis Pearson ..............................................................................................512
Trachylejeunea kusaiensis Inoue et H.A.Mill. ...............................................................................512
Treubia insignis K.I.Goebel ...........................................................................................................43
Treubia insignis subsp. bracteata (Steph.) R.M.Schust. et G.A.M.Scott .........................................43
Treubia insignis subsp. caledonica R.M.Schust. et G.A.M.Scott .....................................................43
Treubia insignis subsp. vitiensis R.M.Schust. et G.A.M.Scott ........................................................43
Treubia lacunosoides T.Pfeif., W.Frey et M.Stech .........................................................................43
Treubia lacunosa (Colenso) Prosk..................................................................................................43
Treubia pygmaea R.M.Schust. .......................................................................................................43
Treubia scapanioides R.M.Schust...................................................................................................43
Treubia tahitensis (Nadeaud) Besch. ..............................................................................................43
Treubia tasmanica R.M.Schust. et G.A.M.Scott ............................................................................43
Triandrophyllum eophyllum (R.M.Schust.) Gradst. ......................................................................141
Triandrophyllum fernandeziense (S.W.Arnell) Grolle ex Fulford et Hatcher .................................141
Triandrophyllum heterophyllum (Steph.) Grolle ...........................................................................141
Triandrophyllum subtriidum (Hook.f. et Taylor) Fulford et Hatcher ..........................................141
Triandrophyllum subtriidum var. triidum (Gottsche) Solari........................................................141
Triandrophyllum symmetricum J.J.Engel ......................................................................................141
Trichocolea argentea Herzog ........................................................................................................257
Trichocolea breviissa Steph. .........................................................................................................257
Trichocolea comptonii Pearson .....................................................................................................257
Trichocolea ilicaulis Steph. ..........................................................................................................257
Trichocolea loccosa Herzog et Hatcher ........................................................................................257
Trichocolea geniculata Pearson .....................................................................................................257
Trichocolea gracillima Austin .......................................................................................................257
Trichocolea hatcheri E.A.Hodgs. ..................................................................................................257
Trichocolea iriomotensis T.Katag. .................................................................................................257
Trichocolea japonica T.Katag. ......................................................................................................257
Trichocolea magna T.Katag. ........................................................................................................257
Trichocolea minutifolia Steph. .....................................................................................................257
Trichocolea mollissima (Hook.f. et Taylor) Gottsche ....................................................................257
Trichocolea pluma (Reinw., Blume et Nees) Mont.......................................................................258
Trichocolea rigida R.M.Schust. ....................................................................................................258
Trichocolea rudimentaris Steph. ...................................................................................................258
Trichocolea sprucei Steph. ............................................................................................................258
Trichocolea tomentella (Ehrh.) Dumort. ......................................................................................258
Trichocolea udarii D.K.Singh ......................................................................................................258
Trichocolea wattsiana Steph. ........................................................................................................258
Trichocoleopsis sacculata (Mitt.) S.Okamura ................................................................................434
Tricholepidozia chaetocarpa (Pearson) E.D.Cooper ......................................................................179
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Tricholepidozia fernandeziensis (Steph.) E.D.Cooper ...................................................................179
Tricholepidozia ferruginea (J.J.Engel et G.L.Merr.) E.D.Cooper ..................................................179
Tricholepidozia issifolia (Steph.) E.D.Cooper..............................................................................179
Tricholepidozia jowettiana (H.A.Mill.) E.D.Cooper ....................................................................179
Tricholepidozia kaindina (Grolle) E.D.Cooper ............................................................................179
Tricholepidozia kogiana (Steph.) E.D.Cooper ..............................................................................179
Tricholepidozia lawesii (Steph.) E.D.Cooper ...............................................................................179
Tricholepidozia leratii (Steph.) E.D.Cooper ................................................................................180
Tricholepidozia lindenbergii (Gottsche) E.D.Cooper ...................................................................180
Tricholepidozia lindenbergii var. complanata (J.J.Engel et G.L.Merr.) E.D.Cooper ......................180
Tricholepidozia lindenbergii var. mellea (J.J.Engel et G.L.Merr.) E.D.Cooper ..............................180
Tricholepidozia lindenbergii var. papillata (J.J.Engel et G.L.Merr.) E.D.Cooper ..........................180
Tricholepidozia marginata (J.J.Engel et G.L.Merr.) E.D.Cooper .................................................180
Tricholepidozia martinii (E.A.Hodgs.) E.D.Cooper.....................................................................180
Tricholepidozia melanesica (H.A.Mill.) E.D.Cooper ....................................................................180
Tricholepidozia neesii (Lindenb.) E.D.Cooper .............................................................................180
Tricholepidozia octoloba (Del Ros.) E.D.Cooper ..........................................................................180
Tricholepidozia plumulosa (Lehm. et Lindenb.) E.D.Cooper .......................................................180
Tricholepidozia pulcherrima (Steph.) E.D.Cooper .......................................................................180
Tricholepidozia pulcherrima var. mooreana (Steph.) E.D.Cooper .................................................180
Tricholepidozia quadriseta (Steph.) E.D.Cooper ..........................................................................181
Tricholepidozia remotifolia (E.A.Hodgs.) E.D.Cooper .................................................................181
Tricholepidozia semperiana (Steph.) E.D.Cooper .........................................................................181
Tricholepidozia tetradactyla (Hook.f. et Taylor) E.D.Cooper .......................................................181
Tricholepidozia trichocoleoides (Herzog) E.D.Cooper ...................................................................181
Trichotemnoma corrugatum (Steph.) R.M.Schust. .......................................................................136
Trilophozia quinquedentata (Huds.) Bakalin .................................................................................81
Trilophozia quinquedentata var. asymmetrica (Horik.) L.Söderstr. et Váňa.....................................82
Tritomaria exsecta (Schmidel) Schifn. ex Loeske ...........................................................................82
Tritomaria exsecta subsp. novaezelandiae J.J.Engel .........................................................................82
Tritomaria exsectiformis (Breidl.) Schifn. ex Loeske ......................................................................82
Tritomaria exsectiformis subsp. arctica R.M.Schust. .......................................................................82
Tritomaria exsectiformis subsp. camerunensis S.W.Arnell ex Váňa ..................................................82
Tritomaria ferruginea (Grolle) Váňa ..............................................................................................82
Tritomaria koreana Bakalin, S.S.Choi et B.Y.Sun .........................................................................82
Tritomaria mexicana Bakalin.........................................................................................................82
Tritomaria scitula (Taylor) Jørg.....................................................................................................82
Tuyamaella angulistipa (Steph.) R.M.Schust. et Kachroo ............................................................350
Tuyamaella borneensis Tixier .......................................................................................................350
Tuyamaella hattorii Tixier ...........................................................................................................350
Tuyamaella jackii (Steph.) Tixier ................................................................................................350
Tuyamaella molischii (Schifn.) S.Hatt. .......................................................................................350
Tuyamaella molischii var. brevistipa P.C.Wu et P.J.Lin ................................................................350
Tuyamaella molischii var. taiwanensis R.L.Zhu et M.L.So............................................................350
Tuyamaella serratistipa S.Hatt. ....................................................................................................350
Tuzibeanthus chinensis (Steph.) Mizut. ........................................................................................414
Vanaea plagiochiloides (Inoue et Gradst.) Inoue et Gradst. ............................................................49
Vandiemenia ratkowskiana Hewson.............................................................................................461
Verdoornia succulenta R.M.Schust. ..............................................................................................456
Verdoornianthus griinii Gradst...................................................................................................414
Verdoornianthus marsupiifolius (Spruce) Gradst. ..........................................................................414
Vetaforma dusenii (Steph.) Fulford et J.Taylor ............................................................................142
World checklist of hornworts and liverworts
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667
Vitalianthus bischlerianus (K.C.Pôrto et Grolle) R.M.Schust. et Giancotti ..................................361
Vitalianthus guangxianus R.L.Zhu, Qiong He et Y.M.Wei..........................................................361
Wettsteinia densiretis (Herzog) Grolle ............................................................................................45
Wettsteinia inversa (Sande Lac.) Schifn.........................................................................................45
Wettsteinia rotundifolia (Horik.) Grolle .........................................................................................45
Wettsteinia schusteriana Grolle.......................................................................................................45
Wiesnerella denudata (Mitt.) Steph..............................................................................................504
Wiesnerella fasciaria C.Gao et K.C.Chang ...................................................................................504
Xenocephalozia navicularis (Steph.) R.M.Schust. .........................................................................217
Xenochila integrifolia (Mitt.) Inoue..............................................................................................253
Xenothallus vulcanicola R.M.Schust. ............................................................................................473
Xylolejeunea aquarius (Spruce) Xiao L.He et Grolle .....................................................................398
Xylolejeunea crenata (Nees et Mont.) Xiao L.He et Grolle ...........................................................398
Xylolejeunea grolleana (Pócs) Xiao L.He et Grolle ........................................................................398
Xylolejeunea muricella Xiao L.He et Grolle ..................................................................................398
Zantenia borneensis (Herzog) Váňa et J.J.Engel .............................................................................56
Zantenia denticulata (Grolle) Váňa et J.J.Engel .............................................................................56
Zantenia karstenii (Schifn.) Váňa et J.J.Engel ...............................................................................56
Zantenia prionophylla (S.Hatt.) Váňa et J.J.Engel ..........................................................................56
Zoopsidella antillana (Steph.) H.Rob...........................................................................................187
Zoopsidella antillana subsp. jamaicensis R.M.Schust. ...................................................................187
Zoopsidella caledonica (Steph.) R.M.Schust. ................................................................................187
Zoopsidella cynosurandra (Steph.) R.M.Schust. ............................................................................187
Zoopsidella integrifolia (Spruce) R.M.Schust................................................................................188
Zoopsidella macella (Spruce) R.M.Schust. ....................................................................................188
Zoopsidella serra (Spruce) R.M.Schust. ........................................................................................188
Zoopsis argentea (Hook.f. et Taylor) Gottsche, Lindenb. et Nees .................................................188
Zoopsis argentea var. lagelliformis (Colenso) R.M.Schust. ............................................................188
Zoopsis bicruris Glenny et E.A.Br. ...............................................................................................188
Zoopsis ceratophylla (Spruce) Hamlin...........................................................................................188
Zoopsis ciliata Colenso ................................................................................................................189
Zoopsis leitgebiana (Carrington et Pearson) Bastow .....................................................................188
Zoopsis liukiuensis Horik. ............................................................................................................188
Zoopsis macrophylla R.M.Schust. .................................................................................................188
Zoopsis martinicensis Steph. .........................................................................................................189
Zoopsis matawaia M.A.M.Renner ...............................................................................................188
Zoopsis nitida Glenny, Braggins et R.M.Schust. ..........................................................................188
Zoopsis setigera K.I.Goebel ..........................................................................................................189
Zoopsis setulosa Leitg....................................................................................................................188
Zoopsis uleana Steph....................................................................................................................189
Acknowledgements
he Early Land Plants Today project has been a community driven and collaborative efort
that has involved numerous dedicated individuals, many institutions, and funding agencies that have helped provide partial funding. he high quality of the data and information
could not have been achieved without rapid responses to our requests for information
regarding publications, PDFs or other forms of bibliographic data from committed individuals. hese include Afroz Alam, Dulip Daniels, Rajendra Lavate, Shuvadeep Majum-
668
Lars Söderström et al. / PhytoKeys 59: 1–828 (2016)
dar, Alexey Potemkin, Ana Séneca, Mafalda Silva, Devendra K. Singh, Sushil K. Singh,
Louis houvenot, Praveen Verma, and Gerhard Winter. We wish to especially thank the
many individuals who have provided a wealth of logistical support that has been broad
ranging from access to data, assistance with translation, access to unpublished molecular
results and information, access to type material or herbaria support, library support, advice
on Latin usage as well as many other forms of support. hese have included Silvia Aranda,
Riccardo Baldini, Fred Barrie, Laura Campos, Tong Cao, Xia-Fang Cheng, Patricia Eckel,
Len Ellis, Lars Hedenäs, Christine Giannoni, Qiong He, Nijole Kalinauskaitė, Lucia Kawasaki, Peter de Lange, Itambi Malombe, John McNeill, David Meagher, Chiara Nepi,
Michelle Price, Qiong Qui, Steven Rogstad, Cecilia Sérgio, Blanka Shaw, Jon Shaw, Akiko
Shinya, Lei Shu, Sorin Ştefănuţ, Alain Vanderpoorten, Anna Vilnet, Yu-Mei Wei, Ying
Yu, and Lan-Ping Zhou. We gratefully acknowledge the Biodiversity Heritage Library as
a resource and the Conservatoire et Jardin botaniques de la Ville de Genève for providing
access to the Franz Stephani database and specimens and Missouri Botanical Garden for
access to the TROPICOS database. hroughout the years, the project has been generously supported in part by the Global Biological Information Facility (GBIF) Seed Money
Award No.2007-41 and supported in part by funding from the Biodiversity Synthesis
Center of the Encyclopedia of Life. Funding from the National Science Foundation (DBI0749762, DBI 1057418, DBI 1458300, DEB- 1145898, EF-0531730, EF-1115116)
has also partially supported aspects of the ELPT project. We also wish to express our
gratitude to the editors and publication team of Phytotaxa that published the important
series of notes that helped lead to the current publication and to the tireless eforts of the
editorial and production team of PhytoKeys that led to the current publication.
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Grolle R (1970a) Radula castlei sp. nov. und Anmerkungen zur Gattung Radula. Bryologist 73
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Grolle R (1970b) Eine neuen Jackiella aus Tasmanien. Journal of the Hattori Botanical Laboratory
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Grolle R (1971a) Die Lebermoose der Crozet-Inseln (Subantarktis). Lindbergia 1 (1/2): 80–82.
Grolle R (1971b) Jamesoniella und Verwandte. Feddes Repertorium 82 (1): 1–100. doi:
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Grolle R (1971c) Lebermoose aus Neuguinea. 9. Jungermannia nivea sp. nov. Miscellanea Bryologica
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Grolle R (1972b) Bazzania in Europa und Makaronesien. Zur Taxonomie und Verbreitung.
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Grolle R (1972c) Zur Kenntnis von Adelanthus Mitt. Journal of the Hattori Botanical Laboratory
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Grolle R (1973a) Miscellanea hepaticologica 131-140. Herzogia 3: 75–82.
Grolle R (1973b) Nephelolejeunea - eine neue Gattung der Tuyamaelloideae. Journal of the
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Grolle R (1974a) Eine neue Taxilejeunea aus Madagaskar und Reunion. Journal of Bryology 8
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Grolle R (1974b) Lebermoose aus Neuguinea. 12. Rhaphidolejeunea. Journal of the Hattori
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Grolle R (1975a) Diplasiolejeunea in Australasien. Feddes Repertorium 86 (1/2): 75–82. doi:
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Grolle R (1975b) Haplolejeunea aus Madagascar – Eine weitere neue Gattung der Tuyamaelloideae.
Journal of the Hattori Botanical Laboratory 39: 203–210.
Grolle R (1975c) Miscellanea hepaticologica 151-160. Lindbergia 3 (1/2): 47–56.
Grolle R (1975d) Miscellanea hepaticologica 141-150. Journal of Bryology 8 (4): 483–492.
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Grolle R (1976a) Verzeichnis der Lebermoose Europas und benachbarter Gebiete. Feddes Repertorium 87 (3/4): 171–279. doi: 10.1002/fedr.19760870303
Grolle R (1976b) Eine weitere Siphonolejeunea –S. elegantissima (Steph.) comb. nov. aus Australien.
Journal of the Hattori Botanical Laboratory 41: 405–409.
Grolle R (1976c) Drepanolejeunea subg. Kolpolejeunea – eine neue Untergattung aus der Palaeotropis.
Journal of the Hattori Botanical Laboratory 40: 191–216.
Grolle R (1977a) Miscellanea hepaticologica 161-170. Journal of Bryology 9 (4): 529–538.
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Grolle R (1977b) Pictolejeunea – eine neue Gattung der Lejeuneoideae aus der Neotropis und
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Grolle R (1978a) Eine neue Diplasiolejeunea-Art aus Sri Lanka. Feddes Repertorium 89 (5/6):
301–305. doi: 10.1002/fedr.19780890502
Grolle R (1978b) Lebermoose aus Neuguinea. 16. Acromastigum. Journal of the Hattori Botanical
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Grolle R (1979a) Lebermoose aus Neuguinea. 17. Harpalejeunea. Journal of the Hattori Botanical
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Grolle R (1979b) Miscellanea hepaticologica 171-180. Journal of Bryology 10 (3): 263–272.
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Grolle R (1979c) Miscellanea hepaticologica 181-190. Journal of the Hattori Botanical Laboratory
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Grolle R (1979d) Miscellanea hepaticologica 191-200. Journal of the Hattori Botanical Laboratory
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Grolle R (1980a) Nephelolejeunea in Australasien. Journal of the Hattori Botanical Laboratory
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Grolle R (1980b) Schusterolejeunea Grolle nom. nov. statt Cladocolea Schust. 1963, non van
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Grolle R (1980c) Miscellanea hepaticologica 201-210. Journal of Bryology 11 (2): 325–334.
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Grolle R (1980d) Zur Kenntnis der Lebermoose von Samoa. I. Wissenschaftliche Zeitschrift
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Grolle R (1981) Miscellanea hepaticologica 211-220. Journal of the Hattori Botanical Laboratory
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Grolle R (1982) Übersicht der Lejeuneaceae in Tasmanien. Wissenschaftliche Zeitschrift der
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Grolle R (1984a) Miscellanea hepaticologica 221-230. Journal of the Hattori Botanical Laboratory
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Grolle R (1984b) Zur Kenntnis der Lejeuneoideae in Cuba (1): Cyclolejeunea. Wissenschaftliche
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Grolle R (1985b) Miscellanea hepaticologica 231-240. Journal of the Hattori Botanical Laboratory 58: 197–202.
Grolle R (1986a) Miscellanea hepaticologica 241-250. Journal of the Hattori Botanical Laboratory
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Grolle R (1986b) Cololejeunea schaeferi spec. nov., ein verkanntes Lebermoos in Makaronesien.
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Grolle R (1987) Miscellanea hepaticologica 251-260. Journal of the Hattori Botanical Laboratory
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Grolle R (1989a) Marsupidium in Guyana. Journal of the Hattori Botanical Laboratory 66:
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Grolle R (1991) Miscellanea hepaticologica 281-290. Journal of the Hattori Botanical Laboratory
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Grolle R (1992a) Zwei neue Arten der Lebermoosgattung Diplasiolejeunea aus der Neotropis.
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Grolle R, Piippo S (1984) Annotated catalogue of Western Melanesian bryophytes. I. Hepaticae
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Grolle R, Reiner-Drehwald ME (1999) Review of the genus Harpalejeunea (Lejeuneaceae)
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Grolle R, Seppelt RD (1986) Seppeltia, a new leafy genus of Metzgeriales from Macquarie
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Schifner V (1900b) Hepaticae massartianae javanicae. Hedwigia 39 (4): 191–208.
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Schifner V (1902) Neue Materialen zur Kenntnis der Bryophyten der atlantischen Inseln.
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Schifner V (1912a) Über eine kritische Form von Riccia sorocarpa und Riccia pseudopapillosa.
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Schumacker R, Váňa J (2005) Identiication keys to the liverworts and hornworts of Europe
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Schuster RM (1950) Notes on nearctic hepaticae. II. he hepaticae in east coast of Hudson
Bay. Bulletin of the National Museum of Canada 122: 1–62.
Schuster RM (1952) Notes on nearctic hepaticae. V. he status of Lophozia gracillima Buch
and its relation to Lophozia longidens, Lophozia porphyroleuca and Sphenolobus ascendens.
Bryologist 55 (3): 173–185. doi: 10.2307/3239846
Schuster RM (1953) Boreal hepaticae. A manual of the liverworts of Minnesota and adjacent
regions. American Midland Naturalist 49 (2): 257–684.
Schuster RM (1956a) North American Lejeuneaceae. V. Schizostipae: Ceratolejeunea. Journal
of the Elisha Mitchell Scientiic Society 72 (2): 292–316.
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Schuster RM (1956b) Notes on American Lejeuneaceae. IV. Paradoxae: Cololejeunea (concl.),
Diplasiolejeunea. Journal of the Elisha Mitchell Scientiic Society 72 (1): 87–125.
Schuster RM (1956c) Aphanolejeunea cornutissima nom. nov. Bryologist 59 (3): 217–218. doi:
10.2307/3239952
Schuster RM (1957a) North American Lejeuneaceae. VI. Lejeunea. Introduction and keys;
subgenus Lejeunea. Journal of the Elisha Mitchell Scientiic Society 73 (1): 122–197.
Schuster RM (1957b) Notes on nearctic hepaticae. XII. Marsupella paroica n. sp. Bryologist 60
(2): 145–151. doi: 10.2307/3240015
Schuster RM (1957c) Notes on Nearctic hepaticae, IX. A study of Plagiochila yokogurensis
Steph. Journal of the Hattori Botanical Laboratory 18: 14–26.
Schuster RM (1957d) North American lejeuneaceae. VI. Lejeunea: subgenus Lejeunea (II, concluded). Journal of the Elisha Mitchell Scientiic Society 73 (2): 388–443.
Schuster RM (1958a) Notes on nearctic hepaticae. VI. Phytogeographical relationships of critical
species in Minnesota and adjacent areas of the Great Lakes. Rhodora 60 (717): 243–256.
Schuster RM (1958b) Notes on nearctic hepaticae. XIII: Tritomaria (Lophoziaceae) in Arctic.
Canadian Journal of Botany 36 (2): 269–288. doi: 10.1139/b58-023
Schuster RM (1959a) A monograph of the nearctic Plagiochilaceae. Part. I. Introduction and
sectio I. Asplenioides. American Midland Naturalist 62 (1): 1–166. doi: 10.2307/2422546
Schuster RM (1959b) A monograph of the nearctic Plagiochilaceae. Part. II. Sectio Zonatae through
Sectio Parallelae. American Midland Naturalist 62 (2): 257–395. doi: 10.2307/2422533
Schuster RM (1959c) Studies on hepaticae. I Temnoma. Bryologist 62 (4): 233–242. doi:
10.2307/3240148
Schuster RM (1960a) Notes on nearctic hepaticae. XIX. he relationships of Blepharostoma,
Temnoma and Lepicolea, with description of Lophochaete and Chandonanthus subg. Tetralophozia, subg. n. Journal of the Hattori Botanical Laboratory 23: 192–210.
Schuster RM (1960b) Studies on hepaticae. II. he new family Chaetophylliopsidaceae. Journal
of the Hattori Botanical Laboratory 23: 68–76.
Schuster RM (1960c) A monograph of the nearctic Plagiochilaceae. Part. III. Sectio Contiguae
to conclusion. American Midland Naturalist 63 (1): 1–130. doi: 10.2307/2422932
Schuster RM (1961a) Studies in Lophoziaceae. 1. he genera Anastrophyllum and Sphenolobus
and their segregates. Revue Bryologique et Lichénologique 30 (1/2): 55–73.
Schuster RM (1961b) Notes on nearctic hepaticae. XVIII. New Lophoziaceae from the arctic
archipelago of Canada. Canadian Journal of Botany 39 (4): 965–992. doi: 10.1139/b61-081
Schuster RM (1962a) North American Lejeuneaceae. VIII. Lejeunea subgenera Microlejeunea
and Chaetolejeunea. Journal of the Hattori Botanical Laboratory 25: 1–80.
Schuster RM (1962b) Notes on American Lejeuneaceae. VII. Lejeunea (Lejeunea) blomquistii
sp. nov. Journal of the Elisha Mitchell Scientiic Society 78 (1): 64–68.
Schuster RM (1963a) An annotated synopsis of the genera and subgenera of Lejeuneaceae. I.
Introduction; annotated keys to subfamilies and genera. Beihefte zur Nova Hedwigia 9:
1–203.
Schuster RM (1963b) Studies on antipodal hepaticae. I. Annotated keys to the genera of antipodal Hepaticae with special reference to New Zealand and Tasmania. Journal of the
Hattori Botanical Laboratory 26: 185–309.
World checklist of hornworts and liverworts
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Schuster RM (1963c) Studies on hepaticae XI-XIII. On Temnoma, Vetaforma and Lophochaete
(Blepharostomaceae; Hepaticae). Nova Hedwigia 5: 27–46.
Schuster RM (1964a) Studies on antipodal hepaticae. IV. Metzgeriales. Journal of the Hattori
Botanical Laboratory 27: 183–216.
Schuster RM (1964b) Studies on hepaticae. XXII-XXV. Pleurocladopsis Schust., gen. n., Eoisotachis Schust., gen. n., Grollea Schust., gen. n., with critical notes on Anthelia Dumort.
Nova Hedwigia 8 (3/4): 275–296.
Schuster RM (1965a) North American Lejeuneaceae. IX. Taxilejeunea. Journal of the Elisha
Mitchell Scientiic Society 81 (1): 32–50.
Schuster RM (1965b) Studies on hepaticae. XXVI. he Bonneria-Paracromastigum-Pseudocephalozia-Hyalolepidozia-Zoopsis-Pteropsiella complex and its allies – a phylogenetic study, part
1. Nova Hedwigia 10 (1/2): 19–61.
Schuster RM (1965c) Studies on antipodal hepaticae. II. Archeophylla Schuster and Archeochaete Schuster, new genera of Blepharostomataceae. Transactions of the British Bryological Society 4 (5): 801–817. doi: 10.1179/006813865804811975
Schuster RM (1966a) Studies on hepaticae, VII-X. On Adelanthus Mitten and Calyptrocolea
Schuster, gen. n. Revue Bryologique et Lichénologique 34 (3/4): 676–703.
Schuster RM (1966b) Studies in Lophoziaceae. 1. he genera Anastrophyllum and Sphenolobus
and their segregates. 2. Cephalolobus gen. n., Acrolophozia gen. n. and Protomarsupella gen. n.
Revue Bryologique et Lichénologique 34 (1/2): 241–287.
Schuster RM (1966c) A memoir on the family Blepharostomataceae, II. Candollea 21 (2):
241–355.
Schuster RM (1966d) he hepaticae and anthocerotae of North America. I. Columbia University
Press, New York, 802 pp.
Schuster RM (1966e) A memoir on the family Blepharostomataceae, I. Candollea 21 (1): 59–136.
Schuster RM (1966f) Studies on hepaticae, XXVIII. On Phycolepidozia, a new, highly reduced
genus of Jungermanniales of questionable ainity. Bulletin of the Torrey Botanical Club
93 (6): 437–449. doi: 10.2307/2483417
Schuster RM (1967a) North American Lejeuneaceae. X. Harpalejeunea, Drepanolejeunea, and
Leptolejeunea. Journal of the Elisha Mitchell Scientiic Society 83 (4): 192–229.
Schuster RM (1967b) A note on the genus Gymnocolea Dum. Bryologist 70 (1): 111–112. doi:
10.2307/3241147
Schuster RM (1967c) Studies on hepaticae. XV. Calobryales. Nova Hedwigia 13 (1/2): 1–76.
Schuster RM (1967d) Studies on antipodal hepaticae. IX. Phyllothalliaceae. Transactions of
the British Bryological Society 5 (2): 283–288. doi: 10.1179/006813867804804296
Schuster RM (1968a) Studies on antipodal hepaticae. X. Subantarctic Scapaniaceae, Balantiopsidaceae and Schistochilaceae. Bulletin of the National Science Museum, Tokyo (n.ser.)
11 (1): 13–31.
Schuster RM (1968b) Studies on the hepaticae, XXIX-XLIV. A miscellany of new taxa and new
range extensions. Nova Hedwigia 15: 437–529.
Schuster RM (1969a) Studies on hepaticae XLVI-XLVII. On Alobiella (Spr.) Schifn. and Alobiellopsis Schust. Bulletin of the National Science Museum, Tokyo (n.ser.) 12 (3): 659–
683.
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Schuster RM (1969b) he hepaticae and anthocerotae of North America. II. Columbia University
Press, New York, 1062 pp.
Schuster RM (1969c) Problems of the antipodal distribution in lower land plants. Taxon 18
(1): 46–91. doi: 10.2307/1218591
Schuster RM (1970a) Studies on antipodal hepaticae, III. Jubulopsis Schuster, Neohattoria
Kamimura and Amphijubula Schuster. Journal of the Hattori Botanical Laboratory 33:
266–304.
Schuster RM (1970b) Studies on hepaticae, XLIX-LIII. New Lejeuneaceae from Dominica and
Jamaica. Bulletin of the Torrey Botanical Club 97 (6): 336–352. doi: 10.2307/2483854
Schuster RM (1971a) he ecology and distribution of hepaticae in a mahogany hammock in
tropical Florida. Castanea 36 (2): 90–111.
Schuster RM (1971b) Studies on Cephaloziellaceae. Nova Hedwigia 22: 121–265.
Schuster RM (1971c) Studies of antipodal Schistochilaceae and Scapaniaceae. Bulletin of the
National Science Museum, Tokyo (n.ser.) 14 (4): 609–660.
Schuster RM (1971d) Two new antipodal species of Haplomitrium (Calobryales). Bryologist
74 (2): 131–143. doi: 10.2307/3241827
Schuster RM (1972) Phylogenetic and taxonomic studies in Jungermanniidae. Journal of the
Hattori Botanical Laboratory 36: 321–405.
Schuster RM (1974) he hepaticae and anthocerotae of North America. III. Columbia University
Press, New York, 880 pp.
Schuster RM (1978a) Studies on Venezuelan hepaticae. I. Phytologia 39 (4): 239–251.
Schuster RM (1978b) Studies on Venezuelan hepaticae. II. Phytologia 39 (6): 425–432.
Schuster RM (1980a) Studies on hepaticae. LIV-LVIII. Kurzia v. Mart. [Microlepidozia (Spr.)
Joerg.], Megalembidium Schust., Psiloclada Mitt., Drucella Hodgs. and Isolembidium
Schust. Journal of the Hattori Botanical Laboratory 48: 337–421.
Schuster RM (1980b) New combinations and taxa of hepaticae I. Phytologia 45 (5): 415–437.
Schuster RM (1980c) he hepaticae and anthocerotae of North America. IV. Columbia University Press, New York, 1334 pp.
Schuster RM (1981a) Austral hepaticae. VIII. Tuyamaelloideae. Phytologia 47 (4): 301–308.
Schuster RM (1981b) Evolution and speciation in Pellia, with special reference to the Pellia
megaspora-endiviifolia complex (Metzgeriales). I. Taxonomy and distribution. Journal of
Bryology 11 (3): 411–431. doi: 10.1179/jbr.1981.11.3.411
Schuster RM (1982) Studies on hepaticae, LIX. On Sandeothallus Schust., gen. n. and the classiication of the Metzgeriales. Nova Hedwigia 36: 1–16.
Schuster RM (1983a) Phytogeography of the Bryophyta. In: Schuster RM (Ed.) New Manual
of Bryology, vol. 1. Hattori Botanical Laboratory, Nichinan, 463–626.
Schuster RM (1983b) Notes on nearctic hepaticae. XVI. New taxa of Frullania from eastern
North America. Phytologia 53 (5): 364–366.
Schuster RM (1984) Diagnoses of some new taxa of hepaticae. Phytologia 56 (2): 65–74.
Schuster RM (1985a) Studies in Porellineae: New taxa of Jubulaceae. Phytologia 57 (5): 369–373.
Schuster RM (1985b) Some new taxa of hepaticae. Phytologia 57 (6): 408–414.
Schuster RM (1985c) Austral hepaticae. XIX. Some taxa new to New Zealand and New Caledonia.
Phytologia 56 (7): 449–464.
World checklist of hornworts and liverworts
787
Schuster RM (1986) Gymnocolea borealis (Frisvoll & Moen) Schust. [Lophozia (Leiocolea) borealis
Frisvoll & Moen] in North America. Lindbergia 12 (1): 5–8.
Schuster RM (1987a) Phylogenetic studies on Jungermanniidae. II. Mastigophoraceae and
Chaetophyllopsidaceae. Memoirs of the New York Botanical Garden 45: 733–748.
Schuster RM (1987b) Preliminary studies on anthocerotae. Phytologia 63 (3): 193–201.
Schuster RM (1987c) On Aureolejeunea Schust. and Brachiolejeunea paramicola Herzog. Phytologia
61 (7): 445–447.
Schuster RM (1987d) Studies on Metzgeriales: I. North American Aneuraceae. Journal of the
Hattori Botanical Laboratory 62: 299–329.
Schuster RM (1989) Studies on the hepatic lora of Prince Edward Islands. I. Aneuraceae.
Journal of the Hattori Botanical Laboratory 67: 59–108.
Schuster RM (1991a) On neotenic species of Radula. Journal of the Hattori Botanical Laboratory
70: 51–62.
Schuster RM (1991b) Diagnoses of new taxa of hepaticae. I. Jungermanniidae. Journal of the
Hattori Botanical Laboratory 70: 143–150.
Schuster RM (1992a) he oil-bodies of the hepaticae. II. Lejeuneaceae (part 2). Journal of the
Hattori Botanical Laboratory 72: 163–359.
Schuster RM (1992b) he hepaticae and anthocerotae of North America. V. Columbia University
Press, New York, 854 pp.
Schuster RM (1992c) Studies on Marchantiales. I-III. Journal of the Hattori Botanical Laboratory
71: 267–287.
Schuster RM (1992d) he hepaticae and anthocerotae of North America. VI. Columbia University Press, New York, 937 pp.
Schuster RM (1993a) Studies on hepaticae, LXII-LXIV. Lepidoziaceae subfamily Zoopsidoideae (1). Nova Hedwigia 56 (1/2): 35–59.
Schuster RM (1993b) On Cephalozia pachycaulis sp. nov. and the perimeters of Cephalozia.
Bryologist 96 (4): 619–625. doi: 10.2307/3243994
Schuster RM (1994) Studies on Lejeuneaceae. I. Preliminary studies on new genera of Lejeuneaceae. Journal of the Hattori Botanical Laboratory 75: 211–235.
Schuster RM (1995a) Phylogenetic and taxonomic studies of Jungermanniidae. III. Calypogeiaceae.
Fragmenta Floristica et Geobotanica 40 (2): 825–888.
Schuster RM (1995b) Studies on Cephaloziellaceae. III. On Cephalomitrion Schust., gen. n.
Nova Hedwigia 61 (3/4): 547–559.
Schuster RM (1995c) he hepaticae of Prince Edward Islands. II. On Gymnocoleopsis (Schust.)
Schust., Lophozia cylindriformis (Mitt.) Steph. and the subgeneric classiication of the genus Lophozia Dumort. Journal of the Hattori Botanical Laboratory 78: 119–135.
Schuster RM (1995d) On a new species of Gymnomitrion, G. mucrophorum Schust., sp. n.
Bryologist 98 (2): 242–245. doi: 10.2307/3243310
Schuster RM (1996a) Studies on antipodal hepaticae. XII. Gymnomitriaceae. Journal of the
Hattori Botanical Laboratory 80: 1–147.
Schuster RM (1996b) On Olgantha Schust., gen. n. Isophylly and evolution of Jungermanniales.
Nova Hedwigia 63 (3/4): 529–543.
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Schuster RM (1996c) Studies on Lejeuneaceae II. Neotropical taxa of Drepanolejeunea (Spr.)
Schifn. Nova Hedwigia 62 (1/2): 1–46.
Schuster RM (1996d) Studies on Cephaloziellaceae. IV. On New Zealand taxa. Nova Hedwigia
63 (1/2): 1–61.
Schuster RM (1997a) On Bragginsella, a new genus of Jungermanniales from New Zealand.
Bryologist 100 (3): 362–367. doi: 10.2307/3244506
Schuster RM (1997b) On a new, microphyllous New Caledonean Acromastigum (Lepidoziaceae).
Nova Hedwigia 64 (3/4): 613–620.
Schuster RM (1998a) On Lejeunea (Papillolejeunea) pocsii Schust., sp. n. of New Zealand.
Journal of the Hattori Botanical Laboratory 85: 83–87.
Schuster RM (1998b) On the genus Scaphophyllum (Jungermanniaceae). Bryologist 101 (3):
428–434. doi: 10.2307/3244182
Schuster RM (1999a) Studies on hepaticae. LXVII-LXVIII. Lepidoziaceae subfamily Zoopsidoideae (4): Monodactylopsis and Pteropsiella. Nova Hedwigia 69 (3/4): 517–540.
Schuster RM (1999b) Studies on hepaticae LXVI. Lepidoziaceae subfamily Zoopsidoideae (3):
Zoopsidella. Nova Hedwigia 69 (1/2): 101–150.
Schuster RM (1999c) Harpalejeunea (Spr.) Schifn. I. Studies on a new Andean species of Harpalejeunea. Journal of the Hattori Botanical Laboratory 87: 287–294.
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Stephani F (1887) Ueber einige Lebermoose Portugals. Hedwigia 26 (1): 1–6.
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Stephani F (1888b) Westindische Hepaticae II. Hepaticae ex insulis St. Domingo et Dominica,
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Stephani F (1899b) Species hepaticarum 1. Bulletin de l’Herbier Boissier 7 (3): 198–225. doi:
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Stephani F (1902d) Species hepaticarum 2. Bulletin de l’Herbier Boissier (sér. 2) 2 (2): 157–179.
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Stephani F (1902g) Species hepaticarum 2. Bulletin de l’Herbier Boissier (sér. 2) 2 (10): 857–888.
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Stephani F (1902h) Species hepaticarum 2. Bulletin de l’Herbier Boissier (sér. 2) 2 (12): 969–987.
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Stephani F (1903a) Species hepaticarum 2. Bulletin de l’Herbier Boissier (sér. 2) 3 (11): 959–974.
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Stephani F (1903b) Species hepaticarum 2. Bulletin de l’Herbier Boissier (sér. 2) 3 (2): 98–129.
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Stephani F (1903c) Species hepaticarum 2. Bulletin de l’Herbier Boissier (sér. 2) 3 (4): 326–341.
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Stephani F (1903d) Species hepaticarum 2. Bulletin de l’Herbier Boissier (sér. 2) 3 (6): 522–537.
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Stephani F (1903e) Species hepaticarum 2. Bulletin de l’Herbier Boissier (sér. 2) 3 (7): 596–611.
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Stephani F (1903f) Species hepaticarum 2. Bulletin de l’Herbier Boissier (sér. 2) 3 (10): 873–888.
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Stephani F (1904a) Species hepaticarum 2. Bulletin de l’Herbier Boissier (sér. 2) 4 (10): 973–988.
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Stephani F (1904b) Species hepaticarum 2. Bulletin de l’Herbier Boissier (sér. 2) 4 (6): 586–601.
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Stephani F (1904c) Species hepaticarum 2. Bulletin de l’Herbier Boissier (sér. 2) 4 (8): 775–790.
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Stephani F (1904e) Species hepaticarum 2. Bulletin de l’Herbier Boissier (sér. 2) 4 (4): 345–360.
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Stephani F (1904f) Species hepaticarum 2. Bulletin de l’Herbier Boissier (sér. 2) 4 (2): 153–168.
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