Fungal Diversity (2020) 105:17–318
https://doi.org/10.1007/s13225-020-00462-6
Refined families of Dothideomycetes: orders and families incertae
sedis in Dothideomycetes
Sinang Hongsanan1,2,6 · Kevin D. Hyde2,3,7 · Rungtiwa Phookamsak2,3,4,5,8,35 · Dhanushka N. Wanasinghe3,4,5,35 ·
Eric H. C. McKenzie9 · V. Venkateswara Sarma10 · Robert Lücking12 · Saranyaphat Boonmee2,11 ·
Jayarama D. Bhat13 · Ning‑Guo Liu2,14,15 · Danushka S. Tennakoon2,11 · Dhandevi Pem2,11 ·
Anuruddha Karunarathna2,3,4,6,50 · Shu‑Hua Jiang17 · Gareth E. B. Jones18,19 · Alan J. L. Phillips20 ·
Ishara S. Manawasinghe2,7,21 · Saowaluck Tibpromma3,4,5,35 · Subashini C. Jayasiri2 · Diana Sandamali2,11,50 ·
Ruvishika S. Jayawardena2,11 · Nalin N. Wijayawardene22 · Anusha H. Ekanayaka2 · Rajesh Jeewon23 ·
Yong‑Zhong Lu24 · Chayanard Phukhamsakda2 · Asha J. Dissanayake16 · Xiang‑Yu Zeng2,25,26 · Zong‑Long Luo27 ·
Qing Tian2 · Kasun M. Thambugala28 · Dongqin Dai22 · Milan C. Samarakoon2,48 · K. W. Thilini Chethana2,11 ·
Damien Ertz29,30 · Mingkwan Doilom3,4,5,35 · Jian‑Kui (Jack) Liu16 · Sergio Pérez‑Ortega31 · Ave Suija32 ·
Chanokned Senwanna2,6 · Subodini N. Wijesinghe2,11,33 · Mekala Niranjan10,36 · Sheng‑Nan Zhang2,6 ·
Hiran A. Ariyawansa34 · Hong‑Bo Jiang2,3,4,11 · Jin‑Feng Zhang2,37 · Chada Norphanphoun2 · Nimali I. de Silva3,48 ·
Vinodhini Thiyagaraja2,3,4,6 · Huang Zhang38 · Jadson D. P. Bezerra39 · Ricardo Miranda‑González40 ·
André Aptroot41 · Hiroyuki Kashiwadani42 · Dulanjalee Harishchandra2,11,21 · Emmanuël Sérusiaux43 ·
Pranami D. Abeywickrama2,11,21 · Dan‑Feng Bao2,6,27 · Bandarupalli Devadatha2,10 · Hai‑Xia Wu44 ·
Kwang Hee Moon45 · Cecile Gueidan46 · Felix Schumm47 · Digvijayini Bundhun2,6 · Ausana Mapook2 ·
Jutamart Monkai2 · Chitrabhanu S. Bhunjun2,11 · Putarak Chomnunti11 · Satinee Suetrong49 · Napalai Chaiwan2,3 ·
Monika C. Dayarathne33 · Jing Yang2,15 · Achala R. Rathnayaka2,11,50 · Jian‑Chu Xu3,4,5,35 · Jiesheng Zheng1 ·
Gang Liu1 · Yao Feng51 · Ning Xie1
Received: 9 June 2020 / Accepted: 10 September 2020 / Published online: 24 December 2020
© The Author(s) 2020
Abstract
Numerous new taxa and classifications of Dothideomycetes have been published following the last monograph of families of
Dothideomycetes in 2013. A recent publication by Honsanan et al. in 2020 expanded information of families in Dothideomycetidae and Pleosporomycetidae with modern classifications. In this paper, we provide a refined updated document on
orders and families incertae sedis of Dothideomycetes. Each family is provided with an updated description, notes, including
figures to represent the morphology, a list of accepted genera, and economic and ecological significances. We also provide
phylogenetic trees for each order. In this study, 31 orders which consist 50 families are assigned as orders incertae sedis in
Dothideomycetes, and 41 families are treated as families incertae sedis due to lack of molecular or morphological evidence.
The new order, Catinellales, and four new families, Catinellaceae, Morenoinaceae Neobuelliellaceae and Thyrinulaceae
are introduced. Seven genera (Neobuelliella, Pseudomicrothyrium, Flagellostrigula, Swinscowia, Macroconstrictolumina,
Pseudobogoriella, and Schummia) are introduced. Seven new species (Acrospermum urticae, Bogoriella complexoluminata,
Dothiorella ostryae, Dyfrolomyces distoseptatus, Macroconstrictolumina megalateralis, Patellaria microspora, and Pseudomicrothyrium thailandicum) are introduced base on morphology and phylogeny, together with two new records/reports
and five new collections from different families. Ninety new combinations are also provided in this paper.
Electronic supplementary material The online version of this
article (https://doi.org/10.1007/s13225-020-00462-6) contains
supplementary material, which is available to authorized users.
Extended author information available on the last page of the article
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Fungal Diversity (2020) 105:17–318
Keywords Abrothallales · Ascrospermales · Asterinales · Botryosphaeriales · Catinellales · Cladoriellales ·
Collemopsidiales · Dyfrolomycetales · Eremithallales · Eremomycetales · Jahnulales · Kirschsteiniotheliales ·
Lembosinales · Lichenotheliales · Microthyriales · Minutisphaerales · Monoblastiales · Murramarangomycetales ·
new family · new genera · new order · new species · Parmulariales · Patellariales · Phaeotrichales · Phylogeny ·
Stigmatodiscales · Strigulales · Superstratomycetales · Taxonomy · Trypetheliales · Tubeufiales · Valsariales · Venturiales ·
Zeloasperisporiales
Table of contents and contributors
Abrothallales
Lichenoconiaceae (S Pérez-Ortega, A Suija, S Hongsanan, R Jeewon)
Ascospermales
Ascospermaceae (D Pem, D Bundhun, S Hongsanan, KD
Hyde)
Asterinales
Asterinaceae (XY Zeng, S Hongsanan, KD Hyde, R
Jeewon)
Asterotexaceae (S Hongsanan, KD Hyde, EHC
McKenzie)
Hemigraphaceae (DQ Dai, S Hongsanan, EHC
McKenzie)
Lembosiaceae (D Sandamali, KD Hyde, S Hongsanan,
EHC McKenzie)
Melaspileellaceae (S Hongsanan, Q Tian, KD Hyde, R
Jeewon)
Morenoinaceae (S Hongsanan, KD Hyde, EHC McKenzie)
Neobuelliellaceae (S Hongsanan, KD Hyde, EHC
McKenzie)
Stictographaceae (DQ Dai, S Hongsanan, R Jeewon)
Botryosphaeriales
Aplosporellaceae (AH Ekanayaka, AJL Phillips, KD
Hyde)
Botryosphaeriaceae (IS Manawasinghe, AJ Dissanaya,
A Phillips)
Melanopsaceae (S Hongsanan, AJL Phillips, KD Hyde)
Phyllostictaceae (AJ Dissanaya, P Abeywickrama, AJL
Phillips, S Hongsanan, KD Hyde)
Planistromellaceae (J Monkai, AJL Phillips, KD Hyde)
Saccharataceae (AJ Dissanaya, AJL Phillips, S Hongsanan KD Hyde)
Catinellales
Catinellaceae (AH Ekanayaka, HA Ariyawansa, JS
Zheng, KD Hyde)
Cladoriellales
Cladoriellaceae (S Hongsanan, DJ Bhat, EHC McKenzie,
KD Hyde)
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Collemopsidiales
Xanthopyreniaceae (S Pérez-Ortega, A Suija, S Hongsanan, EHC McKenzie)
Dyfrolomycetales
Pleurotremataceae (JF Zhang, VV Sarma, S Hongsanan,
EBG Jones, KD Hyde)
Eremithallales
Melaspileaceae (V Thiyagaraja, D Ertz, S Hongsanan,
KD Hyde)
Eremomycetales
Eremomycetaceae (D Pem, S Boonmee, KD Hyde, S
Hongsanan, EHC McKenzie)
Jahnulales
Aliquandostipitaceae (ZL Luo, DF Bao, EBG Jones, KD
Hyde)
Manglicolaceae (S Suetrong, S Preedanon, EBG Jones,
KD Hyde)
Kirschsteiniotheliales
Kirschsteiniotheliaceae (ZL Luo, DF Bao, R Jeewon, KD
Hyde)
Lembosinales
Lembosiaceae (S Hongsanan, KD Hyde, EHC McKenzie)
Lichenotheliales
Lichenotheliaceae (V Thiyagaraja, S Hongsanan, R Jeewon, KD Hyde)
Microthyriales
Microthyriaceae (XY Zeng, S Hongsanan, DJ Bhat, KD
Hyde, EHC McKenzie)
Minutisphaerales
Agrogenosporaceae (J Yang, DJ Bhat, EHC McKenzie,
KD Hyde)
Minutisphaeraceae (KM Thambugala, HA Raja, DJ Bhat,
KD Hyde)
Fungal Diversity (2020) 105:17–318
19
Monoblastiales
Monoblastiaceae (R Lücking, A Aptroot, H Kashiwadani,
KH Moon, C Gueidan, F Schumm, R Phookamsak)
Valsariales
Valsariaceae (C Senwanna, D Pem, S Hongsanan, EHC
McKenzie, KD Hyde)
Murramarangomycetales
Murramarangomycetaceae (S Hongsanan, DJ Bhat, KD
Hyde, EHC McKenzie)
Venturiales
Sympoventuriaceae (D Sandamali, KD Hyde, R Jeewon)
Venturiaceae (D Sandamali, KD Hyde, S Hongsanan, R
Jeewon)
Muyocopronales
Muyocopronaceae (N Chaiwan, S Hongsanan, C Senwanna, KD Hyde)
Natipusillales
Natipusillaceae (S Hongsanan, ZL Luo, KD Hyde, EHC
McKenzie)
Parmulariales
Parmulariaceae (DQ Dai, S Hongsanan, R Jeewon, KD
Hyde)
Patellariales
Patellariaceae (AH Ekanayaka, S Hongsanan, R Jeewon,
KD Hyde)
Phaeotrichales
Phaeotrichaceae (D Sandaruwan, JS Zheng, KD Hyde, S
Hongsanan, EHC McKenzie)
Stigmatodiscales
Stigmatodiscaceae (S Hongsanan, A Rathnayaka, EHC
McKenzie, KD Hyde)
Strigulales
Strigulaceae (SH Jiang, R Lücking, E Sérusiaux)
Tenuitholiascaceae (SH Jiang, R Lücking)
Superstratomycetales
Superstratomycetaceae (P Abeywickrama, S Hongsanan)
Trypetheliales
Polycoccaceae (S Hongsanan, D Ertz, EHC McKenzie)
Trypetheliaceae (R Lücking, SN Zhang, R MirandaGonzález, A Aptroot)
Tubeufiales
Bezerromycetaceae (YZ Lu, J Bezerra, EHC McKenzie,
KD Hyde)
Tubeufiaceae (YZ Lu, J Bezerra, EHC McKenzie, KD
Hyde)
Wiesneriomycetaceae (C Senwanna, J Bezerra, EHC
McKenzie)
Zeloasperisporiales
Zeloasperisporiaceae (S Hongsanan, DJ Bhat R Jeewon,
KD Hyde)
Dothideomycetes, families incertae sedis
Alinaceae (S Hongsanan, S Boonmee, EHC McKenzie)
Argynnaceae (S Boonmee, S Hongsanan, S Konta)
Ascoporiaceae (S Boonmee, EHC McKenzie)
Aulographaceae (D Sandamali, S Hongsanan)
Balladynaceae (S Hongsanan, D Pem, S Boonmee)
Cleistosphaeraceae (S Hongsanan, D Pem, S Boonmee)
Coccoideaceae (S Boonmee, EHC McKenzie, KD Hyde)
Cookellaceae (S Boonmee, EHC McKenzie, KD Hyde)
Dimeriaceae (S Boonmee, KD Hyde, EHC McKenzie,
S Konta)
Dubujianaceae (D Pem, S Hongsanan, KD Hyde)
Dysrhynchisceae (S Boonmee, KD Hyde, S Konta)
Endosporiaceae (D Pem, KD Hyde)
Englerulaceae (HX Wu, S Hongsanan, EHC McKenzie,
KD Hyde)
Homortomycetaceae (KM Thambugala, EHC McKenzie,
KD Hyde)
Hyalomeliolinaceae (S Boonmee, S Konta)
Leptopeltidaceae (R Phookamsak, EHC McKenzie, KD
Hyde)
Macrovalsariaceae (D Pem, KD Hyde)
Meliolinaceae (XY Zeng, S Hongsanan, EHC McKenzie
Mesnieraceae (P Chomnunti, S Hongsanan, S Boonmee)
Naetrocymbaceae (S Hongsanan, DQ Dai, KD Hyde)
Nematotheciaceae (S Hongsanan, S Boonmee, KD Hyde)
Neoparodiaceae (S Hongsanan, S Boonmee, KD Hyde)
Palawaniaceae (A Mapook, S Hongsanan, EHC
McKenzie)
Paranectriellaceae (S Boonmee, EHC McKenzie, KD
Hyde)
Parodiellaceae (S Hongsanan, P Chomnunti, KD Hyde)
Perisporiopsidaceae (S Boonmee, EHC McKenzie, KD
Hyde)
Phaeodimeriellaceae (S Hongsanan, S Boonmee, KD
Hyde)
Pododimeriaceae (S Hongsanan, S Boonmee KD Hyde)
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Pododimeriellaceae (S Hongsanan, D Sandamali, S
Boonmee, KD Hyde)
Polyclypeolinaceae (S Hongsanan, S Boonmee, KD Hyde)
Polystomellaceae (D Pem, S Boonmee, KD Hyde)
Protoscyphaceae (S Boonmee, EHC McKenzie, KD
Hyde)
Pseudoperisporiaceae (S Boonmee, EHC McKenzie, KD
Hyde)
Pseudorobillardaceae (S Tibpromma, WJ Li, S Hongsanan, EHC McKenzie)
Pyrenidiaceae (S Hongsanan, N Huanraluek, D Ertz, KD
Hyde)
Seynesiopeltidaceae (D Pem, S Boonmee, EHC McKenzie, KD Hyde)
Stomatogeneceae (S Hongsanan, S Boonmee, EHC
McKenzie, KD Hyde)
Toroaceae (S Hongsanan, S Boonmee, KD Hyde, EHC
McKenzie)
Thyrinulaceae (XY Zeng, S Hongsanan, KD Hyde, EHC
McKenzie)
Trichopeltinaceae (D Sandamali, S Hongsanan, KD
Hyde)
Trichothyriaceae (XY Zeng, HX Wu, KD Hyde)
Vizellaceae (HX Wu, S Hongsanan, S Boonmee, KD Hyde)
Introduction
Dothideomycetes were previously known as Loculoascomycetes, and is the largest class of ascomycetes (Nannfeldt
1932; Luttrell 1955; Janex-Favre 1971; Barr 1979a, b; Eriksson 1981; Reynolds 1971; Tehler 1990, 1995; LetrouitGalinou et al. 1994; Barr and Huhndorf 2001; Liu and Hall
2004; Hyde et al. 2013; Hongsanan et al. 2020). Members
of this class mostly have bitunicate, fissitunicate asci (Nannfeldt 1932; Luttrell 1955; Eriksson 1981; Barr and Huhndorf 2001; Hyde et al. 2013; Hongsanan et al. 2020). Hyde
et al. (2013) provided descriptions, notes and discussed
taxonomic placement of families in Dothideomycetes. They
also confirmed that Dothideomycetes comprises two subclasses viz. Dothideomycetidae and Pleosporomycetidae,
while other uncertain orders and families were treated as
incertae sedis in Dothideomycetes. Based on morphology
and phylogeny, Dothideomycetidae comprised Capnodiales, Dothideales, and Myrangiales, and Pleosporomycetidae comprised Gloniales, Hysteriales, Mytilinidiales, and
Pleosporales (Hyde et al. 2013, Hongsanan et al. 2020).
Hongsanan et al. (2020) provided a monograph of families
in Dothideomycetidae and Pleosporomycetidae based on
‘Families of Dothideomycetes’ (Hyde et al. 2013) which was
the latest update and the most complete monograph of families in Dothideomycetes, included in Wijayawardene et al.
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Fungal Diversity (2020) 105:17–318
(2017a) who provided ‘Notes for genera: Ascomycota’ and
Wijayawardene et al. (2020) who provided ‘Outline of Fungi
and fungus-like taxa’. Hongsanan et al. (2020) provided an
updated description, notes, including figures to represent
the morphology, list of accepted genera, and economic and
ecological significance of families in Dothideomycetidae
and Pleosporomycetidae. They accepted three orders with
25 families in Dothideomycetidae and four orders with 94
families in Pleosporomycetidae (Hongsanan et al. 2020).
Liu et al. (2017) established guidelines of using divergence
estimates in the classification of Dothideomycetes, and this
was followed by Hongsanan et al. (2020) who provided an
updated divergence estimation of families in Dothideoycetes
using two representative strains from each family. In our
paper, we used the divergence time estimation from Hongsanan et al. (2020) to discuss orderal and familial status of
incertae sedis groups.
The aim of this paper is to bring together data of orders
and families which were not assigned to Dothideomycetidae nor Pleosporomycetidae due to their uncertain phylogenetic placement or lack of sequence data. We included all
published data that we could locate in 2019 and have also
included some data that we were aware of up to the date of
publication. This work refined account of the families of
Dothideomycetes and will provide a baseline for research on
Dothideomycetes over the next decade. As an online database, dothideomycetes.org (Pem et al. 2019b) and fungalgenera.org (Monkai et al. 2019), the contents will be be continually refined and updated and stay abreast of classification
changes as new data and additional evidence are published.
Materials and methods
Layout of the paper
Each family which is currently not assigned to Dothideomycetidae and Pleosporomycetidae is introduced with descriptions and important notes. Accepted genera including the
type species, and notes are provided with basionyms (other
synonyms can be found in Index Fungorum 2020). Faces
of Fungi and Index Fungorum numbers are provided as in
Jayasiri et al. (2015) and Index Fungorum (2020). Estimated
numbers of species for each genus is provided mainly based
on data in Species Fungorum (2020), with species transferred to other genera or species not being included. The
number of species with sequence data available in GenBank
is provided. The morphology of each family is illustrated
by representative photographic plates from type herbarium
specimens, or new species/new collections with sequence
data. We provide drawing for families which we were unable
to obtain herbarium specimens or fresh collections. Notes on
the ecological and economic significance are also provided.
Fungal Diversity (2020) 105:17–318
Molecular phylogeny
Phylogenetic trees of each order are provided, except for
orders where too little sequence data is available. Trees for
each order include representatives from all genera that have
sequence data (Supplemental material 1). The methods
below were applied for most of the phylogenetic trees shown
in this paper. Other additional methods are mentioned in the
legend of each analysis.
Gene regions used in each analysis were selected based
on the most recent study or the availability of sequence data.
Datasets for each partition were aligned using MAFFT (Katoh
et al. 2019), and adjusted manually using Bioedit (Hall 2004).
All missing genes were coded as missing data. Phylogenetic
trees based on individual gene partitions (data not shown)
were congruent with the combined data sets. The phylogenetic
analyses of the combined dataset were performed using maximum likelihood algorithms in RAxML and Bayesian analyses.
Maximum likelihood analysis (ML) was carried out by
using raxmlGUIv.0.9b2 (Silvestro and Michalak 2012). The
search strategy was set to bootstrapping and the analysis performed using the GTRGAMMAI model. The number of replicates was inferred using the stopping criterion (Pattengale
et al. 2009). The bootstrap values expressed from 1,000 repetitions by RAxML analysis which are equal or greater than
70% are given on each node. The best fit model of evolution
was selected by MrModeltest 2.2 (Nylander et al. 2008).
Bayesian phylogenetic analyses, posterior probabilities (PP)
were set by MCMC sampling in MrBayes v3.1.2 (Huelsenbeck and Ronquist 2001; Zhaxybayeva and Gogarten 2002),
following the details in Cai et al. (2008). The first 20% were
discarded, and the remaining trees were used for calculating
posterior probabilities. Posterior probabilities values (PP)
from Bayesian analysis which are equal or greater than 0.90
are given on each node. Phylogenetic tree was viewed in
FigTree v.1.4.0 (Rambaut 2014).
Order of Dothideomycetes
For the subclasses Dothideomycetidae and Pleosporomycetidae see Hongsanan et al. (2020) and Dothideomycetes
genera incertae sedis see Wijayawardene et al. (2020).
Abrothallales Pérez-Ort. & Suija
Lichenoconiaceae Diederich & Lawrey (= Abrothallaceae
Pérez-Ort. & Suija) (57)
Abrothallus De Not.
Lichenoconium Petr. & Syd.
Acrospermales Minter et al.
Acrospermaceae Fuckel
Acrospermum Tode
Gonatophragmium Deighton
Oomyces Berk. & Broome
Pseudovirgaria H.D. Shin et al.
21
Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss. (=
Asterotexales Firmino et al.)
Asterinaceae Hansf.
Asterina Lév.
Asterinella Theiss.
Asterolibertia G. Arnaud
Asterostomella Speg.
Batistinula Arx
Cirsosia G. Arnaud
Dothidasteromella Höhn.
Echidnodella Theiss. & Syd.
Halbania Racib.
Meliolaster Höhn.
Parasterinopsis Bat.
Platypeltella Petr.
Prillieuxina G. Arnaud
Pycnocarpon Theiss.
Schenckiella Henn.
Trichasterina G. Arnaud
Trichopeltospora Bat. & Cif.
Uleothyrium Petr.
Vizellopsis Bat. et al.
Asterotexaceae Firmino et al.
Asterotexis Arx
Hemigraphaceae D.Q. Dai & K.D. Hyde
Hemigrapha (Müll. Arg.) D. Hawksw.
Lembosiaceae Luttrell ex P.M. Kirk et al.
Lembosia Lév.
Melaspileellaceae D.Q. Dai & K.D. Hyde
Melaspileella (P. Karst.) Vain.
Morenoinaceae Hongsanan & K.D. Hyde
Morenoina Theiss.
Neobuelliellaceae Hongsanan & K.D. Hyde
Neobuelliella Hongsanan & K.D. Hyde
Stictographaceae D.Q. Dai & K.D. Hyde
Buelliella Fink
Karschia Körb.
Labrocarpon Etayo & Pérez-Ort.
Melaspileopsis (Müll. Arg.) Ertz & Diederich
Stictographa Mudd
Asterinales genera incertae sedis
Andamanomyces Hosag.
Caribaeomyces Cif.
Caudella Syd. & P. Syd.
Discopycnothyrium Hongsanan & K.D. Hyde
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Fungal Diversity (2020) 105:17–318
Hazslinszkya Körb.
Inocyclus Theiss. & Syd.
Melanographa Müll. Arg.
Pirozynskiella S. Hughes
Vishnumyces Hosag.
Botryosphaeriales C.L. Schoch et al.
Aplosporellaceae Slippers et al.
Alanomyces Sharma
Aplosporella Speg.
Botryosphaeriaceae Theiss. & H. Syd.
Alanphillipsia Crous & M.J. Wingf.
Barriopsis A.J.L. Phillips et al.
Botryobambusa Phook. et al.
Botryosphaeria Ces. & De Not.
Cophinforma Doilom et al.
Diplodia Fr.
Dothiorella Sacc.
Endomelanconiopsis Rojas & Samuels
Eutiarosporella Crous
Lasiodiplodia Ellis & Everh.
Macrophomina Petr.
Marasasiomyces Crous
Mucoharknessia Crous et al.
Neodeightonia C. Booth
Neofusicoccum Crous et al.
Neoscytalidium Crous & Slippers
Oblongocollomyces Tao Yang & Crous
Phaeobotryon Theiss. & Syd.
Sakireeta Subram. & K. Ramakr.
Sardiniella Linaldeddu et al.
Sphaeropsis Sacc.
Tiarosporella Höhn.
Melanopsaceae Phillips et al.
Melanops Nitschke ex Fuckel
Phyllostictaceae Fr.
Phyllosticta Pers.
Pseudofusicoccum Mohali et al.
Planistromellaceae M.E. Barr
Kellermania Ellis & Everh.
Mycosphaerellopsis Höhn.
Planistroma A.W. Ramaley
Umthunziomyces Crous & M.J. Wingf.
Saccharataceae Slippers et al.
Pileospora Tanney & Seifert
Saccharata Denman & Crous (= Neoseptorioides Crous
et al.)
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Septorioides Quaedvl. et al.
Botryosphaeriales genera incertae sedis
Auerswaldiella Theiss. & Syd.
Coccostromella Petr.
Leptoguignardia E. Müll.
Metameris Theiss. & Syd.
Phyllachorella Syd.
Pilgeriella Henn.
Sivanesania W.H. Hsieh & C.Y. Chen
Vestergrenia Rehm
Catinellales Ekanayaka et al.
Catinellaceae Ekanayaka et al.
Catinella Boud.
Cladoriellales Crous
Cladoriellaceae Crous
Cladoriella Crous
Collemopsidiales Perez-Ortega et al.
Xanthopyreniaceae Zahlbr.
Collemopsidium Nyl.
Didymellopsis (Sacc.) Clem. & Shear
Frigidopyrenia Grube
Rhagadodidymellopsis Fdez.-Brime et al.
Xanthopyrenia Bachm.
Zwackhiomyces Grube & Hafellner
Zwackhiomacromyces Etayo & van den Boom
Dyfrolomycetales K.L. Pang et al.
Pleurotremataceae Walt. Watson (= Dyfrolomycetaceae
K.D. Hyde et al.)
Dyfrolomyces K.D. Hyde et al.
Melomastia Nitschke ex Sacc.
Pleurotrema Müll. Arg.
Eremithallales Lücking & Lumbsch
Melaspileaceae W. Watson (= Eremithallaceae Lücking &
Lumbsch)
Encephalographa A. Massal.
Melaspilea Nyl.
Eremomycetales Crous et al.
Eremomycetaceae Malloch & Cain
Eremomyces Malloch & Cain
Rhexothecium Samson & Mouch.
Eremomycetales genus incertae sedis
Arthrographis G. Cochet ex Sigler & J.W. Carmich.
Fungal Diversity (2020) 105:17–318
Jahnulales Pang et al.
Aliquandostipitaceae Inderbitzin
Aliquandostipite Inderbitzin
Brachiosphaera Nawawi
Jahnula Kirschst.
Megalohypha A. Ferrer & Shearer
Neojahnula W. Dong et al.
Pseudojahnula W. Dong et al.
Xylomyces Goos et al.
Manglicolaceae Suetrong & E.B.G. Jones
Manglicola Kohlm. & E. Kohlm.
Kirschsteiniotheliales Hern.-Restr. et al.
Kirschsteiniotheliaceae Boonmee & K.D. Hyde
Kirschsteiniothelia D. Hawksw.
Kirschsteiniotheliales genus incertae sedis
Brachysporiella Bat. (Brachysporiella sensu lato)
Taeniolella S. Hughes sensu lato
Lembosinales Crous
Lembosinaceae Crous
Lembosina Theiss.
Lichenotheliales K. Knudsen et al.
Lichenotheliaceae Henssen
Lichenothelia D. Hawksw.
Endococcus Nyl.
Microthyriales G. Arnaud
Microthyriaceae Sacc.
Arnaudiella Petr.
Calothyriopsis Höhn.
Chaetothyriothecium Hongsanan & K.D. Hyde
Hamatispora L.T.H. Yen et al.
Microthyrium Desm.
Neoanungitea Crous
Paramicrothyrium H.X. Wu & K.D. Hyde
Pseudomicrothyrium X.Y. Zeng et al.
Pseudopenidiella Crous & Koukol
Seynesiella G. Arnaud
Tumidispora Hongsanan & K.D. Hyde
Microthyriales genera incertae sedis
Heliocephala V. Rao et al.
Mitopeltis Speg.
Neoscolecobasidium Crous
Parazalerion Madrid et al.
Thyriodictyella Cif.
Tothia Bat.
23
Minutisphaerales Raja et al.
Acrogenosporaceae Jayasiri & K.D. Hyde.
Acrogenospora M.B. Ellis
Minutisphaeraceae Raja et al
Minutisphaera Shearer et al.
Monoblastiales Lücking et al.
Monoblastiaceae Walt. Watson (= Eriomycetaceae Huanraluek & K.D. Hyde)
Acrocordia A. Massal.
Anisomeridium (Müll. Arg.) M. Choisy
Caprettia Bat. & H. Maia
Eriomyces Huanraluek, Thambugala & K.D. Hyde
Funbolia Crous & Seifert
Heleiosa Kohlm. et al.
Megalotremis Aptroot
Monoblastia Riddle
Phellinocrescentia Crous & Decock
Pseudopassalora Crous
Trypetheliopsis Asahina
Murramarangomycetales Crous
Murramarangomycetaceae Crous
Phaeothyriolum Syd.
Muyocopronales Mapook et al.
Muyocopronaceae K.D. Hyde
Arxiella Papendorf
Leptodiscella Papendorf
Muyocopron Speg.
Mycoleptodiscus Ostaz.
Neocochlearomyces Pinruan et al.
Neomycoleptodiscus Hern.-Restr. et al.
Paramycoleptodiscus Crous & M.J. Wingf.
Pseudopalawania Mapook & K.D. Hyde
Setoapiospora Mapook & K.D. Hyde
Natipusillales Raja et al.
Natipusillaceae Raja et al.
Natipusilla A. Ferrer
Parmulariales D.Q. Dai & K.D. Hyde
Parmulariaceae E. Müll. & Arx ex M.E. Barr
Aldona Racib.
Aldonata Sivan. & A.R.P. Sinha
Antoniomyces Inácio
Aulacostroma Syd. & P. Syd.
Campoa Speg.
Cirsosiopsis Butin & Speer
13
24
Fungal Diversity (2020) 105:17–318
Cocconia Sacc.
Cycloschizon P. Henn.
Cyclostomella Pat.
Dothidasteroma Höhn.
Ferrarisia Sacc.
Hysterostomella Speg.
Kiehlia Viégas
Mintera Inácio & P.F. Cannon
Pachypatella Theiss. & Syd.
Palawaniella Doidge
Parmularia Lév.
Parmulariopsella Sivan.
Parmulariopsis Petr.
Parmulina Theiss. & Syd.
Placoasterella Sacc. ex Theiss. & Syd
Placosoma Syd.
Placostromella Petr.
Pleiostomellina Bat. et al.
Polycyclina Theiss. & Syd.
Polycyclus Höhn.
Protothyrium G. Arnaud
Pseudolembosia Theiss.
Rhagadolobiopsis Guatim. & R.W. Barreto
Rhagadolobium P. Henn. & Lindau
Rhipidocarpon (Theiss.) Theiss. & Syd.
Symphaeophyma Speg.
Syrropeltis Bat. et al.
Thallomyces H.J. Swart
Viegasiella Inácio & P.F. Cannon
Patellariales D. Hawksw. & O.E. Erikss.
Patellariaceae Corda
Baggea Auersw.
Banhegyia L. Zeller & Tóth
Colensoniella Hafellner
Endotryblidium Petr.
Glyphium Nitschke ex F. Lehm.
Haematomyxa Sacc
Holmiella Petrini et al.
Hysteropatella Rehm
Hysteropeltella Petr.
Lahmiomyces Cif. & Tomas.
Lecanidiella Sherwood
Lirellodisca Aptroot
Murangium Seaver
Patellaria Fr.
Poetschia Körb.
Pseudoparodia Theiss. & Syd.
Rhizodiscina Hafellner
Rimula Velen.
Schrakia Hafellner
Stratisporella Hafellner
Tryblidaria (Sacc.) Sacc.
13
Phaeotrichales Ariyaw. et al.
Phaeotrichaceae Cain
Echinoascotheca Matsush.
Phaeotrichum Cain & M.E. Barr
Trichodelitschia Munk
Stigmatodiscales Voglmayr & Jaklitsch
Stigmatodiscaceae Voglmayr & Jaklitsch
Stigmatodiscus Voglmayr & Jaklitsch
Strigulales Lücking et al.
Strigulaceae Zahlbr. (= Phyllobatheliaceae Bitter & F.
Schill.)
Dichoporis Clem.
Flagellostrigula Lücking et al.
Flavobathelium Lücking et al.
Phyllobathelium (Müll. Arg.) Müll. Arg
Phyllocharis Fée,
Phyllocraterina Sérus. & Aptroot
Phylloporis Clem.
Puiggariella Speg.
Raciborskiella Höhnel
Racoplaca Fée
Serusiauxiella S.H. Jiang et al.
Strigula Fr.
Swinscowia S.H. Jiang et al.
Tenuitholiascaceae S.H. Jiang et al.
Tenuitholiascus S.H. Jiang et al.
Superstratomycetales van Nieuwenhuijzen et al.
Superstratomycetaceae van Nieuwenhuijzen et al.
Superstratomyces van Nieuwenhuijzen et al.
Trypetheliales Lücking et al.
Polycoccaceae Ertz et al.
Clypeococcum D. Hawksw.
Polycoccum Saut. ex Körb.
Trypetheliaceae Zenker
Alloarthopyrenia Phukhams. et al.
Aptrootia Lücking & Sipman
Architrypethelium Aptroot
Astrothelium Eschw.
Bathelium Ach.
Bogoriella Zahlbr.
Constrictolumina Lücking et al.
Dictyomeridium Aptroot et al.
Macroconstrictolumina Lücking et al.
Marcelaria Aptroot
Nigrovothelium Lücking et al.
Polymeridium (Müll. Arg.) R.C. Harris
Polypyrenula D. Hawksw.
Fungal Diversity (2020) 105:17–318
Pseudobogoriella Lücking et al.
Pseudopyrenula Müll. Arg.
Schummia Lücking et al.
Trypethelium Sprengel
Viridothelium Lücking et al.
Tubeufiales Boonmee & K.D. Hyde
Bezerromycetaceae J.D.P. Bezerra et al.
Bezerromyces J.D.P. Bezerra et al.
Neorhamphoria Boonmee et al.
Xiliomyces J.D.P. Bezerra et al.
Tubeufiaceae M.E. Barr
Acanthohelicospora Boonmee & K.D. Hyde
Acanthophiobolus Berl.
Acanthostigma De Not.
Acanthostigmina Höhn.
Acanthotubeufia Y.Z. Lu & K.D. Hyde
Aquaphila Goh et al.
Artocarpomyces Subram.
Berkleasmium Zobel
Bifrontia Norman
Boerlagiomyces Butzin
Camporesiomyces D.P. Wei & K.D. Hyde
Chaetosphaerulina I. Hino
Chlamydotubeufia Boonmee & K.D. Hyde
Dematiohelicoma Y.Z. Lu et al.
Dematiohelicomyces Y.Z. Lu et al.
Dematiohelicosporum Y.Z. Lu et al.
Dematiotubeufia Y.Z. Lu et al.
Dictyospora Brahamanage et al.
Discotubeufia Jayasir et al.
Helicangiospora Boonmee et al.
Helicoarctatus Y.Z. Lu et al.
Helicodochium J.S. Monteiro et al.
Helicohyalinum Y.Z. Lu et al.
Helicoma Corda
Helicomyces Link
Helicosporium Nees
Helicotruncatum Y.Z. Lu et al.
Helicotubeufia Y.Z. Lu & J.K. Liu
Kamalomyces R.K. Verma et al.
Kevinhydea N.G. Liu et al.
Manoharachariella Bagyan. et al.
Muripulchra Z.L. Luo et al.
Neoacanthostigma Boonmee et al.
Neochlamydotubeufia Y.Z. Lu et al.
Neohelicoma Y.Z. Lu et al.
Neohelicomyces Z.L. Luo et al.
Neohelicosporium Y.Z. Lu et al.
Neotubeufia Chaiwan et al.
Pleurohelicosporium Y.Z. Lu et al.
Podonectria Petch
25
Pseudohelicomyces Y.Z. Lu et al.
Pseudohelicoon Y.Z. Lu & K.D. Hyde
Tamhinispora Rajeshkumar & Rahul Sharma
Thaxteriella Petr.
Thaxteriellopsis Sivan. et al.
Tubeufia Penz. & Sacc.
Wiesneriomycetaceae Suetrong et al.
Parawiesneriomyces Crous & M.J. Wingf.
Phalangispora Nawawi & J. Webster
Pseudogliophragma Phadke & V.G. Rao
Setosynnema D.E. Shaw & B. Sutton
Speiropsis Tubaki
Wiesneriomyces Koord.
Valsariales Jaklitsch et al.
Valsariaceae Jaklitsch et al.
Bambusaria Jaklitsch et al.
Myrmaecium Nitschke ex Fuckel
Valsaria Ces. & De Not.
Venturiales Y. Zhang ter et al.
Sympoventuriaceae Y. Zhang ter et al.
Acroconidiellina M.B. Ellis
Clavatispora Boonmee & K.D. Hyde
Echinocatena R. Campb. & B. Sutton
Fusicladium Bonord.
Matsushimaea Subram.
Mycosisymbrium Carris
Ochroconis de Hoog & Arx
Scolecobasidiella M.B. Ellis
Scolecobasidium E.V. Abbott
Sympoventuria Crous & Seifert
Veronaeopsis Arzanlou & Crous
Verruconis Samerp. et al.
Yunnanomyces Tibpromma & K.D. Hyde
Venturiaceae E. Müll. & Arx ex M.E. Barr
Apiosporina Höhn.
Atopospora Petr.
Caproventuria U. Braun
Coleroa (Fr.) Rabenh.
Cylindrosympodium W.B. Kendr. & R.F. Castañeda
Dimeriella Speg.
Dimerosporiopsis Henn.
Magnohelicospora R.F. Castañeda et al.
Metacoleroa Petr.
Neocoleroa Petr.
Protoventuria Berl. & Sacc.
Pseudoanungitea Crous
Pseudoparodiella F. Stevens
Sympodiella W.B. Kendr.
Tyrannosorus Unter. & Malloch
13
26
Fungal Diversity (2020) 105:17–318
Venturia Sacc.
Venturiales genus incertae sedis
Cylindrosympodioides Crous & M.J. Wingf.
Lasiobotrys Kunze
Zeloasperisporiales Hongsanan & K.D. Hyde
Zeloasperisporiaceae Crous
Zeloasperisporium R.F. Castañeda
Dothideomycetes families incertae sedis
Alinaceae Boonmee & K.D. Hyde
Alina Racib.
Argynnaceae Shearer & J.L. Crane
Argynna Morgan
Lepidopterella Shearer & J.L. Crane
Ascoporiaceae Kutorga & D. Hawksw.
Ascoporia Samuels & A.I. Romero
Aulographaceae Luttr. ex P.M. Kirk et al.
Aulographum Lib.
Echidnodes Theiss. & Syd.
Lembosiella Sacc.
Thyriopsis Theiss. & Syd.
Balladynaceae Boonmee & K.D. Hyde
Balladyna Racib.
Balladynocallia Bat.
Balladynopsis Theiss. & Syd.
Cleistosphaeraceae Boonmee & K.D. Hyde
Cleistosphaera Syd. & P. Syd.
Coccoideaceae P. Henn. ex Sacc. & D. Sacc.
Coccoidea P. Henn.
Coccoidella Höhn.
Englerodothis Theiss. & Syd.
Cookellaceae Höhn. ex Saccardo & Trotter
Cookella Sacc.
Pycnoderma Syd. & P. Syd.
Dimeriaceae E. Müll. & Arx ex Arx & E. Müll.
Dimerium (Sacc. & P. Syd.) McAlpine
Dubujianaceae D. Pem et al.
Dubujiana D.R. Reynolds & G.S. Gilbert
Dysrhynchisceae Boonmee & K.D. Hyde
Dysrhynchis Clem.
13
Endosporiaceae D. Pem
Endosporium Tsuneda
Englerulaceae P. Henn.
Allosoma Syd.
Digitosarcinella S. Hughes
Englerula P. Henn.
Goosia B. Song
Parenglerula Höhn.
Rhytidenglerula Höhn.
Schiffnerula Höhn.
Thrauste Theiss.
Homortomycetaceae Thambug. et al.
Homortomyces Crous & M.J. Wingf.
Hyalomeliolinaceae Boonmee & K.D. Hyde
Hyalomeliolina F. Stevens
Leptopeltidaceae Höhn. ex Trotter
Dothiopeltis E. Müll.
Leptopeltis Höhn.
Ronnigeria Petr.
Staibia Bat. & Peres
Macrovalsariaceae D. Pem et al.
Macrovalsaria Petr.
Meliolinaceae S. Hughes
Briania D.R. Reynolds
Meliolina Syd. & P. Syd.
Mesnieraceae Arx & E. Müll.
Bondiella Piroz.
Mesniera Sacc. & P. Syd.
Stegasphaeria Syd. & P. Syd.
Naetrocymbaceae Höhn. ex R.C. Harris
Bonaria Bat.
Jarxia D. Hawksw.
Leptorhaphis Körb.
Naetrocymbe Körb.
Tomasellia A. Massal.
Nematotheciaceae Boonmee & K.D. Hyde
Nematothecium Syd. & P. Syd.
Nematostigma Syd. & P. Syd.
Ophioparodia Petr. & Cif.
Neoparodiaceae Boonmee & K.D. Hyde
Neoparodia Petr. & Cif.
Fungal Diversity (2020) 105:17–318
27
Palawaniaceae Mapook & K.D. Hyde
Palawania (Niessl) Syd. & P. Syd.
Stomatogeneceae Boonmee & K.D. Hyde
Stomatogene Theiss.
Paranectriellaceae Boonmee & K.D. Hyde
Paranectriella (Henn. ex Sacc. & D. Sacc.) Magnus. (=
Araneomyces Höhn.)
Puttemansia Henn.
Thyrinulaceae X.Y. Zeng et al.
Blastacervulus H.J. Swart
Paraopeba V.P. Abreu et al.
Thyrinula Petr. & Syd.
Parodiellaceae Theiss. & H. Syd. ex M.E. Barr
Parodiella Speg.
Toroaceae Boonmee & K.D. Hyde
Toroa Syd.
Perisporiopsidaceae E. Müll. & Arx ex R. Kirschner & T.A.
Hofm. (= Parodiopsidaceae Toro)
Asteronia (Sacc.) Henn.
Byssocallis Syd.
Chevalieropsis G. Arnaud
Parodiellina Henn. ex G. Arnaud
Perisporiopsis Henn.
Trichopeltinaceae Bat. et al.
Acrogenotheca Cif. & Bat.
Brefeldiella Speg.
Saccardinula Speg.
Trichopeltella Höhn.
Trichopeltheca Bat.
Trichopeltina Theiss.
Trichothyrinula Petr.
Phaeodimeriellaceae Boonmee et al.
Phaeodimeriella Speg.
Pododimeriaceae Boonmee & K.D. Hyde
Chaetoscutula E. Müll.
Pododimeria E. Müll.
Polyclypeolinaceae Boonmee & K.D. Hyde
Polyclypeolina Bat. & I.H. Lima
Polystomellaceae Theiss. & H. Syd.
Dermatodothella Viégas
Dothidella Speg.
Munkiella Speg.
Parastigmatea Doidge
Protoscyphaceae Kutorga & D. Hawksw.
Protoscypha Syd.
Pseudoperisporiaceae Toro
Bryomyces Döbbeler
Eudimeriolum Speg.
Lasiostemma Theiss.
Nematostoma Syd. & P. Syd.
Pseudorobillardaceae Crous
Pseudorobillarda M. Morelet
Pyrenidiaceae Zahlbr.
Pyrenidium Nyl.
Seynesiopeltidaceae K.D. Hyde
Seynesiopeltis F. Stevens & R.W. Ryan
Trichothyriaceae Theiss.
Lichenopeltella Höhn.
Macrographa Etayo
Pachythyrium G. Arnaud ex Spooner & P.M. Kirk
Trichothyrium Speg.
Vizellaceae H.J. Swart
Acarella Syd.
Blasdalea Sacc. & P. Syd.
Vizella Sacc.
Abrothallales Pérez-Ort. & Suija.
= Lichenoconiales Diederich, Lawrey & K.D. Hyde.
Index Fungorum number: IF 805297; Facesoffungi number: FoF 08045.
Abrothallales was established by Pérez-Ortega and Suija
(in Pérez-Ortega et al. 2014) based on its monophyletic
placement and morphological distinctness within the class
Dothideomycetes. Diederich et al. (in Hyde et al. 2013) had
introduced Lichenoconiales, which Liu et al. (2017) recovered as a sister clade of Abrothallales, raising the question
of whether both orders should be maintained. Diederich
et al. (2018) synonymized Lichenoconiales with Abrothallales and tentatively Lichenoconiaceae with Abrothallaceae.
Pérez-Ortega and Suija agreed with Diederich et al. (2018)
to use the name Abrothallales since Abrothallus is the most
outstanding, easily recognizable and better-known genus of
the group. Lichenoconiaceae is the older name, therefore, we
retain Lichenoconiaceae over Abrothallaceae. Abrothallales
contains species which are lichenicolous, parasymbiontic
or parasitic on macrolichens from a single family Abrothallaceae (Pérez-Ortega et al. 2014; Wijayawardene et al.
13
28
Fungal Diversity (2020) 105:17–318
2017a). The divergence time for Abrothallales is estimated
as 204 MYA (stem age, Hongsanan et al. 2020) (Fig. 1).
Accepted families: Lichenoconiaceae.
Lichenoconiaceae Diederich & Lawrey, in Hyde et al., Fungal Diversity 63: 131 (2013).
= Abrothallaceae Pérez-Ort. & Suija, in Pérez-Ortega,
Suija, Crespo & Ríos, Fungal Diversity 64(1): 303 (2014).
Index Fungorum number: IF 803667; Facesoffungi number: FoF 08046, 58 species.
Parasitic or parasymbiotic on lichen thalli. Vegetative
hyphae immersed in lichen thallus, usually hyaline, I+ violet in some species. Sexual morph: Ascomata apothecioid,
solitary or in groups, superficial, rounded, convex to almost
globose, rarely flattened, black or dark brown, some species
with greenish pruina, at least in young stages. Proper exciple
thin, usually disappearing when mature, composed of radiating hyphae. Hymenium hyaline or greenish in upper part,
covered with light to dark brown or reddish granules dissolving in KOH. Hypothecium light to dark brown, composed
Fig. 1 Phylogram generated
from maximum likelihood
analysis (RAxML) of Abrothallales based on combined ITS,
LSU and tef1 sequence data.
Maximum likelihood bootstrap
values equal or above 70%,
Bayesian posterior probabilities
equal or above 0.90 (MLBS/
PP) are given at the nodes. An
original isolate number is noted
after the species name. The tree
is rooted to Patellaria quercus
(CPC 27232 and BHI F828a).
The ex-type strains are indicated
in bold. Hyphen (-) represents
support values below 70%
MLBS and 0.90 PP
Abrothallus erioder AB74
Abrothallus erioder LG AB60
Abrothallus secedens Perez Ortega 3162 (MA)
98/1.0 Abrothallus secedens SPO305
Abrothallus secedens Perez Ortega 3156 (MA)
Abrothallus secedens Perez Ortega 3161 (MA)
95/0.96
Abrothallus secedens Perez Ortega 3158 (MA)
Abrothallus secedens Perez Ortega 3175 (MA)
Abrothallus canariensis van den Boom 37849 (MA)
100/1.0 Abrothallus canariensis van den Boom 37849 (MA)
Abrothallus kamchatica Yakovchenko (MA)
Abrothallus nephromatis Perez Ortega 3159 (MA)
Abrothallus nephromatis Diederich 12530 BR LICH 7794 34
Abrothallus nephromatis Spribille 10581 (hb. Spribille)
Abrothallus nephromatis LE 260303
Abrothallus nephromatis Ertz 12514 (BR-LICH-6515-16)
100/1.0
81/0.99
100/1.0
Abrothallus boomii (MA) van den Boom 43228
Abrothallus ertzii Ertz 12517 BR LICH 6512 13
Abrothallus welwitschii AB61 Serusiaux LG
Abrothallus welwitschii Perez Ortega 3169 (MA)
Abrothallus welwitschii Ertz 1083 BR LICH 1639 87
100/1.0
80/0.98
Abrothallus welwitschii Perez Ortega 3176 (MA)
Abrothallus welwitschii Perez Ortega 3171 (MA)
Abrothallus welwitschii Perez Ortega 3168 (MA)
87/0.97
Abrothallus welwitschii Spribille 24949 (hb. Spribille)
Abrothallus etayoi Perez Ortega 2003 (MA)
Abrothallus doliiformis Perez-Ortega 2004 (MA)
Abrothallus usneae AB20
99/1.0
Abrothallus suecicus SPO304
Abrothallus acetabuli SPO308
100/1.0
Abrothallus parmeliarum AB36
Abrothallus acetabuli von Brackel 5661
93/0.97 Abrothallus parmeliarum AB57
Abrothallus parmotrematis AB1
Abrothallus buellianus SPO303
96/1.0
100/1.0
Abrothallus hypotrachynae SPO302
Abrothallus granulatae Perez Ortega 3166 (MA)
96/0.98 Abrothallus usneae SPO306
Abrothallus suecicus AB56
Abrothallus cladoniae AB53
Lichenoconium
erodens CBS 128028
73/0.9
Lichenoconium aeruginosum CBS 129239
97/0.99
Lichenoconium usneae CBS 128708
83/0.92
Lichenoconium erodens CBS 128704
100/1.0 Patellaria quercus CPC 27232
Patellaria quercus BHI F828a
92/1.0
99/1.0
Abrothallales
73/-
0.04
13
of isodiametric cells (textura angularis type). Hamathecium
comprising septate, anastomosed and irregularly dichotomously branching interascal filaments. Asci 4–8-spored,
thick-walled, bitunicate, functionally fissitunicate, ovate
to clavate, apedicellate, with a distinct ocular chamber, I-.
Ascospores 2-seriate, irregularly or 2-seriately arranged
within the ascus, ellipsoid to broadly ellipsoid, light to dark
brown, 1- to 3-septate, asymmetric and soleiform, splitting
into part-spores within the ascus in some species, wall finely
ornamented to verrucose. Asexual morph: Coelomycetous.
Vouauxiomyces-type, pycnidium immersed, semi-immersed
or almost superficial, black, with a small ostiole. Pycnidial
wall thick-walled, comprises isodiametric cells (textura
angularis-type). Conidiophores reduced. Conidiogenous
cells percurrently proliferating, ampulliform to lageniform,
lining the cavity of the pycnidium, hyaline, smooth-walled.
Conidia holoblastic, clavate to obpyriform or almost roundish, hyaline to dark brown, smooth to slightly echinulate,
usually embedded in mucilage.
Type: Lichenoconium Petr. & Syd.
Outgroup
Fungal Diversity (2020) 105:17–318
Notes: Lawrey et al. (2011) studied the phylogenetic affinities of Lichenoconium, a genus of lichenicolous fungi known
only in its asexual morph. They found that Lichenoconium
is also a member of Dothideomycetes, and subsequently
Hyde et al. (2013) introduced a new order Lichenoconiales
to accommodate it. Pérez-Ortega et al. (2014) introduced
Abrothallaceae and the new order Abrothallales for the lichenicolous genus Abrothallus. The classification of this genus had
long been debated. This is one of the few Dothideomycetes
genera having apothecioid ascomata, but the bitunicate asci
and the shape of interascal filaments clearly segregates it from
other similar groups (Pérez-Ortega et al. 2014). Pérez-Ortega
et al. (2011) established the connection between the sexual
morph Abrothallus and the asexual morph Vouauxiomyces.
Vouauxiomyces and Lichenoconium share many features, such
as the mode of conidiogenesis and shape of conidia (detailed
descriptions in Hawksworth 1981b), although the conidia are
hyaline in Abrothallus (except in Abrothallus kamchatica) and
dark brown in Lichenoconium. Recent phylogenetic analyses
recovered Abrothallus and Lichenoconium as sister genera,
with a relatively young split (9 MYA) raising the question of
the need to keep two separate orders (Liu et al. 2017). Diederich et al. (2018) in their review of lichenicolous fungi treated
Lichenoconiales as a synonym of Abrothallales, and Lichenoconiaceae as a possible synonym of Abrothallaceae.
Lichenoconium Petr. & Syd., Beih. Reprium nov. Spec.
Regni veg. 42(1): 432 (1927) [1926].
Index Fungorum number: IF 8772; Facesoffungi number:
FoF 08047; 16 morphological species (Species Fungorum
2020), 4 species with molecular data.
Type species: Lichenoconium lichenicola (P. Karst.) Petr.
& Syd. [as ‘lichenicolum’], Beih. Reprium nov. Spec. Regni
veg. 42(1): 432 (1927) [1926].
≡ Dactylium dendroides subsp. lichenicola P. Karst.,
Meddn Soc. Fauna Flora fenn. 14: 107 (1887).
Notes: Lichenoconium species are distributed worldwide
as lichenicolous taxa on a variety of lichen hosts (Lawrey
et al. 2011; Hyde et al. 2013; Wijayawardene et al. 2017a;
Diederich et al. 2018). Many are host-specific, while others
occur on disparate genera (Lawrey et al. 2011). The asexual
morph is characterized by coelomycetous, immersed to
sessile, dark brown pycnidia, an undifferentiated pycnidial
ostiole, simple conidia, globose to ellipsoid, basally often
truncate, thick-walled, brown and commonly verrucose and
conidiogenous cells that are hyaline or poorly pigmented,
holoblastic, elongate and annellidic (Lawrey et al. 2011).
The sexual morph has not been recorded.
Other genera included
Abrothallus De Not. Giorn. Bot. Ital. 1: 194 (1846).
= Vouauxiomyces Dyko & D. Hawksw., Lichenologist
11(1): 57 (1979).
29
Index Fungorum number: IF 4; Facesoffungi number:
FoF 08048; – 42 morphological species (Species Fungorum
2020), 20 species with molecular data.
Type species: Abrothallus bertianus De Not. Giorn. Bot.
Ital. 1: 194 (1846).
Notes: De Notaris described Abrothallus as a lichenized
taxon (De Notaris 1846, 1849). Tulasne (1852) and Lindsay
(1857) unequivocally established its lichenicolous habit.
Suija et al. (2018) fixed the nomenclatural problems concerning the exact date of publication, confirming A. bertianus as the type species. They also reviewed the material
described by Giuseppe De Notaris, Søren Christian Sommerfelt, and Ignaz Kotte lectotypifying Abrothallus species
described by these authors (Suija et al. 2018). These authors
also introduced the combination A. santessonii (≡ Vouauxiomyces santessonii), providing an updated description for this
taxon. Pérez-Ortega et al. (2011) established the connection
between the sexual and asexual morphs. The number of species is tentative since there are numerous single collections
occurring on unusual hosts that may represent new species
and there are only a few collections from certain regions,
e.g. from Africa and Asia. Recent studies focused on the taxonomy of species growing on Peltigerales (Suija et al. 2011,
2015). However, species delimitation, especially those on
specimens growing on Parmeliaceae, is difficult and in need
of a thorough revision (Suija et al. 2018). For morphology of
the type species, see Suija et al. (2018) (Fig. 2).
Economic and ecological significance
Species in Abrothallaceae are not harmful to plants or animals. However, their lichenicolous, parasymbiontic or parasitic lifestyles on macrolichens are interesting for biodiversity and ecological research.
Acrospermales Minter, Peredo & A.T. Watson.
Index Fungorum number: IF 90786; Facesoffungi number: FoF 06407.
Acrospermales was introduced by Minter (2007) to
accommodate Acrospermaceae. The ordinal position of
Acrospermaceae was previously unresolved and was referred
to various orders by many authors. First, it was placed in Hysteriales or Dothideales in the 1st and 2nd editions of Ainsworth & Bisby’s Dictionary of the Fungi (Kirk et al. 2001).
In the 3rd to 5th editions it was transferred to the Dothideales
and in the 6th and 7th it was placed in the Ostropales and Clavicipitales, respectively. The family was also placed, rather
hesitantly, in Pyrenulales by Eriksson (1982). Barr (1990)
argued its ordinal position in more detail, and transferred
it to Xylariales. Minter et al. (2007) introduced a new species in Acrospermaceae and discussed the ordinal position
of Acrospermaceae with the introduction of Acrospermales.
The divergence time for Acrospermales is estimated as 156
MYA (stem age, Hongsanan et al. 2020) (Fig. 3).
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Fungal Diversity (2020) 105:17–318
Fig. 2 Morphological and anatomical features of Abrothallaceae. a
Habitus Abrothallus welwitschii, mature apothecia. b Habitus A. welwitschii, young apothecia with greenish pruina. c Cross-section of A.
parmotrematis on Parmotrema sp. d Interascal elements from A. bertianus. e One-septate ascospore of A. welwitschii, with surface showing ornamentation. f Ascus of A. boomii. g Three-septate ascospores
Accepted families: Acrospermaceae.
Acrospermaceae Fuckel, Jb. nassau. Ver. Naturk. 23–24: 92
(1870) [1869–70].
13
of A. suecicus. h Scanning electron micrograph of A. welwitschii
ascospores. i Ascospore of A. buellianus showing occasional splitting
at the septum. j A. caerulescens asexual stage (pycnidium, Vouauxiomyces-type). k Conidia from A. boomii. l Lichenoconium sp. on A.
santessonii ascoma. Scale bars: a–b = 0.5 mm, c = 50 µm, d–h, k =
10 µm; i = 5 µm, j = 30 µm, l = 20 µm
Index Fungorum number: IF 80430; Facesoffungi number: FoF 06380, 52 species.
Saprobic, epiphytic or symbiotic on herbaceous plants.
Sexual morph: Ascomata solitary or in groups, superficial
or immersed in stromata, erect, elongate, with smooth or
Fungal Diversity (2020) 105:17–318
100/1.00
-/1.00
78/0.75
85/1.00
Pseudovirgaria hyperparasitica CPC 10753
Pseudovirgaria grisea CPC 19132
Acrospermum compressum M 151
Acrospermum graminum M 152
71/0.97
Acrospermum adeanum M 133
80/1.00
Gonatophragmium triuniae CBS 138901
84/1.00
Acrospermales
Fig. 3 Phylogram generated
from maximum likelihood
analysis based on combined
LSU and SSU sequence data
representing Acrospermaceae.
Strigula nemathora (MPN72)
(Strigulaceae, Pleosporales)
is used as the outgroup taxon.
Bootstrap values for maximum
likelihood (ML) equal to or
greater than 70% and clade
credibility values greater than
0.90 (the rounding of values to
2 decimal proportions) from
Bayesian-inference analysis are
labeled on the nodes. Ex-type
strains are in bold and black, the
new isolate is indicated in bold
and blue
31
Acrospermum urticae MFLU 18-1666
Acrospermum graminum MFLUCC 13-0886
Acrospermum longisporium MFLU 17-2849
Strigula nemathora MPN72
Outgroup
0.02
sometimes rough surface, dark brown to black, flattened,
club-shaped to conoid, with a short stipe, swelling when
moist, ostiolate. Peridium comprising two or three layers, an
outer layer composed of dark brown cells of textura angularis, a central layer, composed of hyaline, sometimes pale
brown tissue of elongated cells intertwined, and an inner
layer composed of dense tissue of small, light brown cells.
Hamathecium comprising narrow, long, hyaline, filiform
pseudoparaphyses. Asci typically 8-spored, bitunicate, long,
narrowly cylindrical, pedicellate, apically rounded with an
ocular chamber. Ascospores fasciculate, filiform, hyaline,
multi-septate, almost as long as the asci, smooth-walled, not
fragmenting, without sheath, typically intertwined in a fascicle within the ascus. Asexual morph: Hyphomycetous. Conidiophores micronematous, pale brown, septate, branched
or unbranched. Conidiogenous cells holoblastic, sympodial
with denticles, pale brown, smooth-walled. Conidia cylindrical, long ellipsoid, pale yellow, 1–3-septate, smooth-walled.
Type: Acrospermum Tode.
Notes: Acrospermaceae was introduced with a single genus
Acrospermum by Fuckel (1870) and since then its higher taxonomic placement has undergone various changes (Saccardo
1883; Rehm 1887; Ellis and Everhart 1892; Ainsworth et al.
1973; Barr 1990). The family was previously placed in the
class Dothideomycetes family incertae sedis, due to its uncertain position (Hyde et al. 2013). Two genera, Acrospermum
and Oomyces are currently accepted (Lumbsch and Huhndorf
2010; Wijayawardene et al. 2018). Acrospermum is characterised by erect, elongate, usually brown, superficial ascomata
that are more or less club-shaped and are solitary or in small
groups, with long paraphyses which resemble ascospores.
Oomyces is considered to be clavicipitalean (Diehl 1950) as
it has conoid, yellowish white, multi-locular stromata, bitunicate asci and lacks pseudoparaphyses (Eriksson 1981). No
monograph of the genus is available. Riddle (1920) reviewed
Acrospermum and introduced A. maxoni and A. graminum var.
foliicolum based on asexual morphs. Presently, the asexual
morphs of Acrospermaceae comprise members of Dactylaria
and Gonatophragmium (Wijayawardene et al. 2018). Dactylaria was shown to be polyphyletic by Bussaban et al. (2005)
and is heterogenous (Seifert et al. (2011). Gonatophragmium
was also found to be an asexual morph of Acrospermum by
Kirk et al. (2008), but Seifert et al. (2011) did not assign it to
any taxonomic rank. A Blast search of an LSU sequence of
Gonatophragmium triuniae showed closed hits to Acrospermum adeanum (Crous et al. 2014). We therefore, agree with
Wijayawardene et al. (2018) and include Gonatophragmium
in Acrospermaceae until further data becomes available. The
hyphomycetous genus Pseudovirgaria was introduced by
Shin et al. in Arzanlou et al. (2007), with P. hyperparasitica
as type species. Pseudovirgaria is assigned to Capnodiales,
genera incertae sedis in Index Fungorum (2020), while it
was mentioned as Dothideomycetes genera incertae sedis in
Wijayawardene et al. (2018). In our phylogenetic analysis,
two species of Pseudovirgaria clustered in Acrospermales.
Pseudovirgaria is a hyphomycetous genus and morphologically resembles Gonatophragmium in having cylindric-clavate,
thin-walled conidia. Therefore, we include Pseudovirgaria in
Acrospermaceae based on phylogenetic evidence, and morphological resemblance to asexual genera of Acrospermaceae.
13
32
Descriptions and illustrations of the asexual morphs of Acrospermaceae can be seen in previous studies (i.e. Crous et al.
2014; Berger et al. 2015; Shamsi et al. 2017)
Acrospermum Tode, Fung. mecklenb. sel. (Lüneburg) 1: 8
(1790).
Index Fungorum number: IF 54; Facesoffungi number:
FoF 06381; 28 morphological species (Species Fungorum
2020), 8 species with molecular data.
Type species: Acrospermum compressum Tode, Fung.
mecklenb. sel. (Lüneburg) 1: 8 (1790).
Notes: Tode (1790) introduced Acrospermum with A. compressum as the type species based on fruiting body and ostiole
type. Acrospermum is characterised by superficial, club-like
ascomata, bitunicate asci and fasciculate, filiform, hyaline,
multi-septate ascospores (Riddle 1920). Species of Acrospermum are mostly saprobic and are distributed worldwide.
The specimens of A. compressum were found on dry stems
of Heracleum sphondylium in Germany. Acrospermum compressum can also be observed on dead stems of Urtica dioica,
and A. graminum on grass culms. Acrospermum adeanum is
a necrotrophic parasite and has been observed on 32 different
moss species from 22 different genera, most of which belong
to the pleurocarpous superorder Hypnanae (Döbbeler 1979;
Bell and Newton 2004). The asexual morph of Acrospermum
is hyphomycetous (Wijayawardene et al. 2018).
Acrospermum urticae D. Pem, Camporesi & K.D. Hyde,
sp. nov.
Index Fungorum number: IF 556687; Facesoffungi number: FoF 06382; Fig. 4
Etymology: Name reflects the host from which the fungus
is isolated.
Holotype: MFLU 18-1666.
Saprobic on dead stem of Urtica dioica. Sexual morph:
Ascomata 940–1057 high × 301–345 µm diam. ( x̄ = 1021
× 318 µm), solitary or in groups, superficial, club-shaped
to conoid, erect, uni-locular, brown to blackish when dry,
with a short stipe or sessile, flattened when dry, swelling
when moist, ostiole large, apex rounded. Peridium 11–12
µm in vertical section comprising three layers, an outer layer
comprising dark brown cells of textura angularis, a central
thick layer, comprising pale brown to hyaline tissue of gelatinized hyphae with elongated cells, and an inner layer comprising dense tissue of small, hyaline cells. Hamathecium
comprising narrow, long, pseudoparaphyses. Asci 195–319
× 6.2–6.7 μm ( x̄ = 252.5 × 6.4 µm), 8-spored, bitunicate,
narrowly cylindrical, pedicellate, with an ocular chamber.
Ascospores 122–170 × 1.1–1.2 μm ( x̄ = 146.2 × 1.2 µm),
fasciculate, filiform, hyaline, multi-septate, nearly as long
as the asci, smooth-walled. Asexual morph: undetermined
Material examined: Italy, Ravenna [RA], San Cassiano di Brisighella, on dead aerial stem of Urtica dioica
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Fungal Diversity (2020) 105:17–318
(Urticaceae), 13 August 2018, Erio Camporesi (IT 3999,
holotype; MFLU 18-1666, isotype).
GenBank numbers: LSU: MN597994, SSU: MN597996.
Notes: Acrospermum urticae differs from Acrospermum
longisporium by its smaller ascomata (940–1057 high ×
301–345 µm diam. v.s. 1500–2000 high × 400–500 µm
diam.) and wider ascospores (122.1–175.3 × 1.1–1.2 μm v.s.
150–170 × 0.5–1 μm). Phylogenetic analyses of a combined
LSU, SSU sequence dataset show that A. urticae forms a
distinct lineage in Acrospermaceae with strong ML and
BYPP support (80% ML, 1.0 BYPP; Fig. 3). Therefore, we
introduce Acrospermum urticae as a new species.
Other genera included
Gonatophragmium Deighton, in Cejp & Deighton, Mycol.
Pap. 117: 13 (1969).
Index Fungorum number: IF 8376; Facesoffungi number:
FoF 06486; – 17 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Gonatophragmium mori (Sawada)
Deighton 1969, in Cejp & Deighton, Mycol. Pap. 117: 13
(1969).
≡ Spondylocladium mori Sawada, Spec. Bull. Agric. Exp.
Station Formosa 19: 665 (1919).
Notes: Gonatophragmium was described in Cejp and
Deighton (1969) with G. mori as the type species, a combination based on Spondylocladium mori. Gonatophragmium
mori is a tropical-subtropical leaf-spotting species and is
found on a wide range of hosts. Takahashi and Teramine
(1986) considered Acrospermum viticola to be the sexual
morph of this species, however this association was not
proven by molecular data. Gonatophragmium is distinct in
having pigmented, branched conidiophores formed as erect
to decumbent threads with terminal and intercalary conidiogenous cells, which are regularly swollen around fertile
portions with mostly numerous noticeable conidiogenous
loci. The sexual morph of this genus is undetermined.
Oomyces Berk. & Broome, Ann. Mag. nat. Hist., Ser. 2 7:
185 (1851).
Index Fungorum number: IF 8376; Facesoffungi number:
FoF 06488; – 7 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Oomyces carneoalbus (Lib.) Berk. &
Broome, Ann. Mag. Nat. Hist., Ser. 2 7: 185 (1851).
≡ Sphaeria carneoalba Lib., Pl. crypt. Arduenna, fasc.
(Liège) 3(nos 201–300): no. 241 (1834).
Notes: Oomyces (= Coscinaria Ellis & Everh. 1886
fide Species Fungorum 2017) was introduced by Berk and
Broome (1851), with O. carneoalbus as the type species. The
genus is characterized by conoid, yellowish white, multi-locular stromata and lacks pseudoparaphyses (Eriksson 1981).
Oomyces was described as similar to the egg of some insects,
Fungal Diversity (2020) 105:17–318
33
Fig. 4 Acrospermum urticae (IT 3999, holotype). a–d Ascomata on
host surface. e Ascoma in vertical section f Peridium. g–i Narrowly
cylindrical asci. j–k Filiform ascospores Scale bars: a = 2000 µm, b,
e = 500 µm, c, d = 300 µm, f, g = 100 µm, h = 25 µm, i = 30 µm, j =
50 µm, k = 40 µm
such as Crioceris, because its perithecia are visible as little
dimples in the truncate apex of the fruiting body.
Pseudovirgaria H.D. Shin, U. Braun, Arzanlou & Crous,
in Arzanlou et al., Shin & Crous, Stud. Mycol. 58: 87 (2007)
Index Fungorum number: IF504564; Facesoffungi number: FoF 06487; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Pseudovirgaria hyperparasitica H.D.
Shin, U. Braun, Arzanlou & Crous, in Arzanlou et al., Stud.
Mycol. 58: 87 (2007)
Notes: Phylogenetic analyses indicated that two species of
Pseudovirgaria clustered within Acrospermaceae (Hudson
et al. 2019, this study). Morphologically, it resembles
Gonatophragmium in having cylindric-clavate, thin-walled
conidia. Therefore, we agree with Hudson et al. (2019) to
accept this genus in Acrospermaceae.
Economic and ecological significance
Species of Acrospermaceae are parasitic or endophytic and
may play a negative role by infecting ferns. In Mexico, several species of Terpsichore (T. subtilis and T. taxifolia) are
infected by Acrospermum, typically as black clavate stromata on the hosts.
Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss.
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34
Index Fungorum number: IF 90461; Facesoffungi number: FoF 07605.
Asterinales contains epifoliar fungi which have superficial mycelium forming a network on host plants, 1-celled
appressoria, with a star-like opening to the thyriothecium.
The order was revised by Hongsanan et al. (2014b), however, its classification is unclear due to insufficient sequence
data. Sequence data of Asterinales clustered in two unrelated clades in phylogenetic trees (Ertz et al. 2016; Hyde
et al. 2016b; Liu et al. 2017). The clade containing the type
species Asterotexis cucurbitacearum was treated as Asterotexales (Ertz et al. 2016). However, Hyde et al. (2016b) synonymized it under Asterinales sensu stricto because most
Asterinales strains cluster in this clade. The other clade,
including species of Parmulariaceae were treated as Asterinales sensu lato (Liu et al. 2017). Parmulariaceae was transferred to its own order Parmulariales by Dai et al. (2018).
Although the type species Asterina melastomatis and a
few other Asterinales-like taxa fell within the clade Asterinales sensu lato (Ertz et al. 2016; Hyde et al. 2016b), we
did not include these sequence data in our analysis. This is
because Asterinales-like taxa forming in the Asterinales sensu
lato need to be rechecked since other hyphomycetous strains
were added into this clade, thus, there is possibility that these
sequence data are not Asterinales. Thyrinulaceae is introduced
to accommodate a clade sister to Parmulariales (= Asterinales sensu lato) with Thyrinula as generic type based on the
rules of nomenclatural priority. Hongsanan et al. (2014b) synonymised Lembosiaceae under Asterinaceae, however, adding more sequence data for Lembosia (Fig. 5) indicates that
Lembosia should be raised to a family in Asterinales.
By considering phylogenetic trees (Fig. 5), we retain Lembosiaceae in Asterinales and introduce Morenoinaceae and
Neobuelliellaceae to accommodate the clades of Morenoina
and Neobuelliella, respectively. Based on morphology and
phylogeny, the current Asterinales comprises eight families.
The classification of Asterinales is questionable and needs
more morphological and molecular data to clarify its phylogenetic relationship. The divergence time for Asterinales
is estimated as 221 MYA (stem age, Hongsanan et al. 2020).
Accepted families: Asterinaceae, Asterotexaceae, Hemigraphaceae, Lembisiaceae, Melaspileellaceae, Morenoinaceae, Neobuelliellaceae and Stictographaceae.
Asterinaceae Hansf., Mycol. Pap. 15: 188 (1946).
Index Fungorum: IF 80492; Facesoffungi number: FoF
06726, >1000 species.
Colonies epiphyllous or hypophyllous. Hyphae superficial,
straight to substraight, dark brown, reticulate, with appressoria.
Appressoria 1-celled, mostly lateral, alternate to unilateral. Sextual morph: Thyriothecia superficial, flattened, with stellate or
longitudinal dehiscence. Upper walls brown, comprising radial,
septate cells of textura prismatica. Asci 8-spored, bitunicate,
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Fungal Diversity (2020) 105:17–318
ellipsoid, usually thickened at the apex. Hamathecium cellular
pseudoparaphyses present or absent. Ascospores 2–5-seriate or
fasciculate or conglobate, fusoid to ellipsoid, hyaline to dark
brown, mostly 1-septate, smooth-walled or toughened. Asexual
morph: Coelomycetous states with pycnidia or pycnothyria,
and hyphomycetous states without conidiomata or sporodochia
then gelatinous, pale. Hyphae brown, superficial, with appressoria. Conidiomata pycnothyria, flattened, dimidiate, radiate,
orbicular, stellately dehisced at the centre. Conidiophores
branched or unbranched, hyaline or brown. Conidiogenous cells
monoblastic or percurrent, hyaline or brown. Conidia ovate,
pyriform, angular, or wall straight to sinuate, brown.
Type: Asterina Lév.
Notes: Asterinaceae was established as a member of Microthyriales by Hansford (1946). Members of the family typically
have upper walls comprising radiating cells with star-like or
longitudinal splits and dark brown hyphae with appressoria.
There are 18 genera in this family based on morphology (Hongsanan et al. 2014b; Guatimosim et al. 2015; Wijayawardene
et al. 2017a; Dai et al. 2018). Although Wijayawardene et al.
(2017a) included Echidnodes in Asterinaceae, we exclude it
from Asterinaceae as Hongsanan et al. (2014b) transferred this
genus to Aulographaceae. Phylogenetic studies have several different interpretations of this family. In this study, we reappraise
the phylogenetic relationship of Asterinaceae and related families based on all available sequence data and previous studies.
Asterina Lév., Annls Sci. Nat., Bot., sér. 3 3: 59 (1845).
Index Fungorum number: IF 409; Facesoffungi number:
FoF 06727; >1000 morphological species (Species Fungorum 2020), 9 species with molecular data.
Type species: Asterina melastomatis Lév., Annls Sci.
Nat., Bot., sér. 3 3: 59 (1845).
Notes: Asterina was introduced as a member of Sphaeriaceae with A. azarae, A. compacta, A. pulla and the type
A. melastomatis. It is the largest genus in Asterinaceae, but
only nine species have sequence data available in GenBank
due to its unculturable character. Members of the genus have
circular thyriothecia with stellate dehiscence, lateral appressoria, globose asci, and dark brown, 1-septate ascospores.
Asterina magnoliae X.Y. Zeng, T.C. Wen & K.D. Hyde, in
Hyde et al., Mycosphere 9(2): 349 (2018).
Index Fungorum number: IF 554238; Facesoffungi number: FoF 04089; Fig. 6
Description: see Hyde et al. (2018).
Material examined: Thailand, Chiang Mai, Mae Taeng,
Pa Pae, Bahn Pa Deng, Mushroom Research Centre, 128
Moo 3, on living leaves of Magnolia odora (Magnoliaceae),
8 July 2015, Xiang-Yu Zeng (MFLU 16-0071).
GenBank number: LSU: MN629745.
Notes: Our new collection of Asterina magnoliae is identified by morphological characters (Fig. 6) and phylogenetic
Fungal Diversity (2020) 105:17–318
35
Asterina cynometrae MFLU 13 0373
99/1.0
Asterina cestricola TH 591
Asterostomella grewiae MFLU 13 0629
83/Asterina siphocampyli ppMP 1324
Asterina phenacis TH 589
Asterina weinmanniae TH 592
98/1.0
Asterina zanthoxyli TH 561
Asterina fuchsiae TH 590
Asterina magnoliae MFLU 16-0071
100/1.0
Asterina magnoliae MFLU 16-0072
Asterotexis cucurbitacearum VIC 42814
97/1.0
Parmularia styracis VIC 42447
Cladoriella eucalypti CPC 10953
Cladoriella rubrigena CBS 124760
Venturia inaequalis ATCC 60070
Venturia populina CBS 256.38
100/1.0
100/1.0
0.06
Lembosiaceae
Hemigraphaceae
Neobuelliellaceae
Melaspileellaceae
genus incertae sedis
Stictographaceae
Morenoinaceae
Thyrinulaceae
Incertae sedis
Stictographa lentiginosa Ertz 17447
Labrocarpon canariense Ertz 16308
84/0.99
Melaspileopsis cf diplasiospora Ertz 16247 BR
Morenoina palmicola MFLUCC 15-0284
70/0.94
Morenoina calamicola MFLUCC 14-1162
100/1.0
Blastacervulus robbenensis CBS 120138
-/0.95 Blastacervulus eucalypti CBS 124759
-/0.93
Blastacervulus eucalyptorum CPC 29450
Thyrinula uruguayensis CBS 120122
Thyrinula eucalyptina CPC 35990
93/1.0 Thyrinula dunnii CPC 12977
Thyrinula parasitica CBS 120088
96/1.0
Thyrinula eucalypti CPC 12986
100/1.0
Paraopeba schefflerae 1620
Asterotexaceae
Asterinales incertae sedis
Asterinales
Inocyclus angularis VIC 39747
99/1.0 Lembosia mimusopis MFLU 19-1225
88/1.0
100/1.0
Lembosia mimusopis MFLU 19-0724
66/0.94
Lembosia xyliae MFLU 14-0004
100/1.0
Lembosia albersii MFLU 13-0377
100/1.0
Hemigrapha atlantica Ertz 14014 BR
Neobuelliella poetschii Ertz 18115 BR
Melaspileella proximella G M 2015 04 29
88/0.95
Pirozynskiella laurisilvatica FMR 13133
71/Karschia talcophila Ertz 16749
96/1.0
100/1.0
Karschia cezannei Ertz 19186
Buelliella minimula Lendemer 35969
Melaspilea lekae Ertz 17325
Buelliella physciicola Ertz 18113
Asterinaceae
Parmulariales
Cladoriellales
Outgroup
Fig. 5 Phylogram generated from maximum likelihood analysis
(RAxML) of Asterinales based on LSU sequence data. Maximum
likelihood bootstrap values equal to or greater than 70%, Bayesian
posterior probabilities equal to or greater than 0.90 (MLBS/PP) are
given at the nodes. Isolate numbers are noted after each species name.
The tree is rooted to Venturia inaequalis (ATCC 60070) and Venturia
populina (CBS 256.38). Newly sequence data generated in this study
are in blue. Ex-types are indicated in bold. Hyphen (-) represents support values less than 70% MLBS and 0.90 PP
evidence (Fig. 5). Hyde et al. (2018) provided a full description of this species.
2020), molecular data available for an unnamed species in
the genus.
Type species: Asterinella puiggarii (Speg.) Theiss., Brotéria, sér. bot. 10(2): 116 (1912).
≡ Asterina puiggarii Speg., Anal. Soc. cient. argent.
12(3): 99 (1881).
Notes: Asterinella was introduced as a member of Microthyriaceae. It is characterised by superficial hyphae with
Other genera included
Asterinella Theiss., Annls mycol. 10(2): 160 (1912).
Index Fungorum number: IF 411; Facesoffungi number:
FoF 06729; – 35 morphological species (Species Fungorum
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Fungal Diversity (2020) 105:17–318
Fig. 6 Asterina magnoliae (MFLU 16-0071). a Host leaves. b, c Colonies on leaf surface. d Thyriothecium. e Pycnidioma. f Young asci. g
Mature asci. h Ascospores. i Pycnidia. Scales bars: b–c = 500 μm, d–e = 50 μm, f–g = 20 μm, h–i = 10 μm
intercalary appressoria and thyriothecia with a stellate ostiole. Wu et al. (2014) transferred it to Asterinaceae based on
morphology.
Asterolibertia G. Arnaud 1918, Annals d’École National
d’Agric. de Montpellier, Série 2 16(1–4): 165 (1918) [1917].
Index Fungorum number: IF 421; Facesoffungi number:
FoF 06731; – 35 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Asterolibertia couepiae (Henn.) G. Arnaud,
Annals d’École National d’Agric. de Montpellier, Série 2
16(1–4): 165 (1918) [1917].
≡ Asterina couepiae Henn., Hedwigia 34: 104 (1895).
Notes: Asterolibertia is characterised by intercalary
appressoria. Hongsanan et al. (2014b) synonymised Asterolibertia under Asterina, while Firmino et al. (2016) questioned the intercalary appressoria in Asterolibertia as not
homologous to the lateral appressoria in Asterina. However,
both justifications are based on morphology.
Asterostomella Speg., Anal. Soc. cient. argent. 22(4): 198
(1886).
Index Fungorum number: IF 7271; Facesoffungi number:
FoF 06730; – 87 morphological species (Species Fungorum
2020), 1 species with molecular data.
13
Type species: Asterostomella paraguayensis Speg., Anal.
Soc. cient. argent. 22(4): 198 (1886).
Notes: Asterostomella is a coelomycetous genus characterised by brown, ovoid, aseptate pycnidia, sometimes with a
non-pigmented band in the middle. The genus is considered
as a member of Asterinaceae based on its scutellate conidiomata with stellate dehiscence, which is similar to Asterina.
Batistinula Arx, Publicações Inst. Micol. Recife 287: 4
(1960).
Index Fungorum number: IF 523; Facesoffungi number:
FoF 06732; – 1 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Batistinula gallesiae Arx, Publicações Inst.
Micol. Recife 287: 6 (1960).
Notes: Batistinula typically has 3-septate ascospores. A
specimen that shares the same morphological and biometric
characteristics of the type was collected and sequenced by
Guatimosim et al. (2015). However, that fresh collection was
found on a different host family, which may contradict the
host-specificity of Asterinaceae.
Cirsosia G. Arnaud, Annals d’École National d’Agric. de
Montpellier, Série 2 16(1–4): 127 (1918) [1917].
Fungal Diversity (2020) 105:17–318
Index Fungorum number: IF 1065; Facesoffungi number:
FoF 06734; – 15 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Cirsosia manaosensis (Henn.) G. Arnaud
[as ‘manaoensis’], Annals d’École National d’Agric. de
Montpellier, Série 2 16(1–4): 127 (1918) [1917].
≡ Lembosia manaosensis Henn. [as ‘manaoensis’], Hedwigia 43(4): 265 (1904).
Notes: Cirsosia is mostly similar to Lembosia in having a
linear fissure, but has intercalary appressoria.
37
Index Fungorum number: IF 3102; Facesoffungi number:
FoF 06738; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Meliolaster clavisporus (Pat.) Höhn., Ber.
dt. bot. Ges. 35(10): 701 (1918).
≡ Meliola clavispora Pat., J. Bot., Paris 4: 61 (1890).
Notes: Thyriotheica in Meliolaster are composed of radially arranged of cells, which open by star-like fissures when
mature. Meliolaster is similar to Batistinula and Halbania
in having 3-septate ascospores, but differs in the presence
and shape of appressoria on the hyphae.
Dothidasteromella Höhn., Sber. Akad. Wiss. Wien, Math.naturw. Kl., Abt. 1 119: 421 (1910).
Index Fungorum number: IF 1692; Facesoffungi number:
FoF 06236; – 11 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Dothidasteromella sepulta (Berk. & M.A.
Curtis) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl.,
Abt. 1 119: 421 (1910).
≡ Asterina sepulta Berk. & M.A. Curtis, Proc. Amer.
Acad. Arts & Sci. 4: 129 (1860).
Notes: Dothidasteromella is mostly similar to Echidnodella, Halbania and Uleothyrium in lacking appressoria,
but has Y-shaped dehiscence, 1-septate ascospores, and lack
pseudoparaphyses.
Parasterinopsis Bat., Atas Inst. Micol. Univ. Recife 1: 327
(1960).
Index Fungorum number: IF 3722; Facesoffungi number:
FoF 06739; – 3 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Parasterinopsis sersalisiae (Hansf.) Bat.,
Atas Inst. Micol. Univ. Recife 1: 327 (1960).
≡ Patouillardina sersalisiae Hansf., Proc. Linn. Soc. London 156: 117 (1944) [1943–44].
Notes: Parasterinopsis is retained in Asterinaceae based
on its thyriothecia with irregular fissures and the superficial
hyphae with appressoria. However, it typically has cylindrical, 1–4-septate ascospores.
Echidnodella Theiss. & Syd., Annls mycol. 15(6): 422
(1918) [1917].
Index Fungorum number: IF 1731; Facesoffungi number:
FoF 06761; – 33 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Echidnodella linearis (Syd. & P. Syd.)
Syd., Annls mycol. 15(6): 422 (1918) [1917].
≡ Morenoella linearis Syd. & P. Syd., Annls mycol.
15(3/4): 250 (1917).
Notes: Echidnodella is similar to Lembosia and Cirsosia
in having linear fissures, but lacks appressoria and has cellular pseudoparaphyses.
Platypeltella Petr., in Sydow & Petrak, Annls mycol.
27(1/2): 62 (1929).
Index Fungorum number: IF 4178; Facesoffungi number:
FoF 06741; – 3 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Platypeltella smilacis Petr., Annls mycol.
27(1/2): 62 (1929).
Notes: Platypeltella was introduced as a member of
Microthyriaceae. Wu et al. (2014) included it in Asterinaceae based on the superficial hyphae with intercalary
capitate appressoria. Platypeltella is similar to Asterinella,
but differs in having paraphyses and a round ostiole.
Halbania Racib., Crypt. Par. Java: no. 89 (1889).
Index Fungorum number: IF 2201; Facesoffungi number:
FoF 06735; – 3 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Halbania cyathearum Racib., Crypt. Par.
Java: no. 89 (1889).
Notes: Halbania is mostly similar to Batistinula in
having 3-septate ascospores, but lacks appressoria on the
hyphae. Sequence data is needed to confirm its phylogenetic
placement.
Prillieuxina G. Arnaud, Annals d’École National d’Agric.
de Montpellier, Série 2 16(1–4).
Index Fungorum number: IF 4365; Facesoffungi number:
FoF 06742; – 62 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Prillieuxina winteriana (Pazschke) G.
Arnaud, Annals d’École National d’Agric. de Montpellier,
Série 2 16(1–4): 162 (1918) [1917].
≡ Asterina winteriana Pazschke, Hedwigia 31(3): 104
(1892).
Notes: Prillieuxina is typical of Asterinaceae in having
stellate fissures and brown, 1-septate ascospores. However,
appressoria are very rare in Prillieuxina, and the cells of
the upper walls are radially arranged as in Microthyrium.
Meliolaster Höhn., Ber. dt. bot. Ges. 35(10): 701 (1918).
13
38
Guatimosim et al. (2015) provided the only sequence for
this genus.
Pycnocarpon Theiss., Abh. K.K. Zool.-Bot. Ges. Wien 7(3):
31 (1913).
Index Fungorum number: IF 4564; Facesoffungi number:
FoF 07606; – 4 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Pycnocarpon magnificum (Syd., P. Syd. &
E.J. Butler) Theiss., Abh. K.K. Zool.-Bot. Ges. Wien 7(3):
31 (1913).
≡ Asterina magnifica Syd., P. Syd. & E.J. Butler, Annls
mycol. 9(4): 391 (1911).
Notes: Wijayawardene et al. (2018) accepted this genus in
Dothideomycetes genera incertae sedis. However, Doilom
et al. (2018) included Pycnocarpon in Asterinaceae based
on its superficial, web-like hypha, flattened thyriothecia,
opening by radiating star-like or longitudinal splits, saccate
asci, and conglobose, hyaline to brown, 1-septate ascospores
strongly constricted at the septum (Doilom et al. 2018). We
accept Pycnocarpon in Asterinaceae but note that the upper
wall of ascomata of Pycnocarpon differs from members of
Asterinaceae in having radially arranged, subglobose cells
instead of cells of textura prismatica. Thus, sequence data
is needed to confirm its placement.
Schenckiella Henn., Bot. Jb. 17: 523 (1893).
Index Fungorum number: IF 4885; Facesoffungi number:
FoF 06743; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Schenckiella marcgraviae Henn., Bot. Jb.
17: 523 (1893).
Notes: This is a very unusual genus with a unique combination of characters, with surface hyphae lacking appressoria, Asterina-like thyriothecia, elongated clavate asci, brown,
cellular pseudoparaphyses, which are rarely observed in the
Dothideomycetes, and 4–5-septate brown ascospores.
Trichasterina G. Arnaud, Annals d’École National d’Agric.
de Montpellier, Série 2 16(1–4): 172 (1918) [1917].
Index Fungorum number: IF 5544; Facesoffungi number:
FoF 06744; – 11 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Trichasterina styracis (Theiss.) G. Arnaud,
Annals d’École National d’Agric. de Montpellier, Série 2
16(1–4): 172 (1918) [1917].
≡ Asterina styracis Theiss., Abh. K.K. Zool.-Bot. Ges.
Wien 7(3): 41 (1913).
Notes: Thyriothecia shares almost the same morphology
as Asterina, but with setae on the hyphae. Whether such a
character difference justifies separate genera should be tested
using molecular data.
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Fungal Diversity (2020) 105:17–318
Trichopeltospora Bat. & Cif., in Batista, Costa & Ciferri,
Publicações Inst. Micol. Recife 90: 17 (1958) [1957].
Index Fungorum number: IF 5569; Facesoffungi number:
FoF 06745; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Trichopeltospora pipericola Bat., Cif. &
C.A.A. Costa, in Batista, Costa & Ciferri, Publicações Inst.
Micol. Recife 90: 17 (1958) [1957].
Notes: Trichopeltospora was introduced as a member of
Microthyriaceae. Wu et al. (2011b) transferred Trichopeltospora to Asterinaceae based on its irregular ostiole and
appressoria on the hyphae. This transfer needs to be confirmed by phylogenetic analyses.
Uleothyrium Petr., Annls mycol. 27(5/6): 388 (1929).
Index Fungorum number: IF 5661; Facesoffungi number:
FoF 06762; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Uleothyrium amazonicum Petr., Annls
mycol. 27(5/6): 388 (1929).
Notes: Uleothyrium is similar to Platypeltella in having
rounded ostioles, which are distinct from any other genera
in Asterinaceae. However, Uleothyrium lacks appressoria,
while Platypeltella has intercalary appressoria.
Vizellopsis Bat., J.L. Bezerra & T.T. Barros, Publicações
Inst. Micol. Recife 637: 5 (1969).
Index Fungorum number: IF 5748; Facesoffungi number:
FoF 06746; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Vizellopsis grevilleae Bat., J.L. Bezerra &
T.T. Barros, Publicações Inst. Micol. Recife 637: 5 (1969).
Notes: Vizellopsis was introduced as a member of Microthyriaceae, and Lumbsch and Huhndorf (2010) included this
genus in Dothideomycetes incertae sedis. Dai et al. (2014a)
transferred Vizellopsis to Asterinaceae based on the small,
black thyriothecia, forming below the dark brown mycelium,
comprising radiating cells and a concentrically ridged surface, but strongly thickened and septate hyphae which is
different from other genera of Asterinaceae.
Economic and ecological significance
Asterinaceae species produce haustoria to gain nutrients
from host plants without causing pathogenic damage. They
may reduce photosynthesis by covering the host surface, and
increase the temperature and respiration in those areas as do
other black mildews.
Asterotexaceae Firmino, O.L. Pereira & Crous [as ‘Asterotexiaceae’], in Guatimosim et al., Persoonia 35: 238 (2015).
Index Fungorum number: IF 548079; Facesoffungi number: FoF 07607, 2 species.
Fungal Diversity (2020) 105:17–318
Epiphytes, phytopathogens, forming dark colonies irregular to star-shaped, solitary to confluent. External mycelium growing through ascomatal cavity or fusing with the
host epidermis cells, septate, hyaline, smooth. Appressoria formed underneath the ascomata, solitary or forming
in small clusters, globose, cone-shaped or ovoid to elongate, brown, with a central hyaline penetration peg. Sexual
morph: Ascomata superficial to erumpent, scutellate, dimidiate, brown to black. Scutellum of radially arranged rows of
cells, poorly developed base, opening by numerous irregular
fissures. Hamathecium comprising septate, anastomosing,
cellular pseudoparaphyses, embedded in a gelatinous matrix.
Asci 8-spored, bitunicate, fissitunicate, oblong to cylindrical, with short and rounded pedicel or pedicel sometimes
absent. Ascospores overlapping 2–3-seriate, ellipsoidal, hyaline to slightly yellowish, 1-septate, slightly constricted at
the septum, upper cell broader than lower cell (adapted from
Hongsanan et al. 2014b; Guatimosim et al. 2015). Asexual
morph: Undetermined.
Type: Asterotexis Arx.
Notes: Asterotexaceae was established by Guatimosim
et al. (2015), with the generic type Asterotexis. A phylogenetic tree provided by Guatimosim et al. (2015) showed that
species of Asterotexis formed a distinct clade sister to the
Inocyclus angularis (Incertae sedis clade). They introduced
Asterotexiales to accommodate Asterotexaceae (Guatimosim
et al. 2015). Ertz et al. (2016) indicated that the Asterotexiales clade contains Asterotexis species, Inocyclus angularia
(Parmulariaceae) and some Asterinales species. However,
the Asterotexiales clade in Ertz et al. (2016) was treated as
Asterinaceae sensu stricto (Hyde et al. 2016b). Asterotexales
was synonymized under Asterinales by Liu et al. (2017). In
our phylogenetic analyses (Fig. 5), Asterotexis species cluster with the clade of Asterina species as an unstable clade.
In another analysis, which did not include Lembosia mimusopis (data not shown), Asterotexis clustered with Inocyclus
angularis (Incertae sedis clade). Thus, we retain Asterotexaceae within Asterinales and note that more sequence data
are needed to clarify its phylogenetic placement.
Asterotexis Arx, Fungus, Wageningen 28: 6 (1958).
Index Fungorum number: IF 430; Facesoffungi number:
FoF 06766; 2 morphological species (Index Fungorum
2020), 1 species with molecular data.
Type species: Asterotexis cucurbitacearum (Rehm) Arx.
Notes: Asterotexis is a plant-pathogen found on leaves,
and was identified as a member of Asterinaceae (Inácio
and Cannon 2008; Guerrero et al. 2011; Hongsanan et al.
2014b). Morphologically and phylogenetically, the genus
could not be placed in any family of Asterinales (this study).
Thus, Asterotexis is placed in its own family, Asterotexaceae.
39
Asterotexis cucurbitacearum (Rehm) Arx [as ‘cucurbitarum’], Fungus, Wageningen 28: 6 (1958).
≡ Dothidella cucurbitacearum Rehm, Hedwigia 36(6):
376 (1897).
Index Fungorum number: IF 118911; Facesoffungi number: FoF 07608; Fig. 7
Description: see Hongsanan et al. (2014b).
Material examined: Brazil, Brazilia, Rio de Janeiro, on
Cucurbitaceae, May 1887, E. Ule 676, Ex Herb. Sydow
(SF7565, holotype); COSTA RICA, San José, Finca La
Caja, on surface of leaves of Sechium edule (Cucurbitaceae),
25 March 1927, Det. J.A. Stevensen (S-F220847).
Economic and ecological significance
Species in this family are phytopathogens. The appearance
of colonies on leaves can mainly reduce the photosynthesis,
respiration, disrupt other plants mechanism and can make
host tissues become pale.
Hemigraphaceae D.Q. Dai & K.D. Hyde, in Dai et al.,
MycoKeys 369(2): 67 (2018).
Index Fungorum number: IF 554062; Facesoffungi number: FoF 03910, 9 species.
Biotrophic on lichens. Sexual morph: Ascostromata solitary to gregarious, superficial, stellate, irregularly opening
from the centre to margin, conical in section, coriaceous,
black to dark brown. Peridium comprises two layers, black
and thick-walled cells of textura angularis in outer part, thin
and light brown cells of textura angularis at inner layers.
Hamathecium comprising few, brown, unbranched, filamentous, septate, cellular pseudoparaphyses. Asci 8-spored,
bitunicate, clavate to cylindric-clavate, subglobose, with an
ocular chamber and a short pedicel. Ascospores 3-seriate to
irregularly arranged, ellipsoid, brown, 1-septate, with larger
upper cell and narrower lower cell, smooth-walled. Asexual
morph: Undetermined.
Type: Hemigrapha (Müll. Arg.) R. Sant. ex D. Hawksw.
Notes: Dai et al. (2018) studied the syntype of Hemigrapha asteriscus (≡ Melanographa asteriscus), and concluded that it is different from the family type of Parmulariaceae. Phylogenetically (LSU), H. atlantica forms a distint
lineage within Asterinales (Ertz and Diederich 2015; Dai
et al. 2018; this study). Thus, Hemigraphaceae was established in Asterinales to accommodate a single genus Hemigrapha (Dai et al. 2018).
Hemigrapha (Müll. Arg.) R. Sant. ex D. Hawksw., Kew
Bull. 30(1): 9 (1975).
≡ Melanographa sect. Hemigrapha Müll. Arg., Flora,
Regensburg 65(33): 519 (1882).
Index Fungorum number: IF 2282; Facesoffungi number: FoF 02311; 8 morphological species (Index Fungorum
2020), 1 species with molecular data.
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40
Fungal Diversity (2020) 105:17–318
Fig. 7 Asterotexis cucurbitacearum (a, b, d, f from S-F7565, holotype and c, e, g–n from from S-F22084). a Herbarium specimen. b
Ascomata on substrate. c Ascoma when viewed in squash mount. d
Drawing from von Arx and Müller (1975). e Upper wall of ascoma. f
Asci. g Hamathecium. h–j Asci. k Ocular chamber strained in Melzer’s reagent. l, m 1-septate ascospores. n 2-septate ascospore. Scale
bars: c = 100 μm, e–j = 20 μm, k–n = 10 μm
Type species: Hemigrapha asteriscus (Müll. Arg.) R.
Sant. ex D. Hawksw., Kew Bull. 30(1): 191 (1975).
Notes: Hemigrapha was included in Parmulariaceae
by Lumbsch and Huhndorf (2010), Hyde et al. (2013) and
Wijayawardene et al. (2014a). However, it is dissimilar with
the family type of Parmulariaceae having dark, superficial,
star-shaped ascomata forming on a thallus of lichens (Diederich and Wedin 2000; Dai et al. 2018). Ertz and Diederich
(2015) included H. atlantica in their phylogenetic analysis
and indicated that this genus belongs to Asterinales based
on LSU sequence data, without assigning the family placement. The same result was shown in Dai et al. (2018) and in
this study (Fig. 5).
Index Fungorum number: IF 530383; Facesoffungi number: FoF 02312; Fig. 8
Description: see Dai et al. (2018).
Material examined: Australia, New South Wales, Mount
Kosciuszko, on thallus of Peltigera dolichorrhiza (Lichen).
Müller J. 1882 (G 00292584, syntype).
Hemigrapha asteriscus (Müll. Arg.) R. Sant. ex D.
Hawksw., Kew Bull. 30(1): 191 (1975).
≡ Melanographa asteriscus Müll. Arg., Flora, Regensburg 65(33): 519 (1882).
13
Economic and ecological significance
Members of this family are biotrophic on lichens and appear
as black colonies on the host surface.
Lembosiaceae Hosag, in Hosagoudar et al., J. Mycopathol.
Res. 39(1): 61 (2001).
Index Fungorum number: IF 80503; Facesoffungi numbers: FoF 07609, 160 species.
Epiphytic on living leaves. Superficial hyphae with lateral
appressoria. Sexual morph: Thyriothecia solitary, scattered,
superficial, oval, ellipsoidal, X- or Y-shaped, easily removed
Fungal Diversity (2020) 105:17–318
41
Fig. 8 Hemigrapha asteriscus (G 00292584, syntype). a, b Herbarium material. c, d Appearance of ascostromata on host surface.
d Ascostromata in the water. e Section of ascostroma. f Asci with
hamathecial tissues. g Ascospores. Scale bars: b = 10 mm, c, d = 500
µm, e = 50 µm, f, g = 5 µm
from the host surface, black, opening by a linear fissure,
with basal peridium poorly developed. Upper wall comprising linear, dark cells, which are branched at the margin.
Hamathecium comprising vertical asci inclined upwards,
pseudoparaphyses not observed. Asci bitunicate, fissitunicate dehiscence not observed, subglobose to ovoid, apedicellate, apical region of asci usually with a thick opaque
region, ocular chamber not observed, not staining blue in
IKI. Ascospores overlapping, oblong to obvoid, hyaline to
brown, 1-septate. Asexual morph: Undetermined.
Type: Lembosia Lév.
Notes: Hosagoudar et al. (2001) introduced Lembosiaceae
using the morphological character of elongate thyriothecia with longitudinal or X- or Y-shaped slits (Hongsanan
et al. 2014b). However, Hongsanan et al. (2014b) treated
Lembosiaceae as a synonym of Asterinaceae based on the
first phylogenetic evidence of LSU sequence data of Lembosia albersii. Here, we re-introduced Lembosiaceae using
sequence data currently available in GenBank.
Lembosia Lév., Annls Sci. Nat., Bot., sér. 3 3: 58 (1845).
Index Fungorum number: IF 2724; Facesoffungi number:
FoF 06736, – 160 morphological species (Species Fungorum
2020), 6 species with molecular data
Type species: Lembosia tenella Lév., Annls Sci. Nat.,
Bot., sér. 3 3: 58 (1845).
Notes: Species of Lembosia and Asterina are obligately
biotrophic, having appressoria and similar thyriothecia
(Hosagoudar et al. 2001). However, Lembosia differs from
Asterina in having elongate thyriothecia which dehisce to
13
42
open by a longitudinal or X- or Y-shaped slit (Hosagoudar
1991).
Lembosia xyliae X.Y. Zeng, T.C. Wen & K.D. Hyde, in
Ariyawansa et al., Fungal Diversity: 75: 50 (2015).
Index Fungorum number: IF 551345; Facesoffungi number: FoF 00933, Fig. 9
Description: see Ariyawansa et al. (2015a).
Material examined: Thailand, Chiang Rai, Mae Fah
Luang University, on leaves of Xylia sp. (Fabaceae), 18
January 2014, XY Zeng (MFLU 14-0004, holotype).
Economic and ecological significance
Species in Lembosiaceae are considered as obligatory biotrophs, however, there are no records available to show its
economic significance.
Melaspileellaceae D.Q. Dai & K.D. Hyde, in Dai et al., Phytotaxa 369(2): 70 (2018).
Index Fungorum number: IF 554063; Facesoffungi number: FoF 03911, 1 species.
Saprobic on trees and shrubs. Sexual morph: Ascostromata solitary, superficial, dark to black, coriaceous, small
rounded. Peridium comprises 2 layers, black and thickwalled cells at outer layers, light brown to hyaline cells of
textura angularis of inner layers. Hamathecium comprising
Fig. 9 Lembosia xyliae (MFLU 14-0004, holotype). a, b Colonies
on leaf surface. c Squash mount of ascomata with asci. d Upper
walls and hyphae with hyphopodia. e Immature ascospores. f Mature
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Fungal Diversity (2020) 105:17–318
dense, hyaline, mainly unbranched, filamentous, septate,
cellular pseudoparaphyses around asci, with brownish tips.
Asci 8-spored, bitunicate, broadly clavate to subglobose,
with a rounded apex and a short pedicel. Ascospores 2-seriate to irregularly arranged, ellipsoid, hyaline, 1-septate, with
slightly larger upper cell, with slightly narrower lower cell,
smooth-walled, bearing 2–3 appendages which disappear
when dry. Asexual morph: Undetermined (adapted from
Dai et al. 2018).
Type: Melaspileella (P. Karst.) Vain.
Notes: This family was established by Dai et al. (2018) to
accommodate a single genus Melaspileella based on phylogenetic placement generated from LSU and SSU of M. proximella. The family formed a clade sister to Hemigraphaceae
within Asterinales with high bootstrap support in Dai et al.
(2018), while sister to Pirozynskiella laurisilvatica (FMR
13133) genus incertae sedis in Asterinales in our analyses
(Fig. 5). Thus, we retain Melaspileellaceae in Asterinales.
Melaspileella (P. Karst.) Vain., Ann. Acad. Sci. fenn., Ser.
A 15(no. 6): 317 (1921).
= Mycomelaspilea Reinke, in Pringsheim, Jb. wiss. Bot.
28: 136 (1895).
Index Fungorum number: IF 3095; Facesoffungi number:
FoF 07733; 1 morphological species (Species Fungorum
2020), 1 species with molecular data.
ascospores. Scale bars: a = 400 μm, b = 200 μm, c = 100 μm, d = 50
μm, e–f = 10 μm
Fungal Diversity (2020) 105:17–318
Type species: Melaspileella proximella (Nyl.) Ertz &
Diederich.
Notes: Melaspileella was introduced by Vainio (1921b).
Ertz and Diederich (2015) lectotypified the genus with M.
proximella. They also transferred the genus to Asterinales
without assigning it to any family. A phylogenetic tree based
on LSU and SSU sequence data in Dai et al. (2018) indicated that M. proximella should be assigned in a new family,
Melaspileellaceae.
43
Melaspileella proximella (Nyl.) Ertz & Diederich, Fungal
Diversity 71: 161 (2015).
≡ Arthonia proximella Nyl., Lich. Scand. (Helsinki): 262
(1861).
Index Fungorum number: IF 811374; Facesoffungi number: FoF 07610; Fig. 10
Description: see Zeller and Tóth (1960).
Fig. 10 Melaspileella proximella. a Fruiting body on host
(redrawn from ERD-7986.
Photo: Enrique Rubio). b
Section through ascoma, in
water (redrawn from Finland,
H-NYL 4827, lectotype). c
Ascus with ascospores, in water
(redrawn from Finland, H-NYL
4827, lectotype). d Ascus
with ascospores (Hungary,
on Juniperus, Zeller & Tóth
2835, BP – paratype of Banhegyia setispora). e Ascospore
with polar setulae, in water
(redrawn from Belgium, Ertz
19187). f Ascospore with polar
setulae (redrawn from Finland,
Nylander, H – lectotype). Scale
bars: b = 20 µm, c, d = 10 µm,
e, f = 5 µm
13
44
Economic and ecological significance
Members of this family are saprobic on plants and involved
in recycling organic matter. Melaspileella proximella grows
on trees and shrubs including trunks of Pinus and Tilia, and
twigs of Juniperus (Ertz and Diederich 2015).
Morenoinaceae Hongsanan & K.D. Hyde, fam. nov.
Index Fungorum number: IF 557813; Facesoffungi number: FoF 07611, 26 species.
Saprobic on stem or leaves, forming blackened areas.
Sexual morph: Thyriothecia solitary, aggregated, or gregarious, superficial, easily removed from the host surface,
black, ellipsoid, oblong, curved, opening by X- or Y-shaped
or linear fissures, branched at the margin, from the center
to the outer rim, lacking free hyphae and appressoria at the
margin. Upper wall comprising linear, dark cells, which are
branched at the margin, radiating from the center to the outer
rim. Hamathecium pseudoparaphyses not observed. Asci
8-spored, bitunicate, fissitunicate dehiscence not observed,
subglobose to oblong or saccate to globose, apedicellate,
ocular chamber not observed, or with a distinct, thickened
apical region. Ascospores overlapping 2–3-seriate, oblong
to obvoid, or fusiform, hyaline, 1-septate, constricted at the
septum, with or without guttules, smooth-walled (Hongsanan et al. 2014b; Tibpromma et al. 2017). Asexual morph:
Coelomycetous. “Sirothyriella”, Pycnothyria circular, radially scutellate. Conidiogenous cells holoblastic, simple, hyaline. Conidia cylindrical, hyaline, 1-celled (asexual morph
from Sivanesan 1984; drawing of asexual characters can be
seen in Ellis 1980).
Type: Morenoina Theiss.
Notes: Morenoinaceae resembles Aulographaceae
(Hongsanan et al. 2014b). In our phylogenetic analyses
(Fig. 5), Morenoina calamicola is closely related to Morenoina palmicola (MFLUCC 15-0284) with 67% ML and
0.93 BYPP support as a distinct clade within Asterinales.
Therefore, we introduce Morenoinaceae to accommodate
the Morenoina clade.
Morenoina Theiss., Annls mycol. 11(5): 434 (1913).
= Aulographella Höhn., Ber. dt. bot. Ges. 35: 359 (1917).
Index Fungorum number: IF 3270; Facesoffungi number:
FoF 07612, 26 morphological species (Species Fungorum
2020), 2 species with molecular data.
Type species: Morenoina antarctica (Speg.) Theiss.
Notes: Morenoina was placed in Leptopeltidaceae and
Asterinaceae (von Arx and Müller 1975; Lumbsch and
Huhndorf 2010). Hongsanan et al. (2014b) designated an
epitype and placed this genus in Aulographaceae as Morenoina did not have free hyphae with appressoria as other
members of Asterinaeceae and it has different ascus form
from members of Leptopeltidaceae. Morenoina resembles
Aulographum and differs only in the morphology of the
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Fungal Diversity (2020) 105:17–318
scutellum which comprises inordinately arranged cells and
a hypostroma of subcuticular hyphae beneath the thyriothecium, which is lacking in Morenoina (Ellis 1980). Phylogenetic placement of Aulographaceae is uncertain due to lack
of sequence data. Sequence data of Aulographum hederae
(type species) formed an unstable clade, while two species of Morenoina form a distinct clade within Asterinales
(Fig. 5). Therefore, we place Morenoina in the new family
Morenoinaceae.
Morenoina antarctica (Speg.) Theiss., Annls mycol. 11(5):
434 (1913).
≡ Morenoella antarctica Speg., Boln Acad. nac. Cienc.
Córdoba 11(2): 240 (1887) [1888].
Index Fungorum number: IF 174884; Facesoffungi number: FoF 07613; Fig. 11
Material examined: Argentina, Tierra del Fuego, Ushuaia, south of Paso e Garibaldi, on the stem of Gramineae,
29 October 1989, collected by P.F. Cannon and D.W. Minter,
identified by P.F. Cannon (IMI 349680, epitype).
Economic and ecological significance
Species of this family are saprobic and play a role in recycling organic matter.
Neobuelliellaceae Hongsanan & K.D. Hyde, fam. nov.
Index Fungorum number: IF 558103;
Facesoffungi number: FoF 07614, 1 species.
Lichenicolous. Vegetative hyphae intercellular. Sexual
morph: Ascomata apothecioid, dispersed over the host thallus as black dots, grouped, rarely single, rounded, globose
when young, pressed at the edges, oval when mature, often
bleaching the host thallus, occasionally sunken. Excipulum
dark brown, K+ greenish-brown. Hymenium hyaline to
brownish. Disc olive-brown, I-, KI-. Hamathecium comprising branched, anastomosing, cellular pseudoparaphyses,
capitate at apical and intercalary cells with pigment, thick at
tips, with dark-brown pigment. Asci 4–6-spored, bitunicate,
fissitunicate, cylindrical. Ascospores long, cells rounded, the
upper cell sometimes tapered, hyaline, later ± brown, 1-septate, clearly constricted at the septa, thin-walled (adapted
from Hafellner et al. 2008; Etayo 2010; Yazici and Etayo
2013). Asexual morph: Undetermined.
Type: Neobuelliella Hongsanan & K.D. Hyde
Notes: Neobuelliellaceae is similar to species of Buelliella which are placed in genus incertae sedis in Dothideomycetes. Buelliella minimula (type species) together with
B. physciicola cluster within Stictographaceae (Asterinales) in the phylogenetic analyses of Dai et al. (2018) and
this study. Two strains of Neobuelliella poetschii (≡ Buelliella poetschii) form a distinct clade separately from Stictographaceae, and are sister to Hemigraphaceae (Ertz et al.
Fungal Diversity (2020) 105:17–318
45
Fig. 11 Morenoina antarctica (IMI 349680, epitype). a, b Herbarium and specimen. c, d Appearance of thyriothecia on surface of
plant. e Squash mount of thyriothecium. f Upper wall of thyriothe-
cium. g–h Asci when immature. i Asci at maturity. j–l Ascospores.
Scale bars: e = 100 μm, f = 50 μm, g–l = 20 μm
2015; Dai et al. 2018; this study). Therefore, we introduce
a new family Neobuelliellaceae to accommodate this clade.
et al. 2008; Etayo 2010; Yazici and Etayo 2013). Asexual
morph: Undetermined.
Type species: Neobuelliella poetschii (Hafellner) Hongsanan & K.D. Hyde.
Notes: The genus is introduced to accommodate Neobuelliella poetschii which was known as Buelliella poetschii.
Phylogenetic analyses from Dai et al. (2018) and this study
(Fig. 5) indicate that it does not cluster with the type species
of Buelliella. Thus, we introduce the new genus Neobuelliella to accommodate a single species Buelliella poetschii
(current name is Neobuelliella poetschii).
Neobuelliella Hongsanan & K.D. Hyde, gen. nov.
Index Fungorum number: IF 557815; Facesoffungi number: FoF 07615; 1 morphological species (this study), 1 species with molecular data.
Lichenicolous. Vegetative hyphae intercellular. Sexual
morph: Ascomata apothecioid, dispersed over the host thallus as black dots, grouped, rarely single, rounded, globose
when young, pressed at the edges, oval when mature, often
bleaching the host thallus, occasionally sunken. Hymenium
hyaline to brownish. Disc olive-brown, I-, KI-. Excipulum
dark brown, K+ greenish-brown. Hamathecium comprising
branched, anastomosing, cellular pseudoparaphyses, capitate
at apical and intercalary cells with pigment, thick at tips,
with dark-brown pigment. Asci 4–6-spored, bitunicate, fissitunicate cylindrical. Ascospores long, cells rounded, the
upper cell sometimes tapered, hyaline, later ± brown, 1-septate, clearly constricted at the septa, thin-walled (Hafellner
Neobuelliella poetschii (Hafellner) Hongsanan & K.D.
Hyde, comb. nov.
≡ Buelliella poetschii Hafellner, in Hafellner, Herzog &
Mayrhofer, Mitt. naturw. Ver. Steierm. 137: 187 (2008).
Index Fungorum: IF 557814, Facesoffunginumber: FoF
07616; Fig. 12
Description: see Yazici and Etayo (2013).
13
46
Fungal Diversity (2020) 105:17–318
1-septate ascospores (viz. Karschia, Labrocarpon, Melaspileopsis and Stictographa; Dai et al. 2018). In the phylogenetic tree, these four genera clustered as a distinct clade
within Asterinales (Dai et al. 2018). As the type species of
Buelliella clustered within the clade of Stictographaceae, we
therefore include Buelliella in Stictographaceae.
Fig. 12 Neobuelliella poetschi . a Apothecioid ascomata (redrawn
from Yazici and Etayo 2013). b Ascospores (redrawn from holotype in Hafellner et al. 2008). c Young ascus among hamathecium
(redrawn from Ertz and Diederich 2015)
Notes: Phylogenetically, Buelliella poetschii formed a
distinct lineage within Asterinales, and did not cluster with
B. minimula (type species of Buelliella). Thus, B. poetschii
is synonymized in Neobuelliella.
Economic and ecological significance
Neobuelliellaceae comprises lichenicolous taxa which play
insignificant roles in ecosystems. There is only a single species in this family. Fresh collections are needed to understand the ecological significance of Neobuelliellaceae.
Stictographaceae D.Q. Dai & K.D. Hyde, in Dai et al.,
MycoKeys 369(2): 70 (2018).
Index Fungorum number: IF 554064; Facesoffungi number: FoF 03912, 39 species.
Lichenicolous. Sexual morph: Ascomata solitary, scattered to clustered, superficial, cymbiform to lirelliform, with
slit-like disc, with a slit or star-shaped fissure, black to dark
brown. Peridium thick, outer layers of black to dark brown,
thick-walled cells of textura angularis, inner layers of light
brown to hyaline cells of textura angularis. Hamathecium
comprising dense, filamentous, septate, unbranched or
occasionally branched at upper part around the asci, cellular pseudoparaphyses. Asci 8-spored, bitunicate, clavate to
subglobose, with an ocular chamber, apically rounded, short
pedicellate. Ascospores 2-seriate to irregularly arranged,
ellipsoid, hyaline, becoming light brown to dark brown,
1-septate, with slightly larger upper cell, and narrower lower
cell, smooth-walled. Asexual morph: Undetermined (Dai
et al. 2018).
Type: Stictographa Mudd.
Notes: Stictographaceae was introduced to accommodate
four lichenicolous or saprobic genera which have cymbiform
to lirelliform, or a slit-like discs, black to dark brown ascomata, wide clavate to subglobose asci, and ellipsoid, brown,
13
Stictographa Mudd, Man. Brit. Lich.: 226 (1861).
Index Fungorum number: IF 5238; Facesoffungi number:
FoF 07617; 1 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Stictographa lentiginosa (Lyell ex Leight.)
Mudd.
Notes: Stictographa can be compared with Labrocarpon
in having lichenicolous, black, simple to occasionally short,
branched, lirelliform ascomata with a slit-like opening (Ertz
and Diederich 2015; Diederich et al. 2017). Ertz and Diederich (2015) inferred that Stictographa and Labrocarpon
might be synonyms, however, they do not phylogenetically
cluster together (Ertz and Diederich 2015; Dai et al. 2018).
Thus, Labrocarpon is accepted as a distinct genus. More
collections and species of these genera are needed to clarify
their relationship.
Stictographa lentiginosa (Lyell ex Leight.) Mudd, Man.
Brit. Lich.: 226 (1861).
≡ Opegrapha lentiginosa Lyell ex Leight., Ann. Mag. nat.
Hist., Ser. 2 13: 211 (1854).
Index Fungorum number: IF 406525; Facesoffungi number: FoF 07618; Fig. 13
Description: see Redinger (1938), Sanderson et al.
(2009), Ertz and Diederich (2015).
Other genera included
Buelliella Fink, Lich. Fl. U.S.: 372 (1935).
Index Fungorum number: IF 676; Facesoffungi number:
FoF 03625; 12 morphological species (Species Fungorum
2020, this study), 2 species with molecular data.
Type species: Buelliella minimula (Tuck.) Fink, Lich. Fl.
U.S.: 372 (1935).
≡ Buellia minimula Tuck., Syn. N. Amer. Lich. (Boston)
2: 106 (1888).
Notes: The original diagnosis of this genus was provided
by Fink (1935), and valided by Hafellner (1979). Buelliella
is characterised by a non-lichenized, lichenicolous habit,
initially subsphaerical and closed ascomata, with the upper
ascomatal breaking gradually away to expose the hymenium when mature, becoming apothecioid, with bitunicate,
broadly clavate to subcylindrical asci, and a distinct ocular
chamber, and ellipsoid, 1-septate, pale to medium brown
ascospores. Ertz and Diederich (2015) noted that the genus
is not monophyletic due to B. physciicola and B. poetschii
being distantly related, and sequence data of the type species
Fungal Diversity (2020) 105:17–318
47
Fig. 13 Stictographa lentiginos
(redrawn Ertz and Diederich
2015). a Ascomata growing on
lichen. b Clavate to subglobose
asci and pseudoparaphyses. c
Ascospores. Scale bars: a = 200
µm, b, c = 10 µm
(B. minimula) was unavailable. Thus, it was placed as incertae sedis in Dothideomycetes (Kirk et al. 2008; Ertz and
Diederich 2015; Wijayawardene et al. 2017a). We included
sequence data of B. minimula in our analyses and found that
it forms a lineage within Sticgraphaceae. Buelliella poetschii
is transferred to a new genus, Neobuelliella in this paper.
Karschia Körb., Parerga lichenol. (Breslau) 5: 459 (1865).
Index Fungorum number: IF 2545; Facesoffungi number:
FoF 07619; – 24 morphological species (Species Fungorum
2020), 2 species with molecular data.
Type species: Karschia talcophila (Ach.) Körb., Parerga
lichenol. (Breslau) 5: 460 (1865).
≡ Lecidea talcophila Ach., Lich. univ.: 183 (1810).
Notes: More than 100 species of Karschia are listed in
Index Fungorum (2020), however, most taxa have been
transferred to other genera, such as Buelliella, Cycloschizon, Dothidea and others by Hafellner (1979). Two species
were accepted by Hawksworth et al. (1995), and later four
species were included (Kirk et al. 2008). Karschia is characterized by a lichenicolous life-style, immersed to erumpent,
rounded, black ascomata with a large, irregular opening,
producing pseudoparaphyses with a slightly enlarged and
brown apex, bitunicate, clavate to subcylindrical asci, ellipsoid, brown, smooth ascospores with one septum (Ertz and
Diederich 2015).
Labrocarpon Etayo & Pérez-Ort., in Pérez-Ortega & Etayo,
Lichenologist 42(3): 271 (2010).
Index Fungorum number: IF 515228; Facesoffungi number: FoF 07620; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Labrocarpon canariense (D. Hawksw.)
Etayo & Pérez-Ort. [as ‘canariensis’], in Pérez-Ortega &
Etayo, Lichenologist 42(3): 272 (2010).
≡ Melaspilea canariensis D. Hawksw., Lichenologist
14(1): 84 (1982).
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48
Fungal Diversity (2020) 105:17–318
Lasiodiplodia crassispora A1S3-D20
Lasiodiplodia crassispora CBS 118741
Lasiodiplodia pyriformis CBS 121770
Lasiodiplodia pseudotheobromae CBS 116459
Lasiodiplodia lignicola MFLUCC 11-0656
76/1.00
Lasiodiplodia rubropurpurea CBS 118740
100/1.0 Phaeobotryon negundinis MFLUCC 15-0436
Phaeobotryon rhois CFCC 89662
99/1.0
Barriopsis stevensiana CBS 174.26
83/1.0
Alanphillipsia aloetica CPC 21109
Alanphillipsia euphorbiae CPC 21628
Oblongocollomyces variabilis CBS 121774
89/1.0
73/0.99
Sphaeropsis porosa CBS 110496
Diplodia rosulata CBS 116470
83/1.0
Diplodia cupressi CBS 168.87
Neodeightonia palmicola MFLUCC 10-0822
99/1.0
Tiarosporella corymbiae CBS 142532
Tiarosporella dactylidis MFLUCC 13-0276
97/1.0
Tiarosporella graminis var. karoo CBS 118718
75/0.98
Botryobambusa fusicoccum MFLUCC 11-0143
100/1.0 Dothiorella viticola CBS 117009
97/1.0
Dothiorella citricola ICMP 16828
Dothiorella ostryae MFLU 18-0177
84/0.94
Dothiorella sarmentorum IMI 63581b
97/1.0
Dothiorella iberica CBS 115041
Neofusicoccum luteum CBS 110299
99/1.0 Botryosphaeria fusispora MFLUCC 10-0098
Botryosphaeria agaves MFLUCC 11-0125
72/1.0
Cophinforma atrovirens MFLUCC 11-0425
-/1.0
93/1.0
Macrophomina phaseolina CBS 227.33
Neoscytalidium dimidiatum CBS 145.78
Endomelanconiopsis endophytica CBS 120397
80/0.83 Phyllosticta cussonia CPC14875
99/1.0 Guignardia vaccinii CBS 126.22
75/1.0
93/1.0
Phyllosticta citrichinaensis ZJUCC200956
100/1.0
Phyllosticta hostae CGMCC 3.14355
89/1.0
100/1.0
Phyllosticta hostae JZB3290001
Phyllosticta citricarpa CBS 102374
78/0.97
Phyllosticta citriasiana CBS 120486
100/1.0
71/1.0
Phyllosticta abieticola CBS 112067
Phyllosticta carissicola CPC 25665
Pseudofusicoccum ardesiacum CMW26159
100/1.0 Pseudofusicoccum stromaticum CBS 117448
-/1.0
Kellermania ramaleyae BPI 882823
100/1.0
Kellermania yuccigena BPI 882828
97/1.0
Umthunziomyces hagahagensis CPC 29917
100/1.0 Melanops sp. NJ-2018b isolate 145-1
-/0.86
Melanops tulasnei CBS 116805
Aplosporella longipes CFCC 89661
Aplosporella javeedii CFCC 89657
95/1.0
Aplosporella javeedii HKAS 102118
Aplosporella macropycnidia culture CGMCC 3.17725
99/1.0
99/1.0
Aplosporella africana CMW 25424
Aplosporella yalgorensis MUCC512
100/1.0
-/0.84
Aplosporella prunicola CBS 121167
Alanomyces indica MMI00067
Saccharata capensis CBS 122693
98/1.0
Saccharata hakeicola CPC 29274
99/1.0
Saccharata eucalypti CPC 25529
100/1.0 Saccharata eucalyptorum CPC 29222
100/1.0
Septorioides pini-thunbergii CBS 473.91
100/1.0 Patellaria quercus BHI F828a
Patellaria quercus CPC 27232
100/1.0
13
Botryosphaeriaceae
Phyllostictaceae
Botryosphaeriales
0.05
-/0.75
100/1.00
Planistromellaceae
Melanopsaceae
Aplosporellaceae
Saccharataceae
Outgroup
Fungal Diversity (2020) 105:17–318
◂Fig. 14 Phylogram generated from maximum likelihood analysis
(RAxML) of Botryosphaeriales based on ITS and LSU sequence
data. Maximum likelihood bootstrap values equal to or greater than
70%, Bayesian posterior probabilities equal to or greater than 0.90
(MLBS/PP) are given at the nodes. Isolate numbers are noted after
each species name. The tree is rooted to Patellaria quercus (CPC
27232 and BHI F828a). Newly sequence data generated in this study
are in blue. Ex-type and references strains are indicated in bold.
Hyphen (-) represents support values less than 70% MLBS and 0.90
PP
Notes: Labrocarpon is a monotypic genus which was
introduced by Pérez-Ortega and Etayo (2010) based on morphology of ascomata with periphyses (Ertz and Diederich
2015). Labrocarpon is more or less similar to Stictographa
(Pérez-Ortega and Etayo 2010; Ertz and Diederich 2015).
However, it formed a separate clade from Stictographa
(Ertz and Diederich 2015; Dai et al. 2018). Labrocarpon
is characterized by a lichenicolous life-style, superficial,
rounded to slightly elliptic, black, disk-like ascomata with
a slit-like opening and a well-developed exciple, producing
pseudoparaphyses with a swollen and brown apex, bitunicate, clavate, thick-walled asci with an ocular chamber, and
two-celled ascospores (Ertz and Diederich 2015).
Melaspileopsis (Müll. Arg.) Ertz & Diederich, Fungal
Diversity 71: 161 (2015).
Index Fungorum number: IF 811387; Facesoffungi number: FoF 07621; – 1 morphological species (Dai et al. 2018),
1 species with molecular data.
Type species: Melaspileopsis diplasiospora (Nyl.) Ertz &
Diederich, Fungal Diversity 71: 161 (2015).
≡ Opegrapha diplasiospora Nyl., Acta Soc. Sci. fenn.
7(2): 476 (1863).
Notes: The genus was established by Ertz and Diederich (2015), with the type species M. diplasiospora (≡
Melaspilea diplasiospora in Index Fungorum 2020). Phylogenetically, it does not belong to Melaspileaceae but is
sister to Stictographa within Stictographaceae (Dai et al.
2018). This genus was placed in Melaspileaceae, order
Eremithallales (Wijayawardene et al. 2018). However, we
agree with Dai et al. (2018) to retain Melaspileopsis within
Stictographaceae.
Economic and ecological significance
Stictographaceae accommodates several lichenicolous
and one saprobic genus. Members of this family usually
occur on the thallus of lichens. They are distributed in tropiccal to temperate areas. Stictographa lentiginosa is much
rarer than the host (Phaeographis dendritica) and thus can
act as a useful indicator of the quality of ancient woodlands
in western Europe (Ertz and Diederich 2015).
Botryosphaeriales C.L. Schoch, Crous & Shoemaker.
49
Index Fungorum number: IF 501513; Facesoffungi number: FoF 07659.
Botryosphaeriales was introduced to accommodate a single family Botryosphaeriaceae (Schoch et al. 2006). Previously, nine families (Aplosporellaceae, Botryosphaeriaceae,
Melanopsaceae, Phyllostictaceae, Planistromellaceae, Saccharataceae, Septorioideaceae, Endomelanconiopsidaceae,
Pseudofusicoccaceae) were accepted in Botryosphaeriales
in the Outline of Ascomycetes (Wijayawardene et al. 2018).
However, currently only six families (Aplosporellaceae,
Botryosphaeriaceae, Melanopsaceae, Phyllostictaceae,
Planistromellaceae, Saccharataceae) are accepted in this
order (Phillips et al. 2019), the remaining three families were
synonymized with existing families. Thus, Endomelanconiopsidaceae is considered a synonym of Botryosphaeriaceae,
Pseudofusicoccaceae a synonym of Phyllostictaceae and
Septorioideaceae a synonym of Saccharataceae (Phillips et al. 2019). Species in the order are characterized by
uni- or multilocular ascostromata with multi-layered dark
brown walls, occurring individually or in clusters, frequently
embedded in stromatic tissue. Asci are bitunicate, with a
thick endotunica, pedicellate or sessile, clavate, with a welldeveloped ocular chamber, intermixed with hyaline, septate
pseudoparaphyses, branched or not. Ascospores are hyaline
or pigmented, septate or not, ellipsoid to ovoid, with or
without mucoid appendages or mucilaginous sheath. Conidiomata are pycnidial, uni- to multilocular, frequently embedded in stromatic tissue. Conidiogenous cells are hyaline,
phialidic or annelidic. Conidia are hyaline or pigmented,
septate or not, thin- or thick-walled, with or without mucoid
appendages or sheaths (Phillips et al. 2019). The divergence
time for Botryosphaeriales is estimated as 150 MYA (stem
age, Hongsanan et al. 2020) (Fig. 14).
Accepted families: Aplosporellaceae, Botryosphaeriaceae, Melanopsaceae, Phyllostictaceae, Planistromellaceae, Saccharataceae.
Aplosporellaceae Slippers, Boissin & Crous, Stud. Mycol.
76(1): 41 (2013).
Index Fungorum: IF 805795; Facesoffungi number: FoF
00113, 13 species.
Sexual morph: Stromata pseudothecial, multiloculate
ascomata with multi-layered, dark brown walls, embedded in
stromatic tissue. Hamathecium comprising hyphoid, hyaline
pseudoparaphyses. Asci bitunicate, mostly with a thick endotunica, clavate, with a well-developed ocular chamber and
intermixed with pseudoparaphyses. Ascospores 1–2-seriate,
ellipsoid to ovoid, mostly aseptate, or septate, hyaline to
pigmented, lacking mucoid appendages or sheaths (Slippers et al. 2013; Liu et al. 2012; Thambugala et al. 2014a;
photoplates of sexual characters can be seen in Ekanayaka
et al. 2016). Asexual morph: Coelomycetous. Conidiomata
pycnidial, multilocular, embedded in stromatic tissues.
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Fungal Diversity (2020) 105:17–318
Fig. 15 Asexual morph of Aplosporella javeedii (HKAS 102118).
a Substrate. b Conidiomata on substrate. c Cross section of multiloculate conidioma. d Conidioma wall. e, f Different stages of con-
idiogenesis. g, h Conidia. i Spermatogenous cells. j Spermatia. Scale
bars: c = 1.5 mm, d = 100 μm, e–h, j = 10 μm, i = 25 μm
Conidiophores reduced to conidiogenous cells. Conidiogenous cells hyaline, phialidic. Conidia ellipsoid to subcylindrical, initially hyaline, thin-walled and smooth, becoming pigmented, thick-walled and spinulose, aseptate (Sutton
1980).
Type: Aplosporella Speg.
Notes: Aplosporellaceae was introduced by Slippers
et al. (2013) to accommodate Aplosporella and Bagnisiella.
Currently two genera (Aplosporella and Alanomyces) are
accepted in this family. This family is phylogenetically close
to Botryosphaeriaceae. Liu et al. (2017), using evolutionary evidence, questioned whether this family was introduced
unnecessarily and should be regarded as a genus. However,
based on morphology, phylogeny and evolutionary divergence times, Phillips et al. (2019) regarded Aplosporellaceae
as a distinct family.
Index Fungorum number: IF 7191; Facesoffungi number:
FoF 01427; 12 morphological species (botryosphaeriales.org
2020), 10 species with molecular data.
Type species: Aplosporella chlorostroma Speg.
Notes: Aplosporella was introduced by Spegazzini (1880).
This genus has frequently been incorrectly referred to as
“Haplosporella” (Tilak and Rao 1964; Petrak and Sydow
1927; Tai 1979; Wei 1979). About 340 epithets for Aplosporella are listed in Index Fungorum (July 2019). This
genus is characterised by multiloculate pycnidia, brown,
aseptate, spinulose conidia, hyaline, aseptate spermatia and
fusiform paraphyses (Du et al. 2017). Its sexual morph is
not common (Ekanayaka et al. 2016). Slippers et al. (2013)
indicated that Aplosporella should be separate from Botryosphaeriaceae as a distinct family, which they named
Aplosporellaceae, and confirmed that Aplosporella is wellresolved with species described from living culture. They
also found a close genetic relationship between the genera
Aplosporella and Bagnisiella and considered that Bagnisiella represents the sexual morph and should be reduced
to synonymy with Aplosporella (Schoch et al. 2009a; Slippers et al. 2013). Ekanayaka et al. (2016) introduced a new
Aplosporella Speg., Anal. Soc. cient. argent. 10(4): 157
(1880).
= Bagnisiella Speg., Anal. Soc. cient. argent. 10(4): 146
(1880).
13
Fungal Diversity (2020) 105:17–318
51
species Aplosporella thailandica from Thailand with morphology corresponding to Bagnisiella. Phillips et al. (2019)
formally placed Bagnisiella as a synonym of Aplosporella.
The most recent phylogenetic tree of species in Aplosporella
is provided by Hyde et al. (2020a).
Some taxa in Aplosporellaceae are recorded as plant
pathogens. Aplosporella prunicola was recorded as a probable tree pathogen on gymnosperms (Aylward et al. 2017).
Aplosporella beaumontiana was recorded as a major pathogen associated with a disease of okra (Yan et al. 2018).
Aplosporella javeedii Jami, Gryzenh., Slippers & M.J.
Wingf., Fungal Biology 118(2): 174 (2013).
Index Fungorum number: IF 803637; Facesoffungi number: FoF 07383; Fig. 15
Description: see Phillips and Alves (2009).
Saprobic on dead stems. Sexual morph: Undetermined.
Asexual morph: Conidiomata 1–2 mm diam., erumpent
from bark surface, separate, discoid, dark-brown to black,
multi-locular. Peridium composed cells of textura angularis,
4–10 layers, thick-celled, dark-brown to black cells at outer
layers, thin-walled and hyaline cells at inner layers. Conidiophores reduced to conidiogenous cells. Conidiogenous cells
hyaline, holoblastic, smooth, cylindrical. Conidia 18–25 ×
9–13 μm ( x̄ = 20 × 11 μm, n = 20) ellipsoidal to subcylindrical, with rounded ends, initially light brown, slightly
granulated walls, becoming dark brown (black in mass),
aseptate. Spermatophores reduced to Spermatogenous cells,
occurring intermingled among conidiogenous cells in same
conidioma, subcylindrical, hyaline, smooth. Spermatia 4–7
× 1–3 μm ( x̄ = 6 × 2 μm, n = 20) subcylindrical, straight or
slightly curved, hyaline, smooth-walled, granular.
Material examined: China, Yunnan Province, Kunming,
Kunming Institute of Botany, Botanical Garden, on dead
stems, 24 May 2018, A.H. Ekanayaka (HKAS 102118).
GenBank number: ITS: MW136694, LSU: MW142386,
SSU: MW127178
Notes: This collection is morphologically and phylogenetically similar to the type specimen of Aplosporella javeedii
from South Africa (Jami et al. 2014). Comparison of ITS
and LSU sequence data of this collection with sequence data
of A. javeedii (CMW38165) has 99% similarity. However,
spermatia were found in HKAS 102118.
Botryosphaeriaceae Theiss. & Syd. [as ‘Botryosphaeriacae’], Annls mycol. 16(1/2): 16 (1918).
Index Fungorum number: IF 80530; Facesoffungi number: FoF 00116, 279 species.
Endophytic, pathogenic and saprobic on a wide range
of hosts worldwide. Sexual morph: Ascomata uniloculate,
solitary or clustered, becoming fully or partially erumpent
with maturity, wall multi-layered, dark brown, infrequently
embedded in stromatic tissues. Asci 8-spored, bitunicate,
fissitunicate, with well-developed ocular chamber, thick
endotunica, short pedicellate, clavate. Pseudoparaphyses
cellular, intermixed with asci, hyaline, septate, frequently
constricted at septa, hyphae-like, branched or not, deliquescing with maturity. Ascospores 2–3-seriate, fusoid to ellipsoid or ovoid, hyaline or pigmented, smooth to verruculose,
septate or not, mucoid sheath mostly absent (photoplates of
sexual characters can be seen in Phillips et al. 2013, 2019).
Asexual morph: Coelomycetous. Conidiomata uni or rarely
multiloculate, pycnidial, infrequently embedded in stromatic
tissue. Conidiophores mostly reduced to conidiogenous
cells. Conidiogenous cells hyaline, phialidic, proliferating
percurrently or internally giving rise to periclinal thickenings, or proliferating percurrently forming annelations, without collarettes. Conidia hyaline or pigmented, aseptate or
one or multi-septate, sometimes muriform, smooth or striate,
thin- or thick-walled, without mucoid sheaths or appendages. Spermatogonia similar to conidiomata in anatomy.
Spermatogenous cells ampulliform to lageniform or subcylindrical, hyaline smooth, phialidic. Spermatia developing
in conidiomata or spermatogonia, hyaline, smooth, granular,
subcylindrical or dumbbell-shaped, with rounded ends. (Liu
et al. 2012; Slippers et al. 2013; Phillips et al. 2013).
Type: Botryosphaeria Ces. & De Not.
Notes: Botryosphaeriaceae was introduced by Theissen
and Sydow (1918) for three genera, Botryosphaeria, Phaeobotryon and Dibotryon. Over decades of taxonomic revisions and updates based on morphology, the family became
increasingly complex. However, based on LSU sequence
data Crous et al. (2006a) revealed ten lineages within Botryosphaeriaceae, which they considered to represent individual genera. Phillips et al. (2008) defined, introduced
and reinstated a further five genera in Botryosphaeriaceae.
Aplosporella was shown to reside in Botryosphaeriaceae by
Damm et al. (2007). Endomelanconiopsis was assigned to
this family by Rojas et al. (2008) while Phillips and Alves
(2009) considered that Melanopsis is another genus in Botryosphaeriaceae. Two new genera (Botryobambusa and
Other genus included
Alanomyces Roh. Sharma, Phytotaxa 297(2): 170 (2017).
Index Fungorum number: IF 804153; Facesoffungi number: FoF 07623; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Alanomyces indica Roh. Sharma, Phytotaxa 297(2): 172 (2017).
Notes: Alanomyces, introduced by Sharma et al. (2017), is
characterised by rapidly spreading dark greenish black colonies. Pycnidia are multilocular with a long neck and cylindrical, hyaline, guttulate, single celled spermatia (Sharma
et al. 2017).
Economic and ecological significance
13
52
Cophinforma) were introduced by Liu et al. (2012). Phillips
et al. (2013) resolved Botryosphaeriaceae into 17 genera and
110 species known from culture. Thereafter six new genera
and 85 new species/species combinations have been introduced (Dissanayake et al. 2016). According to a recent study
by Phillips et al. (2019) currently 22 genera are accepted
in Botryosphaeriaceae. In earlier studies Dothiorella and
Spencermartinsia were regarded as two separate genera in
the family (Phillips et al. 2008, 2013). However, Yang et al.
(2017) synonymized Spencermartinsia under Dothiorella.
These two genera are morphologically distinguishable only
in sexual morphs by the hyaline apiculi in Spencermartinsia.
So far, these characters are reported only in S. viticola but no
other species in this genus have been observed with asexual
morphs. Furthermore, phylogenetic distance is also small
between these two genera. Therefore, in this paper we accept
that Spencermartinsia is a synonym of Dothiorella. Phyllostictaceae was re-instated for Phyllosticta (Wikee et al.
2013a). Another three families were introduced in Botryosphaeriales by Slippers et al. (2013), namely Saccharataceae for Saccharata, Aplosporellaceae for Aplosporella and
Melanopsaceae for Melanops. Wyka and Broders (2016)
introduced Septorioideaceae for Septorioides while Yang
et al. (2017) raised Endomelanconiopsis and Pseudofusicoccum to familial status as Endomelanconiopsidaceae and
Pseudofusicoccaceae respectively. Taking into account morphology, phylogeny and evolutionary divergence times, Phillips et al. (2019) synonymised Endomelanconiopsidaceae
with Botryosphaeriaceae, Pseudofusicoccaceae with Phyllostictaceae, and Septorioideaceae with Saccharataceae. In
addition to that, Pilgeriella which fit within Botryosphaeriaceae, but do not have cultures or DNA (Wijayawardene
et al. 2017a). Therefore, we place this genus incertae sedis
in Botryosphaeriales.
Botryosphaeria Ces. & De Not., Comm. Soc. crittog. Ital.
1(fasc. 4): 211 (1863).
Index Fungorum number: IF 635; Facesoffungi number:
FoF 00141; 283 morphological species (Index Fungorum
January, 2020), 13 species with molecular data.
Type species: Botryosphaeria dothidea (Moug.) Ces. &
De Not.
Notes: Botryosphaeria was introduced by Cesati and De
Notaris (1863). Species in Botryosphaeria, based on the type
species B. dothidea, are characterised by ascospores that
are hyaline and aseptate, although they can become pale
brown and septate with age (Shoemaker 1964; Sivanesan
1984; Denman et al. 2000; Alves et al. 2004; Phillips et al.
2005; Jayawardena et al. 2018).
Other genera included
Alanphillipsia Crous & M.J. Wingf., in Crous et al., Persoonia 31: 197 (2013).
13
Fungal Diversity (2020) 105:17–318
Index Fungorum number: IF 805816; Facesoffungi number: FoF 01417; – 5 morphological species (Species Fungorum 2020), 5 species with molecular data.
Type species: Alanphillipsia aloes Crous & M.J. Wingf.,
in Crous et al. Persoonia 31: 197 (2013).
Notes: Alanphillipsia was introduced by Crous et al.
(2013) to accommodate species that are aplosporella-like
in morphology, but have conidia with a hyaline outer layer.
Barriopsis A.J.L. Phillips, A. Alves & Crous (2008).
Index Fungorum number: IF 511712; Facesoffungi number: FoF 01679; – 6 morphological species (Species Fungorum 2020), 6 species with molecular data.
Type species: Barriopsis stevensiana A.J.L. Phillips &
Pennycook, in Wijayawardene et al., Fungal Diversity 86:
56 (2017).
Based on Barriopsis fusca A.J.L. Phillips, A. Alves &
Crous, in Phillips, Alves, Pennycook, Johnston, Ramaley,
Akulov & Crous, Persoonia 21: 39 (2008).
Notes: Barriopsis is characterised by aseptate, brown
ascospores without apiculi (Phillips et al. 2008). Absence
of apiculi on the ascospores differentiates this genus from
Sphaeropsis.
Botryobambusa Phook., Jian K. Liu & K.D. Hyde, in Liu
et al., Fungal Diversity 57(1): 166 (2012).
Index Fungorum number: IF 801313; Facesoffungi number: FoF 02408; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Botryobambusa fusicoccum Phook., Jian
K. Liu & K.D. Hyde, in Liu et al., Fungal Diversity 57(1):
166 (2012).
Notes: Botryobambusa was introduced by Liu et al.
(2012) with a single species with both sexual and asexual
morphs. Phylogenetically this genus is clearly distinguished
from Botryosphaeria. However, it is similar to Botryosphaeria except for smaller asci and ascospores that are surrounded by a mucilagenous sheath.
Cophinforma Doilom, Jian K. Liu & K.D. Hyde, in Liu
et al., Fungal Diversity 57(1): 174 (2012).
Index Fungorum number: IF 801315; Facesoffungi number: FoF 07624; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Cophinforma eucalypti Doilom, Jian K.
Liu & K.D. Hyde, in Liu et al., Fungal Diversity 57(1): 174
(2012).
Notes: Cophinforma was introduced by Liu et al. (2012)
as a monotypic genus for C. eucalypti. Phillips et al. (2013),
based on a multi-gene analysis, introduced two species to
this genus that were previously included in Botryosphaeria. Cophinforma is phylogenetically distinguished from
Fungal Diversity (2020) 105:17–318
Botryosphaeria despite being similar, except for conidia that
are longer than any known species in Botryosphaeria.
Diplodia Fr., in Montagne, Annls Sci. Nat., Bot., sér. 2 1:
302 (1834).
Index Fungorum number: IF 8047; Facesoffungi number:
FoF 00147; – 397 morphological species (Species Fungorum 2020), 30 species with molecular data (Jayawardena
et al. 2019b, c).
Type species: Diplodia mutila (Fr. : Fr.) Fr., Summa Veg.
Scand. 2: 417. 1849.
≡ Sphaeria mutila Fr. : Fr., Syst. Mycol. 2: 424. 1823.
≡ Physalospora mutila (Fr. : Fr.) N.E. Stevens, Mycologia28: 333. 1936.
≡ Botryosphaeria stevensii Shoemaker, Can. J. Bot. 42:
1299. 1964.
Notes: Diplodia comprises two morphological groups
that are supported by two distinct phylogenetic lineages.
In one type the conidia are initially hyaline and aseptate,
later becoming brown and 1-septate. Pigmentation is often
delayed and in some species dark conidia are never seen. In
the second type the conidia become pigmented at an early
stage, even while they are enclosed within the pycnidia, they
only rarely become septate.
Dothiorella Sacc., Michelia 2(no. 6): 5 (1880).
Index Fungorum number: IF 8098; Facesoffungi number:
FoF 00148; – 257 morphological species (Species Fungorum 2020), 44 species with molecular data.
Type species: Dothiorella pyrenophora Berk. ex Sacc.,
Michelia 2(no. 6): 5 (1880)
Notes: Dothiorella has often been confused with Diplodia in morphological characterisations. The type species,
Dothiorella pyrenophora differs from Diplodia by conidia
that are brown and 1-septate early in their development,
while they are still attached to the conidiogenous cells.
Sexual morphs of Dothiorella have pigmented, septate
ascospores. However, the sexual stage of the species is rarely
found in nature and has never been reported in culture.
Dothiorella ostryae Manawasinghe, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number IF556874, Facesoffungi number: FoF 06578; Fig. 16
Etymology: Name reflects the host genus of Ostrya
carpinifolia.
Holotype: MFLU 18-0177.
Saprobic on dead aerial branch of Ostrya carpinifolia. Sexual morph Undetermined. Asexual morph Conidiomata pycnidial, stromatic, solitary, dark brown, semiimmersed, unilocular, globose papillate with a short neck,
wall 5–7 cell layers, outer layers composed of light-brown
cells of textura angularis, inner layer hyaline and thin-walled
53
Conidiophores reduced to conidiogenous cells. Conidiogenous cells holoblastic, hyaline, cylindrical, proliferating
percurrently. Conidia 15–21 × 7–10 ( x̄ = 19 × 9 µm, n =
30) μm, ellipsoidal initially hyaline and aseptate, becoming
pale brown while attached to the conidiogenous cells, finally
dark brown or sepia, 1-septate, with 2 cells of equal length,
thick-walled.
Culture characteristics: Colonies on PDA reach 70 mm
after 5 d in the dark at 25 °C.
Material examined: Italy, Passo delle Forche - Galeata
(province of Forlì-Cesena), dead aerial branch of Ostrya
carpinifolia L. (Betulaceae), 9 April 2018, Erio IT Camporesi (MFLU 18-0177, holotype), ex-type living culture,
JZB3150026.
GenBank number: ITS: MN533805, tef1 MN537429.
Notes: The species identified in this study is similar to
other species in Dothiorella (Phillips et al. 2013). In phylogenetic analysis (Fig. 14), Dothiorella ostryae is closely
related to the clade that contains Dothiorella citricola
(ICMP 16828) and D. viticola (CBS 117009). There are four
Dothiorella species associated with the host genus Ostrya,
namely D. iberica, D. omnivora, D. parva (Farr and Rossman) and D. ostryae (this study).
Endomelanconiopsis Rojas & Samuels, Mycologia 100:
770. (2008).
Index Fungorum number: IF 555482; Facesoffungi number: FoF 07625; – 3 morphological species (Species Fungorum 2020), 3 species with molecular data.
Type species: Endomelanconiopsis endophytica Rojas &
Samuels, Mycologia 100: 770 (2008).
Notes: This genus was introduced by Rojas et al. (2008)
with two species. Tibpromma et al. (2018a) introduced a
third species, Endomelanconiopsis freycinetiae, bringing the
total to three species. Endomelanconiopsis is characterized
by eustromatic conidiomata and holoblastically produced,
brown, non-apiculate, unicellular conidia, each with a longitudinal germ slit (Rojas et al. 2008).
Eutiarosporella Crous, in Crous et al., Phytotaxa 202(2):
85 (2015).
Index Fungorum number: 811248; Facesoffungi number:
FoF 07365; – 7 morphological species (Species Fungorum
2020), 7 species with molecular data.
Type species: Eutiarosporella tritici (B. Sutton & Marasas) Crous, in Crous et al., Phytotaxa 202(2): 85 (2015).
≡ Tiarosporella tritici B. Sutton & Marasas, Trans. Br.
mycol. Soc. 67(1): 74 (1976)
Notes: This genus was introduced by Crous et al. (2015a)
to accommodate tiarosporella-like taxa with long conidiomatal necks and holoblastic conidiogenesis. Eutiarosporella
differs from Marasasiomyces by developing clustered
conidiomata.
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54
Fungal Diversity (2020) 105:17–318
Fig. 16 Dothiorella ostryae (MFLU 18-0177, holotype). a Material
examined. b Submerged conidiomata on host. c, d Cross section of
conidiomata on host. e Developing conidia while attached to the pyc-
nidial wall on host. f, g Conidia on host. h Conidiogenous cells in
culture. 1 Upper view of colony on PDA. j reverse view of 7 days old
culture on PDA. Scale bars: a, b = 2 mm, c–h = 10 µm
Lasiodiplodia Ellis & Everh., Bot. Gaz. 21: 92 (1896).
Index Fungorum number: IF 8708; Facesoffungi number:
FoF 00151; – 64 morphological species (Species Fungorum
2020), 35 species with molecular data.
Type species: Lasiodiplodia theobromae (Pat.) Griff. &
Maubl. Bull. trimest. Soc.Mycol. Fr. 25: 57 (1909).
≡ Botryodiplodia theobromae Pat., in Patouillard &
Lagerheim, Bull. Soc. mycol. Fr. 8(3): 136 (1892).
Notes: Lasiodiplodia was introduced by Ellis in 1894
with L. tubericola as the type species. Ellis did not provide
a description of the genus and species, but this was provided by Clendenin (1896) who attributed both the genus
and species to Ellis and Everhart. Griffon and Maublanc
(1909) considered that Botryodiplodia theobromae was
more suitably accommodated in Lasiodiplodia. Since the
epithet theobromae (1892) is older than tubericola (1896),
L. theobromae should be regarded as the type species of
Lasiodiplodia. In earlier works on Lasiodiplodia and Diplodia it was considered that Lasiodiplodia could represent a
possible synonym of Diplodia (Denman et al. 2000). However, in phylogenetic analyses these two genera were clearly
separated in two distinct clades (Zhou and Stanosz 2001;
Slippers et al. 2004a, b; Phillips et al. 2008). Lasiodiplodia
is distinguished from Diplodia by striations on the conidia.
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Fungal Diversity (2020) 105:17–318
Conidiomatal paraphyses distinguishes Lasiodiplodia from
Neodeightonia. Although Barriopsis has striate conidia, they
are unique in Botryosphaeriaceae because they are also present on immature, hyaline conidia.
Macrophomina Petr., Annls mycol. 21(3/4): 314 (1923).
Index Fungorum number: IF 8814; Facesoffungi number:
FoF 07626; – 4 morphological species (Species Fungorum
2020), 3 species with molecular data.
Type species: Macrophomina philippinensis Petr., Annls
mycol. 21(3/4): 314 (1923).
Other synonyms can be seen in Index Fungorum.
Notes: Macrophomina comprises two species with
cultures and moleular data. This genus is similar to Tiarosporella in having apical mucoid appendages (Sutton and
Marasas 1976). Apart from the phylogeny, these two species
can be distinguished, since Macrophomina has percurrently
proliferating conidiogenous cells (Nag-Raj 1993; Crous et al.
2006a) and conidia that become dark brown at maturity, and
microsclerotia (Phillips et al. 2013).
Marasasiomyces Crous, in Crous et al., Phytotaxa 202(2):
86 (2015).
Index Fungorum number: IF 811252; Facesoffungi number: FoF 07440; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Marasasiomyces karoo (B. Sutton & Marasas) Crous, Crous et al., Phytotaxa 202(2): 85 (2015).
≡ Tiarosporella graminis var. karoo B. Sutton & Marasas,
Trans. Br. mycol. Soc. 67(1): 73 (1976).
Notes: This monotypic genus was introduced by Crous
et al. (2015a) for M. karooo to accommodate tiarosporellalike taxa. Marasasiomyces is distinguished from Tiarosporella by conidiomata with long necks covered in brown
setae, and holoblastic conidiogenesis. These characters are
similar to Eutiarosporella, but differ on account of the nonclustered conidiomata.
Mucoharknessia Crous, R.M. Sánchez & Bianchin., in
Crous et al., Phytotaxa 202(2): 86 (2015).
Index Fungorum number: IF 811254; Facesoffungi number: FoF 01651; – 2 morphological species (Species Fungorum 2020), 2 species with with molecular data.
Type species: Mucoharknessia cortaderiae Crous,
R.M. Sánchez & Bianchin Crous, in Crous et al., Phytotaxa
202(2): 85 (2015).
Notes: This genus was introduced by Crous et al. (2015a)
for species resembling Harknessia. The genus is characterised by conidia bearing mucoid appendages. Mucoharknessia comprises two species.
Neodeightonia C. Booth, in Punithalingam, Mycol. Pap.
119: 17 (1970) [1969].
55
Index Fungorum number: IF 3450; Facesoffungi number:
FoF 07627; – 8 morphological species (Species Fungorum
2020), 8 species with with molecular data.
Type species: Neodeightonia subglobosa C. Booth,
Mycol. Pap. (1970).
Notes: Neodeightonia was regarded as a synonym of Botryosphaeria (von Arx and Müller 1975). However, it was
reinstated by Phillips et al. (2008) based on morphological characters including the dark, 1-septate ascospores, and
its phylogenetic placement. Longitudinal striations on the
conidial wall are an additional characteristic feature of this
genus (Crous et al. 2006a; Phillips et al. 2008). Neodeightonia can be differentiated from Lasiodiplodia by the absence
of conidiomatal paraphyses, and conidial striations distinguish it from Diplodia.
Neofusicoccum Crous, Slippers & A.J.L. Phillips, in Crous
et al., Stud. Mycol. 55: 247 (2006).
Index Fungorum number: IF 500870; Facesoffungi number: FoF 00153; – 45 morphological species (Species Fungorum 2020), 43 species with molecular data (Jayawardena
et al. 2019a, c).
Type species: Neofusicoccum parvum (Pennycook &
Samuels) Crous, Slippers & A.J.L. Phillips., in Crous et al.,
Stud. Mycol. 55: 247 (2006).
≡ Fusicoccum parvum Pennycook & Samuels, Mycotaxon 24: 455 (1985).
Notes: Neofusicoccum was introduced by Crous et al.
(2006a) to accommodate species that are similar to Botryosphaeria, but phylogenetically distinct. Dichomera synasexual morph in Neofusicoccum has been used to differentiate
it from Botryosphaeria, even though not all Neofusicoccum
species form such a synasexual morph (Phillips et al. 2005;
Barber et al. 2005). Development of paraphyses can be
observed in all accepted Botryosphaeria species, but have
not been reported in conidiomata of any Neofusicoccum species. However, this character is difficult to use as developing conidiogenous cells can resemble paraphyses. Accepted
species in this genus have been separated on the basis of
conidial dimensions and pigmentation, pigment production
in culture media and ITS sequence data (Phillips et al. 2013).
However, morphological characterisation of these species
remains controversial (Abdollahzadeh et al. 2013). To
resolve the species, multi-gene sequence data are essential.
Species in Neofusicoccum have been reported as endophytes,
pathogens and saprobes on a wide range of hosts.
Neoscytalidium Crous & Slippers, Stud. Mycol. (2006).
Index Fungorum number: IF 500868; Facesoffungi number: FoF 07628; – 3 morphological species (Species Fungorum 2020), 7 species with molecular data.
Type species: Neoscytalidium dimidiatum (Penz.) Crous
& Slippers, in Crous et al., Stud. Mycol. 55: 244 (2006).
13
56
≡ Torula dimidiata Penz., in Saccardo, Michelia 2(no.
8): 466 (1887).
Notes: In earlier studies, species belonging to this genus
was introduced as Hendersonula toruloidea, which have
conidia that become septate with a darker central cell
and formed a Scytalidium-like synanamorph (Crous et al.
2006a). Since the description, the production of both arthroconidial and pycnidial synasexual morphs has been shown
and led to several controversies in the nomenclature. However, the current genus name was introduced by Crous et al.
(2006a) and other relevant epitypifications and synonymies
by Madrida et al. (2009). Species in this genus have been
reported as pathogens on human skin and nails (Campbell
and Mulder 1977) and plants. Diseases reported associated
with this genus tend to be more common in tropical regions
(Polizzi et al. 2009, 2011; Phillips et al. 2013).
Oblongocollomyces Tao Yang & Crous., in Yang et al., Fungal Biology 121: 339 (2016).
Index Fungorum number: IF 817640; Facesoffungi number: FoF 08049; 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Oblongocollomyces variabilis (F.J.J. Van
der Walt, Slippers & G.J. Marais) Tao Yang & Crous, in
Yang et al., Fungal Biology 121: 339 (2016).
≡ Sphaeropsis variabilis F.J.J. Van der Walt, Slippers &
G.J. Marais, in Slippers et al., Persoonia 33: 164 (2014).
Notes: Oblongocollomyces was introduced by Yang et al.
(2017) as a monotypic genus in Botryosphaeriaceae. In earlier studies, this species was identified as Sphaeropsis variabilis and it was different from Sphaeropsis in having long
conidiomatal necks on PNA, aggregated in dense clusters
of conidiomata on OA (Yang et al. 2017). This species has
conidia with up to 3 septa. Phylogenetically Oblongocollomyces close to Alanphillipsia (Fig. 14).
Phaeobotryon Theiss. & Syd., Annls mycol. 13(5/6): 664
(1915).
Index Fungorum number: IF 3892; Facesoffungi number:
FoF 07629; – 8 morphological species (Species Fungorum
2020), 6 species with molecular data.
Type species: Phaeobotryon cercidis (Cooke) Theiss. &
Syd., Annls mycol. (1915).
≡ Dothidea cercidis Cooke, Grevillea 13(no. 67): 66
(1885).
Notes: Phaeobotryon has 2-septate, brown ascospores
with an apiculus at each end.
Sakireeta Subram. & K. Ramakr., J. Indian bot. Soc. 36:
83 (1957).
Index Fungorum number: IF 9775; Facesoffungi number:
FoF 07583; – 1 morphological species (Wijayawardene et al.
2017a), 1 species with molecular data.
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Type species: Sakireeta madreeya Subram. & K. Ramakr., J. Indian bot. Soc. 36: 84 (1957).
Notes: Sakireeta comprises a single species, S. madreeya,
which was previously known as Tiarosporella madreeya.
This genus is distinguished from the closely related Tiarosporella by conidiomata that are aggregated in a stroma,
and being plurilocular whereas, Tiarosporella has solitary
unilocular conidiomata. In addition, Sakireeta has holoblastic conidiogenous cells (Crous et al. 2015a).
Sardiniella Linald., A. Alves & A.J.L. Phillips, Mycosphere
(2016).
Index Fungorum number: IF 817511; Facesoffungi number: FoF 02405; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Sardiniella urbana Linald., A. Alves &
A.J.L. Phillips, Mycosphere (2016).
Notes: Sardiniella was introduced as a monotypic genus
for Sardiniella urbana. This genus is characterised by
thick-walled, oblong conidia that become pigmented and
1-septate, which distinguishes it from Neofusicoccum. This
genus is similar to Diplodia and Dothiorella. However, in
Dothiorella, and in some species of Diplodia the conidia
become pigmented while still attached to the conidiogenous
cell and this character is not seen in Sardiniella. The conidial
wall in species of Diplodia is thicker than in Sardiniella.
Phylogenetically these two genera are clearly separated.
Sphaeropsis Sacc., Michelia 2(no. 6): 105 (1880).
Index Fungorum number: IF 562041; Facesoffungi number: FoF 01704; – 142 morphological species (Species Fungorum 2020), 5 species with molecular data.
Type species: Sphaeropsis visci (Alb. & Schwein.) Sacc.,
Michelia 2(no. 6): 105 (1880).
≡ Dothidea visci Kalchbr., Hedwigia 8: 117 (1869).
Notes: Sphaeropsis was introduced by Saccardo (1880)
to accommodate species of Diplodia with brown, aseptate
conidia. Since then species were described based on host
association, resulting in more than 600 species names. Pycnidial paraphyses distinguish Sphaeropsis from Diplodia.
The striate conidia of Lasiodiplodia differentiate it from
Sphaeropsis, which has smooth-walled conidia.
Tiarosporella Höhn., in Weese, Ber. dt. bot. Ges. (1919).
Index Fungorum number: IF 10233; Facesoffungi number: FoF 00333; – 8 morphological species (Species Fungorum 2020), 5 species with molecular data.
Type species: Tiarosporella paludosa (Sacc. & Fiori)
Höhn., in Weese, Ber. dt. bot. Ges. 37: 159 (1919).
≡ Neottiospora paludosa Sacc. & Fiori, in Sydow, Hedwigia 38(Beibl.): (137) (1899).
Notes: This genus is characterised by conidia formed
from smooth, hyaline conidiogenous cells that lack periclinal
Fungal Diversity (2020) 105:17–318
thickenings and percurrent proliferations. The conidia are
hyaline, subcylindrical to fusiform with irregular mucoid
appendages. There are 22 species names for this genus in
Index Fungorum.
Economic and ecological significance
Botryosphaeriaceae comprises a diverse range of taxa that
are pathogens, endophytes or saprobes on a wide range of
hosts including dicotyledonous monocotyledonous, and
gymnosperms (Schoeneweiss 1981; Manawasinghe et al.
2016). Their distribution is vast, covering all geographic
and climatic regions, except Polar Regions (Crous et al.
2007a; Rodas et al. 2009; Wunderlich et al. 2011; Manawasinghe et al. 2016). At present, their ecological role as
phytopathogens has gained a great attention (Manawasinghe et al. 2016). The pathogenic taxa in Botryosphaeriaceae
cause several important diseases such as leaf spots, cankers,
dieback, fruit rot, gummosis and even plant death on many
economically important crops (Rodas et al. 2009, Hyde et al.
2014). Some species are pathogenic on many hosts (Diplodia seriata, Lasiodiplodia spp., and Neofusicoccum parvum)
in the same geographic area. Pathogenicity of some taxa
(e.g. B. dothidea) varies with the country for the same host
species (Chen et al. 2014; Netto et al. 2014; Linaldeddu et al.
2015). The pathogenic fungal taxa of this family are recognized as “opportunistic plant fungal pathogens” (Chethana
et al. 2016). It is not clear whether changes in the environment triggers pathogenicity or if disease development is due
to weakening of the host defenses (Chethana et al. 2016;
Manawasinghe et al. 2016). Since phytopathogenic botryosphaeriaceous taxa also have an endophytic lifestyle they
may act as latent pathogens. Some species in this family
have been reported as human pathogens (Polizzi et al. 2009,
2011; Phillips et al. 2013).
Melanopsaceae A.J.L. Phillips, Slippers, Boissin & Crous,
Stud. Mycol. 76(1): 43 (2013).
Index Fungorum number: IF 805796; Facesoffungi numbers: FoF 07630, 111 species.
On woody hosts. Sexual morph: Ascostromata pseudothecial, multiloculate, locules at various levels in ascoma,
immersed, partially erumpent at maturity, black, subglobose, thick-walled. Ascomatal wall thick-walled, comprised
of cells of textura angularis. Asci 8-spored, bitunicate, fissitunicate, pedicellate, clavate. Hamathecium comprising
hyaline, thin-walled, hypha-like, septate, not constricted at
septa, cellular pseudoparaphyses. Ascospores overlapping,
ellipsoid to rhomboid, hyaline, aseptate, thin-walled, with a
persistent mucus sheath. Asexual morph: Coelomycetous.
Conidiomata often formed in the same stroma as ascostromata. Paraphyses septate, branched or not, filiform, hyaline, arising from between the conidiogenous cells. Conidiophores 1–2-septate, branched or not, hyaline, smooth, or
57
reduced to conidiogenous cells. Conidiogenous cells subcylindrical, branched or unbranched, discrete, hyaline, formed
from the inner wall of the conidioma, proliferating percurrently at apex, or with periclinal thickening. Conidia fusoid,
hyaline, aseptate, with a persistent mucus sheath, rarely with
minute marginal frill (adapted from Slippers et al. 2013;
Phillips et al. 2019)
Type: Melanops Nitschke ex Fuckel.
Notes: Slippers et al. (2013) established Melanopsaceae
in Botryosphaeriales to accommodate Melanops. Melanops
has a mucilage sheath around the ascospores and in that
respect is identical to Phyllostictaceae. However, Phyllostictaceae has uniloculate ascostromata, while those of Melanops are multiloculate (Slippers et al. 2013). Phylogenetic
analyses supported the placement of Melanops as a distinct
family in Botryosphaeriales (Wikee et al. 2013b; Slippers
et al. 2013; Phillips et al. 2019). Divergence time estimates
for this family also support its familial status (Slippers et al.
2013; Phillips et al. 2019). However, Jiang et al. (2018b)
suggested that the divergence time estimation in the previous
studies might not be accurate estimates due to the fact that
sequence data from only two taxa were used in the phylogenetic placement, an opinion also voiced by Phillips et al.
(2019).
Melanops Nitschke ex Fuckel, Jb. nassau. Ver. Naturk.
23–24: 225 (1870) [1869–70].
Index Fungorum number: IF 3078; Facesoffungi number:
FoF 07442; 111 morphological species (Species Fungorum
2020), 3 species with molecualar data.
Type species: Melanops tulasnei Fuckel.
Notes: Over 100 species names are listed under Melanops in Index Fungorum (2020), but, only three species have
sequence data (Jiang et al. 2018b). We were unable to obtain
fresh collections to represent this genus. Thus, a drawing is
provided. A description of the type species can be seen in
Phillips and Alves (2009).
Melanops tulasnei Fuckel, Jb. nassau. Ver. Naturk. 23–24:
225 (1870) [1869–70].
Index Fungorum number: IF 150956; Facesoffungi number: FoF 07444; Fig. 17
Description: see Phillips and Alves (2009).
Economic and ecological significance
The ecology and distribution of Melanops is poorly known.
Although its appearance is similar to other Botryosphaeriales, it is not clear whether it is pathogenic or endophytic
(Slippers et al. 2013).
Phyllostictaceae Fr. (as ‘‘Phyllostictei’’), Summa vegetabilium Scandinaviae 2: 420 (1849).
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Fig. 17 Melanops tulasnei (redrawn from neotype and epitype specimens in Phillips and Alves 2009). a Stroma erumpent through the
bark. b Stroma cut through horizontally revealing ascomata and conidiomata. c Ascus. d Ascus tip. e Ascospores. f Conidium and conid-
iogenous cell with mucus sheath around the conidium. g Conidium
with mucous sheath. h Conidia. i Conidiogenous with conidia developing on conidiogenous cells. j Paraphyses. Scale bars: b = 250 μm,
c = 20 μm, d, e, g–j = 10 μm, f = 5 μm
Index Fungorum number: IF 81162; Facesoffungi numbers: FoF 02296, >2000 species.
Foliicolous, pathogenic, endophytic or saprobic on
plants. Sexual morph: Ascostromata pseudothecial, uniloculate, separate to gregarious, globose, brown to black,
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with a central ostiole. Asci 8-spored, bitunicate, fissitunicate, clavate to subcylindrical, fasciculate, pedicellate,
with an ocular chamber. Pseudoparaphyses mostly absent
at maturity, when present broad-cylindric, septate, cellular. Ascospores 2–3-seriate, ellipsoid-fusoid to limoniform,
hyaline, aseptate, smooth-walled, usually with mucilaginous
caps, or surrounded by a mucilaginous sheath. Asexual
morph: Coelomycetous. Conidiomata pycnidial globose,
dark brown, separate to aggregated, with a central ostiole;
wall of 3–6 layers of brown cells of textura angularis. Conidiogenous cells lining the inner wall, hyaline, smooth, subcylindrical to ampulliform or doliiform, proliferating percurrently near apex, frequently covered in mucilaginous sheath.
Conidia ellipsoid-fusoid to obovoid or ovoid, hyaline, aseptate, smooth-walled, guttulate or granular, frequently surrounded by a mucilaginous sheath, often bearing a single
mucilaginous apical appendage.
Type: Phyllosticta Pers.
Notes: Phyllostictaceae (as Phyllostictei) was proposed
by Fries (1849). Seaver (1922) used Phyllostictales and
Phyllostictaceae for Phyllosticta. Phyllostictaceae was
accepted by Hawksworth and David (1989). Schoch et al.
(2006) suggested that Phyllosticta belongs to Botryosphaeriaceae and this was accepted by Crous et al. (2006a) and
Liu et al. (2012) who noted that Phyllosticta is distinct from
other genera in Botryosphaeriaceae. Wikee et al. (2013b)
suggested Phyllostictaceae as a distinct family in Botryosphaeriales and Slippers et al. (2013) formally re-instated
the family.
Ascospores in Melanopsaceae and Phyllostictaceae have
a mucilage sheath. Phyllostictaceae differs from Melanopsaceae by uniloculate ascostromata. Though mucoid apical appendages on the conidia have been regarded as characteristic of Phyllosticta, this feature is not necessarily a
characteristic of the family since they are absent in some
species such as P. leucothoicola or P. neopyrolae (Wikee
et al. 2013b).
Phyllosticta Pers., Traite´sur les Champignons Comestibles
(Paris): 147 (1818).
Index Fungorum number: IF 9384; Facesoffungi number:
FoF 00155; >2000 records (Species Fungorum 2020), several species with molecular data.
Type species: Phyllosticta cruenta (Kunze: Fr.) J. Kickx
f.
≡ Sphaeria cruenta Fr., Syst. mycol. (Lundae) 2(2): 581
(1823).
Notes: Phyllosticta is a geographically widespread genus
of plant pathogenic fungi with a diverse host range and a
Guignardia sexual morph (Fig. 18). Seaver (1922) used
Phyllostictales and Phyllostictaceae for Phyllosticta. Phyllosticta was placed in Botryosphaeriales by Schoch et al.
(2006), who proposed that Botryosphaeriaceae contained
59
Fig. 18 Sexual morph of Phyllosticta ellipsoidea (redrawn from
Wulandari et al. 2011, MFLU 10-0475). a Section of ascoma. b Asci.
c Ascospores with polar mucilaginous appendage at each end. Scale
bars: 25 µm
both Botryosphaeria and Phyllosticta, although no support
was obtained for this relationship. Crous et al. (2006a) and
Liu et al. (2012) also classified Phyllosticta in Botryosphaeriaceae. In both studies it was noted that Phyllosticta was
distinct from other genera in Botryosphaeriaceae, and that
these authors eventually expected it to be placed elsewhere.
Wikee et al. (2013b) redefined Phyllosticta, and showed that
it clusters sister to Botryosphaeriaceae, for which the older
family name Phyllostictaceae was resurrected. In moving
away from a dual nomenclature for fungi, the generic name
Phyllosticta was chosen over Guignardia in previous studies.
There are 3214 Phyllosticta epithets listed in Index Fungorum. In this entry we illustrate Phyllosticta hostae (Fig. 19).
Phyllosticta hostae Y.Y. Su & L. Cai, Persoonia 28: 79
(2012).
Index Fungorum number: IF 564904; Facesoffungi number: FoF 07380; Fig. 19
Leaf spots circular to somewhat irregularly rotundate,
pale brown to brown. Sexual morph: Undetermined. Asexual morph: Colonies on PDA, Pycnidia black, sub epidermal, globose, 40–150 µm in diam. dark brown to black,
darker around ostiole. Conidiogenous cells holoblastic,
phialidic, cylindrical, subcylindrical to ampulliform, hyaline, thin-walled, smooth. Conidia 8–15 × 5–9 µm ( x̄ = 10
× 7, n = 20), unicellular, thin- and smooth-walled, ellipsoid,
subglobose to obovoid, with large central guttule, truncate at
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Fig. 19 Phyllosticta hostae (JZB 3290001). a Leaf spot on rose leaf. b Front view of the colony on PDA. c Rear view of the colony on PDA. d
Conidiomata in culture. e–h Conidia. Scale bars: d = 100 µm, e–h = 10 µm
the base when young, later rounded at both ends, enclosed in
a 1–3 µm thick mucilaginous sheath, and bearing a hyaline,
mucoid apical appendage, 4–8 × 1–3 µm, straight to flexuous, unbranched, tapering towards an acute tip.
Material examined: China, Beijing Tongzhou-Yongle Eco
Park, leaf spot on rose (Rosaceae) leaves, 18 August 2018,
Jia Jing Yi, 28-10-1, living culture: JZB 3290001.
GenBank number: ITS: MW130255.
Notes: Phyllosticta is an important genus of plant pathogens and contains quarantinable pathogenic species in some
countries, especially in Europe (Wulandari et al. 2013).
Phyllosticta hostae was introduced by Su and Cai (2012)
from Hosta plantaginia (Liliaceae). This is the first record
of P. hostae causing leaf spots in rose in China.
13
Other genera included
Pseudofusicoccum Mohali, Slippers & M.J. Wingf., Stud.
Mycol. 55: 249 (2006).
Index Fungorum number: IF 555584; Facesoffungi number: FoF 05299; – 8 morphological species (Species Fungorum 2020), 7 species with molecular data.
Type species: Pseudofusicoccum stromaticum (Mohali,
Slippers & M.J. Wingf.) Mohali, Slippers & M.J. Wingf., in
Crous et al., Stud. Mycol. 55: 249 (2006).
≡ Fusicoccum stromaticum Mohali, Slippers & M.J.
Wingf., Mycol. Res. 110(4): 408 (2006).
Notes: The type species of Pseudofusicoccum, P. stromaticum was introduced as Fusicoccum stromaticum and was
distinguished from other species by its large conidiomata,
the ability to grow at 35°C, and thick-walled conidia. Crous
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61
Fig. 20 Morphology of Pseudofusicoccum. a, b Conidiophores and
conidia of P. stromaticum. c–e Conidiomatal wall, conidiophore and
conidia of P. africanum. f, g Conidia becoming brown and septate
when mature and young conidia of P. artocarpi. h–j Conidiogenous
cells, germinating conidia and conidia of P. adansoniae Scale bars: a
= 5 μm, b, c, e, f–j = 10 μm, d = 25 μm
et al. (2006a) recognized that its conidia were covered by
a mucous sheath, which is lacking in all species of Fusicoccium. Therefore, they introduced Pseudofusicoccum as a
new genus to accommodate this taxon. Pseudofusicoccum
adansoniae, P. ardesiacum and P. kimberleyense were proposed and distinguished from others by production of a pigment in culture and the size of conidia (Pavlic et al. 2008).
Later, P. olivaceum was identified as a new species because
of the discrepancy between ITS and tef1 BLAST results
(Mehl et al. 2011). Pseudofusicoccum artocarpi was introduced as its conidia were clearly longer than others (Trakunyingcharoen et al. 2015). Pseudofusicoccum violaceum is
distinctive because of the violet/purple pigment formed in
culture and this has not been observed in any other Pseudofusicoccum species (Mehl et al. 2011). Pseudofusicoccum
africanum was introduced as it is different from its neighbor
P. violaceum by unique fixed alleles in ITS and tef1 (Jami
et al. 2018). Yang et al. (2017) raised the genus to familial
13
62
status as Pseudofusicoccaceae. The phylogenies reported in
Phillips et al. (2019), as well as those of Minnis et al. (2012)
and Liu et al. (2017), place Pseudofusicoccum in the same
clade as Phyllosticta species. In view of this consistent relationship, together with the evolutionary divergence evidence
as well as morphological data presented in Phillips et al.
(2019), Pseudofusicoccum is considered to be an additional
genus within Phyllostictaceae (Fig. 20).
Economic and ecological significance
Phyllosticta species have globally been recorded as endophytes, plant pathogens and saprobes from economically and
ecologically important plant hosts. Some cause important
diseases such as Citrus black spot and tan spot, subjected
to phyto-sanitary legislation in the European Union and the
USA. Further leaf- and fruit-spotting disease of Musa spp.
(freckle disease), leaf spots in turmeric, cashew, ginger,
orchids and black rot of grapes diseases are also caused by
Phyllosticta species.
Planistromellaceae M.E. Barr, Mycotaxon 60: 437 (1996).
Index Fungorum: IF 81919; Facesoffungi number: 06689,
33 species.
Biotrophic, hemibiotrophic or saprobic on leaves and
stems of various plants in terrestrial habitats. Sexual
morph: Ascostromata multi- or uni-loculate, immersed to
erumpent through cracking or splitting of the host tissue, solitary to gregarious, with periphysate ostioles, with or without papilla. Cells of ascostromata thick-walled, composed
of several layers of dark brown cells, arranged in a textura
angularis. Locules ovoid to globose, developing in the same
stroma of the conidiogenous and/or spermatogenous locules,
collapsed with the empty locule, which previously produced
conidia or spermatia or both, ostiole periphysate. Peridium
of locules composed of a few layers of hyaline to light brown
flattened cells. Hamathecium lacking pseudoparaphyses,
interascal cells abundant even at maturity. Asci 8-spored,
bitunicate, fissitunicate, oblong, clavate to nearly cylindrical, with a pedicel and with an ocular chamber, forming
in a basal layer, often interspersed with and covered by
cellular remnants of interthecial tissues. Ascospores overlapping 1–3-seriate, ellipsoid to broadly obovoid, hyaline
or lightly pigmented, yellowish to brownish, aseptate or
1–2-trans-septate, thin-walled, with or without gelatinous
sheath, guttulate. Asexual morph: Coelomycetous. Conidiomata subepidermal, dark, immersed to erumpent, solitary
to gregarious, pycnidia, locules or acervuli in a stroma or
bearing conidia over stroma surface prior to locule formation, ostiolate. Conidiomata walls comprising several layers
with cells of textura angularis, the outer layers composed
of dark thick-walled cells, lighter towards the inner layers
of hyaline cells. Conidiogenous cells short cylindric, conidiogenesis holoblastic, hyaline, smooth. Conidia oblong,
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ellipsoid-cylindric, hyaline to brown, aseptate or 1-multitrans-septate, smooth-walled or verruculose, with or without
one or more apical appendages. Spermatial state developing
in the same or separate locules. Spermatogenous cells discrete or integrated, phialidic, cylindric to elongate-conical,
determinate, hyaline, smooth. Spermatia bacillary, hyaline,
aseptate, smooth-walled.
Type: Kellermania Ellis & Everh.
Notes: Planistromellaceae was introduced by Barr
(1996) and belongs to Botryosphaeriales (Minnis et al.
2012; Monkai et al. 2013). Lumbsch and Huhndorf (2010)
included the genera Comminutispora, Eruptio, Loratospora.,
Microcyclus, Mycosphaerellopsis, Planistroma and Planistromella in Planistromellaceae based on morphology. Phylogenetic analysis based on SSU, ITS and LSU sequence data
by Minnis et al. (2012) showed that Kellermania, Piptarthron, Planistroma and Planistromella should be treated as
a single genus, thus, they synonymized them under Kellermania (oldest name). However, Monkai et al. (2013) indicated that the type species of Kellermania, which is also
the type of Planistromella, clustered separately from Planistroma based on LSU and ITS sequence data. Planistroma
was accepted as a distinct genus. The family was revised to
accommodate Kellermania, Planistroma Mycosphaerellopsis and Umthunziomyces based on morphology and molecular data (Monkai et al. 2013; Phillips et al. 2019).
Kellermania Ellis & Everh., J. Mycol. 1(12):153 (1885).
Index Fungorum number: IF 8668; Facesoffungi number:
FoF 06690; 23 morphological species (Species Fungorum
2020), 18 species with molecular data.
Type species: Kellermania yuccifoliorum A.W. Ramaley.
Notes: Species in this genus are known from the host
genera Agave and Nolina (Asparagaceae). Five species
have been reported with both sexual and asexual morphs in
Ramaley (1993, 1995, 1998) and Barr (1996). Many species of Kellermania were illustrated by Minnis et al. (2012)
and conidia range from 0 to several trans-septa, and with or
without appendages.
Kellermania yuccifoliorum A.W. Ramaley, Mycotaxon 47:
262 (1993).
Index Fungorum: IF 360150; Facesoffungi number: FoF
06691; Figs. 21 and 22
Description: see Monkai et al. (2013).
Material examined: USA, Califonia, San Bernardino
County, Roadside 20 miles east of Baker (Hwy. 91/466),
on leaves of Yucca brevifolia Engelmann, 14 April 1960,
Isabelle Tavares No. 466 (UC 1202973, holotype of Planistromella yuccifoliorum).
Notes: The type species of Kellermania (K. yuccifoliorum
≡ Planistromella yuccifoliorum) differs from other genera
in Planistromellaceae in having 1–2-septate ascospores
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Fig. 21 Kellermania yuccifoliorum (UC 1202973, holotype of P.
yuccifoliorum). a, b Ascostromata on the host surface. c, d Section
of ascostroma. e, f Ascoma. g Immature ascus. h Mature asci. i Cells
of ascostroma. j Immature ascospore. k, l Mature ascospores. Scale
bars: a = 1 cm, b = 1000 µm, c = 200 µm, d = 100 µm, e, f = 50 µm,
g, h = 30 µm, i = 20 µm, j–l = 10 µm
(Ramaley 1993; Barr 1996). The asexual morph of K. yuccifoliorum is characterized by uniloculate conidiomata, which
develops in the same stroma as the ascogenous locules and
2-septate conidia with a unique apical appendage (Ramaley
1993).
Type species: Mycosphaerellopsis myricariae (Fuckel)
Höhn., Annal. Mycol. 16(1/2):157 (1918).
≡ Sphaeria myricariae Fuckel, Jb. nassau. Ver. Naturk.
27–28: 22 (1874) [1873–74].
Notes: Mycosphaerellopsis differs from other genera
in Planistromellaceae in having uniloculate ascomata
and 1-septate, broadly obovoid ascospores. We accept
Mycosphaerellopsis in this family based on its morphology
(Monkai et al. 2013).
Other genera included
Mycosphaerellopsis Höhn., Annal. Mycol. 16(1/2): 157
(1918).
Index Fungorum number: IF 3345; Facesoffungi number:
FoF 06263; – 2 morphological species (Monkai et al. 2013),
molecular data unavailable.
Planistroma A.W. Ramaley, Mycotaxon 42: 69 (1991).
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Fig. 22 Kellermania yuccifoliorum (redrawn from
Ramaley 1993). a Conidioma. b
Microconidia. c Spermatia and
spermatiogenesis. d Conidiogenesis and origin appearance
of apical appendage. e Conidia.
Scale bars: a = 80 µm, b, d =
27 µm, e = 40 µm
Index Fungorum number: IF 25358; Facesoffungi number: FoF 06264; – 7 morphological species (Monkai et al.
2013), 6 species with molecular data.
Type species: Planistroma yuccigenum A.W. Ramaley,
Mycotaxon 42: 69 (1991), MycoBank: MB 358836.
Notes: Planistroma was established by Ramaley (1991),
and is similar to Kellermania in ascostromata characters,
but Planistroma differs on account of its aseptate ascospores
(Barr 1996). The asexual morphs of Planistroma resemble
Kellermania except in having apically appendaged conidia
in the latter genus (Ramaley 1995). Minnis et al. (2012) synonymized Planistroma under Kellermania based on molecular data support. However, we follow Monkai et al. (2013)
who assigned Planistroma as a distinct genus based on both
morphological and molecular data.
Notes: The monotypic genus Umthunziomyces was introduced by Crous et al. (2016a) and is known only from its
asexual morphs. Umthunziomyces is similar to Kellermania
in having septate conidia, but they are longer and narrower
than in the latter genus. Phillips et al. (2019) included Umthunziomyces in Planistromellaceae with evidence from the
phylogenetic analyses of ITS and LSU sequence data.
Umthunziomyces Crous & M.J. Wingf., in Crous et al., Persoonia 37: 315 (2016).
Index Fungorum number: IF 819069; Facesoffungi number: FoF 06718; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Umthunziomyces hagahagensis Crous &
M.J. Wingf., Persoonia 37: 315 (2016).
Saccharataceae Slippers, Boissin & Crous, Studies in
Mycology 76: 41 (2013).
Index Fungorum number: IF 805794; Facesoffungi numbers: FoF 02296, 22 species.
Saprobic, endophytic, or pathogenic on plants. Sexual
morph: Ascostromata pseudothecial, uniloculate, solitary or in clusters, with multi-layered, dark brown walls,
13
Economic and ecological significance
Members in this family usually grow on living or dead leaves
or on stems of various plants, and are mostly saprobes, but
some species are pathogens such as Kellermania agaves, and
Mycosphaerellopsis moravica (Barr 1996; Ramaley 1993,
1995, 1998; Minnis et al. 2012; Crous et al. 2013). Most
species inhabit Asparagaceae.
Fungal Diversity (2020) 105:17–318
65
Fig. 23 Saccharata proteae (redrawn from Slippers
et al. 2013, CBS 121406).
a Immersed ascomata with
clypeus-like structure. b Asci
and ascospores. c–e Conidiogenous cells and paraphyses. f
Conidia. Scale bars: b–f = 10
µm
infrequently embedded in stromatic tissue, with upper
ascomatal layer darkened and thickened, developing under
a very small epidermal clypeus, ascomatal wall continuous
with clypeus. Asci 8-spored, bitunicate, fissitunicate, with a
thick endotunica, pedicellate or sessile, cylindrical, with a
well-developed ocular chamber. Hamathecium comprising
3–5 µm wide, filiform, septate, hypha-like, branched or not,
abundant, intermixed with asci, cellular pseudoparaphyses.
Ascospores 2–3-seriate, ellipsoidal, hyaline to pigmented,
pale golden brown, granular, septate or not, without mucoid
appendages or sheath. Asexual morph: Coelomycetous.
Conidiomata unilocular pycnidial, infrequently embedded
in stromatic tissue with thickened, darkened upper layer.
Conidiophores sparingly branched, hyaline, subcylindrical,
or reduced to conidiogenous cells. Conidiogenous cells hyaline, smooth, phialidic, proliferating via periclinal thickening or percurrent proliferation, with or without collarettes.
Conidia fusoid, hyaline, aseptate, thin-walled, granular.
Synasexual morph formed in separate conidiomata, or in
same conidiomata with asexual morph. Synasexual morph:
Conidia ellipsoid or oval, pigmented, aseptate, thick-walled,
finely verruculose. Spermatogonia similar to conidiomata
in anatomy. Spermatogenous cells ampulliform to lageniform or subcylindrical, hyaline smooth, phialidic. Spermatia
developing in conidiomata or spermatogonia, subcylindrical
or dumbbell-shaped, with rounded ends, hyaline, smooth,
granular.
Type: Saccharata Denman & Crous.
Notes: The narrow, aseptate, filiform pseudoparaphyses in
Saccharataceae are unique in Botryosphaeriales and the pale
golden brown ascospores are also distinctive. Slippers et al.
(2013) accommodated this family in Botryosphaeriales. Saccharataceae grouped separately from all other families that
were basal in the phylogenetic tree, suggesting a long, separate evolutionary history. Saccharataceae has previously
been known only from southern Africa, and is most diverse
on the Proteaceae. Recent research has shown, however, that
it has also been introduced as endophytes in other countries
where South African Proteaceae are now being cultivated
(Marincowitz et al. 2008).
Saccharata Denman & Crous, in Crous et al., CBS Diversity
Ser. (Utrecht) 2: 104 (2004).
Index Fungorum number: IF 28918; Facesoffungi number: FoF 02297; 19 morphological species (Species Fungorum 2020), 15 species with molecular data.
Type species: Saccharata proteae (Wakef.) Denman &
Crous.
≡ Phyllachora proteae Wakef., Bull. Misc. Inf., Kew(5):
164 (1922).
13
66
Notes: Crous et al. (2004) described the first species
of this genus from Proteaceae in the South Western Cape
region of South Africa. Three other species were added to
the genus, two from Proteaceae and one from Encephalartos
(Marincowitz et al. 2008; Crous et al. 2008, 2009). Apart
from its restricted distribution and host range, Saccharata
is also unique in its asexual morphology, which includes
a hyaline, fusicoccum-like and a pigmented diplodia-like
asexual morph. Based on phylogenetic analysis, evolutionary estimates and morphological data, Phillips et al. (2019)
considered Septorioideaceae a synonym of Saccharataceae.
We were unable to obtain fresh collections of Saccharata
or other genera of Saccharataceae, therefore, a drawing of
Saccharata proteae is provided (Fig. 23).
Saccharata proteae (Wakef.) Denman & Crous, in Crous
et al., CBS Diversity Ser. (Utrecht) 2: 104 (2004).
≡ Phyllachora proteae Wakef., Bull. Misc. Inf., Kew(5):
164 (1922).
Index Fungorum number: IF 370531; Facesoffungi number: FoF 07631; Fig. 23
Description: see Slippers et al. (2013).
Other genera included
Pileospora J.B. Tanney & K.A. Seifert, Mycol. Prog. 18:
169 (2019).
Index Fungorum number: IF 824738; MycoBank number:
MB82473; – 1 species with molecular data.
Type species: Pileospora piceae J.B. Tanney & K.A. Seifert, Mycol. Prog. 18: 169 (2019).
Notes: Pileospora is distinct from Septorioides by its
aseptate, ellipsoidal-fusiform conidia with apical appendages versus subcylindrical to fusiform-ellipsoidal, 1–10-septate conidia lacking appendages. Pileospora is similar to
Neofusicoccum, Saccharata, and other genera of Botryosphaeriales having fusicoccum-like asexual states with cooccurring irregularly-shaped conidia. An irregular, mucoid,
apical appendage distinguishes conidia of Pileospora from
above mentioned similar genera.
Septorioides Quaedvl., Verkley & Crous, Stud. Mycol. 75:
383 (2013).
Index Fungorum number: IF 804464; Facesoffungi number: FoF 06278; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Septorioides pini-thunbergii (S. Kaneko)
Quaedvlieg et al. Stud. Mycol. 75: 383 (2013).
≡ Septoria pini-thunbergii S. Kaneko, in Kaneko et al.,
Trans. Mycol. Soc. Japan 30(4): 463 (1989).
Notes: This genus is characterized by acervular conidiomata that open by means of an irregular rupture and paraphyses that are intermingled among conidiogenous cells
(Quaedvlieg et al. 2013). Septorioides pini-thunbergii was
13
Fungal Diversity (2020) 105:17–318
associated with needle blight and the cause of sooty mould
of Pinus thunbergii in Japan (Suto 2000). Later, this species
was isolated as an endophyte from P. densiflora in South
Korea (Yoo and Eom 2012). The trend of raising established genera to familial status continued when Wyka and
Broders (2016) introduced Septorioideaceae within the Botryosphaeriales to accommodate Septorioides species. They
introduced a novel species Septorioides strobi as the second
species of this family. However, Phillips et al. (2019) placed
Septorioides as a genus in Saccharataceae taking into consideration the results of phylogenetic analyses, evolutionary
divergence and morphological data.
Economic and ecological significance
Saccharataceae has previously been known only from southern Africa, and is most diverse on Proteaceae. Research
has shown, however, that it has also been introduced as an
endophyte into other countries where South African Proteaceae are now cultivated (Marincowitz et al. 2008). This
plant family, which has a high endemic richness in southern
Africa, has evolved in the region for more than 100 million
years (Barker et al. 2007). This date allows for the estimated
57 (28–100) MYA (based on rDNA SSU) separation of the
Saccharataceae as a family and to have evolved with these
endemic plant hosts in the region. The species are typically
associated with leaf spots and stem cankers and they appear
to be pathogens. Separate studies have also shown that they
are endophytes (Swart et al. 2000, Taylor et al. 2001), similar
to members of the Botryosphaeriaceae.
Catinellales Ekanayaka, K.D. Hyde & Ariyawansa, ord.
nov.
Index Fungorum number: IF 557327; Facesoffungi number: FoF 07646
Saprobic on dead wood in terrestrial habitats. Sexual
morph: Ascomata apothecial, discoid. Excipulum composed of angularis to globose cells. Hamathecium comprising cylindric, septate, paraphyses. Asci cylindric, clavate,
inoperculate with an ocular chamber, J-. Ascospores ellopsoidal, uni-seriate. Asexual morph: Undetermined.
Type: Catinellaceae Ekanayaka, K.D. Hyde &
Ariyawansa.
Notes: Catinella was established by Boudier (1907).
Several authors have misidentified this fungus and placed it
within Leotiomycetes (Durand 1922). However phylogenetic
study of Greif et al. (2007) strongly supported its placement
within Dothideomycetes. Catinella formed a distinct lineage
within Dothideomycetes in Hongsanan et al. (2020), thus we
introduced this new order to accommodate Catinella and
allied species in the class Dothideomycetes. The divergence
time for Catinellales is estimated as 163 MYA (Hongsanan
et al. 2020).
Fungal Diversity (2020) 105:17–318
67
≡ Peziza olivacea Batsch, Elench. fung. (Halle): 127
(1783).
Notes: Catinella was established by Boudier (1907), with
C. olivacea as the type species. The genus was recognised as
a member in Leotiomycetes based on its morphology. However, sequence data of C. olivacea indicated that this genus
formed a distinct clade within Dothideomycetes (Fig. 24).
Economic and ecological significance
Catinellaceae is saprobic on dead wood and plays a role in
recycling of organic matter.
Fig. 24 Catinella olivacea (a = redrawn from Gamundí 1981; b–e =
redrawn from Akata et al. 2016). a Peridium. b Apothecial ascomata.
c Ascospores. d Asci. e Anchor hyphae. Scale bars: a = 10 μm, b =
20 μm
Accepted families: Catinellaceae.
Catinellaceae Ekanayaka, K.D. Hyde & Ariyawansa, fam.
nov.
Index Fungorum number: IF 557328; Facesoffungi number: FoF 07647, 1 species.
Saprobic on dead wood in terrestrial habitats. Sexual
morph: Ascomata apothecial, discoid, cupulate, flattened
disc. Excipulum composed of subangularis to globose
cells, ectal cells are pigmented. Hamathecium comprising cylindric, septate, swollen apices agglutinate to form
a pseudoepithecium, paraphyses. Asci 8-spored, cylindric,
clavate, inoperculate, unitunicate, J-, with well-developed
ocular chamber. Ascospores ellopsoidal, uni-seriate, darkly
pigmented, one-celled. Asexual morph: Undetermined.
Type genus: Catinella Boud.
Notes: The new family Catinellaceae bears some morphological characteristics similar to Leotiomycetes, i.e. cupulate
apothecia, unitunicate asci, aseptate ascospores. However,
Dothideomycetes also show these characters, although they
are not common within the class (Greif et al. 2007).
Catinella Boud., Hist. Class. Discom. Eur. (Paris): 150
(1907).
Index Fungorum number: IF 845; Facesoffungi number:
FoF 07648; 3 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type: Catinella olivacea (Batsch) Boud.
Cladoriellales Crous.
Index Fungorum number: IF 823435; Facesoffungi number: FoF 07649.
Cladoriellales was established by Crous et al. (2017),
with a single genus Cladoriella. Molecular data (LSU)
confirmed its status as a distinct order within Dothideomycetes. This order contains asexual, cladosporium-like species
which are found as saprobes or pathogens on leaf surface.
In Fig. 25, four species of Cladoriella (Cladoriella eucalypti, C. kinglakensis, C. rubrigena, and C. xanthorrhoeae)
grouped with high support. The divergence time for Cladoriellales is estimated as 183 MYA (stem age, Hongsanan
et al. 2020, Fig. 2).
Accepted families: Cladoriellaceae.
Cladoriellaceae Crous, in Crous et al., Persoonia 39: 417
(2017).
Index Fungorum number: IF 823436; Facesoffungi numbers: FoF 07650, 5 species.
Saprobes or pathogens on leaf surface. Sexual morph:
Undetermined. Asexual morph: Hyphae coiling, branched,
septate, with swollen cells giving rise to conidiophores, with
hyphopodium-like structures at the base, simple, intercalary,
brown to dark brown, thick-walled, smooth to finely verruculose. Conidiophores separate, erect, subcylindrical, straight,
septate, thick-walled, brown to dark brown, smooth to finely
verruculose. Conidiogenous cells terminal or intercalary,
mono- or poly-tretic, sympodial, with 1–2 conspicuous loci,
thickened, darkened, refractive, with a minute central pore.
Conidia remains in long acropetal chains, narrowly ellipsoidal to cylindrical or fusoid, brown, non or 1-septate, thickwalled, finely verruculose, with apical conidium rounded at
the apex; additional conidia with 1–2 truncate, conspicuous
hila; thickened, darkened, refractive, with a minute central
pore. Chlamydospores absent (Crous et al. 2006b, 2017).
Type: Cladoriella Crous.
Notes: Cladoriellaceae was established by Crous
et al. (2017) to accommodate a single genus Cladoriella.
Sequence data for this family are available in GenBank and
it also supports Cladoriellaceae as a family of Cladoriellales
in Dothideomycetes.
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68
Blastacervulus eucalypti CBS 124759
97/0.99
Thyrinula parasitica CBS 120088
100/1.0
Parmularia styracis VIC 42447
Parmularia styracis VIC 42587
-/0.97
-/1.0
100/1.0
97/1.0
Cladoriella rubrigena CBS 124760
Cladoriella xanthorrhoeae CBS 143398
100/1.0
Asterina cestricola TH 591
Asterina zanthoxyli TH 561
Asterina weinmanniae TH 592
100/1.0
0.03
Asterinales
100/1.0
Cladoriellales
Cladoriella eucalypti CPC 10953
78/0.98
Cladoriella kinglakensis CPC 32730
Capnodium coffeae OSC 100414
Outgroup
Leptoxyphium fumago CBS 123.26
Cladoriella Crous, Stud. Mycol. 55: 54 (2006).
Index Fungorum number: IF 500799; Facesoffungi number: FoF 07651; 5 morphological species (Species Fungorum 2020), 4 species with molecular data.
Type species: Cladoriella eucalypti Crous.
Notes: Crous et al. (2006b) introduced Cladoriella to
accommodate cladosporium-like species. This genus is similar to Devriesia, but lacks chlamydospores, and produces a
distinct red pigment on media. Phylogenetic analyses also
support this distinctness as Cladoriella formed a clade apart
from Cladosporium complex (Mycosphaerellaceae), Cladophialophora complex (Herpotrichiellaceae), or Pseudocladosporium complex (Venturiaceae) (Crous et al. 2006b)
(Fig. 26).
Economic and ecological significance
Species in this family are saprobic and probably are plant
pathogenic (Wijayawardene et al. 2017a). They are found on
leaf surfaces, however, the lifestyle of species in Cladoriellaceae is not well-studied.
Collemopsidiales Pérez-Ort.
Index Fungorum number: IF 815627; Facesoffungi number: FoF 07652.
This recently erected order includes a single family, Xanthopyreniaceae. It encompasses species of lichen-forming
13
Parmulariales
100/1.0
Thyrinulaceae
Fig. 25 Phylogram generated
from maximum likelihood analysis (RAxML) of Cladoriellales
based on LSU sequence data.
Maximum likelihood bootstrap
values equal or above 70%,
Bayesian posterior probabilities
equal or above 0.90 (MLBS/
PP) are given at the nodes. An
original isolate number is noted
after the species name. The tree
is rooted to Capnodium coffeae
(OSC 100414) and Leptoxyhium
fumago (CBS 123.26). The
ex-type strains are indicated in
bold. Hyphen (-) represents support values below 70% MLBS
and 0.90 PP
Fungal Diversity (2020) 105:17–318
fungi, lichenicolous or forming loose associations with
cyanobacteria. The diversity of the order has been largely
underestimated and the actual number of species could
exceed 200. The divergence time for Collemopsidiales is
estimated as 230 MYA (stem age, Hongsanan et al. 2020)
(Fig. 27).
Accepted families: Xanthopyreniaceae.
Xanthopyreniaceae Zahlbr., in Engler & Prantl, Nat.
Pflanzenfam., Edn 2 (Leipzig) 8: 91 (1926).
Index Fungorum number: IF 81525; Facesoffungi number: FoF 07653, 53 species.
Lichen-forming fungi, lichenicolous or forming loose
associations with cyanobacteria. Vegetative hyphae usually
hyaline. Sexual morph: Ascomata perithecioid, solitary or
in groups, superficial or immersed in the substrarum, subglobose. Peridium thin to thick, composed by elongated to
roundish, more or less compresed cells of textura angularis.
Involucrelum present in some species, with textura intricata.
Hamathecium comprising thin, septate, highly branched and
anastomosed interascal filaments. Asci 4–8-spored, thickwalled, bitunicate, fissitunicate, ovate to clavate, apedicellate, with a distinct ocular chamber. Ascopores irregularly
arranged within the ascus, ellipsoid to broadly ellipsoid,
hyaline, 1-septate, wall cell smooth, usually with a gelatinous shell. Asexual morph: Coelomycetous. Pycnidium,
Fungal Diversity (2020) 105:17–318
69
with X. tichothecioides, suggesting that the correct generic
name for those species should be Collemopsidium, and
many species of Pyrenocollema were subsequently combined into this genus (Grube and Ryan 2002; Mohr et al.
2004). Pérez-Ortega et al. (2016) studied the phylogenetic
relationships of Xanthopyreniaceae using six molecular
markers. Although they found Xanthopyreniaceae belongs
to Dothiodeomycetes, and described the new order Collemopsidiales, their exact position remained elusive despite
of using different taxa sampling, very likely due to the use
of Arthoniomycetes as outgroup. Subsequent work by Liu
et al. (2017) found high support for the sister relationships
of Collemopsidiales with Monoblastiales. Based on uncertainties by Pérez-Ortega et al. (2016), Tedersoo et al. (2018)
described the new class Collemopsidiomycetes which should
be considered as a synonym of Dothiodeomycetes. Lumbsch
and Huhndorf (2007) included the genera Collemopsidium,
Didymellopsis, Frigidopyrenia, Pyrenocollema, and Zwackhiomyces as members of the Xanthopyreniaceae. Later,
Wijayawardene et al. (2018) excluded Frigidopyrenia and
Pyrenocollema from the family and placed them in Pezizomycotina genera incertae sedis. Unfortunately, no molecular data is available for Didymellopsis, Frigidopyrenia and
Pyrenocollema. Pérez-Ortega et al. (2016) discussed that
the number of species in the family may have been highly
underestimated.
Fig. 26 Conidiophore and conidia of Cladoriella spp. a Cladoriella eucalypti (redrawn from Crous et al. 2006, CBS H-18043). b–d
Cladoriella xanthorrhoeae (redrawn from Crous et al. 2017, CBS
H-23300)
superficial or immersed in the substratum, black, with a
small ostiole. Pycnidial wall thick-walled cells of textura
angularis, composed of ± isodiametric or elongated cells.
Conidiogenous cells ± cylindrical, phialidic. Conidiospores
bacilliform to ellipsoid.
Type: Xanthopyrenia Bachm.
Notes: Xanthopyreniaceae was introduced by Zahlbruckner to accommodate the single species Xanthopyrenia tichothecioides (= Arthopyrenia tichothecioides), a species growing on periodically inundated calcareous rocks. The species
was later combined into Pyrenocollema (P. tichothecioides)
and subsequently many species associated with cyanobacteria, and with perithecioid ascomata, highly branched and
anastomosed interascal filaments and hyaline 1-septate
ascospores were transferred or described in Pyrenocollema
(e.g. Santesson 1992; Coppins et al. 1992; McCarthy and
Kantvilas 1999; Nordin 2002). However, Grube and Ryan
(2002) stated that the type species of the genus, P. tremelloides is in fact a parasite on Nostoc and is not congeneric
Xanthopyrenia Bachm., Nova Acta Acad. Caes. Leop.Carol. German. Nat. Cur. 105(1): 65 (1919).
Index Fungorum number: IF 5810; Facesoffungi number:
FoF 07654; – 2 morphological species (Pérez-Ortega et al.
2016), molecular data unavailable.
Type species: Xanthopyrenia tichothecioides (Arnold)
Bachm.
≡ Arthopyrenia tichothecioides Arnold, Flora, Regensburg 52: 268 (1869).
Notes: The genus was described to accommodate the single species Arthopyrenia tichothecioides, a species occurring with seawater or spray on calcareous rocks (Thüs and
Schultz 2009). This species and X. heardensis, the only other
species described in the genus, were subsequently transferred to Pyrenocollema. Pérez-Ortega et al. (2016) showed
that Collemopsidium as currently circumscribed is paraphyletic, so Xanthopyrenia might be the correct generic name for
non-marine Collemopsidium species (Fig. 28).
Other genera included
Collemopsidium Nyl., Flora, Regensburg 64: 6 (1881).
Index Fungorum number: IF 1182; Facesoffungi number:
FoF 07655; – 16 morphological species (Species Fungorum
2020), 3 species with molecular data (Pérez-Ortega et al.
2016).
13
70
Collemopsidium cf. elegans MA-LICH 18342
-/85/1.0
Collemopsidium cf. halodytes MA-LICH 18333
Collemopsidium cf. foveolatum MA-LICH 18341
100/1.0
Collemopsidium pelvetiae MA-LICH 18370
96/1.0
Collemopsidium cf. sublitorale MA-LICH 18331
96/0.99
Collemopsidium cf. ostrearum MA-LICH 18344
-/-
Collemopsidiales
Zwackhiomyces coepulonus Etayo 25240
Collemopsidium angermannicum MA-LICH 18373
100/1.0
Anisomeridium ubianum MPN94
100/1.0
Anisomeridium phaeospermum MPN539
100/1.0
Acrocordia subglobosa HTL940
Monoblastiales
Fig. 27 Phylogram generated
from maximum likelihood
analysis (RAxML) of Collemopsidiales based on LSU and
SSU sequence data. Maximum
likelihood bootstrap values
equal or above 70%, Bayesian
posterior probabilities equal
or above 0.90 (MLBS/PP) are
given at the nodes. An original
isolate number is noted after the
species name. The tree is rooted
to and Abrothallus cladoniae
(AB53) and A. parmeliarum
(AB36). Hyphen (-) represents
support values below 70%
MLBS and 0.90 PP
Fungal Diversity (2020) 105:17–318
Abrothallus parmeliarum AB36
100/1.0
Outgroup
Abrothallus cladoniae AB53
0.02
Type species: Collemopsidium iocarpum (Nyl.) Nyl.,
Flora, Regensburg 64: 6 (1881).
≡ Pyrenopsis iocarpa Nyl., Not. Sällsk. Fauna et Fl.
Fenn. Förh., Ny Ser. 5: 26 (1861).
Notes: Grube and Ryan (2002) studied the types of Collemopsidium and Pyrenocollema and stated that the former
is the correct generic name for species previously included
in Pyrenocollema. Pérez-Ortega et al. (2016) showed that
the genus as currently understood is paraphyletic and species
occurring in marine and terrestrial and freshwater habitats
belong to different lineages. Collemopsidium iocarpum was
described growing close to the seashore and it is likely the
correct name for the marine clade. Species concepts and
taxonomy have to be thoroughly studied. Pérez-Ortega et al.
(2016) showed that each of the species commonly accepted
in the literature likely encompass dozens of different taxa.
Cyanobacteria associated with Collemopsidium are largely
undetermined, marine species likely belong to Hyella (Swinscow 1965b). Xanthocapsa may be involved in freshwater
symbioses (Grube and Hafellner 1990).
Didymellopsis (Sacc.) Clem. & Shear, Gen. fung., Edn 2
(Minneapolis): 66 (1931).
= Didymella subgen. Didymellopsis Sacc. & D. Sacc.,
Syll. fung. (Abellini) 17: 657 (1905).
Index Fungorum number: IF 1550; Facesoffungi number:
FoF 07656; – 8 morphological species (Species Fungorum
2020), molecular data unavailable.
13
Type species: Didymellopsis latitans (Nyl.) Sacc. ex
Clem. & Shear, Gen. fung., Edn 2 (Minneapolis): 265
(1931).
≡ Obryzum latitans Nyl., Flora, Regensburg 68: 298
(1885).
Notes: All species in Didymellopsis are found on lichens
with cyanobacterial photobionts, but D. perigena which
grows on Catapyrenium squamulosum might be associated with cyanobacteria surrounding the host thallus. Differences between Didymyellopsis and Zwackhiomyces are
subtle (Zhurbenko et al. 2015) and a thorough revision of
the group, including DNA sequence analyses should be carried out.
Frigidopyrenia Grube, Phyton, Horn 45(2): 307 (2005).
Index Fungorum number: IF 521258; Facesoffungi number: FoF 07657; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Frigidopyrenia bryospila (Nyl.) Grube,
Phyton, Horn 45(2): 308 (2005).
≡ Verrucaria bryospila Nyl., Flora, Regensburg 47: 357
(1864).
Notes: Grube (2005) described the monotypic genus
Frigidopyrenia for Verrucaria bryospila occurring on
mosses in arctic-alpine areas, forming distinct squamules
and with cyanobacteria as photobiont. Grube (2005) discussed the morphological differences of the genus with
similar genera such as Collemopsidium, Magmopsis and
Fungal Diversity (2020) 105:17–318
71
Fig. 28 Morphological and anatomical features of Xanthopyreniaceae. a Habitus Collemopsidium foveolatum-morphotype. b C.
halodytes-morphotype. c C. ostrearum-morphotype. d Zwackhiomyces coepulonus. e Ascus from C. angermannicum. F Interascal fila-
ments from C. cf. foveolatum. g Ascospore from C. cf. sublitorale.
h Ascospore from C. angermannicum with gelatinous sheath. Scale
bars: a–c = 1 mm; d = 0.5 mm; e–h = 10 µm
Pyrenocollema, which have different peridial structure. It
is not clear that the genus belongs in Xanthopyreniaceae.
Index Fungorum number: IF 835454; Facesoffungi number: FoF 09130; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Rhagadodidymellopsis endocarpi Fdez.Brime, Gaya, Llimona & Nav.-Ros., Plant and Fungal Systematics 65(1): 177 (2020).
Rhagadodidymellopsis Fdez.-Brime, Gaya, Llimona &
Nav.-Ros., Plant and Fungal Systematics 65(1): 177 (2020).
13
72
Dyfrolomyces thamplaensis MFLUCC 15-0635
100/1.0
Dyfrolomyces distoseptatus NFCCI: 4377
80/1.0
Dyfrolomyces sinensis MFLUCC 17-1344
Dyfrolomyces phetchaburiensis MFLUCC 15-0951
-/0.98
96/0.97
Dyfrolomyces thailandica MFLU 16-1173
Dyfrolomyces rhizophorae BCC 15481
-/1.0
82/-
Dyfrolomyces rhizophorae JK 5456A
Melomastia italica MFLUCC 15-0160
100/1.0
Dyfrolomycetales
Fig. 29 Phylogram generated
from maximum likelihood
analysis (RAxML) of Dyfrolomycetales based on LSU and
SSU sequence data. Maximum
likelihood bootstrap values
equal or above 70%, Bayesian
posterior probabilities equal
or above 0.90 (MLBS/PP) are
given at the nodes. An original
isolate number is noted after
the species name. The tree is
rooted to Anisomeridium phaeospermum (MPN539) and A.
ubianum (94). The ex-type and
reference strains are indicated in
bold. Hyphen (-) represents support values below 70% MLBS
and 0.90 PP
Fungal Diversity (2020) 105:17–318
Melomastis maolanensis GZCC 16-0102
100/1.0
Dyfrolomyces tiomanensis NTOU3636
Acrospermum adeanum M133
100/1.0
Acrospermales
Acrospermum compressum M151
100/1.0
Anisomeridium ubianum 94
Outgroup
Anisomeridium phaeospermum MPN539
0.02
Notes: This recently erected genus, with the only species R. endocarpi growing on Endocarpon pusillum, was
described based on the presence of ascomata occurring in
stromata with uneven wall. It seems to be closely related to
Didymellopsis and Zwackhiomyces (Fernández-Brime et al.
2020). It is here tentatively placed in Collemopsidiales and
Xanthopyreniaceae until molecular data is available.
Zwackhiomyces Grube & Hafellner, Nova Hedwigia 51(3–
4): 305 (1990).
Index Fungorum number: IF 25388; Facesoffungi number: FoF 07658; – 26 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Zwackhiomyces coepulonus (Norman)
Grube & R. Sant., in Grube & Hafellner, Nova Hedwigia
51(3–4): 310 (1990).
≡ Arthopyrenia coepulona Norman, Bot. Notiser: 192
(1868) [1867].
Notes: Zwackhiomyces is one of the most diverse genera
of lichenicolous fungi with 35 known species (Diederich
et al. 2018). Species mostly grow on crustose lichens with
green photobionts, although five species grow on hosts with
cyanobacteria as photobiont. Pérez-Ortega et al. (2016)
sequenced the type species and confirmed the genus is a
member of Xanthopyreniaceae.
Zwackhiomacromyces Etayo & van den Boom, Opuscula
Philolichenum 13: 70 (2014).
13
Index Fungorum number: IF 803963; Facesoffungi number: FoF 08929; – 2 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Zwackhiomacromyces constrictocarpus
Etayo & van den Boom, Opuscula Philolichenum 13: 72
(2014).
Notes: van den Boom and Etayo (2014) established the
new genus based on material from the Iberian Peninsula
growing on Megaspora verrucosa. The new genus differs
from Zwackhiomyces in having lageniform ascomata, with
large and granulose papillae, larger asci and ascospores,
and normally 3-septate smooth-walled ascospores (van den
Boom and Etayo 2014). Later, Etayo and Berger (in Berger
and Zimmerman 2016) combined Pyrenidium hyalosporum
growing in Placopsis gelida into Zwackhiomacromyces
based on ascoma and ascospore characters. This genus tentatively placed in the order Collemopsidiales and Xanthopyreniaceae until molecular data is available.
Economic and ecological significance
Members of the Xanthopyreniaceae are usually inconspicuous and have been often overlooked. Most of the lichenicolous species do not cause harm to the hosts, and many of
the interactions may be considered commensalistic. Marine
species of Collemopsidium with boring capacity may have
relevant ecological significance since they actively produce
bioweathering in littoral rocky substrates, boosting the deteriorative action of waves. Furthermore, those species are
Fungal Diversity (2020) 105:17–318
able to colonize and modified mollusk and barnacle shells,
mediating mimicry in some species (Espoz et al. 1995).
Dyfrolomycetales K.L. Pang, K.D. Hyde and E.B.G. Jones.
Index Fungorum number: IF 805281; Facesoffungi number: FoF 07670.
Dyfrolomycetales with the type genus Dyfrolomyces was
introduced to accommodate marine fungi and now includes
terrestrial, wood-inhabiting taxa which are characterized
by immersed, ostiolate, clypeate, papillate ascomata, bitunicate, cylindrical, short pedicellate asci, with a distinct
ocular chamber along with a ring-like subapical ring, and
overlapping uni-seriate, broadly fusiform, symmetrical, hyaline, multi-septate ascospores. Only the single family Pleurotremataceae is accepted in this order. The divergence time
for Dyfrolomycetales is estimated as 171 MYA (stem age,
Hongsanan et al. 2020) (Fig. 29).
Accepted families: Pleurotremataceae.
Pleurotremataceae Walt. Watson, New Phytol. 28: 113
(1929).
Index Fungorum: IF 81192; Facesoffungi number: FoF
01911, 47 species.
= Dyfrolomycetaceae K.D. Hyde, K.L. Pang, Alias,
Suetrong & E.B.G. Jones, in Pang et al., Cryptog. Mycol.
34(3): 227 (2013).
Saprobic on wood in terrestrial and aquatic habitats.
Sexual morph: Ascomata perithecial, gregarious or solitary, immersed or erumpent throughout the host, dark-brown
to black, ovoid to subglobose, carbonaceous to membranaceous, clypeate, ostiolate, papillate. Peridium comprising of
dark pigmented cells of textura angularis or epidermoidea.
Hamathecium comprising numerous, filamentous, hyaline,
septate, branched or unbranched, narrow cellular pseudoparaphyses. Asci 8-spored, bitunicate, clavate to cylindrical,
short pedicellate, apical ring present, J-. Ascospores uniseriate, ellipsoidal to cylindrical or fusiform, hyaline, septate, smooth-walled, mostly with guttules. Asexual morph:
Undetermined.
Type: Pleurotrema Müll.
Notes: Pleurotremataceae is typified by Pleurotrema with
Pleurotrema polysemum as the type species. Subsequently,
the nomenclature and placement of this family has been
the subject of discussion until Maharachchikumbura et al.
(2015) placed it under Chaetosphaeriales, Sordariomycetes.
Maharachchikumbura et al. (2016) examined the isotype
of P. polysemum and found that Dyfrolomycetaceae (Dyfrolomycetales) is a synonym of Pleurotremataceae, thus
removed Pleurotremataceae from Sordariomycetes to be a
single family of Dyfrolomycetales. This family comprises
three genera, and most members are characterized by perithecial, ostiolate, glabrous, ovoid to subglobose ascomata,
clavate to cylindrical, unitunicate, short pedicellate asci with
73
a J- apical ring, and hyaline, ellipsoidal to cylindrical, septate, guttulate ascospores that are uni-seriately arranged in
the asci.
Pleurotrema Müll. Arg., Bot. Jb. 6: 388 (1885).
Index Fungorum number: IF 4251; Facesoffungi number:
FoF 07671; 14 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Pleurotrema polysemum (Nyl.) Müll. Arg.
≡ Parathelium polysemum Nyl., Bot. Ztg. 20: 279 (1872).
Notes: Pleurotrema was introduced as lichenized by Müller (1885). The placement of this genus has been the subject
of controversy as it has been placed in different families
viz., Pyrenulaceae, Pleurotremataceae and Ascomycota
genera incertae sedis, based on its morphological characters
(Barr 1994; Kirk et al. 2001; Lumbsch and Huhndorf 2010).
Maharachchikumbura et al. (2016) re-examined the isotype
of Pleurotrema polysemum, and synonymized Dyfrolomycetaceae under Pleurotremataceae.
Other genera included
Dyfrolomyces K.D. Hyde, K.L. Pang, Alias, Suetrong &
E.B.G. Jones, in Pang et al., Cryptog. Mycol. 34(3): 227
(2013).
Index Fungorum number: IF 804660; Facesoffungi number: FoF 07672; – 9 morphological species (Species Fungorum 2020), 6 species with molecular data.
Type species: Dyfrolomyces tiomanensis K.L. Pang,
Alias, K.D. Hyde, Suetrong & E.B.G. Jones, in Pang et al.,
Cryptog. Mycol. 34(3): 228 (2013).
Notes: Dyfrolomyces was established based on morphological and molecular analyses. Members of this genus are
characterized by relatively large, immersed, globose or
subglobose, clypeate, ostiolate, papillate ascomata, bitunicate, fissitunicate, cylindrical asci and broadly fusiform,
symmetrical, hyaline, septate ascospores, with or without a
mucilaginous sheath.
Dyfrolomyces distoseptatus M. Niranjan and V.V Sarma,
sp. nov.
Index Fungorum number: IF 556726; Facesoffungi number: FoF 06625; Fig. 30
Etymology: In reference to the distoseptate ascospores.
Saprobic on undetermined decaying twig. Sexual morph:
Ascomata 550–630 high × 450–600 μm wide, perithecial,
immersed in periderm, erumpent neck with pseudoparaphyses, clypeate, ostiolate, papillate. Peridium 40 μm, with
two strata, outer thick, carbonaceous and inner brown and
hyaline cells of textura angularis to epidermoidea cells.
Hamathecium comprising 1.8–2.1 μm wide, filamentous,
septate, unbranched, dense, narrow cellular pseudoparaphyses, longer than asci. Asci (123.1)126.7–146.2 (148.6)
× (4.1)4.7–6.3 (6.5) μm ( x̄ = 136.8 × 5.6, n = 25), 8–spored,
13
74
Fig. 30 Dyfrolomyces distoseptatus (AMH 9984, holotype). a Host
plant. b Papillate necks. c, d, i Vertical section of ascomata. e Peridium. f Pseudoparaphyses. g, h Asci j, k Cultures in Petri plates from
13
Fungal Diversity (2020) 105:17–318
single spore isolations. l–o Ascospores. Scale bars: d = 100 µm, e–i
= 50 µm, l–o = 10 µm
Fungal Diversity (2020) 105:17–318
bitunicate, cylindrical, apical ends obtuse, smooth-walled,
short pedicellate, persistent. Ascospores (19.4)19.7–24.9 ×
(4.1) 4.3–5 μm ( x̄ = 21.9 × 4.7, n = 25), uni-seriate, fusoid,
obtuse ends, hyaline, 3-distoseptate, apical ends slightly
bent. Asexual morph: Undetermined.
Material examined: India, Andaman and Nicobar Islands,
South Andaman, Manjery, (11˚52’25.7”N 92˚64’89.9”E), on
unidentified twig, 10 December 2017, M. Niranjan (AMH
9984, holotype), extype living culture NFCCI: 4377.
GenBank numbers: ITS: MK024391, LSU: MH971236.
Notes: Dyfrolomyces distoseptatus clustered with D.
thamplaensis with strong bootstrap support (100%) in ML
analysis. It also forms a sister relationship clade with D.
sinensis (Fig. 29). However, D. distoseptatus has several
distinguishable characters. It is closely related to D. thamplaensis in having distoseptate ascospores with acute ends,
but the ascospores in the latter have prominent guttules. In
addition, the asci of D. distoseptatus are smaller than D.
thamplaensis (123.1–148.6 × 4.1–6.3 μm vs. 114–160 ×
6–8.5 μm) (Zhang et al. 2017a). Furthermore, D. distoseptatus produces ascospores that are 3-septate when compared
to 6–7 septate ascospores in D. sinensis (Hyde et al. 2018).
Hence, based on the above-mentioned morphological differences (Fig. 30) and molecular sequence analyses (Fig. 29),
a new species, Dyfrolomyces distoseptatus, is introduced.
Melomastia Nitschke ex Sacc., Atti Soc. Veneto-Trent. Sci.
Nat., Padova, Sér. 4 4: 90 (1875).
Index Fungorum number: IF 3118; Facesoffungi number:
FoF 07673; – 24 morphological species (Species Fungorum
2020), 2 species with molecular data.
Type species: Melomastia mastoidea (Fr.) J. Schröt., in
Chon, Krypt. -Fl. Schlesien (Breslau) 3.2(3): 320 (1894)
[1908].
≡ Sphaeria mastoidea Fr., K. svenska Vetensk-Akad.
Handl., ser. 3 38: 267 (1817).
Notes: Melomastia was introduced by Saccardo (1875)
to accommodate the type species M. friesii. Schröter (1894)
synonymized M. friesii under M. mastoidea. Species of
Melomastia are characterized by immersed, black, ostiolate
ascomata, cylindrical, thick-walled, pedicellate asci with a
J- subapical ring, and ovoid, hyaline, 2-septate, guttulate
ascospores (Norphanphoun et al. 2017).
Economic significance
Only three genera are accepted in Pleurotremaceae, and
most of the taxa in this family are saprobes in aquatic, mangrove and terrestrial habitats (Pang et al. 2013; Norphanphoun et al. 2017; Zhang et al. 2017a).
Eremithallales Lücking & Lumbsch.
Index Fungorum number: IF 540500; Facesoffungi number: FoF 07674.
75
Eremithallales was introduced for a single species
Eremithallus costaricensis (Lücking et al. 2008). Previously
Eremithallales was reported close to Lichinomycetes. However, both phylogenetic and morpholocal analysis confirmed
the placement of this order in Dothideomycetes with strong
support and later Melaspileaceae was included (Ertz and
Diederich 2015). In Hongsanan et al. (2020), Eremithallales froms a distinct clade within Dothideomycetes. The
divergence time for Eremithallales is estimated as 238 MYA
(stem age Hongsanan et al. 2020).
Accepted families: Melaspileaceae.
Melaspileaceae Walt. Watson, New Phytol. 28: 94 (1929).
= Eremithallaceae Lücking & Lumbsch, in Lücking,
Lumbsch, Di Stéfano, Lizano, Carranza, Bernecker, Chaves
& Umaña, Symbiosis 46(3): 163 (2008).
Index Fungorum number: IF 80992; Facesoffungi number: FoF 07675, 40 species.
Lichenized. Thallus thin, white, corticolous, with Trentepohlia photobiont. Sexual morph: Ascomata apothecioid,
immersed when young, erumpent to superficial when
mature, with a flat to slightly convex disk and a slightly
elevated margin, often surrounded by marginal lobes from
the substrate, roundish. Peridium reddish brown, K+ olivaceous brown at exciple issue, with a I- hymenium, apically
not enlarged, cellular pseudoparaphyses that are sometimes
branched or anastomosed, anastomosed periphyses arising
from the inner excipular layer. Asci 6–8-spored, elongate
or clavate to subcylindrical, wall apically thickened, with a
distinct ocular chamber, I- and K/I-. Ascospores overlapping,
2-seriate, ellipsoid to oblong, hyaline to brown, 1-septate,
constricted near the septum, smooth-walled, sometimes K/
I+ blue gelatinous sheath. Asexual morph: Undetermined.
Type: Melaspilea Nyl.
Notes: The taxonomic placement of this group was
uncertain due to the absence of molecular data. However,
the first phylogenetic analysis confirmed the placement of
Melaspileaceae under Eremithallales. The same result is
shown in our phylogenetic tree (Fig. 31). Morphologically,
Melaspilea sensu stricto is similar to Eremithallus in ascomata, exciple, hamathecium, ascus and ascospore types, and
being lichenized with a trentepohlioid photobiont (Ertz and
Diederich 2015). Thus, Eremithallus was reduced as a synonym of Melaspilea and Eremithallaceae was synonymized
under Melaspileaceae (Ertz and Diederich 2015). This was
followed by Wijayawardene et al. (2017a) and our study.
Melaspilea Nyl., Act. Soc. linn. Bordeaux 21(4): 416 (1857)
[1856].
= Eremithallus Lücking, Lumbsch & L. Umaña, in Lücking et al., Symbiosis 46(3): 163 (2008).
13
76
100/1.0
Melaspilea costaricensis Lücking 15683 (F)
Melaspilea costaricensis Sipman 54968 (B)
Encephalographa elisae EB 0347
85/0.98
Melaspilea enteroleuca Aptroot 20685 (BR)
100/0.99
100/1.0
Melaspilea enteroleuca Farou (BR)
96/-
Melaspilea sp. Ertz 17341 (BR)
Eremithallales
Dothideomycetes sp. AN13
100/1.0
99/0.98
100/-
Lecanactis abietina Ertz 5068 (BR)
Opegrapha vulgata Ertz 7564 (BR)
98/0.99
Arthonia didyma Ertz 7587 (BR)
92/1.0
Lichenostigma alpinum Ertz 17522 (BR)
Trypethelium nitidiusculum AFTOL-ID 2099
Laurera megasperma AFTOL-ID 2094
Trypetheliales
100/1.0
Arthoniomycetes
Fig. 31 Phylogram generated
from maximum likelihood
analysis (RAxML) of Eremithallales based on LSU and SSU
sequence data. Maximum
likelihood bootstrap values
equal or above 70%, Bayesian
posterior probabilities equal
or above 0.90 (MLBS/PP) are
given at the nodes. An original
isolate number is noted after
the species name. The tree is
rooted to Diploschistes ocellatus (AFTOL 958) and Stictis
radiata (AFTOL 398). The
ex-type and references strains
is indicated in bold. Hyphen (-)
represents support values below
70% MLBS and 0.90 PP
Fungal Diversity (2020) 105:17–318
Stictis radiata AFTOL 398
0.02
100/1.0
Outgroup
Diploschistes ocellatu AFTOL 958
Index Fungorum number: IF 3094; Facesoffungi number:
FoF 07676; 37 morphological species (Species Fungorum
2020), 3 species with molecular data.
Type species: Melaspilea arthonioides (A. Massal.) Nyl.
≡ Catillaria arthonioides A. Massal., Geneac. lich.
(Verona): 19 (1854).
Notes: Melaspilea was introduced for two species M.
deformis and M. arthonioides. Taxa in this genus are distributed worldwide and occur primarily as epiphytes on bark
or wood, rocks and parasites or commensals with lichens
(Ertz and Diederich 2015). Melaspilea is a heterogeneous
group and more collections are needed to resolve the genus
(Ertz and Diederich 2015).
Melaspilea enteroleuca (Ach.) Ertz & Diederich, Fungal
Diversity 71: 151 (2015).
≡ Lecidea enteroleuca Ach., Lich. univ.: 177 (1810).
Index Fungorum number: IF 811372; Facesoffungi number: FoF 07677; Fig. 32
Description and discussions: see epitype in Ertz and
Diederich (2015).
Other genus included
Encephalographa A. Massal., Geneac. lich. (Verona): 13
(1854).
Index Fungorum number: IF 1776; Facesoffungi number:
FoF 07678; – 3 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Encephalographa cerebrina (DC.) A. Massal., Miscell. Lichenol.: 49 (1856).
13
≡ Opegrapha cerebrina DC., in Lamarck & de Candolle,
Fl. franç., Edn 3 (Paris) 2: 312 (1805).
Notes: Encephalographa comprises only a single species (Wijayawardene et al. 2017a). This genus was placed
in Hysteriaceae (Renobales and Aguirre 1990). Ertz and
Diederich (2015) indicated that the type species Encephalographa elisae, belongs in Eremithallales. Presently,
Encephalographa is included in Melaspileaceae (Ertz and
Diederich 2015). The species is characterized by saxicolous,
lirelliform, superficial and black ascomata, lichenized, dark
brown to black exciple, I- hymenium, anastomosed paraphyses, clavate, 8-spored and thick-walled asci, apically thickened with distinct ocular chamber, I- and K/I-, and ellipsoid,
1-septate, smooth to finely ornamented, hyaline to brownish
ascospores, constricted at the septum with thin perispore
(Renobales and Aguirre 1990; Ertz and Diederich 2015).
Economic and ecological significance
Melaspileaceae comprises lichenized, lichenicolous and
saprobic taxa (Ertz and Diederich 2015). Saprobic fungi
play a vital role in decomposing plant and animal debris
(Dix and Webster 1995) while lichens are frequently used as
air pollution indicators, which is a simple and economically
viable approach to detect and monitor air pollution (Lopes
et al. 2019).
Eremomycetales Crous, Spatafora, Haridas & I.V. Grig., in
Haridas et al., Stud. Mycol. 96: 151 (2020)
Index Fungorum number: IF 831891; Facesoffungi number: FoF 07679
Fungal Diversity (2020) 105:17–318
77
Fig. 32 Melaspilea enteroleuca (France, Ertz 19235 (BR)). a Thallus with apothecia. b Section of a single apothecium in water. c Hymenium
and ascomatal margin in water. d Ascus in water. e Ascospores in water. Scale bars: a = 0.5 mm, b = 100 µm, c = 50 µm, d = 10 µm, e = 5 µm
Notes: This order was introduced by Haridas et al. (2020).
Analyses of combined LSU and SSU sequence data (Fig. 33)
reveals that the genera Arthrographis, Eremomyces and
Rhexothecium grouped together, forming a distinct clade
apart from other orders in Dothideomycetes. The divergence
time for Eremomycetales is estimated as 262 MYA (stem
age, Hongsanan et al. 2020).
Accepted families: Eremomycetaceae.
Eremomycetaceae Malloch & Cain, Can. J. Bot. 49: 847
(1971).
Index Fungorum number: IF 80751; Facesoffungi number: FoF 05359, 3 species.
Saprobic, isolated primarily on animal dung or soil. Colonies flocculent, drift white, superficial, dense, and growing
slowly on agar media. Hyphae septate, branch, hyaline to
brown. Sexual morph: Ascomata sphaerical to ellipsoid,
solitary, scattered, superficial on hyphae, or submerged in
the agar, dark brown to black, lacking ostioles, but tending to split open before maturity. Peridium thin, pseudoparenchymatous, composed of large brown cells of textura
angularis. Hamathecium lacking pseudoparaphyses. Asci
irregularly disposed within the ascoma, 8-spored, bitunicate,
not fissitunicate, obovoid to clavate, thin-walled, pedicellate, and evanescent. Ascospores multi-seriate, small, often
fabiform to broadly oblate, hyaline to pale brown, aseptate,
13
78
Arthrographis kalrae CBS 693.77
94/1.0
Arthrographis longispora UTHSC 053220
Arthrographis arxii IFM 52652
94/1.0
Eremomyces bilateralis CBS 781.70
Eremomycetaceae
Eremomycetales
Fig. 33 Phylogram generated
from maximum likelihood
analysis (RAxML) of genera
in Eremomycetales based on
LSU and SSU sequence data.
Maximum likelihood bootstrap
values equal or above 70%,
Bayesian posterior probabilities
equal or above 0.90 (MLBS/
PP) are given at the nodes. An
original isolate number is noted
after the species name. The tree
is rooted to Dyfrolomyces thamplaensis (MFLUCC 15-0635).
The ex-type strains are indicated
in bold. Hyphen (-) represents
support values below 70%
MLBS and 0.90 PP
Fungal Diversity (2020) 105:17–318
100/1.0
Rhexothecium globosum CBS 955.73
Dyfrolomyces thamplaensis MFLUCC 15-0635
Outgroup
0.02
smooth-walled to slightly verrucose. Asexual morph: Coelomycetous or hyphomycetous. Pycnidia globose to subglobose, thin-walled, papillate. Conidiophores filiform, septate,
hyaline to brown. Conidia ellipsoidal, hyaline, aseptate, thinwalled, guttulate.
Type: Eremomyces Malloch & Cain.
Notes: Malloch and Cain (1971) introduced Eremomycetaceae for species having solitary ascomata and hyaline,
aseptate ascospores. Eremomycetaceae consists of three
species: Eremomyces bilateralis, Rhexothecium globosum,
and Pithoascus langeronii, (= Eremomyces langeronii).
Eremomycetaceae are most similar to Pseudeurotiaceae
(Malloch and Sigler 1988), but differ in ascomatal initials
(coiled in Pseudeurotiaceae, pseudoparenchymatous in
Eremomycetaceae). Eremomycetaceae comprises two genera
Eremomyces, the type genus and Rhexothecium (Lumbsch
and Huhndorf 2010). Eremomyces has fabiform and hyaline
ascospores, while Rhexothecium has broadly oblate and pale
yellow to brown ascospores. Eremomyces is more similar to
Trichosporiella (Helotiales) in having subglobose to globose, black conidiomata and scopulariopsis-like hyaline
conidia. Rhexothecium resembles Trichosporiella (Helotiales) in having solitary to catenulate, lateral, and hyaline,
basic truncate conidia. Species of Eremomycetaceae are distributed worldwide and can be commonly found in Egypt,
Kenya, North America and the Netherlands. Wijayawardene
et al. (2018) included Arthrographis in Eremomycetaceae,
while Liu et al. (2017) included Arthrographis in a new family Arthrographaceae. However, whether Arthrographaceae
is valid or not is unclear as there is no mention of this family
in the literature or in Index Fungorum. Members of both
Arthrographis and Eremomyces produce hyaline to subhyaline conidia, however Arthrographis is distinct in producing unicellular arthroconidia and chlamydospores which
are absent in Eremomyces. We include Arthrographis in
13
Eremomycetales, genera incertae sedis until further studies
are carried out.
Eremomyces Malloch & Cain, Can. J. Bot. 49(6): 847
(1971).
= Pithoascina Valmaseda et al., Can. J. Bot. 65(9): 1905
(1987).
Index Fungorum number: IF 1880; Facesoffungi number:
FoF 05360; 2 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Eremomyces bilateralis Malloch & Cain.
Notes: Eremomyces bilateralis, which was isolated on
dung of the North American porcupine. The genus is characterised by pseudoparenchymatous ascomatal initials and
cleistothecia, clavate to ovoid, evanescent short asci and
one-celled hyaline to pale brown ascospores (Hyde et al.
2013). Superficially, Eremomyces resembles some tropical
parasites such as Cleistosphaera and Pilgeriella (Malloch
and Cain 1971). However, it differs from these genera in
having irregularly disposed asci, a cephalothecoid peridium
and in its coprophilous habitat. Eremomyces was compared
to members of Sporormiaceae based on cleistothecial characters and members of Venturiaceae as it produces setose
ascomata (Malloch and Cain 1971). Presently, it is clear
that Eremomyces is distinct from all other genera and it is
included in its own monotypic family Eremomycetaceae
because of apparent lack of close association with any other
group. Cultures and sequence data are available but lacking
for the type.
Eremomyces bilateralis Malloch & Cain, Can. J. Bot. 49(6):
849 (1971).
Index Fungorum number: IF 313945; Facesoffungi number: FoF 05361; Fig. 34
Fungal Diversity (2020) 105:17–318
79
Fig. 34 Eremomyces bilateralis (TRTC 045344, holotype). a Herbarium packet and label. b Hebarium culture c Pycnidium on the upper surface
of agar media. d Squash mount of pycnidium. e Conidia. Scale bars: c = 2000 µm, d = 300 µm, e = 25 µm, f = 50 µm
Saprobic on dung of the North American porcupine. Colonies flocculent, drift white to dark brown, superficial, compact, and growing slowly on agar media. Hyphae septate,
branched, hyaline to brown. Sexual morph: Ascomata solitary, scattered, superficial on hyphae, globose to ellipsoid,
dark brown to black, lacking ostioles. Peridium thin, composed of large brown cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 8-spored, bitunicate,
obovoid, thin-walled, pedicellate, evanescent. Ascospores
multi-seriate, fabiform, hyaline, aseptate, smooth-walled
(adapted from Hyde et al. 2013). Asexual morph: Coelomycetous. Pycnidia in culture brown to dark-brown, globose to
subglobose, thin-walled, papillate. Conidiophores filiform,
septate, hyaline to brown. Conidiogenous cells formed inside
the swollen part. Conidia 3.2–4.8 × 1.7–2.8 μm ( x̄ = 3.85 ×
2.3 μm, n = 20), ellipsoidal, hyaline, aseptate, thin-walled,
guttulate.
Material examined: Canada, Ontario, Leeds Co., East of
Brockville, on porcupine dung in cave, 5 September 1966,
J.C. Krug (TRTC 45344, holotype).
Notes: Eremomyces bilateralis is found on dung of usually sedentary animals, particularly rodents in Canada
(Ontario), Kenya, Tanzania and the USA (California). Colonies are relatively slow growing on most media. Ascomata
tend to be fragmented when immature and lack characteristic cephalothecoid peridium when mature. On the natural
substrate the ascomata are equally conspicuous and produce
short setae compared to long hairs seen in culture.
Other genera included
Rhexothecium Samson & Mouch., Can. J. Bot. 53(16): 1637
(1975).
Index Fungorum number: IF 4699; Facesoffungi number:
FoF 06699; – 1 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Rhexothecium globosum Samson &
Mouch., Can. J. Bot. 53(16): 1637 (1975).
Notes: Rhexothecium globosum was found from desert
soil in Egypt. The genus is characterised by sphaerical
ascomata with a pseudoparenchymatous peridium, clavate
8-spored asci and yellow to yellow-brown sphaerical
ascospores. The asexual morph of Rhexothecium is hyphomycetous and described as trichosporiella-like. Hyde et al.
(2013) accepted Rhexothecium as a genus in Eremomycetaceae. Cultures and sequence data are available but lacking
for the type species.
Economic and ecological significance
Species included in Eremomyces are implicated as opportunistic human pathogens.
Jahnulales K.L. Pang, Abdel-Wahab, El-Shar., E.B.G. Jones
& Sivichai.
Index Fungorum number: IF 90550; Facesoffungi number: FoF 07682.
Jahnulales was introduced in Dothideomycetes based
on phylogenetic analysis of SSU nrDNA sequence data
from Aliquandostipite, Jahnula and Patescospora (Pang
et al. 2002). This order is phylogenetically related to the
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Fungal Diversity (2020) 105:17–318
100/1.0
100/1.0
-/0.91
76/1.0
Jahnula appendiculata BCC11400
Jahnula appendiculata BCC11445
Jahnula bipileata F49-1
Jahnula bipileata AF220-1
100/1.0 Xylomyces chlamydosporus SS0807
91/1.0
Xylomyces chlamydosporus SS2917
Ascagilis seychellensis SS2113.2
Ascagilis seychellensis A492
Ascagilis sunyatsenii UBCF 13876
100/1.0 Jahnula dianchia KUMCC 17-0034
Jahnula dianchia KUMCC 17-0039
Jahnula dianchia S-740
100/1.0
Jahnula dianchia MFLUCC 16-1353
96/Jahnula dianchia MFLUCC 16-0983
100/1.0 100/1.0
Jahnula sangamonensis A482-1B
Jahnula sangamonensis KUMCC 15-0229
100/1.0
Jahnula sangamonensis F81-1
68/0.9
Neojahnula australiensis SS0665
100/1.0
Brachiosphaera tropicalis E192 1
Brachiosphaera tropicalis SS2523
100/1.0 Megalohypha aqua-dulces AF005-2b
71/1.0
Megalohypha aqua-dulces AF005-2a
Pseudojahnula potamophila F111-1
73/1.0
Jahnula rostrata F4-3
Jahnula granulosa SS1567
90/1.0
Jahnula aquatica R68-1
94/0.99 Aliquandostipite crystallinus A514-1
75/1.0
86/-
100/1.0
100/1.0
92/1.0
Aliquandostipite crystallinus F83-1
Aliquandostipite separans CY2787
Aliquandostipite siamensiae SS81-02
Aliquandostipite khaoyaiensis CBS 118232
100/1.0 Manglicola guatemalensis BCC20157
Manglicola guatemalensis BCC20156
94/1.0
97/1.0
100/1.0
100/1.0
-/0.94
100/1.0
Arthrographis arxii IFM 52652
Arthrographis longispora UTHSC 05 3220
Rhexothecium globosum CBS 955.73
100/1.0
Eremomyces bilateralis CBS 781.70
100/1.0 Pseudovirgaria grisea CPC 19134
Pseudovirgaria hyperparasitica CPC 10753
Acrospermum compressum M 151
Acrospermum adeanum M 133
Gonatophragmium triuniae CBS 138901
98/1.0
Acrospermum graminum M 152
100/1.0 Dyfrolomyces tiomanensis NTOU3636
100/1.0
Dyfrolomyces tiomanensis NTOU3636
Dyfrolomyces rhizophorae JK 5456A
Phyllobathelium anomalum 242
99/1.0
Flavobathelium epiphyllum MPN67
Strigula jamesii MPN548
-/0.98
Strigula nemathora MPN72
Hysteropatella elliptica CBS 935.97
Hysteropatella clavispora CBS 247.34
75/1.0
Thyrinulaceae
Acrospermales
72/1.0
Asterotexis cucurbitacearum VIC 24814
Asterotexis cucurbitacearum PMA M 0141224
Paraopeba schefflerae 1620
100/1.0
Thyrinula parasitica CBS 120088
Arthrographis kalrae CBS 693.77
Eremomycetaceae
100/1.0
100/1.0
Manglicolaceae
Asterina cestricola TH 591
Asterina phenacis TH 589
Asterina zanthoxyli TH 561
Asterina fuchsiae TH 590
Asterina weinmanniae TH 592
Asterinales
-/93
100/1.0
Jahnulales
-/1.0
Aliquandostipitaceae
Dyfrolomycetales
Strigulales
Outgroup
Fig. 35 Phylogram generated from maximum likelihood analysis
(RAxML) of Jahnulales based on ITS, LSU and SSU sequence data.
Maximum likelihood bootstrap values equal or above 60%, Bayesian
posterior probabilities equal or above 0.90 (MLBS/PP) are given at
the nodes. Isolate/specimen number is noted after the species name.
The tree is rooted to Hysteropatella clavispora (CBS 247.34) and
H. elliptica (CBS 935.97). The ex-type strains are indicated in bold.
Hyphen (-) represents support values below 70% MLBS and 0.90 PP
Dothideales, Patellariales and Pleosporales (Campbell et al.
2007). Most Jahnulales species occur on rotting or soft
submerged corticated or decorticated wood (Huang et al.
2018). Phylogenetic analyses (Fig. 35) indicate that Jahnulales comprises two major lineages which are designated as
two different families (Aliquandostipitaceae and Manglicolaceae). Jahnula forms as paraphyletic within Aliquandostipitaceae. The divergence time for Jahnulales is estimated as
204 MYA (stem age, Hongsanan et al. 2020).
Accepted families: Aliquandostipitaceae, Manglicolaceae.
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Fungal Diversity (2020) 105:17–318
Aliquandostipitaceae Inderb., in Inderbitzin et al., Am. J.
Bot. 88(1): 54 (2001).
Index Fungorum IF82130; Facesoffungi number: FoF
07683, 36 species.
Saprobic on submerged or trapped decorticated wood
or leaves in freshwater streams, rivers, lakes, peat swamps
and marine habitats. Sexual morph: Ascomata scattered or
rarely in small groups, immersed to erumpent or superficial,
if superficial often attached to substrate by hyphal stalks,
with repent or pendant thick brown hyphal strands, globose
to subglobose, hyaline, pale brown or black, coriaceous to
subcarbonaceous, ostiolate, papillate. Peridium variable,
3–7 cell layers thick, comprising very large, light brown to
brown, thin-walled, cells of textura globulosa to angularis.
Hamathecium comprising 2–7 μm wide, hyaline, septate,
sparsely branched, filamentous, cellular pseudoparaphyses,
anastomosing above the asci. Asci 8-spored, bitunicate, fissitunicate, clavate to cylindrical, pedicel absent or very short,
with or without an ocular chamber. Ascospores 2- to multiseriate, ellipsoid to fusiform, hyaline to pale brown to dark
brown, 1-septate, guttulate, with or without various types
of appendages and/or sheaths. Asexual morph: Hyphomycetous. Conidiophores reduced or un-branched or sparingly
branched, hyaline. Conidiogenous cells monoblastic, thallic
or sympodial or percurrent, monilioid, hyaline or brown.
Conidia phragmosporous or staurosporous, with globose
central cell, with 4-several, radiating septate arms, hyaline
or brown, single or branched or un-branched chains (Seifert
et al. 2011) or tetraradiate with 4–8 appendages.
Type: Aliquandostipite Inderb.
Notes: Aliquandostipitaceae was introduced by Inderbitzin et al. (2001) for taxa characterized by mycelium five
times wider than those of most ascomycetes. Ascomata,
borne either on these thick hyphae or on hyphal stalks are
membranous, have a hamathecium of persistent pseudoparaphyses, bitunicate asci and hyaline to pale brown, 1-septate
ascospores with or without a sheath or with appendages.
Manglicola has also been referred to the Jahnulales and as
the type for Manglicolaceae by Suetrong et al. (2011). Patescospora was synonymised under Aliquandostipite (Suetrong
et al. 2011). The phylogenetic analysis based on ITS and
LSU sequence data showed that the genera Aliquandostipite,
Ascagilis, Brachiosphaera, Jahnula, Megalohypha, Neojahnula Pseudojahnula, and Xylomyces cluster together with
high support (100% ML, 1.00 PP). We follow the previous
studies and accept these eight genera in Aliquandostipitaceae (Dong et al. 2020).
Aliquandostipite Inderb., Am. J. Bot. 88(1): 54 (2001).
Index Fungorum number: IF 28650; Facesoffungi number: FoF 07684; 6 morphological species (Index Fungorum
2020), 6 species with molecular data.
Type species: Aliquandostipite khaoyaiensis Inderb.
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Notes: The genus was introduced for two freshwater lignicolous ascomycetes collected in Thailand and China and it
is characterised by immersed-erumpent or superficial ascomata. Hamathecium consists of pseudoparaphyses. Asci are
bitunicate, and fissitunicate. Hyphae are up to 50 μm wide
(Suetrong et al. 2011; Liu et al. 2015).
Other genera included
Ascagilis K.D. Hyde, Aust. Syst. Bot. 5(1): 109 (1992).
Index Fungorum number: IF 21262; Facesoffungi number: FoF 09187; – 7 morphological species (Dong et al.
2020), 7 species with molecular data.
Type species: Ascagilis bipolaris K.D. Hyde, Aust. Syst.
Bot. 5(1): 111 (1992).
Notes: The type species Ascagilis bipolaris was transferred to Jahnula by Hyde and Wong (1999). Dong et al.
(2020) resurrected Ascagilis to accommodate 7 species
based on its mucilaginous pad of ascospores which is considered as noticeable characteristic of Ascagilis. Phylogenetic analyses supported the placement of Ascagilis in Aliquandostipitaceae (Dong et al. 2020; this study).
Brachiosphaera Nawawi, in Descals et al., Trans. Br. mycol.
Soc. 67: 213 (1976).
Index Fungorum number: IF 7438; Facesoffungi number:
FoF 07685; – 2 morphological species (Species Fungorum
2020), 2 species with molecular data.
Type species: Brachiosphaera tropicalis Nawawi, Transactions of the British Mycological Society 67 (2): 213
(1976).
Notes: The hyphomycetous genus Brachiosphaera is characterised by its effuse colonies, septate hyphae and conidia
frequently formed in chains.
Jahnula Kirschst., Ann. mycol. 34(3): 196 (1936).
Index Fungorum number: IF 2526; Facesoffungi number:
FoF 07686; – 17 morphological species (Huang et al. 2018,
Species Fungorum 2020), 6 species with molecular data.
Type species: Jahnula aquatica (Plötner & Kirschst.)
Kirschst., Ann. Mycol. 34: 196 (1936).
≡ Amphisphaeria aquatica Plöttn. & Kirschst., in
Kirschstner, Verh. bot. Ver. Prov. Brandenb. 48: 52 (1906)
[1907].
Notes: The largest genus within Jahnulales, Jahnula, is
characterised by perithecial, solitary, superficial to subimmersed ascomata, bitunicate, fissitunicate, cylindrical,
pedicellate asci and light brown to dark brown ascospores.
All 17 Jahnula species have been reported from wood or
decorticated wood in freshwater habitats (Huang et al.
2018). The genus is currently polyphyletic, and Jahnula
sensu stricto accommodates J. aquatica, J. granulosa, J.
potamophila and J. rostrata (Suetrong et al. 2011; Huang
et al. 2018).
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82
Fig. 36 Jahnula dianchia (HKAS 92632). a Appearance of ascomata on substrate. b Vertical section of ascoma. c Structure of peridium. d Pseudoparaphyses. e–i Asci. j–o Ascospores. p Germinating
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Fungal Diversity (2020) 105:17–318
ascospore. q, r Colony on MEA. Scale bars: b = 150 μm, c, e–i = 50
μm, j–p = 20 μm, d = 15 μm
Fungal Diversity (2020) 105:17–318
Jahnula dianchia S.K. Huang & K.D. Hyde, Mycol. Progr.
17: 549 (2018).
Index Fungorum number: IF 553200; Facesoffungi number: FoF 03149; Fig. 36
Saprobic on decaying wood submerged in freshwater habitats. Sexual morph: Ascomata 285–390 μm high, 250–350
μm diam, perithecial, solitary, superficial to semi-immersed,
obpyriform to subglobose, black, papillate, ostiolate. Peridium around 40 μm thick, membranous, composed of brown
to hyaline cells of textura angularis. Hamathecium comprising 2–3 μm wide, septate, branched, filiform, cellular
pseudoparaphyses, embedded in a gelatinous matrix. Asci
150–170 µm ( x̄ = 160 µm, SD = 10, n = 15) long, 15–17
μm ( x̄ = 16 µm, SD = 1, n = 15) wide, 8-spored, bitunicate,
fissitunicate, cylindrical, pedicellate, rounded at apex, with
a distinct ocular chamber. Ascospores 27–29 µm ( x̄ = 28
µm, SD = 1, n = 20) long, 11–13 μm ( x̄ = 12 µm, SD = 1,
n = 20) wide, uni-seriate, oval to broadly ellipsoid, slightly
curved, initially pale brown, becoming dark brown at maturity, 1-septate, verruculose, multiguttulate. Asexual morph:
Undetermined.
Material examined: China, Yunnan Province, saprobic on
decaying wood submerged in Erhai Lake, December 2014,
Z.L. Luo, S-364 (HKAS 92632), living culture MFLUCC
16-0983; saprobic on decaying wood submerged in Erhai
Lake, December 2014, H.Y. Su, S-460, living culture
MFLUCC 16-1353; saprobic on decaying wood submerged
in a freshwater stream in Cangshan Mountain, June 2016,
S.M. Tang, S-740.
GenBank numbers: ITS: MH793537, LSU: MH793543
(MFLUCC 16–0983); ITS: MH793538, LSU: MH793544
(MFLUCC 16-1353), LSU: MT797171 (S-740).
Notes: Jahnula dianchia was collected from a freshwater lake in Yunnan Province, China. During our study of
lignicolous freshwater fungi from Northwestern Yunnan
Province, three isolates were obtained from different collections. The morphology of those isolates fit well with Jahnula dianchia, and the phylogenetic analysis showed that
our isolates cluster with this species wih strong support. We
therefore identify our isolates as J. dianchia based on morphology and phylogeny.
Megalohypha A. Ferrer & Shearer, in Ferrer et al., Mycologia 99: 458 (2007).
Index Fungorum number: IF 505741; Facesoffungi number: FoF 07687; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Megalohypha aqua-dulces A. Ferrer &
Shearer, in Ferrer et al., Mycologia 99(3): 456 (2007).
Notes: Megalohypha was introduced based on two collections obtained from submerged wood in tropical forest
streams in Panama and Thailand. Megalohypha is characterized by hyaline, translucent ascomata, with subtending,
83
wide, brown, septate, stoloniferous hyphae, a peridium of
large, thin-walled cells, a hamathecium of septate pseudoparaphyses, clavate, fissitunicate, 8-spored asci and 1-septate, brown, rough-walled ascospores, with longitudinal
sulcate striations (Ferrer et al. 2007).
Neojahnula W. Dong, H. Zhang & K.D. Hyde, Fungal
Divers (revising).
Index Fungorum number: IF 557823; Facesoffungi number: FoF 07688; – 1 morphological species (Dong et al.
2020), 1 species with molecular data.
Type species: Neojahnula australiensis (K.D. Hyde) W.
Dong, H. Zhang & K.D. Hyde, Fungal Divers (revising).
≡ Jahnula australiensis K.D. Hyde, Aust. Syst. Bot. 6(2):
161 (1993).
Notes: The type species, Neojahnula australiensis was
transferred from Jahnula collected from submerged wood in
freshwater (Hyde 1993). Phylogenetically, the species clustered basal to Brachiosphaera within Aliquandostipitaceae
(Prihatini et al. 2008; Hyde et al. 2017; Huang et al. 2018;
Dong et al. 2020). Morphologically, J. australiensis could
not be placed in Jahnula because of the few setae surrounding the ascomata, a thin peridium comprising single row of
cells, asci with an ocular chamber and a faint apical ring
(Dong et al. 2020). Therefore, Dong et al. (2020) introduced
Neojahnula to accommodate this species.
Pseudojahnula W. Dong, H. Zhang & K.D. Hyde, Fungal
Divers (revising).
Index Fungorum number: IF 557824; Facesoffungi number: FoF 07689; – 1 morphological species (Dong et al.
2020), 1 species with molecular data.
Type species: Pseudojahnula potamophila (K.D. Hyde
& S.W. Wong) W. Dong, H. Zhang & K.D. Hyde, Fungal
Divers (revising).
≡ Jahnula potamophila K.D. Hyde & S.W. Wong, Nova
Hedwigia 68(3–4): 499 (1999).
Notes: Pseudojahnula was introduced by Dong et al.
(2020) to accommodate the unstable lineage of Jahnula
potamophila. This genus differs from Jahnula by hyaline to
metallic grey ascomata, a several-layered peridium, comprising hyaline cells and covered with sparse hyaline hairs,
obclavate asci, with an ocular chamber, and ascospores surrounded by a wavy mucilaginous sheath (Hyde and Wong
1999; Dong et al. 2020).
Xylomyces Goos et al., Mycologia 69(2): 282 (1977).
Index Fungorum number: IF 10460; Facesoffungi number: FoF 07690; – 8 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Xylomyces chlamydosporus Goos et al.
[as‘chlamydosporis’], Mycologia 69(2): 282 (1977).
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84
Notes: Xylomyces is characterised by large, dematiaceous,
thick-walled, multi-septate, more or less fusiform chlamydospores that superficially resemble phragmoconidia, however, it has been shown to be polyphyletic and not all species
belong in the Jahnulales.
Ecological significance
Members of Aliquandostipitaceae are saprobes with the ability to decompose lignocellulose in woody litter, and thus
contribute to the recycling of material in aquatic habitats
(Yuen et al. 1998; Bucher et al. 2004).
Manglicolaceae Suetrong & E.B.G. Jones, in Suetrong et al.,
Fungal Divers 51(1): 183 (2011).
Index Fungorum number: IF 563225; Facesoffungi numbers: FoF 07691, 2 species.
Saprobic on intertidal mangrove wood and the brackish water palm Nypa fruticans. Sexual morph: Ascomata
superficial, solitary, or in groups, immersed in the substrate
with a hypostroma, obtuse clavate to fusiform, coriaceous,
olive-brown, pedicellate, ostiolate, epapillate, periphysate.
Peridium thick, 3–5-layered, inner layers hyaline, outer layers pale brown to olive-brown of cells of textura angularis.
Hamathecium comprising numerous, simple, septate, trabeculate pseudoparaphyses. Asci 8-spored, bitunicate, cylindrical, thick-walled, with an ocular chamber and a short pedicel, developed at the base of the ascocarp venter between
the pseudoparaphyses. Ascospores uni-seriate, fusiform,
apiculate, 1-septate, with larger and dark brown upper cell,
with smaller and light brown lower cell, gelatinous appendages covering both ends. Asexual morph: Undetermined.
Type: Manglicola Kohlm. & E. Kohlm.
Notes: The family was introduced to accommodate a
marine ascomycete with large, club-shaped, coriaceous
ascomata, lacking a papilla, and with trabeculate pseudoparaphyses, cylindrical asci and ascospores, with a large apical cell, and a basal turbinate cell with gelatinous appendages at each end. Phylogenetic analyses using LSU and SSU
indicated that Manglicolaceae is related to Aliquandostipitaceae within Jahnulales (100 ML/1.0 PP). This result was
confirmed by many studies, such as Suetrong et al. (2011)
and Hyde et al. (2013). Manglicolaceae is similar to Aliquandostipitaceae in that it has wide hyphae, ascomata
with short stalks, cylindrical asci, and large ascospores with
appendages, and all are saprobic species in aquatic habitats
(Hyde et al. 2013). However, it differs from Aliquandostipitaceae by its marine mangrove habitat, larger ascomata, with
a wide ostiole, surrounded by hyaline, clavate hyphae and
periphysate ostioles, large, unequally, 1-septate, pale brown
ascospores, with a turbinate basal cell and with few asci
per ascoma. Manglicolaceae comprises a monotypic genus,
Manglicola.
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Fungal Diversity (2020) 105:17–318
Manglicola Kohlm. & E. Kohlm., Mycologia 63(4): 840
(1971).
Index Fungorum number: IF 2995; Facesoffungi number:
FoF 07692; 2 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Manglicola guatemalensis Kohlm. & E.
Kohlm.
Notes: Huhndorf (1994) referred Manglicola to the Hypsostromataceae order incertae sedis based on its superficial,
large, elongate ascomata (stalked) with a soft texture, trabeculate pseudoparaphyses, long pedicellate asci attached
in a basal arrangement in the centrum and fusiform, septate ascospores. However, phylogenetic analyses indicated
that Manglicola formed a clade as a family in Jahnulales
(Suetrong et al. 2010, 2011; Hyde et al. 2013), while Hypsostroma saxicola and H. caimitalense (Hypsostromataceae)
are placed in Pleosporales with high bootstrap support (Hyde
et al. 2013). Manglicola contains two species, M. guatemalensis (type species) and M. samuelsii Huhndorf. Manglicola guatemalensis differs from M. samuelsii by its marine
habitat, few asci per ascomata, 1-septate ascospores which
are constricted at the septum, deliquescing appendages
covering both ascospore apices, with an apical cylindrical
appendage and subglobose basal appendage, while M. samuelsii is terrestrial, found on bamboo in lowland rainforest,
has numerous asci per ascomata and 3-septate ascospores
without a sheath or appendages (Jones et al. 2019). Further
studies are required to determine if M. samuelsii is correctly
placed in Manglicolaceae.
Manglicola guatemalensis Kohlm. & E. Kohlm., Mycologia
63(4): 841 (1971).
Index Fungorum number: IF 317136, Facesoffungi number: FoF 06123; Fig. 37
Saprobic on frond bases of Nypa fruticans. Sexual
morph: Ascomata 425–866 µm height, 115–310 µm in diam
around the center, obtusely clavate to fusiform, pedicellate,
solitary or aggregated, superficial, ostiolate, coriaceous,
olive-brown. Ascoma attached by a hypostroma immersed
in the host tissue of N. fruticans. Hamathecium comprising
1.25–2.5 µm in diam., simple, numerous, septate trabeculate,
hyaline, pseudoparaphyses. Asci 300–605 × 30–42.5 µm,
8-spored, cylindrical, bitunicate, thick-walled, with an apical
apparatus. Ascospores 95–112.5 × 20–30 µm, uni-seriate,
fusiform, apiculate, light brown to dark brown, unequally
1-septate, constricted at the septum, apical cell larger, basal
cell, turbinate, light brown, with a gelatinous appendage at
the spore apex. Asexual morph: Undetermined.
Material examined: Thailand, Suratthani Province, Ban
Hui Sap Mangrove Forest, superficial on frond bases of
Nypa fruticans, 8 Jan 2019, S. Preedanon & S. Suetrong
(BBH 45364).
Fungal Diversity (2020) 105:17–318
Fig. 37 Manglicola guatemalensis (BBH 45364). a Superficial ascomata on N. fruticans, partially immersed in mud (MU), composed of
dark ascospores (AS) in mature asci (AC), visible through the thin wall
peridium, spores exuded at the ostiole (OS). b Ascus apex with an apical apparatus. c Trabeculate pseudoparaphyses (PR) numerous, septate,
Notes: The genus has been reported as a saprobe from
bark of dead roots, and from a seedling of Rhizophora mangle in Guatemala (see morphology Suetrong et al. 2010).
Economic and ecological significance
Manglicolaceae contains two species saprobic on woody
material involved in decay of dead woody structures and
likely to be involved in nutrient recycling.
Kirschsteiniotheliales Hernandez-Restrepo, R.F.
Castañeda, Gen. & Crous.
Index Fungorum number: IF 821220; Facesoffungi number: FoF 07693.
Kirschsteiniotheliales was introduced based on phylogenetic analysis. Only a single family Kirschsteiniotheliaceae
is accepted in this order. Phylogenetic analyses indicate that
Kirschsteiniotheliales formed a distinct clade within Dothideomycetes with high support (Fig. 38). The divergence
time for Kirschteiniotheliales is estimated as 221 MYA
(stem age, Hongsanan et al. 2020).
85
simple. d Ascospore in ascus with apical (AA) and basal (BA) appendages. e–i Ascospores. j Germinating ascospore, with germ tube developing laterally from the basal cell. Scale bars: a= 250 µm, b, e–j = 25
µm, c = 15 µm, d = 20 µm
Accepted families: Kirschsteiniotheliaceae.
Kirschsteiniotheliaceae Boonmee & K.D. Hyde, Mycologia
104(3): 705 (2012).
Index Fungorum number: IF 561021; Facesoffungi number: FoF 01737, 24 species.
Saprobic on decaying wood. Sexual morph: Ascomata
superficial, subglobose to globose, dark brown to black,
membranaceous, solitary, with a central papilla. Peridium
composed of cells of textura angularis. Hamathecium comprising numerous, filiform, hyaline, mostly cellular pseudoparaphyses. Asci 8-spored, bitunicate, cylindrical to clavate,
with a long pedicel, apical rounded, with an ocular chamber.
Ascospores 2-seriate, ellipsoidal, septate, slightly curved, dull
green, brown to dark brown at maturity, with median septum or in lower part, some ascospores with secondary septa,
thick-walled, lacking a mucilaginous sheath. Asexual morph:
Hyphomycetous. Conidiophores macronematous, mononematous, erect, apically branched, arising straight or slightly
curved, solitary, elongate and thick-walled, septate, brown to
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Fungal Diversity (2020) 105:17–318
100/1.0 Kirschsteiniothelia aethiops S-1322
Kirschsteiniothelia aethiops S-1406
88/1.00
Kirschsteiniothelia aethiops CBS 109.53
100/1.0 Kirschsteiniothelia aethiops MFLUCC 15-0424
Kirschsteiniothelia aethiops S-783
Kirschsteiniothelia aethiops DAOM 231155
Kirschsteiniothelia lignicola MFLUCC 10-0036
98/1.00 Kirschsteiniothelia rostrata MFLUCC 15-0619
96/1.00 Kirschsteiniothelia rostrata MFLUCC 16-1124
100/1.00
Kirschsteiniothelia tectonae MFLUCC12-0050
100/1.00
Kirschsteiniothelia thujina JF13210
94/1.00
Kirschsteiniothelia submerse S-481
Kirschsteiniothelia submerse MFLUCC 15-0427
100/1.00 Kirschsteiniothelia submerse S-601
100/1.00
100/1.00 Kirschsteiniothelia arasbaranica IRAN2509C
Kirschsteiniothelia arasbaranica IRAN2508C
100/1.00 100/1.00
Kirschsteiniotheliales
Kirschsteiniothelia aethiops MFLUCC 16-1104
100/1.0
Kirschsteiniothelia fluminicola MFLUCC 16-1263
Kirschsteiniothelia phoenicis MFLUCC 18-0216
Kirschsteiniothelia aquatica MFLUCC 17-1685
100/1.00
78/-
Kirschsteiniothelia cangshanensis MFLUCC 16-1350
Acrospermum adeanum M133
100/1.00
100/1.00
83/0.99
Acrospermum compressum M151
Acrospermum gramineum M152
100/1.00
Dyfrolomyces tiomanensis NTOU3636
Dyfrolomyces rhizophorae JK 5456A
Phyllobathelium anomalum MPN242
Phyllobathelium firmum
87/-
Flavobathelium epiphyllum MPN67
100/1.00
Strigula jamesii MPN548
100/1.00
100/1.00
Jahnula bipileata F49-1
-/100/1.00
-/-
Jahnula sangamonensis A402-1B
Jahnula seychellensis SS 2113.2
Aliquandostipite khaoyaiensis CBS 118232
Aliquandostipite crystallinus R76-1
Jahnula aquatica R68-1
Homortomyces tamaricis MFLUCC 13-0280
100/1.00 Homortomyces tamaricis MFLUCC 14-0167
100/1.00
83/1.00
Homortomyces tamaricis MFLUCC 13-0441
Homortomyces combreti CPC 19808
Tubeufia javanica MFLUCC 12-0545
Tubeufia cylindrothecia BCC 3559
100/1.00 Helicomyces roseus CBS 283.51
100/1.00 Helicomyces roseus MFLUCC 15-0343
0.06
100/-
Pleospora herbarum CBS 191.86
Pleospora herbarum MFLUCC 14-0920
HomortoJahnulales mycetaceae Tubeufiales
100/1.00
Dyfrolomycetales
Strigulales
100/1.00
Acrospermales
Outgroup
Fig. 38 Phylogram generated from maximum likelihood analysis (RAxML) of Kirschsteiniotheliales based on ITS, LSU and SSU
sequence data. Maximum likelihood bootstrap values equal or above
70%, Bayesian posterior probabilities equal or above 0.90 (MLBS/
PP) are given at the nodes. Isolate/specimen number is noted after
the species name. The tree is rooted to Pleospora herbarum (CBS
191.86, MFLUCC 14–0920). The ex-type strains are indicated in bold.
Hyphen (-) represents support values below 70% MLBS and 0.90 PP
dark brown, smooth-walled. Conidiogenous cell tretic, terminal, constricted at delimiting septa. Conidia broadly ellipsoid-obovoid, brown to dark brown, constricted and darkly
pigmented at the septa, rounded at both ends, smooth-walled.
Type: Kirschsteiniothelia D. Hawksw., J. Linn. Soc., Bot.
91: 182 (1985).
Notes: Kirschsteiniotheliaceae species are mostly saprobes on dead wood from terrestrial and aquatic habitats
(Boonmee et al. 2012; Hyde et al. 2013; Su et al. 2016).
Taeniolella is an asexual genus with saprobic, endophytic
and lichenicolous life styles. It was found to be a polyphyletic genus distributed between the Dothideomycetes and
the Sordariomycetes (Ertz et al. 2016). The type species
Taeniolella exilis formed a lineage within Kirschsteiniotheliaceae (Ertz et al. 2016), therefore, the genus was placed
in Kirschsteiniotheliaceae by Ertz et al. (2016) and Wijayawardene et al. (2017a). However, Heuchert et al. (2018) and
Dong et al. (2020) suggested that four freshwater species of
Taeniolella are undoubtedly not congeneric with Taeniolella
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Fungal Diversity (2020) 105:17–318
87
Fig. 39 The sexual morph characters of Kirschsteiniothelia spp.
a, b Appearance of ascomata on host surface. c, d Section through
ascoma. e, f Peridium. i, j Immature asci. k, l Asci. m–s Ascospores.
Note: a, c, e, g, i, k, m–p = Kirschsteiniothelia lignicola (holotype);
b, d, f, h, j, l, q–s = Kirschsteiniothelia emarceis (holotype)
sensu lato. Thus, we suggest to exclude this genus from
Kirschsteiniotheliaceae and further studies are needed to
resolve placement of this group.
Kirschsteiniothelia aethiops (Sacc.) D. Hawksw., J. Linn.
Soc., Bot. 91(1–2): 185 (1985).
≡ Amphisphaeria aethiops Sacc., Syll. fung. (Abellini)
1: 722 (1882).
Index Fungorum number: IF 104401; Facesoffungi number: FoF 01738; Fig. 40
Saprobic on decaying wood submerged in freshwater
habitats. Sexual morph: Undetermined. Asexual morph:
Colonies on the natural substrate superficial, effuse, gregarious, hairy, dark brown to black. Mycelium immersed,
composed of septate, branched, thin-walled, smooth, pale
brown hyphae. Coinidiophores 148–228 µm ( x̄ = 188 µm,
SD = 40, n = 20) long, 6–8 μm ( x̄ = 7 µm, SD = 1, n = 20)
wide, mononematous, macronematous, septate, usually with
one or a number of short branches near apex, erect, straight
Kirschsteiniothelia D. Hawksw., J. Linn. Soc., Bot. 91: 182
(1985).
Index Fungorum number: IF 25723; Facesoffungi number: FoF 08040; 24 morphological species (Bao et al. 2018;
Species Fungorum 2020), 11 species with molecular data.
Type species: Kirschsteiniothelia aethiops (Sacc.) D.
Hawksw., Bot. J. Linn. Soc. 91(1–2): 185 (1985).
Notes: Kirschsteiniothelia has little DNA sequence data
for Kirschsteiniothelia species available in GenBank, thus,
more collections and sequence data are needed (Fig. 39).
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88
Fig. 40 Kirschsteiniothelia aethiops (S-1322) a Colonies on natural
substrate. b, c Conidiophores with conidia. d, e Conidiogenous cells.
f–i Conidia. j Germinating conidium. k, l Culture on PDA from sur-
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Fungal Diversity (2020) 105:17–318
face and reverse. Scale bars: b = 90 μm, c = 70 μm, d, e = 40 μm, f–i
= 15 μm, j = 30 μm
Fungal Diversity (2020) 105:17–318
89
Fig. 41 Lembosina aulographoides (redrawn from Crous et al.
2019b, CPC 33049, holotype).
a Hysterothecioid ascomata on
host surface. b, c Young asci. d
Asci viewed from above. e Asci
when mature. f Ascospores.
Scale bars: a = 250 μm, b–f =
10 μm
or flexuous, smooth, brown to dark brown, gradually paler
towards the apex, cylindrical, septate. Conidiogenous cells
25–33 µm ( x̄ = 29 µm, SD = 4, n = 20) long, 5–7 μm ( x̄ =
6 µm, SD = 1, n = 20) wide, tretic, integrated, sometimes
percurrent, terminal, subhyaline to pale brown, doliiform
or lageniform, new cell developing from apical or subapical part of subtending cell. Conidia 41–53 µm ( x̄ = 47 µm,
SD = 6, n = 40) long, 12–16 μm ( x̄ = 14 µm, SD = 2, n =
40) wide, acrogenous, solitary, dry, cylindrical, sometimes
clavate, rounded at the apex, 3–4-septate, constricted and
darkly pigmented at the septa, smooth, brown.
Material examined: China, Yunnan Province, saprobic on
submerged decaying wood in a freshwater stream in Laojunshan Mountain, H.W. Shen, June 2017, S-1322.
Sequence data: ITS MH793539, LSU MH793545, SSU
MH793556.
Notes: Kirschsteiniothelia aethiops is the generic type
of Kirschsteiniothelia. Phylogenetic analysis based on
ITS, LSU and SSU sequence data showed that our strains
(S-1322 and S-1406) are identical to the strains of K. aethiops (Fig. 38). The morphology of our isolates also fit well
with K. aethiops (Su et al. 2016).
Economic and ecological significance
Poch et al. (1992) reported the first chemical examination of
a Kirschsteiniothelia species and extracted five major compounds from cultures of this species. Species of Kirschsteiniotheliales have the potential as important sources of novel
biologically active secondary metabolites. Most Kirschsteiniothelia species are saprobes involved in nutrient cycling.
Lembosinales Crous.
Index Fungorum number: IF 832086; Facesoffungi number: FoF 07694.
Crous et al. (2019b) provided the first molecular data for
Lembosina and found that it formed a distinct lineage next
to Lichenoconiales. Therefore, they introduced Lembosinales to accommodate this lineage. The divergence time
for Lembosinales is estimated as 253 MYA (Hongsanan
et al. 2020).
Accepted families: Lembosinaceae.
Lembosinaceae Crous, in Crous et al., Fungal Systematics
and Evolution 5: 91 (2019) [2020].
Index Fungorum number: IF 832087; Facesoffungi number: FoF 07695, 26 species.
Sexual morph: Ascomata thyriothecial or hysterothecial, linear, rarely y-shaped, solitary, gregarious, superficial,
loose on host surface, black, opening with linear fissures.
Upper wall comprises thin layer of mostly neatly arranged
dark cells, branched at the outer rim, poorly developed at
the base. Hamathecium comprising sparse, filiform, cellular
pseudoparaphyses. Asci 8-spored, bitunicate, subglobose to
oblong. Ascospores 2–4-seriate, hyaline to brown, 1-septate,
upper cell slightly wider and shorter than the lower cell,
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90
Fungal Diversity (2020) 105:17–318
Lichenothelia convexa L1607
Lichenothelia convexa L1606
70/Lichenothelia convexa L1844
99/1.0
Lichenothelia calcarea L1706
Lichenothelia calcarea L1324
90/Lichenothelia arida L2196
70/Lichenothelia arida L2195
96/1.0
Lichenothelia arida L1707
Lichenothelia sp. L984
-/0.9
Lichenothelia rugosa Ertz 16065 BR
Anisomeridium ubianum MPN94
84/1.0
75/Megalotremis verrucose MPN104
100/0.94
Acrocordia subglobosa HTL940
Heleiosa barbatula JK 5548I
74/0.93
Lichenothelia calcarea L1840
Lichenothelia convexa L1852
Lichenothelia calcarea L1799
100/1.0 Cladoriella rubrigena CBS 124760
Cladoriellales
Cladoriella eucalypti CPC 10953
Lichenothelia calcarea L1717
Abrothallus parmeliarum AB36
Abrothallales
99/1.0
Abrothallus cladoniae AB53
100/100/1.0 Patellaria quercus CPC 27232
Patellaria cf. atrata BCC 28877
Hysteropatella prostii H B 9934b
Botryosphaeria dothidea CMW 8000
82/1.0
Lasiodiplodia crassispora CBS 118741
100/1.0
Aplosporella africana CMW 25424
100/1.0
Curvularia lunata CBS 730.96
Outgroup
Pleospora herbarum CBS 191.86
78/100/1.0
Lichenotheliales
s. str.
Monoblastiaceae
Lichenotheliales (1)
Patel- Botryolariales sphaeriales
0.03
Fig. 42 Phylogram generated from maximum likelihood analysis (RAxML) of Lichenotheliales based on LSU, SSU and mtSSU
sequence data. Maximum likelihood bootstrap values equal or above
70%, Bayesian posterior probabilities equal or above 0.90 (MLBS/
PP) are given at the nodes. An original isolate number is noted after
the species name. The tree is rooted to Curvularia lunata (CBS
730.96) and Stemphylium versicarium (CBS 191.86). The ex-type
strains are indicated in bold. Hyphen (-) represents support values
below 70% MLBS and 0.90 PP
with basal protrusion (Crous et al. 2019b). Asexual morph:
Undetermined.
Type: Lembosina Theiss.
Notes: This family was established based on phylogenetic
placement of Lembosina aulographoides. More fresh collections and sequence data are needed to confirm its placement.
Index Fungorum number: IF 2728; Facesoffungi number:
FoF 07696; 26 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Lembosina aulographoides (E. Bommer
et al.) Theiss.
Notes: Lembosina was placed in Leptopeltidaceae and
Asterinaceae by von Arx and Müller (1975) and Hawksworth et al. (1995), respectively. Hyde et al. (2013) placed
this genus in Aulographaceae. However, Crous et al. (2019b)
indicated that Lembosina based on L. aulographoides, does
Lembosina Theiss., Annls mycol. 11(5): 437 (1913).
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91
not belong to any of these families and should be placed in
its own family and order.
Lembosina aulographoides (E. Bommer, M. Rousseau &
Sacc.) Theiss., Annls mycol. 11(5): 437 (1913).
≡ Lembosia aulographoides E. Bommer et al., Bull. Soc.
R. Bot. Belg. 29(1): 238 (1890).
Index Fungorum number: IF 148168; Facesoffungi number: FoF 07697; Fig. 41
Description: see Crous et al. (2019b).
Economic and ecological significance
The life style of Lembosiaceae is unclear, however, they tend
to be saprobes and involved in nutrient recycling.
Lichenotheliales K. Knudsen, Muggia & K.D. Hyde.
Index Fungorum number: IF 805298; Facesoffungi number: FoF 07698.
The order Lichenotheliales comprises non-lichenized
taxa, including saprobic, lichenicolous, epiphytic or endolithic species growing on or within rocks where they are
associated with algae (Hyde et al. 2013). These fungi are
found in hot and cold arid regions worldwide (Muggia et al.
2015). Lichenotheliales comprises only Lichenotheliaceae
which comprises one genus Lichenothelia and Endococcus. Ametrano et al. (2019) indicated that Lichenotheliales formed two clades within Dothideomycetes named
Lichenotheliales sensu stricto and Lichenotheliales (1). The
same result is shown in our analyses (Fig. 42). Lichenotheliales sensu stricto is monophyletic (Ametrano et al. 2019).
The divergence time for Lichenotheliales is estimated as 265
MYA (stem age, Hongsanan et al. 2020).
Accepted families: Lichenotheliaceae.
Lichenotheliaceae Henssen, Syst. Ascom. 5: 137 (1986).
Index Fungorum number: IF 81652; Facesoffungi numbers: FoF 07699, 51 species.
Saprobic or lichenicolous. Endolithic or epilithic thallus
in saxicolous lifestyle. Lichenicolous shows episubstratic or
endokapylic thallus black, dispersed or continuous, areolate or not, frequently producing black superficial or rarely
effigurate, hyphae, branching or not, sometimes connecting
scattered stromata; meristematic growth frequent. Sexual
morph: Ascomata perithecioid and with interascal filaments
or fertile stromata without an ostiole and with asci forming
in pseudoparenchymatous locules. Asci 8-spored, globose
to broadly clavate, bitunicate, sometimes K/I+bluish around
the outer layers or apex of the ascus. Interascal gel amyloid
or not. Ascospores uni- to 2-seriate, partially overlapping,
hyaline or brown, 1-septate to muriform (sometimes variable). Asexual morph: Undetermined.
Type: Lichenothelia D. Hawksw.
Fig. 43 Morphology of Lichenothelia spp. (redrawn from Hawksworth 1981a). a Asci of Lichenothelia scopularia. b Ascospores of
Lichenothelia metzleri
Notes: The family Lichenotheliaceae was introduced to
encompass Lichenothelia and Lichenostigma in Dothideales (Henssen and Jahns 1973; Hyde et al. 2013; Valadbeigi
et al. 2016). Lichenostigma was placed with Lichenothelia
in Lichenotheliaceae based on morphology (Henssen 1987;
Muggia et al. 2015). However, based on phylogenetic analysis Lichenostigma was excluded from Lichenotheliaceae
and placed to Lichenostigmatales in Arthoniomycetes (Ertz
et al. 2014; Hyde et al. 2013; Muggia et al. 2013, 2015).
Presently, Lichenotheliaceae comprises two genera namely
Lichenothelia and Endococcus. Endococcus was formally
referred to Dothideales (Hawksworth 1979). Lumbsch and
Huhndorf (2010) and Hyde et al. (2013) placed Endococcus
in Dothideomycetes genera incertae sedis. However, Jayasiri et al. (2016) included this genus in Lichenotheliaceae
based on morphological similarities to the type species of
Lichenothelia. Lichenotheliaceae is placed in Lichenotheliales based on morphological, habitat and molecular evidence
(Hyde et al. 2013). Both sexual and asexual morphs have
been recorded as saprobic, lichenicolous or lichenized taxa
(Hyde et al. 2013; Valadbeigi et al. 2016; Wijayawardene
et al. 2017a).
Lichenothelia D. Hawksw., Lichenologist 13(2): 142 (1981).
Index Fungorum number: IF 2855; Facesoffungi number
FoF07700; 11 morphological species (Species Fungorum
2020), 9 species with molecular data.
Type species: Lichenothelia scopularia (Nyl.) D.
Hawksw.
≡ Verrucaria scopularia Nyl., Not. Sällsk. Fauna et Fl.
Fenn. Förh., Ny Ser. 3: 85 (1861).
Notes: The cosmopolitan genus Lichenothelia is characterized by fertile ascomata and many species are described
13
92
based on morphological identification (Ametrano et al.
2019). Both sexual and asexual morphs have been recorded
(Valadbeigi et al. 2016; Wijayawardene et al. 2017a) and
the taxa show a wide range of life strategies including rock
inhabiting, lichenized, lichenicolous and saprobic taxa
(Ametrano et al. 2019). Based on modern phylogeny, this
genus is incorporated in the order Lichenotheliales (Hyde
et al. 2013). Broader sampling with informative loci at the
species level is required for species delimitation within this
genus (Ametrano et al. 2019) (Fig. 43).
Endococcus Nyl., Mém. Soc. Sci. Nat. Cherbourg 3: 193
(1855).
Index Fungorum number: IF 1791; Facesoffungi number
FoF 00560; 40 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Endococcus rugulosus Nyl., Mém. Soc.
Imp. Sci. Nat. Cherbourg 3: 193 (1855).
=Verrucaria rugulosa Borrer ex Leight., Brit. Sp. Ang.
Lich.: 47 (1851).
Notes: Endococcus species occur on crustose, foliose, and
fruticose lichens and some are lichenicolous, while others
colonize diverse rock types (Halıcı et al. 2007; Knudsen
2008; Jayasiri et al. Hong). Species are commonly hostspecific, while some species have wide host ranges (Halıcı
et al. 2007; Jayasiri et al. 2016). Endococcus was affiliated
in Dothideales (Hawksworth 1979). However, the taxonomic
placement of this genus has changed over time (Hafellner 2019). Jayasiri et al (2016a) transferred this genus to
Lichenotheliaceae based on the morphological characteristics, such as perithecioid ascomata with widely porate ostioles, globose to broadly clavate asci and hyaline or brown
with 1-septate ascospores. However, Diederich et al (2018)
accepted this genus under Verrucariaceae while Wijayawardene et al (2018) accepted in uncertain placement in
Dothideomycetes. Molecular data are lacking for this genus
and needs critical revision (Hafellner 2019).
Economic and ecological significance
Species such as Lichenothelia scopularia, Lichenothelia arida, L. calcarea and L. convexa are lichenicolous
(Hawksworth 1981a), while other species are reported as
lichenized and non-lichenized (Hyde et al. 2013). Ametrano
et al. (2019) stated that Lichenothelia consists of rock-inhibiting taxa which colonize natural and anthropogenic lithic
substrates.
Microthyriales G. Arnaud.
Index Fungorum number: IF 90485; Facesoffungi number: FoF 07701.
Arnaud (1918) introduced this order, with type family
Microthyriaceae. Microthyriales contained two families
viz. Microthyriaceae and Micropeltidaceae based on their
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Fungal Diversity (2020) 105:17–318
flattened ascomata with poorly developed base. However,
members of Microthyriaceae have brown to black thyriothecia, comprising cuboid or angular cells arranged in parallel
rows, with prominent central ostioles, and usually uniseptate
ascospores (Doidge 1942; Müller and von Arx 1962; Luttrell 1973; Barr 1987b; Hofmann and Piepenbring 2006;
Wu et al. 2011b; Hyde et al. 2013; Hongsanan et al. 2015b;
Hongsanan and Hyde 2017), while Micropeltidaceae has
black-blue or greenish thyriothecia, comprising interwoven
hyphae, with a central ostiole, and multiple trans-septate
ascospores (Clements and Shear 1931; Batista 1959; von
Arx and Müller 1975; Barr 1987b; Wu et al. 2011b; Hyde
et al. 2013; Hongsanan and Hyde 2017; Hongsanan et al.
2015a; Zeng et al. 2019). Hongsanan and Hyde (2017)
excluded Micropeltidaceae from Microthyriales based on
their phylogenetic analyses. This was supported by new
collections of Micropeltidaceae in Zeng et al. (2019).
Therefore, Microthyriales contains a single family, Microthyriaceae based on morphology and phylogeny. Species
of Microthyriales cluster together as a distinct clade within
Dothideomycetes with high support (Fig. 44). The divergence time for Microthyriales is estimated as 184 MYA
(stem age, Hongsanan et al. 2020).
Accepted families: Microthyriaceae.
Microthyriaceae Sacc., Syll. fung. (Abellini) 2: 658 (1883).
Index Fungorum number: IF 81008; Facesoffungi number: FoF 06747, 120 species.
Saprobic on leaves. Hyphae mostly absent, superficial,
brown, septate, branched, reticulate. Sexual morph: Thyriothecia superficial, circular, flattened, dark brown, solitary or
gregarious, with a round, central ostiole, with poorly developed basal plate. Upper wall brown, comprising radially
arranged cells of textura prismatica, often darkened at the
ostiole and tapering at the margin. Hamathecium comprising asci inclined from the base and rim towards the central
ostiole. Pseudoparaphyses present or absent. Asci 8-spored,
bitunicate, fissitunicate, cylindrical to obpyriform, with or
without a pedicel, with or without an ocular chamber at the
tip. Ascospores overlapping uni- to 3-seriate, fusiform to
ellipsoidal, hyaline or brown, usually 1-septate, with cilia
or appendages, smooth-walled. Asexual morph: Mycelium
consisting of hyaline to pale brown, smooth to verruculose, branched, septate hyphae. Conidiophores micronematous or macronematous, mononematous, arising from a
brown stroma or from superficial hyphae, erect, solitary,
unbranched or branched, straight or flexuous, smooth or
roughened, septate, subcylindrical, pale brown to brown;
microconidiophores when present, reduced to conidiogenous cells on hyphae, visible as slight thickenings on hyphal
cells, somewhat erumpent, pale brown, truncate apex. Conidiogenous cells mono- to polyblastic, integrated, terminal,
smooth or verruculose, determinate or sympodial, clavate,
Fungal Diversity (2020) 105:17–318
93
100/1.0
Microthyrium propagulensis IFRD 9037
Sympodiella multiseptata CBS 566.71
90/1.0
100/1.0
99/1.0
Microthyrium microscopicum CBS 115976
Spirosphaera beverwijkiana CBS 469.66
Anungitopsis speciosa CBS 181.95
97/1.0
100/1.0
Neoanungitea eucalypti CBS 143173
Condylospora vietnamensis NBRC 107639
100/1.0
-/0.98
Hamatispora phuquocensis VTCCF1219
Pseudomicrothyrium thailandicum MFLU14-0286
100/1.0
100/1.0
100/1.0
Heliocephala gracilis MUCL 41200
Heliocephala elegans MUCL 39003
Heliocephala natarajanii MUCL 43745
Heliocephala zimbabweensis MUCL 40019
Stomiopeltis betulae CBS 114420
75/0.99
76/1.0
Microthyriaceae
Microthyriales
Microthyrium buxicola MFLUCC 15-0212
Spirosphaera minuta CBS 476.66
86/-
Chaetothyriothecium elegans CPC 21375
Tumidispora shoreae MFLUCC 14-0574
87/0.94
100/1.0
89/0.95
Pseudopenidiella piceae CBS 131453
Pseudopenidiella gallaica CBS 121796
Ochroconis dracaenae CPC 26115
100/1.0
Sympoventuria capensis CBS 120136
74/71/0.98
100/1.0
Venturia inaequalis CBS 594.70
Trichodelitschia bisporula CBS 262.69
Phaeotrichum benjaminii CBS 541.72
78/0.99
100/1.0
Zeloasperisporium siamense IFRDCC 2194
Zeloasperisporium ficusicola MFLUCC 15-0221
Zeloasperisporium hyphopodioides CBS 218.95
100/1.0
Natipusilla naponensis AF217-1
Natipusilla decorospora AF236-1
Kirschsteiniothelia lignicola MFLUCC10-0036
Venturiales
Phaeotrichales
Zeloasperisporiales
Natipusillales
Outgroup
0.1
Fig. 44 Phylogram generated from maximum likelihood analysis
(RAxML) of Microthyriales and related orders based on ITS and
LSU sequence data. Maximum likelihood bootstrap values equal
or above 70%, Bayesian posterior probabilities equal or above 0.90
(MLBS/PP) are given at the nodes. An original isolate number is
noted after the species name. The tree is rooted to Kirschsteiniothelia lignicola (MFLUCC10-0036). The ex-type strains are indicated
in bold. Hyphen (-) represents support values below 70% MLBS and
0.90 PP
subcylindrical, ampulliform or ovoid. Ramoconidia when
present, pale brown, verruculose, subcylindrical to fusoidellipsoid, aseptate, giving rise to branched chains of conidia.
Conidia solitary or in branched chains, sometimes radial on
compact heads, acrogenous or acropleurogenous, verruculose, subcylindrical to ellipsoid, obclavate, pale brown, aseptate to multi-septate, sometimes rostrate; hilum inconspicuous, truncate, sometimes unthickened, sometimes thickened;
conidia in Hamatispora consisting of a main axis and lateral
branches; main axis hook-shaped with a long tail bearing
lateral branches; lateral branches straight,developing from
different cells of the helicoid part of the main axis.
Type: Microthyrium Desm.
Notes: The phylogenetic placement of this family was first
provided by Wu et al. (2011b), represented by Microthyrium
microscopicum. New genera and new species in Microthyriaceae with molecular data were provided to support the
placement of Microthyriaceae. Eleven genera are accepted
in this family based on morphology and phylogeny.
The genera Condylospora, Spirosphaera and Stomiopeltis are not included in the family due to their polyphyletic characters or lack of evidence. Caribaeomyces is
excluded from the family as it is typical of Asterinales by
its hyphopodia and structure of upper walls. Hansfordiella
and Isthmospora are also excluded from the family, as they
are asexual morphs of Trichothyrium (Trichothyriaceae)
(Wijayawardene et al. 2017a). Monorhizina was referred as a
member of Microthyriaceae in some studies. This genus was
synonymised under Lauterbachiella and Rhagadolobium by
Dingley (1972) and Müller and von Arx (1962), respectively.
13
94
Hofmann (2009) accepted Monorhizina in Microthyriaceae,
while Hyde et al. (2013) did not accept this genus. Kirk
et al. (2013) retained Monorhizina as a distinct genus but
did not assign it to any family. Wijayawardene et al. (2017a)
listed Monorhizina as a genus in Microthyriaceae. We do
not accept Monorhizina in Microthyriaceae and note that
the placement of this genus in Microthyriaceae needs future
study.
Microthyrium Desm., Annls Sci. Nat., Bot., sér. 2 15: 137
(1841).
Index Fungorum number: IF 3206; Facesoffungi number:
FoF 06748; 91 morphological species (Species Fungorum
2020), 3 species with molecular data.
Type species: Microthyrium microscopicum Desm.,
Annls Sci. Nat., Bot., sér. 2 15: 138 (1841).
Notes: Microthyrium is characterized by flattened thyriothecia with cells of radial arrangement and hyaline
ascospores with cilia.
Other genera included
Arnaudiella Petr., Annls mycol. 25(3/4): 339 (1927).
Index Fungorum number: IF 311; Facesoffungi number:
FoF 06749; – 10 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Arnaudiella caronae (Pass.) Petr., Annls
mycol. 25(3/4): 339 (1927).
≡ Seynesia caronae Pass., Atti Reale Accad. Lincei, Rendic., Sér. 4 4(2): 63 (1888).
Notes: Arnaudiella is similar to Microthyrium in having flattened thyriothecia and the radial arrangement of
upper wall with a darkened ostiole, but is distinct in brown
ascospores.
Calothyriopsis Höhn., Sber. Akad. Wiss. Wien, Math.naturw. Kl., Abt. 1 128(7–8): 552 (1919).
Index Fungorum number: IF 761; Facesoffungi number:
FoF 06750; – 4 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Calothyriopsis conferta (Theiss.) Höhn.,
Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 128(7–8):
552 (1919).
≡ Microthyrium confertum Theiss., Annls mycol. 7(4):
352 (1909).
Notes: Calothyriopsis is typical of Microthyriaceae in
having flattened thyriothecia and 1-septate ascospores. However, the arrangement of upper wall is different from the
type genus. Therefore, its placement needs to be confirmed
by phylogeny.
Chaetothyriothecium Hongsanan & K.D. Hyde, Phytotaxa
161(2): 161 (2014).
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Fungal Diversity (2020) 105:17–318
Index Fungorum number: IF 804964; Facesoffungi number: FoF 06751; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Chaetothyriothecium elegans Hongsanan
& K.D. Hyde, Phytotaxa 161(2): 161 (2014).
Notes: Chaetothyriothecium was introduced with a single
species C. elegans, which typically has radial setae growing
around a central ostiole. Based on the flattened thyriothecia
with cells in a radial arrangement and phylogenetic analyses,
Chaetothyriothecium is accommodated in Microthyriaceae.
Sequence data of LSU is available for the genus
Hamatispora L.T.H. Yen, K. Yamag. & K. Ando, in Yen
et al., Mycoscience 59(6): 468 (2018).
Index Fungorum number: IF 822829; Facesoffungi number: FoF 06752; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Hamatispora phuquocensis L.T.H. Yen,
K. Yamag. & K. Ando, in Yen et al., Mycoscience 59(6):
468 (2018).
Notes: Hamatispora is a hyphomycetous genus with
staurospores that are question mark-shaped or hook-shaped
with 3 arms developing from each cell on the helicoid part
(Yamaguchi et al. 2018). It is considered as a member of
Microthyriaceae based on the phylogeny.
Neoanungitea Crous, in Crous et al., Persoonia 39: 359
(2017).
Index Fungorum number: IF 823489; Facesoffungi number: FoF 07702; – 2 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Neoanungitea eucalypti Crous, in Crous
et al., Persoonia 39: 359 (2017)
Notes: Neoanungitea was introduced with a single species
that shares features of both Anungitea and Anungitopsis. It
is characterized by conidiogenous cells forming a terminal
rachis with flat-tipped sympodial loci. Crous et al. (2017)
classified Neoanungitea as a member of Microthyriaceae
based on phylogenetic analysis. Sequence data of ITS and
LSU is available for the genus.
Paramicrothyrium H.X. Wu & K.D. Hyde, in Wu, Schoch,
Boonmee, Bahkali, Chomnunti & Hyde, Fungal Diversity
51(1): 204 (2011).
Index Fungorum number: IF 563363; Facesoffungi number: FoF 06754; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Paramicrothyrium chinensis H.X. Wu &
K.D. Hyde [as ‘chinensis’], in Wu et al., Fungal Diversity
51(1): 204 (2011).
Notes: Paramicrothyrium was introduced with a single
species Paramicrothyrium chinensis, which is characterized
by irregular ostioles without a darkened ring. However, the
Fungal Diversity (2020) 105:17–318
95
Fig. 45 Pseudomicrothyrium thailandicum (MFLU 14-0286, holotype). a, b Colony on the surface of host leaf. c Ascoma. d, e Ascoma
in cross section. f Basal layer. g Pseudoparaphyses. h–k Asci from
young state to mature state. l–n Ascospores from young state to
mature state. o Germinating ascospore. p Ascoma with hyphae. Scale
bars: a = 400 μm, b, p =100 μm, c, d = 50 μm, e, f = 20 μm, g–o =
10 μm
sequence data of the type species is questionable, as it does
not cluster with the main clade of Microthyriales.
filiform, narrow cellular pseudoparaphyses. Asci 8-spored,
bitunicate, fissitunicate, ovoid to obpyriform. Ascospores
2–3-seriate, fusiform to ellipsoidal, hyaline, 1-septate,
upper cell slightly wider than lower cell, not constricted at
the septum, with 2–4 apical appendages, usually with 2 large
guttules in the upper cell, 2 small guttules in the lower cell,
smooth-walled. Asexual morph: Undetermined.
Type species: Pseudomicrothyrium thailandicum X.Y.
Zeng, S. Hongsanan & K.D. Hyde.
Notes: Pseudomicrothyrium is similar to Microthyrium in
having hyaline ascospores with cilia around the ostiole, but
differs in superficial, brown, reticulate hyphae. Phylogenetically, the new species described below forms a distinct clade
within Microthyriaceae.
Pseudomicrothyrium X.Y. Zeng, S. Hongsanan & K.D.
Hyde, gen. nov.
Index Fungorum number: IF 556881; Facesoffungi number: FoF 06755; – 1 morphological species (this study), 1
species with molecular data.
Saprobic on dead leaves. Hyphae superficial, pale brown,
septate, branched, loosely reticulate. Sexual morph: Thyriothecia superficial, circular, light brown, solitary or gregarious, slightly carbonaceous, with a colorless basal layer, connected with hyphae, easily separated from the host surface.
Upper wall light brown, comprising cells of radial arrangement of textura prismatica, darken at the central ostiole,
tapering at margin. Hamathecium comprising asci inclined
from the base and rim towards the central ostiole, with thin,
Pseudomicrothyrium thailandicum X.Y. Zeng, S. Hongsanan & K.D. Hyde, sp. nov.
13
96
Etymology: Referring to its occurrence in Thailand.
Index Fungorum: IF 556882; Facesoffungi number: FoF
06756; Fig. 45
Saprobic on dead leaves. Hyphae superficial, pale brown,
septate, branched, loosely reticulate. Sexual morph: Thyriothecia 32–42 high × 140–160 μm diam. ( x̄ = 37 × 148
µm, n = 10), superficial, circular, light brown, solitary or
gregarious, slightly carbonaceous, with a colorless basal
layer, connected with hyphae, easily separated from the host
surface. Upper wall light brown, comprising cells of radial
arrangement of textura prismatica, darken at the central
ostiole, tapering at margin. Hamathecium comprising asci
inclined from the base and rim towards the central ostiole,
with thin, filiform, narrow cellular pseudoparaphyses. Asci
35–40 × 10–15 μm ( x̄ = 37 × 13 µm, n = 10), 8-spored,
bitunicate, fissitunicate, ovoid to obpyriform. Ascospores
13–15 × 4–5 μm ( x̄ = 14 × 5 µm, n = 20), 2–3-seriate, fusiform to ellipsoidal, 1-septate, upper cell slightly wider than
lower cell, not constricted at the septum, hyaline, with 2–4
apical appendages, usually with 2 large guttules in the upper
cell and 2 small guttules in the lower cell, smooth-walled.
Asexual morph: Undetermined.
Material examined: Thailand, Chaing Rai, Mae Fah
Luang University, on dead leaves of unidentified host, 3
March 2014, Xiang-Yu Zeng (MFLU 14-0286, holotype).
GenBank numbers: LSU: MT741680, SSU: MT741682.
Notes: Pseudomicrothyrium is introduced in this paper.
Sequence data of P. thailandicum indicate that it forms a
distinct lineage within Microthyriaceae with high support
(100% ML, 1.0 PP).
Pseudopenidiella Crous & Koukol, in Crous et al., Persoonia 28: 167 (2012).
Index Fungorum number: IF 800382; Facesoffungi number: FoF 06757; – 3 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Pseudopenidiella pini (P.M. Kirk &
Minter) P.M. Kirk, Index Fungorum 120: 1 (2014).
≡ Polyscytalum pini P.M. Kirk & Minter, in Kirk, Trans.
Br. mycol. Soc. 80(3): 462 (1983).
Notes: Pseudopenidiella was introduced as a hyphomycetous genus characterised by the formation of dimorphic
conidiophores with terminal, aseptate ramoconidia producing branched conidial chains and lack of coronate-type scars
on conidia or conidiogenous cells (Crous et al. 2012b). It
is similar to Cladosporium (Capnodiaceae), Digitopodium
and Penidiella (Teratosphaeriaceae), but phylogenetically
related to Microthyriaceae.
Seynesiella G. Arnaud, Annals d’École National d’Agric. de
Montpellier, Série 2 16(1–4): 202 (1918) [1917].
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Fungal Diversity (2020) 105:17–318
Index Fungorum number: IF 5018; Facesoffungi number:
FoF 06758; – 5 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Seynesiella juniperi (Desm.) G. Arnaud,
Annals d’École National d’Agric. de Montpellier, Série 2
16(1–4): 203 (1918) [1917].
≡ Dothidea juniperi Desm., Annls Sci. Nat., Bot., sér. 2
15: 141 (1841)
Notes: Seynesiella was transferred from Venturiaceae to
Microthyriaceae by Barr (1987b) based on the upper walls
with cells of radial arrangement. However, the light brown
ascospores are different from the type genus. Sequence data
are needed to confirm its taxonomic placement.
Tumidispora Hongsanan & K.D. Hyde, in Ariyawansa et al.,
Fungal Diversity 75: 95 (2015)
Index Fungorum number: IF 551375; Facesoffungi number: FoF 00944; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Tumidispora shoreae Hongsanan & K.D.
Hyde, in Ariyawansa et al., Fungal Diversity 75: 95 (2015).
Notes: Tumidispora is similar to Microthyrium in having
flattened thyriothecia with radial arrangement of upper wall
and hyaline, 1-septate ascospores, but differs in the shape
of ostiole, lack superficial hyphae, and lacking appendages
around the ascospores.
Economic and ecological significance
Species of this group are saprobes and poorly studied but
play a role in recycling organic matter.
Minutisphaerales Raja, Oberlies, Shearer & A.N. Mill.
Index Fungorum number: IF 811951; Facesoffungi number: FoF 08064.
Minutisphaerales was established by Raja et al. (2015) to
accommodate Minutisphaeraceae, members of which are
found on submerged wood in freshwater streams. Jayasiri
et al. (2018) included Acrogenosporaceae in this order based
on the morphological similarities and phylogenetic analyses.
The same result is shown in Fig. 46. The divergence time
for Minutisphaerales is estimated as 224 MYA (stem age,
Hongsanan et al. 2020).
Accepted families: Acrogenosporaceae, Minutisphaeraceae.
Acrogenosporaceae Jayasiri & K.D. Hyde, Mycosphere
9(4): 809 (2018).
Index Fungorum number: IF 554451; Facesoffungi number: FoF 04575; 13 species.
Saprobic on bark and wood. Sexual morph: Hysterothecia superficial, laterally compressed, thick-walled, with
a prominent sunken slit, solitary to gregarious, erect and
elevated, presenting an almost pedicellate appearance.
Fungal Diversity (2020) 105:17–318
Hysterium barrianum ANM1495
100/1.0
Hysteriaceae
Hysteriales
Strigulaceae
Strigulales
Hysterium thailandica MFLUCC 16-0338
-/-
Oedohysterium insidens CBS 238.34
98/1.0
Rhytidhysteron neorufulum MFLUCC 13-0216
99/1.0
Rhytidhysteron thailandicum MFLUCC 14-0503
-/0.98
Strigula jamesii MPN548
80/-
Strigula smaragdula LFF016344
81/-
Flavobathelium epiphyllum MPN67
-/0.91
88/1.0
-/100/1.0
Minutisphaera japonica HHUF 30098
Minutisphaera aspera G427 1a
Minutisphaera fimbriatispora L A242 8A
Minutisporaceae
100/1.0
Minutisphaera parafimbriatispora G156 4a
100/1.0
Acrogenospora carmichaeliana MFLU 18-1130
98/0.98
Acrogenospora carmichaeliana FMR11021 Acrogenosporaceae
Minutisporales
Fig. 46 Phylogram generated
from maximum likelihood
analysis (RAxML) of Minutisphaerales based on ITS, LSU,
SSU and tef1 sequence data.
Maximum likelihood bootstrap
values equal or above 70%,
Bayesian posterior probabilities
equal or above 0.90 (MLBS/
PP) are given at the nodes. An
original isolate number is noted
after the species name. The tree
is rooted to Capnodium coartatum (MFLUCC 10-0069) and
Leptoxyphium kurandae (CBS
129530). The ex-type strains are
indicated in bold. Hyphen (-)
represents support values below
70% MLBS and 0.90 PP
97
100/1.0
Acrogenospora thailandica MFLUCC 17-2396
Valsaria insitiva CBS 127882
Valsaria spartii CBS 139070
97/1.0
Munkovalsaria rubra CBS 109505
100/1.0
Capnodium coartatum MFLUCC 10-0069
Valsariaceae
Valsariales
100/1.0
Outgroup
Leptoxyphium kurandae CBS 129530
0.04
Hamathecium comprising persistent cellular, hypha-like,
hyaline, septate, paraphyses. Asci 8-spored, cylindrical to
clavate. Ascospores 1–2-seriate, ellipsoidal to fusiform, hyaline or moderately pigmented, 1- or 2-celled, with the septum near the lower end, hyaline or moderately pigmented.
Asexual morph: Hyphomycetous. Colonies effuse, scattered, black, glistening, hairy. Mycelium mostly immersed,
consisting of septate, subhyaline to pale brown hyphae.
Conidiophores macronematous, mononematous, pale to
mid brown, often slightly paler at the apex, smooth, septate,
unbranched, long cylindrical or slightly tapering towards the
apex, straight or slightly flexuous, erect, solitary or in small
groups. Conidiogenous cells monoblastic, integrated, terminal or intercalary, light brown or hyaline, cylindrical, often
elongated percurrently. Conidia holoblastic, solitary, globose, obovoid to ellipsoidal, asaptate, dark brown to black,
smooth or verrucose, truncate at base.
Type genus: Acrogenospora M.B. Ellis.
Notes: Acrogenosporaceae was introduced by Jayasiri et al. (2018) for a single genus Acrogenospora. Based
on morphological characters and phylogenetic analyses,
Acrogenosporaceae was placed within Minutisphaerales as
a sister clade to the type family Minutisphaeraceae (Jayasiri
et al. 2018). This study further supports the earlier result
(Fig. 46). Descriptions and illustrations of the sexual morphs
of Acrogenosporaceae can be seen in previous studies (i.e.
Greville 1825; Duby 1862; von Arx and Müller 1975).
Acrogenospora M.B. Ellis, Dematiaceous Hyphomycetes
(Kew): 114 (1971).
Index Fungorum number: IF 7036; Facesoffungi number:
FoF 08065; 13 morhological species (Species Fungorum
2020), 3 species with molecular data.
Type species: Acrogenospora sphaerocephala (Berk. &
Broome) M.B. Ellis.
Notes: Acrogenospora altissima was presumed to be
the asexual morph of Farlowiella australis Dennis on host
surfaces (Ellis 1972) while A. megalospora and F. carmichaeliana were linked by cultural studies (Mason 1941).
Molecular data support the synonymy of Acrogenospora and
Farlowiella (Jayasiri et al. 2018; Hyde et al. 2019). Nomenclaturally, Acrogenospora was protected over Farlowiella
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Fungal Diversity (2020) 105:17–318
Fig. 47 Acrogenospora sphaerocephala (MFLU 18-1130). a Colony on substrate. b, c Conidiophores with conidia. d Conidiophore.
e Conidiogenous cell with conidium. f, g Conidiogenous cells. h, i
Conidia. Scale bars: a = 100 μm, b–d = 50 μm, e, g = 30 μm, f, i =
20 μm, h = 15 μm
(Rossman et al. 2015). Acrogenospora was monographed by
Goh et al. (1998b). The genus is similar to Monotosporella
in having macronematous, mononematous conidiophores,
monoblastic conidiogenous cells and brown, sphaerical,
obovoid to ellipsoidal conidia. However, Acrogenospora
has unicellular conidia, while Monotosporella has septate
conidia. With little available sequence data, two strains of
Monotosporella clustered in Melanommataceae (Dothideomycetes) and Pleurotheciales (Sordariomycetes), respectively (Schoch et al. 2009a; Yang et al. 2019), which are
phylogenetically distinct from Acrogenospora.
Material examined: Thailand, Prachuap Khiri Khan Province, near 1230.1950 N, 9931.3500 E, on decaying wood
submerged in a freshwater stream, 25 December 2014, Jaap
van Strien, Site 5-14-2 (MFLU 18-1130).
Acrogenospora cf. sphaerocephala (Berk. & Broome) M.B.
Ellis, Dematiaceous Hyphomycetes (Kew): 114 (1971).
Index Fungorum number: IF 308236; Facesoffungi number: FoF 04687; Fig. 47
Description: see Hyde et al. (2019).
13
Ecological and economic significance
Acrogenosporaceae contains saprobic, lignicolous taxa
found in terrestrial, but mostly in freshwater habitats. They
play an important role in wood decay by the decomposition
of lignocelluloses (Wong et al. 1998; Krauss et al. 2011;
Hyde et al. 2016b). Generally, it is assumed that aquatic
fungi transfer organic matter directly to the higher trophic
levels of aquatic food webs (Christian et al. 2011).
Minutisphaeraceae Raja, Oberlies, Shearer & A.N. Mill.,
Mycologia 107 (4): 854 (2015).
Fungal Diversity (2020) 105:17–318
99
Fig. 48 Minutisphaera parafimbriatispora (ILLS 72341,
holotype). a, e Squash mount
of ascoma. b Asci. d Ascomata
on host surface. f Ascospores.
Scale bars: a, b = 20 μm, c =
10 μm, d, e = 100 μm. f = 50
μm
Index Fungorum number: IF 811062; Facesoffungi numbers: FoF 08066, 4 species.
Saprobic on submerged wood in freshwater habitats.
Sexual morph: Ascomata pseudothecioid or apothecioid,
erumpent to superficial, brown, with an ostiole and irregularly curved, dark brown to black hypha-like structures
around the ostiole. Peridium thin-walled, composed of
cells of textura angularis to textura globosum. Hamathecium comprising septate, cellular pseudoparaphyses with or
without enlarged pigmented tips. Asci 8-spored, fissitunicate, ovoid to obclavate, sessile to short-pedicellate, apically
rounded with or without an ocular chamber. Ascospores
2–4-seriate, clavate, fusiform to ellipsoidal, hyaline or pale
brown septate, multiguttulate, smooth or rough-walled, with
gelatinous sheath, with or without filamentous appendages radiating around the mid-septum. Asexual morph:
Undetermined.
Type: Minutisphaera Shearer, A.N. Mill. & A. Ferrer,
Ferrer et al.
Notes: This monotypic family was established by Raja
et al. (2015) to accommodate Minutisphaera. Raja et al.
(2015) placed Minutisphaeraceae in a newly introduced
order Minutisphaerales, Dothideomycetes based on its distinct morphological characters and multi-gene phylogenies.
The morphological characters such as small, globose to subglobose, ostiolate ascomata, septate, cellular pseudoparaphyses, fissitunicate, ovoid to obclavate asci and hyaline to
brown, 1-septate, ascospores with or without a gelatinous
sheath, suggest that Minutisphaeraceae could be placed in
Pleosporales (Zhang et al. 2012). Nevertheless, the molecular phylogenetic analysis shows that the family has no relationship with the members of Pleosporales (Raja et al. 2015;
Liu et al. 2017). Bao et al. (2019) provided a new species
of Minutisphaera. All the taxa of Minutisphaeraceae are
freshwater fungi occurring on submerged woody substrates.
Minutisphaera Shearer, A.N. Mill. & A. Ferrer, Mycologia
103(2): 415 (2011).
Index Fungorum number: IF 518355; Facesoffungi number: FoF 08067; 5 morphological species (Species Fungorum 2020), 5 species with molecular data.
Type species: Minutisphaera fimbriatispora Shearer,
A.N. Mill. & A. Ferrer.
Notes: Minutisphaera, typified by M. fimbriatispora was
described from submerged wood in freshwater habitats in
the USA (Ferrer et al. 2011). Subsequently, M. japonica was
described in Japan (Raja et al. 2013), and M. aspera and M.
parafimbriatispora in the USA (Raja et al. 2015). The members of this genus are only known by their sexual morphs
and no asexual morph has been reported.
Minutisphaera parafimbriatispora Raja, Oberlies, Shearer
& A.N. Mill., in Raja et al., Mycologia 107(4): 855 (2015).
Index Fungorum number: IF 5811064; Facesoffungi number: FoF 08068; Fig. 48
Description: see Raja et al. (2015).
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100
Material examined: USA, North Carolina, Guilford
County, Piedmont Plateau, Bur-Mil Park, Greensboro,
swampy area behind Lake Brandt, on submerged decorticated wood, 20 October 2011, H.A. Raja G156-1 (ILLS
72341, holotype).
Economic and ecological significance
Members of this family usually inhabit woody substrates in
freshwater habitats as saprobes (Ferrer et al. 2011; Raja et al.
2013, 2015). Some species produce dipeptides and aromatic
polyketides secondary metabolites belonging to two structural classes (Raja et al. 2015).
Monoblastiales Lücking, M.P. Nelsen & K.D. Hyde.
MycoBank number: MB 805299; Index Fungorum number: IF 805299; Facesoffungi number: FoF 08069.
This order comprises lichenized and non-lichenized
fungi. It was introduced by Hyde et al. (2013) to accommodate the single family, Monoblastiaceae. Among the mostly
lichenized orders in Dothideomycetes, Monoblastiales differs from Strigulales in the net-like physes and from most
Trypetheliales in having euseptate ascospores with few septa
and the poor secondary chemistry; from both orders it also
deviates in the macroconidia often arranged in a gelatinous
matrix (Hyde et al. 2013). The divergence time for Monoblastiales is estimated as 230 MYA (stem age, Hongsanan
et al. 2020).
Accepted families: Monoblastiaceae.
Monoblastiaceae Walt. Watson, New Phytologist 28: 106
(1929).
= Eriomycetaceae Huanraluek & Hyde, in Hyde et al.,
Fungal Diversity 100: 146 (2020).
MycoBank number: MB 81020; Index Fungorum number:
IF 81020; Facesoffungi number: FoF 08851; approximately
120 species (this paper).
Lichenized on bark, more rarely on rocks or leaves (supraor rarely subcuticular), more rarely non-lichenized and saprobic on leaves or bark (Eriomyces, Funbolia, Heleiosa,
Pseudopassalora) or hyperparasitic on bracket fungi (Phellinocrescentia); usually in terrestrial, chiefly lowland to
montane tropical to subtropical habitats, with few species
extending into temperate regions and in one case (Heleiosa) growing on leaves of Juncus in salt marshes. Thallus in lichenized species mostly reduced and ecorticate,
white, to distinctly corticate, grey-green to olive brown,
sometimes shiny (Megalotremis, Trypetheliopsis), in nonlichenized species absent. Photobiont in lichenized species
trentepohlioid. Ascomata when present scattered, clustered,
or aggregated, immersed to sessile, mostly black but sometimes covered by thallus, rarely non-carbonaceous, globose
to pear-shaped or conical, ostiolate, ostiole apical or eccentric, with periphyses. Involucrellum present, sometimes
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Fungal Diversity (2020) 105:17–318
only apically, or reduced, usually dark brown to carbonized,
paraplectenchymatous in thin sections. Excipulum dense,
consisting of compressed hyphae, appearing prosoplectenchymatous in thin sections but structure sometimes difficult
to observe when carbonaceous, hyaline to brown or brownblack. Hamathecium usually comprising 0.5–0.7 µm wide
paraphyses (trabeculate pseudoparaphyses according to Harris (1990, 1995), hyaline, straight, branched and anastomosing, rarely (Eriomyces) formed by 1–3 µm wide, branched
cellular pseudoparaphyses. Asci (1–)2–8-spored, rarely polyspored (Eriomyces), bitunicate, fissitunicate, mostly cylindrical to sometimes cylindrical-clavate, shortly pedicellate,
with narrow to broad, non-amyloid ocular chamber and in
part fluorescent ring- or cap-structures. Ascospores uni- to
biseriate or irregularly arranged, ellipsoid-oval to oblong
or sometimes fusiform, hyaline to rarely brown, aseptate or
1–3-septate, with thin to rather thick (Megalotremis, Trypetheliopsis) eusepta and more or less rectangular lumina,
smooth-walled or ornamented or rarely with appendages
(Heleiosa), sometimes slightly constricted at the septa, in
1-septate ascospores the upper cell often distinctly larger
than the lower cell (particularly in Anisomeridium). Conidiomata common, usually pycnidia, very rarely hyphomycetous
(Funbolia, Pseudopassalora); pycnidia immersed to sessile
and usually visible as black dots, but sometimes conspicuous and flask-shaped with a short to long a beak or hair-like
(Anisomeridium, Caprettia) or asymmetrically cup- or earshaped (campylidiiform: Trypetheliopsis). Conidia acrogenous or rarely pleurogenous (Funbolia), either macro- or
microconidia; macroconidia usually aseptate (septate in
Funbolia, Pseudopassalora), (globose to) broadly ellipsoid to oblong-bacillar or guttuliform, (2.5–)5–15(–45) ×
(2–)3–7(–16) µm large, hyaline (brown in Funbolia, Pseudopassalora), often forming clusters embedded in a gelatinous
matrix, these clusters in some species in the form of sacci
or cirri (Anisomeridium, Caprettia); microconidia aseptate,
globose to broadly ellipsoid or fusiform, small, 2–4(–5) ×
1–2(–4) µm large, hyaline. Chemistry: Most species do not
contain secondary substances; lichexanthone and anthraquinone pigments are known from a few taxa.
Type: Monoblastia Riddle.
Notes: For a detailed discussion of the lichenized core
group of the family (Acrocordia, Anisomeridium, Caprettia, Megalotremis, Monoblastia, Trypetheliopsis), including
a critical review of the currently applied genus concept, see
Lücking et al. in Hyde et al. (2013). The scarce molecular
data available support monophyly of Megalotremis and a
close relationship of the latter with Trypetheliopsis, while
Anisomeridium is likely polyphyletic (Fig. 49), a notion
anticipated by Harris (1995). Representatives of all genera
except Caprettia and the type genus, Monoblastia, have been
sequenced, so the position of Caprettia within the family,
based on hamathecium and ascus structure, ascospore type,
Fungal Diversity (2020) 105:17–318
101
Fig. 49 Best-scoring maximum
likelihood tree of currently
available data in Monoblastiales and its only family,
Monoblastiaceae (including
Eriomycetaceae), based on a
combined alignment of five
markers (nuSSU, nuLSU, ITS,
mtSSU, TEF1α), with a length
of 3612 bases (single autapomorphic base calls in gappy
sites removed). The tree was
reconstructed using the universal GTR-Gamma model without
site partitioning, and the final
likelihood was − 10218.109712
and the peculiar conidiomata and conidia, remains tentative. The accurcate circumscription of the family hinges on
the presumed close relationship of Monoblastia with e.g.
Acrocordia, and so far only a single species of the latter has
been sequenced. The taxonomic delimitation between Anisomeridium, Megalotremis, and Trypetheliopsis is discussed
in detail below.
In this updated treatment, we also include several nonlichenized, monospecific genera in the family. None of them
were originally established with a connection to Monoblastiaceae (Kohlmeyer et al. 1996; Crous et al. 2011a, b, 2014;
Hyde et al. 2020b), but two genera (Funbolia, Heleiosa)
were associated with the latter in a subsequent study (Nelsen
et al. 2011a). Recently, Hyde et al. (2020b) established the
new family Eriomycetaceae for the five non-lichenized lineages; however, these authors did not include lichenized representatives of Monoblastiaceae in their analysis and therefore overlooked the close relationship that had previously
been established by Nelsen et al. (2011a). The expanded
phylogeny presented here does not resolve Monoblastiaceae
and Eriomycetaceae as reciprocally monophyletic, but renders the non-lichenized lineages as a moderately supported
clade nested within a paraphyletic backbone comprising the
lichenized lineages (Fig. 49).
Given this topology, one could theoretically retain
Eriomycetaceae by splitting a Acrocordiaceae from
Monoblastiaceae, the latter then corresponding to the clade
containin Anisomeridium sensu lato, Megalotremis, and Trypetheliopsis (Fig. 49). However, this solution is not appropriate, for various reasons. First, the type of Monoblastiaceae,
Monoblastia, has not yet been sequenced. While its relationship with the other lichenized genera is supported by morphological and anatomical characters (Lücking et al. in Hyde
et al. 2013), it is unclear whether it is more closely related
to Acrocordia or to the Anisomeridium-Megalotremis-Trypetheliopsis clade, thus creating potentially unstable nomenclature regarding the application of the name Monoblastiaceae.
Second, while the non-lichenized lineages partly deviate
from the lichenized taxa anatomically, the genus Heleiosa
is very similar to the latter: its ascoma anatomy, including
the thin, anastomosing paraphyses and the cylindrical asci
with broad ocular chamber (Kohlmeyer et al. 1996), is in line
with that of Acrocordia and Monoblastia. Also the pycnidial
genus Phellinocrescentia agrees with lichenized taxa, in particular species of Anisomeridium, in the guttuliform conidia
(Crous et al. 2014). Thus, even if Eriomyces differs in its
peculiar asci and the rather thick pseudoparaphyses (Hyde
et al. 2020b), and Funbolia and Pseudopassalora deviate from the other genera by their hyphomycetous asexual
morphs (Crous et al. 2011a, b), there are at least two genera
in the former Eriomycetaceae that agree phenotypically with
lichenized Monoblastiaceae.
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102
The inclusion of lichenized and non-lichenized lineages
in a single family, even with such disparate morphological
and anatomical features, is not entirely surprising. The situation is analogous to Graphidaceae and Trypetheliaceae,
which feature some non-lichenized or borderline lichenized
lineages as early diverging clades (Nelsen et al. 2009, 2011a,
2014; Lücking et al. 2016; Cáceres et al. 2020; MirandaGonzález et al. 2020). Several lineages of lichenized fungi
produce hyphomycetous asexual morphs, particularly in
the order Arthoniales (Arthoniomycetes), e.g. in the genera Reichlingia and Tylophoron (Frisch et al. 2014; Van
den Broeck et al. 2018), so the relationship of Funbolia and
Pseudopassalora with the other lineages in Monoblastiaceae
is also not entirely unexpected, although without molecular
evidence such relationships would not have been predicted.
Monoblastia Riddle, Mycologia 15: 70 (1923).
MycoBank number: MB 3251; Index Fungorum number:
IF 3251; Facesoffungi number: FoF 08070; 12 morphologically defined species (Lücking et al. 2017; Harada 2018),
molecular data thus far unavailable.
Lichenized on bark or rarely on limestone, in terrestrial,
chiefly lowland to montane tropical to subtropical habitats,
with few species extending into temperate regions. Thallus usually ecorticate, white. Photobiont Trentepohlia.
Ascomata scattered or rarely clustered, erumpent to sessile,
mostly black, carbonaceous, ostiolate, ostiole round. Involucrellum present or reduced, carbonized. Excipulum dense,
consisting of compressed hyphae, appearing prosoplectenchymatous in thin, bleached sections, hyaline to brown or
brown-black. Hamathecium comprising 0.5–0.7 µm wide
paraphyses, hyaline, straight, branched and anastomosing.
Asci (2–)8-spored, bitunicate, fissitunicate, cylindrical, short
pedicellate, with broad, non-amyloid ocular chamber and
fluorescent cap-structures. Ascospores mostly uni-seriate,
ellipsoid-oval, hyaline, aseptate, ornamented with warts,
short ridges or spines. Pycnidia rare. Conidia acrogenous,
microconidia, aseptate, oblong to ellipsoid to fusiform,
small, hyaline. Chemistry: Secondary substances absent.
Type species: Monoblastia palmicola Riddle, Mycologia
15: 70 (1923).
Notes: See Lücking et al. in Hyde et al. (2013) for discussion and illustrations.
Other genera:
Acrocordia A. Massal., Geneac. Lich. (Verona): 17 (1854).
MycoBank number: MB 46; Index Fungorum number: IF
46; Facesoffungi number: FoF 08071; ten morphologically
defined species (Lücking et al. 2017), molecular data available for one species.
Lichenized on bark or rarely on bryophytes or on limestone, in temperate to tropical montane regions. Thallus ecorticate, white. Photobiont Trentepohlia. Ascomata
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scattered or rarely clustered, erumpent to sessile, black,
carbonaceous, ostiolate, ostiole apical or lateral. Involucrellum present or reduced, carbonized. Excipulum dense,
consisting of compressed hyphae, appearing prosoplectenchymatous in thin, bleached sections, hyaline to brown or
brown-black. Hamathecium comprising 0.5–0.7 µm wide
paraphyses, hyaline, straight, branched and anastomosing.
Asci (2–)8-spored, bitunicate, fissitunicate, cylindrical, short
pedicellate, with broad, non-amyloid ocular chamber and
fluorescent cap-structures. Ascospores mostly uni-seriate,
rarely biseriate, ellipsoid-oval to rarely oblong, hyaline,
1-septate, usually granular ornamented. Pycnidia rare.
Conidia acrogenous, microconidia, aseptate, oblong to
ellipsoid to fusiform, small, hyaline. Chemistry: Secondary
substances absent.
Type species: Acrocordia garovaglii A. Massal., Geneac
Lich (Verona): 17 (1854) [= A. conoidea (Fr.) Körb. var.
conoidea, Syst Lich Germ: 358 (1855)].
Notes: This genus was not treated in detail by Lücking
et al. in Hyde et al. (2013). Acrocordia resembles both Anisomeridium and Monoblastia in general habit. It differs from
the first in the broad ocular chamber of the asci and from the
second in the 1-septate vs. aseptate ascospores (Fig. 50). The
septum usually has characteristing endospore thickenings
that make it appear laterally somewhat bulging (Fig. 50).
Arthopyrenia endobrya (≡ Anisomeridium endobryum;
Megalotremis endobrya) is an unusual lichen that grows
intracellularly within dead bryophytes, known from tropical montane forest in Brazil (type; Döbbeler and Poelt 1981)
and Costa Rica (Aptroot et al. 2008). It is characterized by
sessile, basally constricted, black perithecia and 1-septate
ascospores 35–50 × 10–15 µm in size. Harris (1995) recognized its position within Monoblastiaceae and placed it in
Anisomeridium. Because of its slight septal endospore thickenings, it was later recombined into Megalotremis (Aptroot
et al. 2008), although the ascospore size range (see below
Megalotremis for further discussion) fits Anisomeridium better, Given the ecorticate thallus, the fully exposed perithecia, the asci with broad ocular chamber, and the thin-walled
ascospores lacking crystals and with the septa slightly bulging (Fig. 51), the species is actually best referred to Acrocordia, where several other species have sessile, subglobose
perithecia, such as A. subglobosa. The biseriately arranged
ascospores apparently lacking granular ornamentation deviate from most of the other species in that genus, but such
variation is also accepted in Anisomeridium (see below). The
necessary combination is introduced below.
Acrocordia endobrya (Döbbeler & Poelt) Lücking & Aptroot, comb. nov.
MycoBank number: MB 836365; Index Fungorum number: IF 836365; Facesoffungi number: FoF 08072.
Fungal Diversity (2020) 105:17–318
103
Fig. 50 Morphology, asci and ascospores in Acrocordia species.
a–d A. conoidea (a, b Germany, Schumm 70; c, d Germany, Schumm
12542). e–h A. gemmata (e–f Portugal, Schumm 11491; g Portugal,
Schumm 11547; h, i Germany, Schumm 11586). j, k A. salweyi (Por-
tugal, Aptroot & Schumm 14215). Scale bars: a, c, j = 1 mm, b, d, k
= 10 µm. For further illustrations of species of this genus, see also
Schumm (2008, 2011) and Schumm and Aptroot (2013)
Bas.: Arthopyrenia endobrya Döbbeler & Poelt, Pl Syst
Evol 138: 276 (1981); Anisomeridium endobryum (Döbbeler
& Poelt) R.C. Harris, More Florida Lichens: 146 (1995);
Megalotremis endobrya (Döbbeler & Poelt) Aptroot in Aptroot et al., Biblioth Lichenol 97: 68 (2008).
MycoBank number: MB 201; Index Fungorum number:
IF 201; Facesoffungi number: FoF 08073; approximately 60
morphologically defined species (this paper; Harada 2019),
molecular data thus far available for four species.
≡ Arthopyrenia sect. Anisomeridium Müll. Arg., Flora
66: 290 (1883)
MycoBank number: MB 700075; Index Fungorum number: IF 700075; Facesoffungi number: FoF 08852.
Anisomeridium (Müll. Arg.) M. Choisy, Icon Lich Univ 1:
24 (1928).
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Fungal Diversity (2020) 105:17–318
Fig. 51 Morphology, asci and ascospores of Acrocordia endobrya
(Costa Rica, Sipman 51790). a, b Thallus on dead bryophytes with
perithecia and perithecia enlarged. c–e Mature and immature asci
showing broad ocular chamber. f–h Ascospores. Scale bars: a, b = 1
mm, c, d = 50 µm; e, f, g = 10 µm
Lichenized on bark or more rarely on leaves or rocks,
possibly lichenicolous; in lowland to montane tropical to
subtropical habitats, with few species extending into temperate regions. Thallus usually ecorticate or sometimes with
thin cortex, white or shades of grey-green to green. Photobiont Trentepohlia. Ascomata scattered or rarely clustered,
erumpent to sessile, mostly black, carbonaceous, ostiolate,
ostiole apical or lateral. Involucrellum present or reduced,
carbonized. Excipulum dense, consisting of compressed
hyphae, appearing prosoplectenchymatous in thin, bleached
sections, hyaline to brown or brown-black. Hamathecium
comprising 0.5–0.7 µm wide paraphyses, hyaline, straight,
branched and anastomosing. Asci (2–)8-spored, bitunicate,
fissitunicate, cylindrical, short pedicellate, with narrow, nonamyloid ocular chamber and fluorescent cap- or ring-structures. Ascospores uni- or biseriate to irregularly arranged,
ellipsoid-oval to broadly fusiform, small to medium-sized
[(10–)15–45(–55) × 4.5–15(–20) µm], hyaline to very rarely
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Fungal Diversity (2020) 105:17–318
brown, 1(–3)-septate, with thin eusepta and rectangular
lumina, smooth-walled or granular ornamented, sometimes
slightly constricted at the septa, in 1-septate ascospores the
upper cell often distinctly larger than the lower cell (very
rarely the opposite). Pycnidia common, immersed to sessile,
visible as black dots but sometimes conspicuous and flaskshaped. Conidia acrogenous, either macro- or microconidia;
macroconidia usually aseptate, (globose to) broadly ellipsoid to bacillar, (2.5–)5–15(–45) × (2–)3–7(–16) µm large,
hyaline, often forming clusters embedded in a gelatinous
matrix, these clusters in some species in the form of sacci or
cirri; microconidia aseptate, globose to broadly ellipsoid or
fusiform, small, 2–4(–5) × 1–2(–4) µm large, hyaline. Chemistry: Most species do not contain secondary substances;
lichexanthone present in some taxa.
Type species: Arthopyrenia xylogena Müll.Arg., Flora
66: 290 (1883) [= Anisomeridium subnectendum (Nyl.)
R.C.Harris, More Florida Lichens: 150 (1995)].
Notes: Lücking et al. (in Hyde et al. 2013) did not treat
this genus in detail. Citing Riedl (1963), Harris (1995)
considered Arthopyrenia xylogena Müll. Arg. the ‘lectotype’ of the genus. However, Müller (1883) only included
a single species in Arthopyrenia sect. Anisomeridium, viz.
Arthopyrenia xylogena, which is then by default the (holo-)
type (although the terms ‘holo-’ and ‘lectotype’ technically
do not apply to names above species [ICN Art. 10.1, Note
1]. Riedl’s (1963: 270) statement that “… xylogena muss als
Typusart der Sektion Anisomeridium Müll. Arg. betrachtet
werden.” [“… xylogena is to be considered the type species
of section Anisomeridium Müll. Arg.”] refers to the differences between Müller’s protologue of the species description
and the features observed by Riedl himself, casting doubt as
to Müller properly described the taxon at hand. However,
this only affects typification of the species itself, not typification of the genus, which is fixed by the single species name
listed in the protologue.
The taxonomic delimitation between Anisomeridium and
Megalotremis is elaborated in detail below under the latter
genus (see below). Lücking et al. (2017) gave the number of
species in Anisomeridium as 200, but revision of published
literature revealed that only 57 species are currently to be
accepted in the genus, after transferral of several taxa to
Megalotremis (see below).
Caprettia Bat. & H. Maia, Atas Inst Micol Univ Pernambuco
2: 377 (1965).
MycoBank number: MB 813; Index Fungorum number:
IF 813; Facesoffungi number: FoF 08074; 8 morphologically defined species (this paper), molecular data thus far
unavailable.
Lichenized on leaves, in tropical regions. Thallus ecorticate, greenish, subcuticular (subgenus Caprettia) or supracuticular (subgenus Porinula). Photobiont trentepohloid.
105
Ascomata scattered (absent in subgenus Caprettia), sessile,
subglobose, yellowish to red-brown or brown-black, carbonaceous or not, glabrous or with short to conspicuous,
horizontal setae, ostiolate, ostiole apical. Involucrellum
reduced, carbonized or not. Excipulum dense, consisting of
compressed hyphae, appearing prosoplectenchymatous in
thin sections, hyaline to brown. Hamathecium comprising
0.5–0.7 µm wide paraphyses, hyaline, straight, branched and
anastomosing. Asci (2–)4–8-spored, bitunicate, fissitunicate,
cylindrical, short pedicellate, with broad, non-amyloid ocular chamber and fluorescent cap-structures. Ascospores uniseriate, ellipsoid-oval to sometimes curved, hyaline, 1-septate, smooth. Pycnidia frequent, typically hair-like (subgenus
Caprettia) or long-beaked with often slightly inflated base
(subgenus Porinula). Conidia acrogenous, microconidia,
aseptate, ellipsoid, small, hyaline, within pycnidial tube
aggregated into gelatinous, rectangular sacci extruded from
pycnidial beak as single diaspores. Chemistry: Secondary
substances absent.
Type species: Caprettia amazonensis Bat. & H. Maia,
Atas Inst Micol Univ Pernambuco 2: 378 (1965).
Notes: For illustrations of selected species, see Lücking
et al. in Hyde et al. (2013). This genus includes several phenotytically disparate elements. The type species, C. amazonensis, grows subcuticularly and produces black, hair-like,
straight pycnidia (no ascomata). Subgenus Porinula Lücking & Sérus. presents supracuticular growth and includes
two distinct groups. The type species, C. tanzanica, as well
as C. ornata, C. setifera, C. goderei, C. neotropica, and C.
nyssogenoides, all frequently produce long-beaked pycnidia
and intergrade from very pale and glabrous to black and distinctly setose ascomata. A further species, C. confusa, differs
in the absence of pycnidia and the large ascospores, resembling those of Trypetheliopsis (Sérusiaux and Lücking 2003;
Lücking 2008; Yeshitela et al. 2009). In lieu of molecular
data, these taxa are here kept in a single genus, although it
is unclear whether they form a monophyletic group and to
what extend they are related to Anisomeridium sensu lato
and Trypetheliopsis (see also Harris 1995).
Sérusiaux and Lücking (2003) introduced Caprettia subgenus Porinula as a combination based on Porinula [non
Porinula (Nyl.) Flagey], overlooking that an illegitimate
later homonym cannot serve as base for a legitimate combination. The name must therefore be considered a replacement name, Caprettia subgenus Porinula [ICN Art. 58.1].
At the genus level, however, the replacement name Porinella
R. Sant. has priority.
Eriomyces Huanraluek, Thambugala & K.D. Hyde in Hyde
et al., Fungal Diversity 100: 146 (2020).
MycoBank number: MB 556538; Index Fungorum number: IF 556538; Facesoffungi number: FoF 06151; one
13
106
phylogenetically and morphologically defined species (Hyde
et al. 2020b).
Saprobic on dead twigs of Hevea brasiliensis in terrestrial, lowland to montane tropical to habitats. Ascomata
scattered to somewhat aggregate, immersed to erumpent,
brown-black to greyish black, applanately wart-shaped to
drop-shaped or angular, ostiolate, ostiole more or less eccentric, with periphyses. Involucrellum present apically, dark
brown to carbonized, paraplectenchymatous in thin sections.
Excipulum prosoplectenchymatous, hyaline to pale brown.
Hamathecium comprising 1–3 µm wide, branched cellular
pseudoparaphyses. Asci polyspored, bitunicate, fissitunicate,
clavate, long-pedicellate, with indistinct tholus and ocular
chamber. Ascospores densely arranged, oblong, sometimes
slightly curved, hyaline, aseptate, smooth-walled. Conidiomata not observed.
Type species: Eriomyces heveae Huanraluek, Thambugala & K.D. Hyde in Hyde et al., Fungal Diversity 100: 147
(2020).
Notes: This monospecific genus is characterized by the
unusual, polyspored, clavate asci with long stipe, deviating
from the other genera in this family in which ascomata are
known. In the original description, the ostioles were given as
central but based on the published photographic plate (Hyde
et al. 2020b: Fig. 92) they appear to be irregularly excentric,
also shown by the irregular shape of the ascomata.
Funbolia Crous & Seifert in Crous et al., Persoonia 26: 115
(2011).
MycoBank number: MB 560161; Index Fungorum number: IF 560161; Facesoffungi number: FoF 08075; 1 phylogenetically and morphologically defined species (Crous
et al. 2011a).
Type species: Funbolia dimorpha Crous & Seifert in
Crous et al., Persoonia 26: 115 (2011).
Notes: For description and illustrations see Crous et al.
(2011a).
Heleiosa Kohlm., Volkm.-Kohlm. & O.E. Erikss., Can J Bot
74: 1830 (1996).
MycoBank number: MB 27767; Index Fungorum number:
IF 27767; Facesoffungi number: FoF 08076; 1 phylogenetically and morphologically defined species (Kohlmeyer et al.
1996).
Type species: Heleiosa barbatula Kohlm., Volkm.Kohlm. & O.E. Erikss., Can J Bot 74: 1830 (1996).
Notes: For description and illustrations see Kohlmeyer
et al. (1996).
Megalotremis Aptroot, Biblioth Lichenol 44: 124 (1991).
MycoBank number: MB 26295; Index Fungorum number: IF 26295; Facesoffungi number: FoF 08077; 16
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Fig. 52 Ascospore size variation in Anisomeridium and Megalotremis, highlighting taxa of uncertain affinity to either genus and their
current status. Axes denote maximum length and width reported for
each species (in µm). Note the formation of three smaller groups in
Anisomeridium
morphologically defined species (this paper); molecular data
available for one species.
Lichenized on bark, in tropical regions. Thallus usually
with thin, cartilaginous cortex, rarely ecorticate, greenish
grey to white. Photobiont Trentepohlia. Ascomata scattered, usually erumpent, black but at least partly covered
by thalline layer, typically carbonaceous, ostiolate, ostiole
apical or lateral. Involucrellum present, carbonized. Excipulum dense, consisting of compressed hyphae, appearing
prosoplectenchymatous in thin, bleached sections, brown to
brown-black. Hamathecium comprising 0.5–0.7 µm wide
paraphyses, hyaline, straight, branched and anastomosing.
Asci (1–)2–8-spored, bitunicate, fissitunicate, cylindrical,
short pedicellate, with broad, non-amyloid ocular chamber and fluorescent cap-structures. Ascospores uni-seriate
to irregularly arranged, ellipsoid-fusiform, rather large
[50–130(–225) × (18–)20–45(–60) µm], sometimes with the
upper cell enlarged, hyaline, 1-septate, with internal, needleshaped crystals. Pycnidia rare. Conidia acrogenous, either
macro- or microconidia; macroconidia usually aseptate,
(globose to) ellipsoid, (4–)5–9(–15) × 2–5(–15) µm large,
hyaline; microconidia aseptate, globose to broadly ellipsoid
or fusiform, small, 2–7(–7) × 1.5–2(–4) µm large, hyaline;
macro- and/or microconidia in some species aggregated into
conglutinate masses sometimes deposited outside the pycnidia. Chemistry: Secondary substances absent.
Fungal Diversity (2020) 105:17–318
Type species: Megalotremis verrucosa (Makhija & Patw.)
Aptroot, Biblioth. Lichenol. 44: 126 (1991).
Notes: In the protologue, Aptroot (1991) did not compare
Megalotremis with Anisomeridium but placed the new genus
in Trypetheliaceae because of the endospore thickenings;
he also pointed out the ‘internal folds’ of the ascospores
which were subsequently interpreted as crystals (Aptroot
et al. 2008). Harris (1995) synonymized Megalotremis under
Anisomeridium, stating that corticate thalli, one of the perceived differences, also occur in the latter genus. Harris
(1995) further suspected that the second species of Megalotremis included in the protologue, M. biocellata, was related
to Arthopyrenia sensu lato. Aptroot et al. (2008) differentiated Megalotremis from Anisomeridium by the corticate
thallus and large ascospores with endospore thickenings,
and from Trypetheliopsis (as Musaespora) by the absence
of campylidioid asexual morphs.
Given that the currently available molecular data support
this separation (Fig. 49), we analysed the species included
under Anisomeridium sensu lato by Harris (1995) in terms
of thallus morphology and ascospore size (and endospore
thickenings). Of the 72 species included in the world-wide
key, plus an additional eight species of Megalotremis not
included or established subsequently, 55 have endoperidermal, ecorticate thalli with erumpent to prominent, usually exposed, black perithecia, and ascospores with rather
thin walls and septa up to 55 × 20 µm, but in most cases
smaller than 30 × 10 µm (Fig. 52), with smooth or granular
ornamented walls. Notably, these species cluster in three
ascospore size groups (Fig. 52). Fifteen species have epiperidermal, corticate thalli with immersed-erumpent perithecia
partly covered by thallus, and ascospores with thickened
walls and septa larger than 55 × 20 µm and up to 130(–225)
× 45(–60) µm, typically with internal, needle-shaped crystals (or in two cases lack ascomata but have peculiar, tubeshaped pycnidia). These correspond to the current concept
of Megalotremis, and of the eight species treated by Harris (1995), the following four are currently included in that
genus: M. infernale, M. megalospora, M. nemorosa, and M.
verrucosa (type of Megalotremis). Another four species are
recombined into Megalotremis below.
Several species do not fit into the above scheme and
deserve attention. A group of at least four species has corticate thalli but fully exposed, prominent, black perithecia and
ascospores typical of Anisomeridium, including A, glaucescens, A. indicum, A. stromaticum, and A. truncatum. These
taxa superficially resemble certain species of Lithothelium,
although they differ fundamentally in anatomy, and require
further study. Their thalli, although corticate, are opaque
and differ from the usually shiny thalli of Megalotremis.
The transference of Anisomeridium endobryum (≡ Megalotremis endobrya) to Acrocordia is discussed above under
that genus.
107
Anisomeridium excellens produces large ascospores
(55–85 × 20–28 µm) fitting the range of Megalotremis and
far above typical Anisomeridium, but the thallus is clearly
endoperidermal and ecorticate, thus fitting Anisomeridium
morphologically. Specimens identifiable with this species
are not uncommon in the Caribbean, and usually also have
at least partly clustered ascomata, and never any superficial
thallus. The unusually large ascospores are granular ornamented, a rare feature in bitunicate lichenized Ascomycota
and largely known from the genera Anisomeridium and
Bogoriella (Trypetheliaceae); they also lack internal crystals, another feature pointing to Anisomeridium rather than
Megalotremis. Thus, A. excellens is currently retained in
Anisomeridium (see also further discussion under M. biocellata below), but represents an oddball due to its ascospore
size and requires molecular study.
Anisomeridium megalosporum (≡ Megalotremis megalospora) was described by Vainio (1923) almost as an afterthought in his work on Thaxter’s lichen collections from
Trinidad, as he considered it non-lichenized. Aptroot (1995)
found it to be lichenized, with the ascospores containing
needle-shaped crystals, and therefore proposed its inclusing in Megalotremis. The only obstacle to this classification is the thallus morphology, keyed out as endoperidermal and ecorticate by Harris (1995), who also saw the type.
Notably, Harris (1995) describes the ascospores as granular
ornamented, not mentioning any crystals. In that respect,
great similarities exist with A. excellens, which in Harris’s
key strongly overlaps with his concept of A. megalosporum
except for 2–4-spored vs. 2-spored asci. It is possible that
part of Harris’s A. megalosporum represents a 2-spored form
of A. excellens, whereas A. megalosporum sensu stricto is
a different taxon. For the time being, we retain the latter in
Megalotremis as an exception analogous to the case of A.
excellens in Anisomeridium.
The presumably large-spored Anisomeridium planiusculum revealed a complex situation. Nylander (1858) described
the ascospores as 1-septate, hyaline, and 58–78 × 20–30 µm.
Harris (1995) reported the ascospores as 1-septate, hyaline,
and 62–82 × (22–)27–33 µm, i.e. congruent with Nylander’s
protologue, giving the diminutive specimen in H (H-NYL
732; H9505212) as holotype. Said holotype features a portion of an apparently endoperidermal, whitish thallus with
erumpent ascomata not sharply delimited from the surrounding thallus. Harris’s drawings along with that material agree
with his published measurements and depict the ascospore
walls as uniform. However, the same type has an annotation and drawing by P. G. Patwardhan & U. Makhija date
1977, which shows secondary endospore constrictions (as
in Constrictolumina) and give the size of the ascospores
as considerably smaller, namely 35–65 × 13–20 µm. All
this would point to similarities of the presumed ‘holotype’
with M. biocellata (see below). Two more collections from
13
108
Fée’s herbarium with the name Verrucaria planiuscula are
in G, one with three large pieces and (G00290329) one with
a small portion of the same species abundantly present on
the large pieces (G00290330), both featuring a lichen with
well-developed, corticate thallus and sharply delimited,
black perithecia. The large pieces feature a drawing of two
ascospores with the size annotation “58–78 × 20–30 µm”,
in Nylander’s handwriting and with an additional annotation by Müller “= jun!”. The small piece has two ascospore
drawings by Müller, one depicting a 1-septate and the other
a muriform ascospore, with the size annotation “65–75 ×
23–37 µm”, and the name annotation “Anthracothecium
planiusculum Müll. Arg.’ Müller (1888) indeed published
the combination A. planiusculum (Nyl.) Müll.Arg., stating
that the 1-septate, hyaline ascospores are young and the
mature ascospores become muriform and brown. While this
sounds unlikely, precisely such ascospore ontogeny has been
demonstrated e.g. for Aptrootia terricola in the Trypetheliaceae (Sweetwood et al. 2012; Aptroot and Lücking 2016).
Given the agreement of the ascospore size annotation on
the specimen with the three large pieces in G (G00290329)
with Nylander’s (1858) protologue, that specimen must be
selected as lectotype: Verrucaria planiuscula Nyl., Expos.
Synopt. Pyrenocarp.: 58 (1858); Pyrenula planiuscula
(Nyl.) Tuck., Proc. Amer. Acad. Arts & Sci. 7: 233 (1868);
Anthracothecium planiusculum (Nyl.) Müll. Arg., Flora 71:
208 (1888); Arthopyrenia planiuscula (Nyl.) Zahlbr., Cat.
Lich. Univers. 1: 310 (1921); Anisomeridium planiusculum (Nyl.) R.C. Harris, More Florida Lichens: 149 (1995).
Type. Jamaica, on Quassia excelsa (= Q. amara), s.dat.,
s.col. (G G00290329, lectotype, here selected, MycoBank
MBT393283). The initially 1-septate ascospores and the
morphology and substrate make a relationship of this taxon
with Aptrootia unlikely and suggest correct placement in
the genus Anthracothecium. Based on ascospore size, it
would then largely agree with A. prasinum, although that
species has 8-spored asci (Aptroot 2012), whereas Harris (1995) gives the asci of Anisomeridium planiusculum
as 2–4-spored. The only other species of Anthracothecium
with solitary ascomata and apical ostioles, A. macrosporum,
agrees with the presumably 2–4-spored asci of V. planiuscula but has much larger ascospores. However, Nylander
(1858) described the asci of V. planiuscula as “… 4nae
(8nae?) …” and Müller (1888) as “… 8-nae …”. Notably,
on the isolectotype material of Verrucaria prasina in G
(G00290128), which Müller (1884) renamed Anthracothecium eschweileri, because he considered the thallus not to be
green (Latin: prasinus), Müller depicted a similar ascospore
development as for Verrucaria planiuscula, with 1-septate
ascospores next to muriform ones. We can therefore rather
safely assume that Verrucaria planiuscula Nyl. is another
synonym of Anthracothecium prasinum and not a species
of Anisomeridium or Megalotremis.
13
Fungal Diversity (2020) 105:17–318
Anisomeridium tarmugliense (≡ Ditremis tarmugliensis)
was described as having “vertically elongated” ascomata
with wide open ostioles that are white inside (Makhija and
Patwardhan 1990). The illustration shows an irregularly
lobed ascoma circumference. These characters contradict
placement in Anisomeridium or Monoblastiaceae. Except
for the ascospores (the hamathecial filaments were not
described), the morphology points to Stictidaceae, e.g.
Ostropa or Robergea. We regard the position of this taxon
as unresolved, but it is unlikely to be a lichenized bitunicate
ascomycete.
Megalotremis biocellata Aptroot is characterized by
an endoperidermal, ecorticate thallus and rather large
ascospores with secondary endospore invaginations in each
cell. Both the thallus and the ascospore type are characteristic for tropical, lichenized species of Arthopyrenia sensu
lato currently placed in the genus Constrictolumina in Trypetheliaceae (Aptroot and Lücking 2016). Harris (1995)
also related M. biocellata to Arthopyrenia sensu lato and
suspected that it could be the same as Tomasellia dispora.
The latter was described as having 12-locular ascospores
(Müller 1887, p. 427), but this is surely a lapsus for 2-locular, as Müller drew the ascospores on the holotype material (G-G00294870) as 1-septate. Harris (1995) annotated
the holotype of T. dispora as synonymous with A. malaccitula (≡ Constrictolumina malaccitula), although he did
not list that name as synonym of the latter. Instead, he gave
A. bifera and Didymella gigantea as synonyms. Examination of the type of M. biocellata supports its placement in
Constrictolumina, but it was initially unclear whether it is
actually conspecific with C. malaccitula or represents a
closely related taxon with larger ascospores. Aptroot (1991)
described the ascospores of M. biocellata as 47–53 × 17–21
µm in size. Harris (1995) studied the holotype of Verrucaria
malaccitula (H-H9505214, H-NYL 1706) and, contrary
to Nylander’s (in Nylander and Crombie 1884) measurements (26–36 × 12–16 µm), gave the ascospores as 37–48 ×
15–16 µm, a range adopted by Aptroot and Lücking (2016).
A specimen from Papua New Guinea (Aptroot 19093)
included under C. malaccitula in the latter work had mature
ascospores measuring up to 49 × 17 µm. The type material of T. dispora reportedly has ascospores 38–48 × 14–16
µm (Müller 1887), and that of A. bifera 42–45 × 15–16 µm
(Zahlbruckner 1935), whereas Didymella gigantea, with
smaller ascospores, does not seem to belong in this complex also on account of its morphology. Specimens identified as M. biocellata from Costa Rica (Aptroot et al. 2008)
have ascospores 40–45 × 16–20 µm in size [due to a lapsus,
the ascospores of Costa Rican material of that species (FH,
Brenes 188; see Aptroot et al. 2008: 71, Fig. 11A–B) were
due to a mixup of images erroneously also illustrated as
those of C. cinchonae in Aptroot and Lücking 2016: 919,
fig. 45L]. All this would point to a continuous range of about
Fungal Diversity (2020) 105:17–318
40–50 × 15–20 µm large ascospores for a single species,
with malaccitula as the oldest available epithet. We also discovered a lapsus in the protologue of M. biocellata (Aptroot
1991: 125): whereas in the description the ascospores were
given as 47–53 × 17–21 µm in size, in the corresponding
illustration (Aptroot 1991: fig. 48) they were depicted as
67–68 × 20–21 µm. Re-examination of the type material
showed an ascospore size of 45–50 × 15–20 µm. Thus, we
consider M. biocellata, another synonym of C. malaccitula,
besides T. dispora and A. bifera (see Lücking et al. in Hongsanan et al. 2020, this volume: Trypetheliaceae).
Megalotremis elegans (R.C. Harris) Lücking & Aptroot,
comb. nov.
MycoBank number: MB 836366; Index Fungorum number: IF 836366; Facesoffungi number: FoF 08078.
Bas.: Anisomeridium elegans R.C. Harris, More Florida
Lichens: 145 (1995).
Megalotremis holopolia (Nyl.) Lücking & Aptroot, comb.
nov.
MycoBank number: MB 836367; Index Fungorum number: IF 836367; Facesoffungi number: FoF 08079.
Bas.: Verrucaria holopolia Nyl., Bull. Soc. Linn. Normandie, Sér. 2(2): 131 (1868); Anisomeridium holopolium
(Nyl.) R.C. Harris, More Florida Lichens: 147 (1995).
Megalotremis immersa (Makhija & Patw.) Lücking & Aptroot, comb. nov.
MycoBank number: MB 836368; Index Fungorum number: IF 836368; Facesoffungi number: FoF 08080.
Bas.: Ditremis immersa Makhija & Patw., Biovigyanam
16: 17 (1990); Anisomeridium immersum (Makhija & Patw.)
R.C. Harris, More Florida Lichens: 147 (1995).
Megalotremis monospora (Makhija & Patw.) Lücking &
Aptroot, comb. nov.
MycoBank number: MB 836369; Index Fungorum number: IF 836369; Facesoffungi number: FoF 08081.
Bas.: Ditremis monospora Makhija & Patw., Biovigyanam 16: 20 (1990); Anisomeridium monosporum (Makhija
& Patw.) R.C. Harris, More Florida Lichens: 148 (1995).
Phellinocrescentia Crous & Decock in Crous et al., Persoonia 33: 235 (2014)
MycoBank number: MB 810600; Index Fungorum number: IF 810600; Facesoffungi number: FoF 08082; one phylogenetically and morphologically defined species (Crous
et al. 2014).
Type species: Phellinocrescentia guianensis Crous &
Decock in Crous et al., Persoonia 33: 235 (2014).
Notes: For description and illustrations see Crous et al.
(2014).
109
Pseudopassalora Crous in Crous et al., Persoonia 27: 41
(2011).
MycoBank number: MB 560570; Index Fungorum number: IF 560570; Facesoffungi number: FoF 08083; one phylogenetically and morphologically defined species (Crous
et al. 2014).
Type species: Pseudopassalora gouriqua Crous in Crous
et al., Persoonia 27: 41 (2011).
Notes: For description and illustrations see Crous et al.
(2011b).
Trypetheliopsis Asahina, J. Jap. Bot. 13: 319 (1937).
MycoBank number: MB 5627; Index Fungorum number:
IF 5627; Facesoffungi number: FoF 08084; 8 morphologically defined species (Harada 2017; Lücking et al. 2017),
molecular data available for one species.
= Musaespora Aptroot & Sipman, Lichenologist 25(2):
123 (1993).
MycoBank number: MB 22430; Index Fungorum number:
IF 22430; Facesoffungi number: FoF 08085.
Lichenized on bark or leaves, in tropical regions. Thallus
corticate, olive-green to brownish. Photobiont Trentepohlia.
Ascomata scattered or rarely pseudostromatic, erumpent to
prominent, brown-black and carbonaceous or rarely with
orange-red pigment, ostiolate, ostiole apical. Involucrellum present, carbonized. Excipulum dense, consisting of
compressed hyphae, appearing prosoplectenchymatous in
thin, bleached sections, brown to brown-black. Hamathecium comprising 0.5–0.7 µm wide paraphyses, hyaline,
straight, branched and anastomosing, usually inspersed. Asci
8-spored, bitunicate, fissitunicate, cylindrical, short pedicellate, with broad, non-amyloid ocular chamber and fluorescent cap-structures. Ascospores irregularly arranged, ovaloblong, typically slightly curved, hyaline, 1-septate, large
(70–120 × 12–25 µm), with internal, needle-shaped crystals.
Conidiomata campylidiiform, brown black or rarely with
orange-red pigment. Conidia acrogenous, microconidia,
aseptate, drop-shaped, small, hyaline, aggregated in gelatinous mass. Chemistry: Secondary substances absent except
for unidentified red pigment in the type and in T. coccinea.
Type species: Trypetheliopsis boninensis Asahina, J. Jap.
Bot. 13: 319 (1937).
Notes: Trypetheliopsis was described as a monospecific
genus for a corticolous lichen with pseudostromatic ascomata producing a bright, orange-red pigment and with large,
1-septate, fusiform ascospores with markedly submedian
septum (Asahina 1937; Santesson 1970). The genus was
largely forgotten, until Kashiwadani et al. (2009) resurrected it, considering it an earlier name for Musaespora,
based on similarities (in particular the orange-red pigment)
with Musaespora coccinea.
13
110
Fungal Diversity (2020) 105:17–318
Fig. 53 Morphology, asci and ascospores in Trypetheliopsis boninensis. a–c Thallus with ascomata, pycnidia, and campylidia (Japan, Kashiwadani 52239 & Moon). d–r Hamathecium, asci, and ascospores (holotype). Scale bars: d = 0.5 µm, e, f, h–n = 40 µm, g, o–r = 10 µm
Trypetheliopsis boninensis Asahina, J. Jap. Bot. 13(5): 319
(1937).
MycoBank number: MB 411966; Index Fungorum number: IF 411966; Facesoffungi number: FoF 08086; Fig. 53.
13
Lichenized on bark in tropical regions. Thallus corticate,
olive-green. Photobiont Trentepohlia. Ascomata pseudostromatic, prominent, with orange-red pigment, ostiolate,
ostiole apical. Involucrellum present, carbonized. Excipulum dense, consisting of compressed hyphae, appearing
Fungal Diversity (2020) 105:17–318
prosoplectenchymatous in thin, bleached sections, brown.
Hamathecium comprising 0.5–0.7 µm wide paraphyses, hyaline, straight, branched and anastomosing, usually inspersed.
Asci 8-spored. Ascospores irregularly arranged, oval-oblong,
fusifurm, 1-septate, with constrictions at septum, 40–50 ×
15–20 µm, with internal, needle-shaped crystals. Conidiomata of two type, pycnidia and campylidiiform, with orangered pigment. Conidia acrogenous, aseptate, drop-shaped,
small, hyaline, aggregated in gelatinous mass. Chemistry:
Several orange-red pigments, identified as skyrin, oxyskyrin
and skyrinol (Santesson 1970).
Material examined: JAPAN, Ryukyu, Pref. Okinawa, Iriomote-jima, around Kanpirei Fall, Taketomi-cho, Yaeyamagun, 24°21’17″ N, 123°48’29″ E, alt. c. 70 m, on bark, 9
March 2018, Kashiwadani 52239 & Moon, TNS).
Notes: We were able to restudy the type and other material in detail (Fig. 53). The ascomata are pseudostromatic,
with 5–10 perithecia per pseudostroma, rather closely
resembling the pseudostromata of Trypethelium sensu stricto
(Aptroot and Lücking 2016). The ascospores are distinctly
fusiform with a marked submedian septum, resembling
those of certain Megalotremis species. Recently collected
material (Kashiwadani 52239 & Moon, TNS) shows two
types of conidiomata: red pycnidia and dark red campylidia
(Fig. 53). Trypetheliopsis boninensis is thus clearly congeneric with Musaespora and differs from the type of the latter,
M. corticola (= T. gigas) largely in the sessile vs. prominent, strongly pigmented ascomata. The close relationship
between these taxa is also illustrated by T. coccinea, which
features a similar pigment as T. boninensis and differs from
the latter in the foliicolous growth habit, distinctly verrucose
thallus and erumpent to prominent, solitary ascomata.
111
6.
6.
7.
7.
8.
8.
9.
9.
10.
10.
11.
11.
Key to genera of Monoblastiaceae:
1.
1.
2.
2.
3.
3.
4.
4.
5.
5.
Ascomata absent; conidiomata present………………2
Ascomata present……………………………………6
Conidiomata hyphomycetous; conidia brown………3
Conidiomata coelomycetous (pycnidia, frequently
beaked or campylidiiform)……………………………4
Conidia pleurogenous, dimorphic, broadly ellipsoid
and 1-septate to oblong-bacillar and tapering and
(2–)3(–7)-septate, walls smooth; on bark……Funbolia
Conidia acrogenous, uniform, oblong-fusiform,
aseptate to 1-septate, walls finely verruculose; on
leaves………………………………Pseudopassalora
Non-lichenized; growing on bracket fungi (Phellinus)
……………………………………Phellinocrescentia
Lichenized; growing on bark or leaves………………5
Conidiomata campylidiiform…………Trypetheliopsis
Conidiomata beaked pycnidia or entirely hair-like
………………………………………………………6
12.
12.
13.
13.
14.
Pycnidia with distinctly inflated base and short beak;
on bark…………………………Anisomeridium p.p.
Pycnidia with slightly inflated base or entirely hairlike; on leaves………………………………………7
Thallus subcuticular, with Cephaleuros-like photobiont or appearing non-lichenized; ascomata
unknown; pycnidia hair-like, very thin, straight, rather
dense…………………Caprettia subgenus Caprettia
Thallus supracuticular, with Phycopeltis photobiont;
ascomata frequent; pycnidia setiform, thicker, often
bent, usually scattered…………………………………
…………………………Caprettia subgenus Porinula
Non-lichenized; growing on leaves; either asci clavate
with long stipe or ascospores with appendages………9
Lichenized; growing on bark or rarely on rock; asci
cylindrical, with short stipe, ascospores lacking appendages…………………………………………………10
Asci distinctly clavate, with long stipe, polyspored;
ascospores aseptate, oblong, without appendages;
growing on dead twigs and leaves…………Eriomyces
Asci cylindrical, with short stype, 8-spored; ascospores
1-septate, ellipsoid, with appendages; growing on
senescent leaves of Juncus in salt marshes…Heleiosa
Ascospores aseptate, with granular or peg-like ornamentation………………………………Monoblastia
Ascospores 1–3-septate, smooth or with granular ornamentation……………………………………………11
Ascospores small to medium-sized (less than 50 × 20
µm) and with thin walls and septa or septa slightly
buldging laterally; thallus mostly thin and ecorticate or
with thin cortex and opaque…………………………12
Ascospores large (more than 50 × 20 µm) and with
thickened walls and/or septa and often with internal,
needle-shaped crystals; thallus usually well-developed, usually with cartilaginous cortex and often with
metallic glance, very rarely endoperidermal and ecorticate………………………………………………14
Asci with broad ocular chamber; ascospores oblong,
usually granular ornamented; perithecia often sessile;
in temperate or tropical montane regions…Acrocordia
Asci with narrow ocular chamber; ascospores ellipsoid to fusiform, smooth to more rarely granular
ornamented; perithecia usually immersed to prominent, rarely in sessile pseudostromata; mostly tropical…………………………………………………13
Pycnidia hair-like or setiform; ascomata often non-carbonaceous; on leaves………………………Caprettia
Pycnidia wart-shaped or at best flask-shaped with short
beak; ascomata carbonaceous; usually on bark, rarely
on leaves………………………………Anisomeridium
Pycnidia campylidiiform; ascospores usually slightly
curved…………………………………Trypetheliopsis
13
112
100/1.0
-/0.90
90/0.99
100/1.0
Phaeothyriolum corymbiae CPC 35789
Phaeothyriolum corymbiae CBS 143434
Phaeothyriolum amygdalinum CPC 36084
Phaeothyriolum eucalyptorum CBS H-24021
Murramarangomycetales
Fig. 54 Phylogram generated
from maximum likelihood
analysis (RAxML) of Murramarangomycetales based on ITS,
LSU, and SSU sequence data.
Maximum likelihood bootstrap
values equal or above 70%,
Bayesian posterior probabilities
equal or above 0.90 (MLBS/
PP) are given at the nodes. An
original isolate number is noted
after the species name. The tree
is rooted to Patellaria quercus
(CPC 27232). The ex-type
strains are indicated in bold.
Hyphen (-) represents support
values below 70% MLBS and
0.90 PP
Fungal Diversity (2020) 105:17–318
Phaeothyriolum dunnii CPC 35847
Acrospermum compressum M151
100/1.0
Acrospermales
Acrospermum longisporium MFLU 17-2849
100/1.0
Natipusilla limonensis PE3-2a
Natipusilla decorospora ILL AF236-1
Natiousillales
Zeloasperisporium siamense
100/1.0
Zeloasperisporium wrightiae MFLUCC 15-0225
100/1.0
Zeloasperisporiales
Venturia inaequalis CBS 815.69
Venturia inaequalis CBS 594.70
Patellaria quercus CPC 27232
Venturiales
Outgroup
0.04
14. Pycnidia variously shaped but not campylidiiform
(or pycnidia absent); ascospores straight or slightly
curved………………………………………………15
15. On leaves; ascospores slightly curved…………………
……………………………………Caprettia (confusa)
15. On bark; ascospores straight………………………16
16. Thallus epiperidermal, corticate and usually shiny;
ascospores with internal, needle-shaped crystals……
…………………………………………Megalotremis
16. Thallus endoperidermal, ecorticate…………………17
17. Ascospores with internal, needle-shaped crystals
…………………………Megalotremis (megalospora)
17. Ascospores granular ornamented……………………
……………………………Anisomeridium (excellens)
Economic and ecological significance
Monoblastiaceae (including Eriomycetaceae) comprises
both lichenized and non-lichenized ascomycetes. No economic significance has been reported for lichenized members of the family, but foliicolous representatives have been
classified according to their potential to monitor environmental health of tropical forest ecosystems. Thus, Anisomeridium foliicola, Trypetheliopsis epiphylla and T. kalbii
were listed as characteristic of mostly closed forest (Lücking 1997). Corticolous Monoblastiaceae also appear to be
largely confined to closed forest (Rivas Plata et al. 2008),
although Lucheta et al. (2019) characterized Anisomeridium
13
leptospermum as urban lichen. The anthraquinone pigments
in Trypetheliopsis have been chemically studied (Santesson
1970) but their potential applications are unknown. In studies in the semi-arid vegetation of northeastern Brazil, species of Anisomeridium were shown to form part of the diet
of termites (Barbosa-Silva and Vasconcellos 2019; BarbosaSilva et al. 2019).
The non-lichenized lineages formerly separated in Eriomycetace are apparently mostly saprobic and hence function
as decomposers of plant and fungal organic matter, but their
exact ecology has not yet been assessed (Kohlmeyer et al.
1996; Crous et al. 2011a, b, 2014; Hyde et al. 2020b).
Murramarangomycetales Crous.
Index Fungorum number: IF 823433; Facesoffungi number:
FoF 08087.
Murramarangomycetales was introduced by Crous et al.
(2017) to accommodate Murramarangomycetaceae. The
family formed a distinct lineage which could not be assigned
to any orders in Dothideomycetes (Crous et al. 2017; Hongsanan et al. 2020; this study, Fig. 54).
Accepted families: Murramarangomycetaceae.
Murramarangomycetaceae Crous, in Crous et al., Persoonia
39: 385 (2017).
Index Fungorum number: IF 823434; Facesoffungi numbers: FoF 08088, 7 species.
Fungal Diversity (2020) 105:17–318
113
to solitary conidia. Conidia in slimy mass, aseptate, hyaline,
smooth, bacilliform (adapted from Crous et al. 2017, 2019c).
Type: Phaeothyriolum Syd.
Notes: This family was introduced by Crous et al. (2017)
to accommodate the monotypic genus, Murramarangomyces
which contains a single species, M. corymbiae. Phylogenetic
analyses indicated that Murramarangomycetaceae is different from other families and orders within Dothideomycetes,
therefore, Crous et al. (2017) established a new order to
accommodate this family. However, Murramarangomyces
was synonymised under Phaeothyriolum by Crous et al.
(2019c). Murramarangomycetaceae and Murramarangomycetales accommodates Phaeothyriolum base on the Phaeothyriolum clade not clustering with any other families and
orders in Dothideomycetes. Seqeuence data of the type species of Phaeothyriolum is needed.
Fig. 55 Phaeothyriolum corymbiae (redrawn from Crous et al. 2017,
CBS H-23281, holotype). a Mycelium becoming fertile conidiogenous cells. b, c Conidigenous cell aggregated in cauliflower-like
form. d Conidia
Leaf spots, associated with several fungi in a leaf spot
colony, confined to one side of the leaf. Mycelium interepidermal, brown, septate, branched, giving leaf spots a redpurple appearance without any superficial mycelium being
present, mycelium aggregating in substomatal cavity. Sexual
morph: Thyrothecium superficial, radial orientation or clusters, circular, flattened, with lobed, brown, smooth to irregular margin of cells of textura epidermoidea or angularis.
Hamathecium comprising hypha-like, dissolving at maturity,
or persistent, hyaline, septate, anastomosing or not, pseudoparaphyses. Asci 8-spored, ellipsoid-obclavate, pedicellate,
with small ocular chamber. Ascospores 3- to multi-seriate,
fusoid-ellipsoid to obovoid, 1-septate, guttulate, granular,
slightly constricted at median septum or not, encased in
prominent mucoid sheath which is absent at maturity, hyaline, becoming brown once discharged onto agar. Asexual
morph: Hyphomycetous. Hypophyllous, circular, dark
brown. Mycelium in vitro consisting of hyaline, smooth
hyphae, irregular, constricted at septa. Conidiogenous cells
thick-walled cells that become fertile conidiogenous cells,
aggregated in cauliflower-like clusters of brown, doliiform
to globose, brown conidiogenous cells, phialidic, giving rise
Phaeothyriolum Syd., Annls mycol. 36(4): 305 (1938).
= Murramarangomyces Crous, in Crous et al., Persoonia
39: 385 (2017).
Index Fungorum number: IF 3962; Facesoffungi number:
FoF 08089; 7 morphological species (Species Fungorum
2020), 4 species with molecular data.
Type species: Phaeothyriolum eucalyptinum Syd.
Notes: Phaeothyriolum was placed in genera incertae
sedis in Dothideomycetes in Wijayawardene et al. (2017a).
Sequence data of the type species is unavailable. Murramarangomyces based on a single species M. corymbiae was
found to be aexual morph of Phaeothyriolum corymbiae.
Thus, Murramarangomyces was synonymized under Phaeothyriolum (Crous et al. 2019c). Crous et al. (2019c) indicated that four species of Phaeothyriolum clustered together
within Dothideomycetes, but its placement was not assigned.
We were unable to obtain a fresh collection of species in this
family, so a drawing is provided (Fig. 55)
Economic and ecological significance
Species in this family cause leaf spots either on living leaves,
or on leaf litter (Crous et al. 2019c), confined to one side of
the leaf.
Muyocopronales Mapook, Boonmee & K.D. Hyde.
Index Fungorum number: IF 551615; Facesoffungi number: FoF 01886.
This order was established by Mapook et al. (2016b)
to accommodate a single family, Muyocopronaceae. The
order formed a distinct clade within Dothideomycetes
and was related to Acrospermales and Dyfrolomycetales
(Mapook et al. 2016b; this study). Muyocopronales differs
from Acrospermales and Dyfrolomycetales by superficial,
flattened, carbonaceous, brittle ascomata, cellular pseudoparaphyses that are longer than the asci and ellipsoidal to
ovate, unicellular ascospores (Hyde et al. 2013; Mapook
13
114
Fungal Diversity (2020) 105:17–318
Muyocopron garethjonesii MFLU 16-2664a
75/1.0
Muyocopron dipterocarpi MFLUCC 14-1103
-/1.0
Muyocopron unilateralis IMI 324533
100/1.0
Muyocopron coloratum CBS 720.95
100/1.0
Muyocopron castanopsis MFLUCC 14-1108
Muyocopron castanopsis MFLU CC 10-0042
Muyocopron lithocarpi MFLU CC 10-0041
100/1.0
100/1.0
Muyocopron lithocarpi MFLUCC 14-1106
Pseudopalawania siamensis MFLUCC 17-1476a
Pseudopalawania siamensis MFLUCC 17-1476b
Mycoleptodiscus terrestris CBS 231.53
Mycoleptodiscus endophytica MFLUCC 17-0545
Neomycoleptodiscus venezuelense CBS 100519
72/-
100/1.0
Neocochlearomyces chromolaenae culture BCC THA 68252
Paramycoleptodiscus albizziae CPC 27552
Arxiella terrestris CBS 268.65
100/1.0
Muyocopronales
100/1.0
Arxiella dolichandrae CBS 138853
77/0.90
100/1.0
Leptodiscella rintelii CBS 144927
Leptodiscella africana CBS 400.65
86/0.94
Setoapiospora thailandica MFLUCC 17-1426
100/1.0
100/1.0
Palawania thailandensis MFLUCC 14-1121
Palawania thailandensis MFLU 16-1873
100/1.0
100/1.0
Stigmatodiscus tamaricis L114
Stigmatodiscus oculatus AP161116
100/1.0
Dyfrolomyces rhizophorae BCC15481
Dyfrolomyces tiomanensis NTOU3636
100/1.0
0.04
Acrospermum gramineum M152
Acrospermum compressum M151
Palawaniaceae
fam. incertae sedis
Stigmatodis
cales
Dyfrolomycetales
Outgroup
Fig. 56 Phylogram generated from maximum likelihood analysis
(RAxML) of Muyocopronales based on ITS, LSU, rpb-2, SSU and
tef1 sequence data. Maximum likelihood bootstrap values equal
or above 70%, Bayesian posterior probabilities equal or above 0.90
(MLBS/PP) are given at the nodes. An original isolate number is
noted after the species name. The tree is rooted to Acrospermum compressum (M151) and A. gramineum (M152). The ex-type strains are
indicated in bold. Hyphen (-) represents support values below 70%
MLBS and 0.90 PP
et al. 2016b). Our phylogenetic analyses indicate that the
order comprises a single family, Muyocopronaceae. Palawaniaceae is sister to the clade of Muyocopronaceae (81%
ML, Fig. 56). Thus, we do not include this family within
Muyocopronales. Currently there is one family with seven
accepted genera in this order (this paper). The divergence
time for Muyocopronales is estimated as 171 MYA (stem
age, Hongsanan et al. 2020).
Accepted families: Muyocopronaceae.
Muyocopronaceae K.D. Hyde, in Hyde et al., Fungal Diversity 63: 164 (2013).
Index Fungorum number: IF 804506; Facesoffungi numbers: FoF 08090, 68 species.
Saprobic, common on the surface of dried twigs, stems
and less common on leaves, appearing as black spots, mycelium absent. Sexual morph: Ascomata superficial on host
surface, circular, scattered, flattened, rarely coalescing, carbonaceous and coriaceous, black, with a poorly developed
basal layer, with a central irregular ostiole, lenticular when
section. Upper wall irregularly arranged radiating cells,
13
Fungal Diversity (2020) 105:17–318
115
Fig. 57 Muyocopron dipterocarpi (MFLU 18-1385). a Ascomata on host substrate. b Section through ascostroma. c Peridium d Pseudoparaphyses. e–h Asci. i–m Ascospores. n Germinated spores. o Appressoria. Scale bars a = 200 µm, b = 100 µm, c, d, i–o = 10 µm, e–h = 30 µm
branched at margin cells. Peridium 2-layered, an outer layer
comprises brown to black pseudoparenchymatous of compact thick-walled textura epidermoidea cells, light brown
of textura angularis cells at inner layer. Hamathecium comprising dense, septate, cellular pseudoparaphyses which are
longer than the asci and immersed in mucilage and inclined
towards the centre. Asci 8-spored, bitunicate, fissitunicate,
pedicellate, with small ocular chamber. Ascospores 2–3-seriate, ellipsoidal to ovate with obtuse ends, hyaline, aseptate,
with or without granular appearance. Asexual morph: See
Papendorf (1967), Crous et al. (2016b, 2018a), HernándezRestrepo et al. (2019).
Type: Muyocopron Speg.
Notes: Muyocopronaceae was invalidly introduced by
Luttrell (1951), and was synonymized under Microthyriaceae (Wu et al. 2011a, b). Hyde et al. (2013) accepted and
formally introduced Muyocopronaceae as a distinct family,
as well as provided an English diagnosis (Latin diagnosis
no longer required). Phylogenetic analyses indicated that the
family, Muyocopronaceae comprises nine genera Arxiella,
Leptodiscella, Mycoleptodiscus, Muyocopron, Neocochlearomyces, Neomycoleptodiscus, Paramycoleptodiscus,
Setoapiospora, and Pseudopalawania (Mapook et al 2020;
this study, Fig. 56)
13
116
Muyocopron Speg., Anal. Soc. cient. argent. 12(3): 113
(1881).
Index Fungorum number: IF 3294; Facesoffungi number:
FoF 01887; 48 morphological species (Species Fungorum
2020), 5 species with molecular data.
Type species: Muyocopron corrientinum Speg.
Notes: Muyocopron was introduced by Spegazzini (1881),
with the type species M. corrientinum. Saccardo (1883)
re-described this genus and placed it in Microthyriaceae.
Muyocopron was synonymized under Ellisiodothis by von
Arx and Müller (1954). Later, it was transferred to Botryosphaeriaceae and Microthyriaceae (von Arx and Müller 1975; Lumbsch and Huhndorf 2007, 2010). However,
Muyocopron differs from members of Microthyriaceae by
pseudothyriothecia with the peridial wall comprising two
layers, pseudoparaphyses longer than the asci, and aseptate
ascospores (Hyde et al. 2013; Mapook et al. 2016b). Molecular studies indicated that Muyocopron is distantly related
to the Microthyriaceae (Wu et al. 2011a, b; Mapook et al.
2016b, and Fig. 56). The genus clustered with Dyfrolomycetaceae (Dyfrolomycetales), but as a distinct order (Hyde
et al. 2013; Mapook et al. 2016b).
Muyocopron dipterocarpi Mapook, Doilom, Boonmee &
K.D. Hyde, Phytotaxa 265 (3): 223 (2016).
Index Fungorum number: IF 551617; Facesoffungi number: FoF 01889, Fig. 57
Description: see Senwanna et al. (2019).
Material examined: Thailand, Phayao Province, Mueang
District, Wiang Subdistrict, on dried twig (attached on
tree) of Hevea brasiliensis, 2 October 2016, C. Senwanna,
RBPY036 (MFLU 18-1385).
Other genera included
Arxiella Papendorf, Trans. Br. mycol. Soc. 50(1): 73 (1967).
Index Fungorum number: IF 7233; Facesoffungi number:
FoF 08091; – 3 morphological species (Species Fungorum
2020), 3 species with molecular data.
Type species: Arxiella terrestris Papendorf, Trans. Br.
mycol. Soc. 50(1): 73 (1967).
Notes: Species of Arxiella have been reported with
asexual morphs (Papendorf 1967; Crous et al. 2014). Arxiella terrestris and A. lunata have hyphomycetous characters, while A. dolichandrae has coelomycetous characters.
Coelomycetous Arxiella is characterized by sporodochial,
brown conidiomata on agar surface, with globose to somewhat elongated, phialidic, brown conidiogenous cells, and
reniform, medianly 1-septate conidia, with guttulate, hornlike appendages at the ends (Crous et al. 2014). Hyphomycetous Arxiella is characterized by septate, branched, hyaline to slightly olivaceous hyphae, with 1- to multi-celled,
all cells conidiiferous conidiophores, and reniform, ends
obliquely cornute, 1-septate, hyaline to slightly coloured
13
Fungal Diversity (2020) 105:17–318
conidia, on short or elongate sterigma-like projections, single or in short chains or in small irregular groups (Papendorf 1967). Arxiella was treated as incertae sedis by Crous
et al. (2014). Crous et al. (2018a) showed the placement of
Arxiella within Muyocopronaceae. Subsequently, this was
supported by Hernández-Restrepo et al. (2019) who provided a phylogenetic tree from four loci (ITS, LSU, rpb-2,
and tef1) and concluded that this genus should be placed in
Muyocopronaceae.
Leptodiscella Papendorf, Trans. Br. mycol. Soc. 53(1): 146
(1969).
Index Fungorum number: IF 8743; Facesoffungi number:
FoF 08092; – 5 morphological species (Species Fungorum
2020), 5 species with molecular data.
Type species: Leptodiscella africana (Papendorf) Papendorf, Trans. Br. mycol. Soc. 53(1): 146 (1969).
≡ Leptodiscus africanus Papendorf, Trans. Br. mycol.
Soc. 50(4): 687 (1967).
Notes: Leptodiscella was described based on hyphomycetous characters, producing conidia, long appendages,
chlamydospores forming long or short chains or without
chlamydospores present, ellipsoid to oblong-oval or obovoid appressoria or absent. Conidiogenous cells are solitary
or aggregated, mono- or polyblastic, sympodial (Papendorf
1969; Udagawa and Toyazaki 1985; Madrid et al. 2012).
Papendorf (1967) reported that Leptodiscella was isolated
from leaf litter and soil of an Acacia karroo Heyne community in South Africa, but molecular data is unavailable.
Madrid et al. (2012) provided sequence data of Leptodiscella
based on phenotypic comparisons and presented taxonomic
position. Crous et al. (2018a) used ITS region in their phylogenetic analysis and showed the placement of Leptodiscella
within Muyocopronaceae.
Mycoleptodiscus Ostaz., Mycologia 59(6): 970 (1968)
[1967].
Index Fungorum number: IF 9029; Facesoffungi number:
FoF 08094; – 7 morphological species (Hernández-Restrepo
et al. 2019), 3 species with molecular data.
Type species: Mycoleptodiscus terrestris (Gerd.) Ostaz.,
Mycologia 59(6): 970 (1968) [1967].
≡ Leptodiscus terrestris Gerd., Mycologia 45(4): 552
(1953).
Notes: The type species was included in the invalid
genus Leptodiscus (Gerdemann 1953) which was placed
in Microthyriaceae (von Arx and Müller 1975; Lumbsch
and Huhndorf 2007). Thongkantha et al. (2009) provided
the first phylogenetic analysis of a Mycoleptodiscus species
(M. coloratus) based on LSU sequence data, and placed
it in Magnaporthales (Sordariomycetes). However, Crous
et al. (2018a) indicated that Mycoleptodiscus, based on phylogenetic placement of the type species, M. terrestris, is a
Fungal Diversity (2020) 105:17–318
member of Muyocopronales. More discussion can be seen
in Hernández-Restrepo et al. (2019). The sexual morph of
this genus has not been described, except for M. affinis, it
however was introduced as Omnidemptus affinis by Cannon
and Alcorn (1994). The major characters are restricted to
species characterized by cylindrical conidia with appendages
at one or both ends. Conidiogenous cells lack a collarette,
are solitary or aggregated, mono- or polyblastic, sympodial, often denticulate or phialidic, globose or ampulliform.
Conidia are cylindrical, the appressoria have a visible pore
surrounded by dark radial lines, and sclerotia (HernándezRestrepo et al. 2019)
117
with a filamentous appendage at each end, produced in a
mucous pale yellow to brown mass (Hernández-Restrepo
et al. 2019). The only species in this genus was found on leaf
litter of Gyranthera caribensis in Venezuela. In the phylogenetic analyses of Hernández-Restrepo et al. (2019), this
genus formed a lineage within Muyocopronaceae and sister
to the strain of Mycoleptodiscus endophyticus (MFLUCC
17-0545) with good support (71% ML, 0.98 BYPP). In our
analysis (Fig. 56), the genus is related to the clade containing Mycoleptodiscus endophyticus and Mycoleptodiscus
terrestris with moderate support. However, both analyses
showed that Neomycoleptodiscus is a distinct genus in
Muyocopronaceae.
Neocochlearomyces Pinruan, Sommai, Suetrong, J.Z. Groenew. & Crous, in Crous et al., Persoonia 41: 381 (2018).
Index Fungorum number: IF 828085; Facesoffungi number: FoF 08095; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Neocochlearomyces chromolaenae Pinruan, Sommai, Suetrong, J.Z. Groenew. & Crous, in Crous
et al., Persoonia 41: 381 (2018).
Notes: The phylogenetic analyses from ITS, LSU, rpb2, and tef1 (Hernández-Restrepo et al. 2019) showed that
Neocochlearomyces formed a distinct lineage within Muyocopronaceae. The type species was sister to Neomycoleptodiscus venezuelense (CBS 100519) and Mycoleptodiscus
endophyticus (MFLUCC 17-0545) in the analysis of Hernández-Restrepo et al. (2019). While, it is related to Paramycoleptodiscus albizziae in our analyses but as a distinct genus
(Fig. 56). The genus is characterized by macronematous and
mononematous, subcylindrical, septate, thick-walled conidiophores, with terminal and intercalary conidiogenous cells
on one side of the swollen fan-like structure, and falcate,
aseptate conidia that are equilateral, convex, obtuse to subobtusely rounded, forming a slimy spore mass (Crous et al.
2018a).
Paramycoleptodiscus Crous & M.J. Wingf., in Crous et al.,
Persoonia 36: 371 (2016).
Index Fungorum number: IF 817042; Facesoffungi number: FoF 08093; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Paramycoleptodiscus albiziae Crous &
M.J. Wingf. [as ‘albizziae’], in Crous et al., Persoonia 36:
371 (2016).
Notes: The genus is characterised by mycelium consisting
of hyaline to brown, branched, septate hyphae, chlamydospore-like cells that form a stroma, sporodochium of densely
aggregated stroma, solitary or aggregated, subglobose to
slightly ampulliform conidiogenous cells, and mono- or
polyblastic, sympodial, aseptate conidia that are hyaline,
smooth, granular, and base with well defined fusarium-like
foot cell (Crous et al. 2016b). The phylogenetic analyses of
Crous et al. (2018a) indicated that the type species of this
genus is closely related to Mycoleptodiscus terrestris, but
morphological characters are different. Paramycoleptodiscus
differs from Mycoleptodiscus by aseptate conidia and basal
appendage in the form of a foot cell, while Mycoleptodiscus
has septate conidia with apical appendages.
Neomycoleptodiscus Hern.-Restr., J.D.P. Bezerra & Crous,
in Hernández-Restrepo et al., Persoonia 42: 221 (2019).
Index Fungorum number: IF 829829; Facesoffungi number: FoF 08096; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Neomycoleptodiscus venezuelense Hern.Restr., J.D.P. Bezerra & Crous, in Hernández-Restrepo et al.,
Persoonia 42: 221 (2019).
Notes: Neomycoleptodiscus was established by Hernández-Restrepo et al. (2019) to accommodate a single species
Neocochlearomyces venezuelense. The genus has broadly
lunate conidia and variable production of conidial appendages, smooth, hyaline to pale brown hyphae, sporodochiumlike, brown conidiomata, with ampulliform to doliiform,
angular, smooth, with a circular aperture situated conidiogenous cells, and one septate, cylindrical, guttulate conidia,
Pseudopalawania Mapook. & K.D. Hyde, Biomolecules,
10: 569 (2020).
Mycobank number: MB834934; Facesoffungi number:
FoF 08097; – 1 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Pseudopalawania siamensis Mapook &
K.D. Hyde, Biomolecules, 10: 569 (2020).
Notes: The genus has sub-carbonaceous to superficial,
flattened, carbonaceous ascomata, poorly developed basal
layer and an irregular margin, with an ostioles, branching
pseudoparaphyses, cylindric-clavate asci, with an ocular
chamber observed clearly when immature, and broadly fusiform to inequilateral, 1-septate, hyaline ascospores, guttulate when immature, with thin gelatinous sheath (Mapook
et al. 2020). Although, morphology of Pseudopalawania
is typical of Palawania, molecular analyses confirmed its
13
118
Natipusilla limonensis ILL AF286-1
99/1.0
100/1.0
96/0.92
Natipusilla limonensis PE3 2a
Natipusilla decorospora ILL AF236-1
Natipusillalceae
100/1.0
100/1.0
Natipusilla naponensis ILL AF217-1
Natipusilla bellaspora ILL PE91 1a
Zeloasperisporiaceae
77/1.0
Zeloasperisporium wrightiae MFLUCC 15-0225
Zeloasperisporiales
Zeloasperisporium siamense IFRDCC 2194
100/1.0
Natipusillales
Fig. 58 Phylogram generated
from maximum likelihood
analysis (RAxML) of Natipusillales based on LSU and SSU
sequence data. Maximum likelihood bootstrap values equal or
above 70%, Bayesian posterior
probabilities equal or above
0.90 (MLBS/PP) are given at
the nodes. Isolate/specimen
number is noted after the species name. The tree is rooted
to Astrothelium variolosum
(MPN43), Trypthelium eluteriae
(111). The ex-type strains are
indicated in bold. Hyphen (-)
represents support values below
70% MLBS and 0.90 PP
Fungal Diversity (2020) 105:17–318
Trypethelium eluteriae 111
Outgroup
Astrothelium variolosum MPN43
0.03
placement within Muyocopronaceae (Mapook et al. 2020;
this study Fig. 56).
Setoapiospora Mapook & K.D. Hyde, Fungal Diversity 100:
5–277 (2020).
Index Fungorum number: IF 556905; Facesoffungi number: FoF 06793; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Setoapiospora thailandica Mapook & K.D.
Hyde, in Fungal Diversity 100: 5–277 (2020).
Notes: The genus is characterised by appearing as dark
brown to black spots on branches of wood, carbonaceous
ascomata, with dark brown setae, peridium composed of
cells of textura epidermoidea, pseudoparaphyses, 8-spored
asci, with an ocular chamber, and 1-septate ascospores, constricted at the septum, with a small lower cell and a large
upper cell, tapering towards ends, with granular appearance.
(Hyde et al. 2020b). In the phylogenetic analyses indicated
that this genus formed a distinct lineage within Muyocopronaceae (Hyde et al. 2020b, this study Fig. 56).
Economic and ecological significance
Species of Muyocopronaceae are commonly found as
saprobes on dead twigs and stems, thus playing roles in
decomposition.
Natipusillales Raja, Shearer, A.N. Mill. & K.D. Hyde.
Index Fungorum number: IF 805300; Facesoffungi number: FoF 08110.
Natipusillales was established to accommodate members
of the freshwater ascomycetes belonging to Natipusillaceae
based on LSU and SSU sequence data (Hyde et al. 2013).
Phylogenetic analyses placed this order in Dothideomycetes
(Hyde et al. 2013; Liu et al. 2017). Natipusillales was related
13
to Zeloasperisporiales which is not freshwater fungi and has
very different morphological characters (Hongsanan et al.
2015b). There is a single family with one genus accepted in
this order (Fig. 58). The divergence time for Natipusillales
is estimated as 180 MYA (stem age, Hongsanan et al. 2020).
Accepted families: Natipusillaceae.
Natipusillaceae Raja, Shearer & A.N. Mill., Mycologia
104(2): 570 (2012).
Index Fungorum number: IF 561948; Facesoffungi number: FoF 08111, 4 species.
Saprobic on decaying wood. Sexual morph: Ascomata
globose to subglobose, erumpent to superficial, hyaline to
light brown. Peridium membranous, composed of pseudoparenchyma cells of textura angularis in surface view.
Hamathecium comprising septate, sparse, cellular pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, globose,
subglobose or obclavate. Ascospores multi-seriate, fusiform to cylindrical, hyaline becoming brown with age, one
to several septate, multiguttulate or not. Asexual morph:
Undetermined.
Type: Natipusilla A. Ferrer, A.N. Mill. & Shearer.
Notes: Natipusillaceae comprises the freshwater genus
Natipusilla with four species (Ferrer et al. 2011; Raja et al.
2012). Natipusillales was established to accommodate members of Natipusillaceae based on LSU and SSU sequence
data (Hyde et al. 2013).
Natipusilla A. Ferrer, A.N. Mill. & Shearer, Mycologia
103(2): 417 (2011).
Index Fungorum number: IF 518365; Facesoffungi number: FoF 08112; 4 morphological species (Species Fungorum 2020), 4 species with molecular data.
Fungal Diversity (2020) 105:17–318
119
Fig. 59 Morphology of Natipusilla spp. (a–d = redrawn of Natipusilla decorospora from holotype AF236-1; e, f = redrawn of N. limonensis
from holotype AF286-1). a Ascomata. b, e Asci. c, d, f Ascospores. Scale bars: a–f = 10 µm
Type species: Natipusilla decorospora A. Ferrer, A.N.
Mill. & Shearer.
Notes: Species of Natipusilla have small ascomata, none
to few pseudoparaphyses, globose asci and mostly one-septate ascospores (Ferrer et al. 2011; Raja et al. 2012). There
are four species accepted in this genus. We were unable to
obtain a fresh collection of species in this family, so a drawing is provided (Fig. 59).
from Asterinales. Taxonomic details can be seen in Dai et al.
(2018). Only a single from Parmulariales has sequence data.
The clade of Parmulariales is sister to the new family Thyrinulaceae with high boostrap support (Figs. 5, 60), and this
result is similar to that found by Dai et al. (2018). The divergence time for Parmulariales is estimated as 115 MYA (stem
age, Hongsanan et al. 2020).
Accepted families: Parmulariaceae.
Economic and ecological significance
As typical lignicolous freshwater taxa, species of Natipusillales play an important role in nutrient and carbon cycling,
biological diversity and ecosystem functioning of freshwater
ecosystems with their ability to decompose lignocellulose
in woody litter, softening the wood and releasing nutrients
(Palmer et al. 1997; Wong et al. 1998; Yuen et al. 1998;
Bucher et al. 2004; Hyde et al. 2016a).
Parmulariaceae E. Müll. & Arx ex M.E. Barr, Mycologia
71(5): 944 (1979).
Index Fungorum number: IF 81109; Facesoffungi numbers: FoF 02296, 149 species.
Epiphytic or parasitic on living leaves of plants and
ferns or thallus of lichens. Colonies foliicolous, superficial,
dark brown to black. Hyphae spreading out from ascomata
border, brown, septate, branching, sinuous, with or lacking
appressoria. Sexual morph: Ascostromata solitary to gregarious, or in groups, superficial to immersed, shield-like to
star-shaped, elliptical to boat-shaped, dark brown to black,
carbonaceous to membranaceous, strongly flated, or longitudinal slits slightly protuberant, and contain numerous asci.
Locules immersed in ascostromata or in longitudinal ridges
Parmulariales D.Q. Dai & K.D. Hyde.
Index Fungorum number: IF 554065; Facesoffungi number: FoF 03913.
The order was established by Dai et al. (2018) based on its
phylogeny and morphological characters which are different
13
120
Parmularia styracis VIC 42587
100/1.00
Parmularia styracis VIC 42450
Parmularia styracis VIC 42447
87/0.95
93/0.90
Thyrinula parasitica CBS 120088
100/1.0
92/0.93
Blastacervulus eucalypti CBS 124759
Thyrinulaceae
Paraopeba schefflerae AMG-2017
Parmulariales
Fig. 60 Phylogram generated
from maximum likelihood
analysis (RAxML) of Parmulariales based on LSU sequence
data. Maximum likelihood
bootstrap values equal or above
70%, Bayesian posterior probabilities equal or above 0.90
(MLBS/PP) are given at the
nodes. Isolate/specimen number
is noted after the species name.
The tree is rooted to Venturia
inaequalis (ATCC 60070) and
V. populina (CBS 256.38). The
ex-type strains are indicated in
bold. Hyphen (-) represents support values below 70% MLBS
and 0.90 PP
Fungal Diversity (2020) 105:17–318
Cladoriellales
Cladoriella rubrigena CBS 124760
100/1.0
Cladoriella eucalypti CPC 10953
Venturia populina CBS 256.38
Outgroup
100/1.0
Venturia inaequalis ATCC 60070
0.03
radiating from the centre. Stromata wall thin to thick, composed of black, large cells of textura prismatica. Peridium
of locules thin to thick, composed of dark brown to hyaline
smaller cells of textura angularis. Hamathecium comprising
dark brown to hyaline, septate, pseudoparaphyses present or
absent. Asci 8-spored, thick-walled, bitunicate, long ovate,
broad cylindrical, broad-clavate, or obclavate, globose to
subglobose, short pedicellate, with a distinct ocular chamber.
Ascospores 2–multi-seriate, oblong to ellipsoid, ellipsoid to
ovate, hyaline to dark brown, 1-septate, wall smooth to verrucose. Asexual morph: Undetermined.
Type: Parmularia Lév.
Notes: Parmulariaceae was invalidly introduced by Müller and von Arx (1962) for taxa with colonies growing on the
surface of living plant tissues, mostly leaves, and later was
validated by Barr (1979a). The family contains a range of
ascomata types, asci and ascospores forms (Inácio and Cannon 2008). von Arx and Müller (1975) accepted 27 genera
in this family and Lumbsch and Huhndorf (2007) included
34 genera. Inácio and Cannon (2008) excluded Chaetaspis
and Kentigia, with 34 genera accepted. Hyde et al. (2013)
accepted 30 genera including a new genus Antoniomyces,
which was erected by Inácio et al. (2012). Placostromella
was re-described by Thambugala et al. (2014b) and transferred to this family. Placoasterella and Placosoma were
transfered to Parmulariaceae, as they were similar to taxa
in this family (Hongsanan et al. 2014b). Pleiostomellina
13
was referred by Ariyawansa et al. (2014). Dictyocyclus was
moved to Myriangiaceae based on morphological similarity (Dissanayake et al. 2014). Wijayawardene et al. (2014a)
accepted 34 genera in Parmulariaceae. Wu et al. (2014)
re-examined the holotype specimen of the type species of
Cirsosiopsis and transferred the genus to Parmulariaceae
based on its morphology. A new genus Rhagadolobiopsis
was introduced by Guatimosim et al. (2014) and Jayasiri
et al. (2016) re-examined herbarium of type species of Cocconia Sacc. and confirmed its placement within Parmulariaceae. Dai et al. (2018) excluded Hemigrapha, introducing Hemigraphaceae to accommodate it and re-examined
Inocyclus, moving it to Asterinales genera incertae sedis
based on the morphology and phylogenetic analysis. Doilom et al. (2018) transferred Syrropeltis to Parmulariaceae
based on morphology. Thus, 34 genera are accepted in Parmulariaceae. Parmulariaceae is close to Asterinaceae, but
differs in having ascostromata formed by several layers of
pigmented cells, pseudoparaphyses, broadly ellipsoidal to
obclavate asci, and lacks appressoria (Inácio and Cannon
2008; Inácio et al. 2012). Species of Asterinaceae have
superficial hyphae mostly with appressoria, thyriothecia
with a thin upper wall layer and poorly developed base and
subglobose asci (Pirozynski and Shoemaker 1970; Inácio
et al. 2012; Hongsanan et al. 2014b).
Parmularia Lév., Annls Sci. Nat., Bot., sér. 3 5: 286 (1846).
Fungal Diversity (2020) 105:17–318
Index Fungorum number: IF 3744; Facesoffungi number:
FoF 02297; 12 morphological species (Species Fungorum
2020), 1 species with molecular data, 34 predicted number
species.
Type species: Parmularia styracis Lév.
Notes: Parmularia typified by Parmularia styracis, was
introduced by Léveillé (1846). Parmularia is similar to
Mintera in having elongate locules radiating from a central
point. However, Mintera has mycelium with appresoria, and
black ascostroma with a single locule which aggregate into
a star shape. In Parmularia single ascostroma contain many
locules. Guatimosim et al. (2015) re-collected the type species and provided ITS, LSU and tef1-α sequence data. For
further morphological details of type species, see Dai et al.
(2018). Parmularia is characterized by large, black, shieldlike, flattened ascostromata with ridges radiating from the
centre to the outer rim, and the ridges comprising elongated,
radiating locules, filamentous pseudoparaphyses with brown
and verrucose apices and ellipsoid, 1-septate ascospores
(Inácio and Cannon 2008; Hyde et al. 2013; Dai et al. 2018)
Other genera included
Aldona Racib., Parasit. Alg. Pilze Java’s (Jakarta) 1: 19
(1900).
Index Fungorum number: IF 115; Facesoffungi number:
FoF 00309; – 4 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Aldona stella-nigra Racib., Parasit. Alg.
Pilze Java’s (Jakarta) 1: 19 (1900).
Notes: Aldona, typifided by Aldona stella-nigra was
introduced by Raciborski (1900). The type was originally
collected on living leaves of Pterocarpus indicus in Sumatra, western Indonesia. Aldona was initially placed in Hysteriaceae, then transferred to Phacidiaceae, and finally to
Parmulariaceae (Saccardo 1904; von Höhnel 1917; Müller and Patil 1973; von Arx and Müller 1975). The genus
was reported as pathogenic occurring on terrestrial habitats
and bearing an asexual ceolomycetous morph (for morphology see Tian et al. 2015; Wijayawardene et al. 2018, 2020).
Aldona is characterized by semi-immersed to erumpent,
black, linear, radial or star-shaped, branching, coriaceous,
shiny ascomata opening by longitudinal slits, black, shiny,
globose or irregular spermatogonia growing on the spot, surrounded by ascomata and elongate-clavate, hyaline, transseptate ascospores with upper cells larger and wider, basal
cells short and narrow (Tian et al. 2015).
Aldonata Sivan. & A.R.P. Sinha, Mycol. Res. 92(2): 248
(1989)
Index Fungorum number: IF 25242; Facesoffungi number: FoF 00311; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.
121
Type species: Aldonata pterocarpi Sivan. & A.R.P. Sinha,
Mycol. Res. 92(2): 249 (1989)
Notes: Aldonata was reported as a pathogen on living leaves of Pterocarpus draco in the Andaman Islands
(Sivanesan and Sinha 1989). The asexual morph of this
genus has not yet been reported (Tian et al. 2015). Aldonata
is characterized by semi-immersed, globose to subglobose,
black, shiny, ascomata appearing as flexuous lines on the
leaf spot surface with a clearly defined margin, solitary to
gregarious, sub-immersed, black and shiny, carbonaceous,
globose to irregular, spermatogonia mostly growing around
the central of greyish white spot and ellipsoid to fusiform,
muriform, hyaline ascospores with up to 8 transverse and
longitudinal septa (Tian et al. 2015).
Antoniomyces Inácio, Mycol Progress 11: -6 (2012).
Index Fungorum number: IF 563246; Facesoffungi number: FoF 02299; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Antoniomyces loranthicola Inácio, Mycol
Progress 11: -6 (2012).
Notes: This genus was introduced by Inácio et al. (2012)
according to its similarity of morphology with Parmulariaceae. This was followed in Wijayawardene et al. (2017a).
Antoniomyces is characterized by solitary to gregarious,
superficial, black, circular, ascostromata opening by longitudinal slits and forming in the central area of mycelium,
clavate to subglobose asci and cylindric-ellipsoidal or ellipsoidal to narrowly ovoid, 1-septate ascospores (Inácio et al.
2012).
Aulacostroma Syd. & P. Syd., Philipp. J. Sci., C, Bot. 9:
176 (1914).
Index Fungorum number: IF 456; Facesoffungi number:
FoF 02301; – 4 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Aulacostroma palawanense Syd. & P. Syd.,
Philipp. J. Sci., C, Bot. 9(2): 176 (1914).
Notes: Aulacostroma was reported as pathogenic on Pandanus merrillii in Philippines (see morphology in Inácio
and Cannon 2008; Wijayawardene et al. 2018, 2020). Aulacostroma is characterized by stromatic, solitary to gregarious, superficial, black ascomata, broadly clavate, globose
to subglobose, obovoid asci and brown ellipsoid, 1-septate
ascospores (Inácio and Cannon 2008).
Campoa Speg., Boln Acad. nac. Cienc. Córdoba 25: 90
(1921).
Index Fungorum number: IF 778; Facesoffungi number:
FoF 02303; – 4 morphological species (Species Fungorum
2020), molecular data unavailable.
13
122
Type species: Campoa pulcherrima Speg., Boln Acad.
nac. Cienc. Córdoba 25: 90 [no. 173, reprint pages 92]
(1921).
Notes: Campoa comprises five species, which were collected from Phillipines, Chile, Japan and the Dominican
Republic (Inácio and Cannon 2008). The genus was reported
as pathogenic on living leaves of Temu divaricatum (Inácio
and Cannon 2008; Wijayawardene et al. 2018, 2020). Campoa is characterized by superficial, flat, black ascostromata
with filamentous pseudoparaphyses, cylindrical to cylindricclavate asci and ellipsoidal ascospores (Inácio and Cannon
2008).
Cirsosiopsis Butin & Speer, Sydowia 31(1–6): 10 (1979)
[1978].
Index Fungorum number: IF 1068; Facesoffungi number:
FoF 08113; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Cirsosiopsis violacescens Butin & Speer,
Sydowia 31(1–6): 10 (1979) [1978].
Notes: The monotypic genus was introduced by Butin and
Speer (1978) with its family placement in Microthyriaceae.
Wu et al. (2014) re-examined the holotype of type species C.
violacescens and transferred it to Parmulariaceae based on
its morphological similarity with Parmularia. This genus is
characterized by black and discoid ascostromata opening by
irregular disintegration at the centre, globose to subglobose
asci and 1-septate, ellipsoid to ovate ascospores (Wu et al.
2014).
Cocconia Sacc., Syll. fung. (Abellini) 8: 738 (1889).
Index Fungorum number: IF 1149; Facesoffungi number:
FoF 00558; – 14 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Cocconia placenta (Berk. & Broome)
Sacc., Syll. fung. (Abellini) 8: 738 (1889).
≡ Rhytisma placenta Berk. & Broome, J. Linn. Soc., Bot.
14(no. 74): 131 (1873) [1875].
Notes: Jayasiri et al. (2016) re-examined the holotype and
confirmed its family level placement within Parmulariaceae.
The genus was reported as pathogenic on living leaves of
Symplocos spicata (see morphology in Inácio and Cannon
2008; Jayasiri et al. 2016; Wijayawardene et al. 2018, 2020).
Cocconia is characterized by superficial, shield-like, dark
brown to black, carbonaceous, flated ascostromata producing thick-walled, cylindrical asci and ellipsoidal,1-septate,
brown ascospores (Jayasiri et al. 2016).
Cycloschizon Henn., Bot. Jb. 33: 39 (1902).
Index Fungorum number: IF 1376; Facesoffungi number:
FoF 02305; – 13 morphological species (Species Fungorum
2020), molecular data unavailable.
13
Fungal Diversity (2020) 105:17–318
Type species: Cycloschizon brachylaenae (Rehm) Henn.,
Bot. Jb. 33: 39 (1902).
≡ Schneepia brachylaenae Rehm, Hedwigia 40(Beibl.):
(173) (1901).
Notes: The type species Cycloschizon brachylaenae was
collected on living leaves of Brachylaena neriifolia from
South Africa. Index Fungorum (2019) lists 14 species,
whereas Inácio and Cannon (2008) compared 12 taxa by
their hosts, distributions and main morphological characters. Cycloschizon is characterized by clustered, superficial,
black, circular, flat ascomata opening by one circumferential,
producing broadly clavate to subglobose, thick-walled asci
and ellipsoidal ascospores with one septum (Inácio and Cannon 2008). For a drawing and other details, see Inácio and
Cannon (2008), Wijayawardene et al. (2018, 2020).
Cyclostomella Pat., Bull. Herb. Boissier 4: 656 (1896).
Index Fungorum number: IF 1378; Facesoffungi number:
FoF 02307; – 3 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Cyclostomella disciformis Pat., Bull. Herb.
Boissier 4: 656 (1896).
Notes: Cyclostomella comprises four species (Inácio and
Cannon 2008). The genus was reported as pathogenic on
living leaves, usually in tropical regions such as Costa Rica,
Cuba and Brazil. A comparison of morphological features
of four taxa of Cyclostomella was provided by Inácio and
Cannon (2008). Cyclostomella is characterized by superficial, circular to elliptical, disc-like, black, rather flat ascostromata producing cylindric to clavate to subglobose asci
and ellipsoid, dark brown, 1-septate ascospores (Inácio and
Cannon 2008).
Dothidasteroma Höhn., Sber. Akad. Wiss. Wien, Math.naturw. Kl., Abt. 1 118: 1509 (1909).
Index Fungorum number: IF 1691; Facesoffungi number:
FoF 08114; – 4 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Dothidasteroma pterygotae (Berk. &
Broome) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
Kl., Abt. 1 118: 1510 (1909).
≡ Rhytisma pterygotae Berk. & Broome, J. Linn. Soc.,
Bot. 14(no. 74): 131 (1873) [1875].
Notes: Dothidasteroma was erected by von Höhnel
(1909a) with D. pterygotae as type species. This genus
is characterized by fruiting bodies forming a 1-cell layer,
superficial, flat ascostromata with irregular locules. Inácio
and Cannon (2008) accepted three species in Dothidasteroma. Inácio et al. (2011) introduced a new species. Thus,
four species are accepted in Dothidasteroma.
Ferrarisia Sacc., Atti Inst. Veneto Sci. lett., ed Arti 10: 61
(1919).
Fungal Diversity (2020) 105:17–318
Index Fungorum number: IF 1986; Facesoffungi number:
FoF 02309; – 8 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Ferrarisia philippina Sacc., Atti Soc.
Veneto-Trent. Sci. Nat., Padova 23: 61 (1917).
Notes: Ferrarisia philippina was synonymized under F.
ipomoeae. Thus, the current name for the generic type is F.
ipomoeae (Inácio and Cannon 2008). Ferrarisia jasmini was
transfered to Palawaniella by von Arx and Müller (1975).
Eight species are accepted in Ferrarisia by Inácio and Cannon (2008). Ferrarisia was reported as pathogenic on living
leaves and usually occurrs in tropics (Inácio and Cannon
2008; Wijayawardene et al. 2018, 2020). Ferrarisia is characterized by small, superficial, circular to elliptical, black
ascostromata with subglobose asci and ellipsoid, brown
ascospores with one septum (Inácio and Cannon 2008).
Hysterostomella Speg., Anal. Soc. cient. argent. 19(6): 260
(1885).
Index Fungorum number: IF 2476; Facesoffungi number:
FoF 02313; – 25 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Hysterostomella guaranitica Speg., Anal.
Soc. cient. argent. 19(6): 260 [no. 305] (1885).
Notes: This genus is widely distributed throughout the
tropics and is pathogenic on living leaves of palms (Inácio
and Cannon 2008; Wijayawardene et al. 2018, 2020). Hysterostomella is characterized by circular or elliptical, flat, black
ascostromata opening by irregular slits, with filamentous
pseudoparaphyses above the asci, which are subglobose,
obovoid, and clavate (Inácio and Cannon 2008).
Kiehlia Viégas, Bragantia 4(1–6): 156 (1944).
Index Fungorum number: IF 2567; Facesoffungi number:
FoF 08115; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Kiehlia obscura Viégas, Bragantia 4(1–6):
156 (1944).
Notes: Kiehlia obscurawas originally collected on living
leaves of bamboo from Brazil. Inácio and Cannon (2008) reexamined two varieties of the second species K. bambusina,
and provided illustrations of sexual and asexual morphs.
Index Fungorum (2020) listed K. bambusina as Phyllachora bambusina, hence only the type species is accepted
in this genus. Kiehlia is characterized by elongate, immersed
stromata and conidiomata growing in a same locule, clavate
asci and ovoid, hyaline ascospores with one septum (Viégas
1944).
Mintera Inácio & P.F. Cannon, Mycol. Res. 107(1): 86
(2003).
123
Index Fungorum number: IF 28710; Facesoffungi number: FoF 02317; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Mintera reticulata (Starbäck) Inácio & P.F.
Cannon, Mycol. Res. 107(1): 86 (2003).
≡ Parmularia reticulata Starbäck, Ark. Bot. 5(no. 7): 6
(1905).
Notes: This genus was reported as pathogenic on living
leaves with only sexual morph found. Mintera is characterized by stromatic, gregarious, black ascomata grouping
into a pyrethrum-like shape on mycelium, cylindric-clavate,
subglobose asci and ellipsoid, brown, 1-septate ascospores
(Inácio and Cannon 2008).
Pachypatella Theiss. & Syd., Annls mycol. 13(3/4): 228
(1915).
Index Fungorum number: IF 3670; Facesoffungi number:
FoF 02319; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Pachypatella alsophilae (Racib.) Theiss. &
Syd., Annls mycol. 13(3/4): 228 (1915).
≡ Hysterostomella alsophilae Racib., Parasit. Alg. Pilze
Java’s (Jakarta) 2: 22 (1900).
Notes: Pachypatella was reported as pathogenic on living
leaves of ferns. Two species including the type are described
(morphology see Inácio and Cannon 2008; Wijayawardene
et al. 2018, 2020). Pachypatella is characterized by superficial, circular or elliptical, flat ascostromata, becoming irregular in shape, with clavate, or cylindrical to cylindric-clavate
asci and cylindric-ellipsoidal to ellipsoidal or occasionally
oblong ascospores (Inácio and Cannon 2008).
Palawaniella Doidge, Bothalia 1(1): 16 (1921).
Index Fungorum number: IF 3683; Facesoffungi number:
FoF 02321; – 6 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Palawaniella eucleae Doidge, Bothalia
1(1): 16 (1921).
Notes: Doidge (1942) synonymized P. eucleae under P.
orbiculata. Currently, seven taxa are placed in this genus. It
was reported as pathogenic occurring on living leaves from
South Africa and Brazil (morphology see Inácio and Cannon 2008; Wijayawardene et al. 2018, 2020). Palawaniella
is characterized by stromatic ascomata that are gregarious
in the centre of black spots, becoming flattened and with a
single locule (Inácio and Cannon 2008).
Parmulariopsella Sivan., Trans. Br. mycol. Soc. 55(3): 509
(1970).
Index Fungorum number: IF 3746; Facesoffungi number:
FoF 02323; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
13
124
Fungal Diversity (2020) 105:17–318
Fig. 61 Parmulariopsis pulchella (W No: 10820, holotype). a Type
material. b, c Appearance of black ascostromata on the host. d Section of ascostroma. e Peridium. f–i Asci with ascospores. j Interthe-
cial, dark brown filamentous pseudoparaphyses. k–o Ascospores.
Scale bars: a = 10 mm, b, c = 1 mm, d, e = 100 µm, f–j = 10 µm,
k–o = 5 µm
Type species: Parmulariopsella burseracearum Sivan.,
Trans. Br. mycol. Soc. 55(3): 509 (1970).
Notes: This monotypic genus was collected on Santiria
trimera from Africa. This genus is similar to Parmularia,
but differs as the ascomata are covered by dark hyphae (Inácio and Cannon 2008). Parmulariopsella was reported as
pathogenic on living leaves (morphology see Inácio and
Cannon 2008; Wijayawardene et al. 2018, 2020). This genus
is characterized by solitary ascomata growing in the centre
of the black mycelium, globose to subglobose, obovoid asci
and ellipsoid, brown, 1-septate ascospores (Inácio and Cannon 2008).
13
Parmulariopsis Petr., Sydowia 8(1–6): 186 (1954).
Fungal Diversity (2020) 105:17–318
Index Fungorum number: IF 3747; Facesoffungi number:
FoF 02325; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Parmulariopsis pulchella Petr., Sydowia
8(1–6): 186 (1954).
Notes: This genus forms large, black ascostromata on
living leaves and is only known from the type collection
from Malaysia (Inácio and Cannon 2008). Parmulariopsis
is characterized by its shield-like ascostromata with locules
radiating from the centre. It looks similar to Parmularia,
but Parmulariopsis has elliptical to oblong, or long fusiform
stromata, whereas Parmularia has rounded stromata.
Parmulariopsis pulchella Petr., Sydowia 8(1–6): 186
(1954).
Index Fungorum number: IF 302295; Facesoffungi number: FoF 02326, Fig. 61
Parasitic on lower surface of leaves. Sexual morph:
Ascostromata 2–20 long × 1–4 mm wide, solitary to gregarious, superficial, elliptical to oblong, or long fusiform,
or irregular, shield-like, black, carbonaceous, flated, with
ridges radiating from the centre to the outer rim; ridges
containing elongated locules, which open by a longitudinal slit and contain numerous asci, cells of ascostromata
thick-walled and elongate and composed of black, amorphous tissues in the stromatic crust. Locules in vertical section 70–150 diam. × 65–85 µm high, subglobose, immersed
in ascostromata. Peridium of ascostromata 30–40 µm thick,
comprises of several black layers, composed of dark brown
cells of textura prismatica. Cells between locules 30–55 µm
thick, composed of brown to hyaline cells of texture angularis. Hamathecium comprising interthecial, hyaline to brown,
septate, filamentous pseudoparaphyses, 50–65 × 1.5–2.5 µm,
with swollen and brown tips. Asci 40–70 × 20–30 µm ( x̄ =
52.4 × 23.3 µm, n = 20), 8-spored, thick-walled, bitunicate,
clavate to subglobose, short-pedicellate, with a distinct ocular chamber. Ascospores 19–25 × 8.5–9 µm ( x̄ = 19.7 × 8.7
µm, n = 20), irregularly arranged, ellipsoidal, with larger
and rounder upper cells, with narrow and longer lower cells,
hyaline, 1-septate, constricted at the septum, smooth-walled.
Asexual morph: Undetermined.
Material examined: Malaysia, Sabah, Mt. Kinabalu, Tenmopok, 5000 ft., on leaves of Joinvillea sp., 9 March 1932,
J. & M. S. Clemens (W No: 10820, holotype).
Notes: Parmulariopsis was typified by P. pulchella, without designating any type specimens. Inácio and Cannon
(2008) re-examined the isotypes from S and K. In the present study, a specimen deposited in W labeled with the same
collecting information as in the original article is thought to
be the holotype.
Parmulina Theiss. & Syd., Annls mycol. 12(2): 194 (1914).
125
Index Fungorum number: IF 3748; Facesoffungi number:
FoF 08116; – 5 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Parmulina exsculpta (Berk.) Theiss. &
Syd. [as ‘exculpta’], Annls mycol. 12(1): 45 (1914).
≡ Dothidea exsculpta Berk., Hooker’s J. Bot. Kew Gard.
Misc. 6: 233 (1854).
Notes: Inácio and Cannon (2008) re-examined a specimen
from K, and designated it as neotype, as the authors considered the holotype was missing. This genus was reported
as pathogenic on living leaves (morphology see Inácio and
Cannon 2008; Wijayawardene et al. 2018, 2020). Parmulina
is characterized by superficial, circular to irregular, rather
flat, black ascostromata (Inácio and Cannon 2008).
Placoasterella Sacc. ex Theiss. & Syd., Annls mycol.
13(3/4): 236 (1915).
Index Fungorum number: IF 4121; Facesoffungi number:
FoF 08117; – 4 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Placoasterella schweinfurthii (Henn.) Theiss. & Syd., Annls mycol. 13(3/4): 236 (1915).
≡ Asterella schweinfurthii Henn., Bull. Herb. Boissier 1:
118 (1893).
Notes: Placoasterella was erected with two species.
Another two species were introduced with their family
placement in Asterinaceae (Müller and von Arx 1962).
Hongsanan et al. (2014b) re-examined holotype of P. schweinfurthii and transferred the genus to Parmulariaceae
based on morphology. Placoasterella is characterized by
superficial, circular, flat, black, carbonaceous ascostromata
with small, solitary, subglobose locules, lacking pseudoparaphyses, ovoid asci and ellipsoid to fusoid ascospores with
one septum (Hongsanan et al. 2014b).
Placosoma Syd., Annls mycol. 22(3/6): 302 (1924).
Index Fungorum number: IF 4139; Facesoffungi number:
FoF 08118; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Placosoma nothopanacis Syd., Annls
mycol. 22(3/6): 303 (1924).
Notes: Placosoma was introduced by Sydow (1924) with
a single species. Prasad et al. (1961) described a second
species P. salvadorae. Thus, two species are accepted in this
genus. Placosoma was placed in Asterinaceae by Lumbsch
and Huhndorf (2010). However, Hongsanan et al. (2014b)
re-examined the type species P. nothopanacis and suggested
that Placosoma is typical of Parmulariaceae. Placosoma is
characterized by superficial, circular, flattened, blackened,
carbonaceous ascostromata, subglobose to oblong asci and
broadly ellipsoidal ascospores with one septum (Hongsanan
et al. 2014b).
13
126
Placostromella Petr., Sydowia 1(1–3): 9 (1947).
Index Fungorum number: IF 4141; Facesoffungi number:
FoF 08119; – 3 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Placostromella macrospora Petr., Sydowia
1(1–3): 9 (1947).
Notes: Placostromella was intoduced by Petrak (1947d)
with Placostromella macrospora as type species. Later P.
amazonensis and P. castanopsis were transferred to Placostromella (Inácio et al. 2005). Three taxa including the type
are accepted in Placostromella (morphology see Thambugala et al. 2014b; Wijayawardene et al. 2018, 2020). This
genus is characterized by subcuticular, carbonaceous, black,
rounded or broadly ellipsoid ascostromata with a single locule, cylindrical to clavate asci and cylindric-ellipsoidal or
oblong to obovate ascospores (Thambugala et al. 2014b).
Pleiostomellina Bat., J.L. Bezerra & H. Maia, in Batista &
Bezerra, Portug. acta biol., Sér. B 7(4): 373 (1964).
Index Fungorum number: IF 4204; Facesoffungi number:
FoF 08120; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Pleiostomellina pernambucensis Bat., J.L.
Bezerra & Cavalc., in Batista & Bezerra, Portug. acta biol.,
Sér. B 7(4): 374 (1964).
Notes: The monotypic genus was introduced by Batista
and Bezerra (1964). Ariyawansa et al. (2014) referred Pleiostomellina to Parmulariaceae based on its similarities with
Parmularia. Pleiostomellina is characterized by superficial,
flattened carbonaceous ascostromata with numerous locules
in a circle, subglobose asci without a pedicel and 2–3-seriate, ellipsoidal, brown, 1-septate ascospores (Ariyawansa
et al. 2014).
Polycyclina Theiss. & Syd., Annls mycol. 13(3/4): 212
(1915).
Index Fungorum number: IF 4310; Facesoffungi number:
FoF 02327; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Polycyclina rhytismoides (Speg.) Theiss.
& Syd., Annls mycol. 13(3/4): 212 (1915).
≡ Hysterostomella rhytismoides Speg., Boln Acad. nac.
Cienc. Córdoba 11(4): 580 (1889).
Notes: The monotypic genus Polycyclina is parasitic on
living leaves of Lomaria imperalis (drawing see Thambugala
et al. 2014b; Wijayawardene et al. 2018, 2020). Polycyclina
is characterized by solitary, superficial, circular to elliptical
to irregular, disc-like, black, rather flat ascostromata, clavate
to cylindric-clavate asci and ellipsoid to cylindric-ellipsoid,
dark brown, aseptate ascospores (Thambugala et al. 2014b).
Polycyclus Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
Kl., Abt. 1 118: 1542 [82 repr.] (1909).
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Fungal Diversity (2020) 105:17–318
Index Fungorum number: IF 4312; Facesoffungi number:
FoF 02329; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Polycyclus andinus (Pat.) Theiss. & Syd.,
Annls mycol. 13(3/4): 210 (1915).
≡ Hysterostomella andina Pat., in Patouillard & Lagerheim, Bull. Herb. Boissier 3(1): 73 (1895).
Notes: Polycyclus was introduced by von Höhnel (1909a),
however, the type species was selected by Theissen and
Sydow (1915). Inácio and Cannon (2008) compared P. andinus and P. marginalis, and considered they were possibly
conspecific. This genus was reported as pathogenic on living
leaves of ferns (morphology see Inácio and Cannon 2008;
Wijayawardene et al. 2018, 2020). Polycyclus is characterized by superficial, circular to elliptical, disc-like, black, flat
ascostromata with the fissured centre raised up to the wart
shape, and with 1–5 locules disposed as a ring around a
central column, opening by longitudinal to irregular fissures
(Inácio and Cannon).
Protothyrium G. Arnaud, C. r. hebd. Séanc. Acad. Sci.,
Paris 164: 574 (1917).
Index Fungorum number: IF 4395; Facesoffungi number:
FoF 02331; – 3 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Protothyrium salvadorae (Cooke) G.
Arnaud, Annals d’École National d’Agric. de Montpellier,
Série 2 16(1–4): 101 (1918) [1917].
≡ Phyllachora salvadorae Cooke, Grevillea 13(no. 67):
65 (1885).
Notes: Four species are accepted in Protothyrium (Inácio and Cannon 2008). Species were reported as pathogenic
on living leaves (morphology see Inácio and Cannon 2008;
Wijayawardene et al. 2018, 2020). Protothyrium is characterized by superficial, circular or elliptical, flat, black ascostromata, becoming irregular in shape, opening by irregular slits
(Inácio and Cannon 2008).
Pseudolembosia Theiss., Annls mycol. 11(5): 432 (1913).
Index Fungorum number: IF 4448; Facesoffungi number:
FoF 02333; – 4 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Pseudolembosia geographica (Massee)
Theiss., Annls mycol. 11(5): 432 (1913).
≡ Lembosia geographica Massee, Bull. Misc. Inf., Kew:
181 (1899).
Notes: Pseudolembosia was introduced by Theissen
(1913) with P. geographica as the type species. Index Fungorum (2020) listed five epithets, but only four taxa are
accepted by Inácio and Cannon (2008). This genus was
reported as pathogens on living leaves (morphology see
Inácio and Cannon 2008; Wijayawardene et al. 2018, 2020).
Pseudolembosia is characterized by superficial, circular to
Fungal Diversity (2020) 105:17–318
elliptical, disc-like, black, rather flat ascostromata raised up
to Y-shaped to irregular near the center (Inácio and Cannon
2008).
Rhagadolobiopsis Guatim. & R.W. Barreto, Mycologia
106(2): 277 (2014).
Index Fungorum number: IF 802297; Facesoffungi number: FoF 08121; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Rhagadolobiopsis thelypteridis Guatim. &
R.W. Barreto, Mycologia 106(2): 277 (2014).
Notes: Rhagadolobiopsis was erected by Guatimosim
et al. (2014), with a single species, R. thelypteridis. This
species is reported as a pathogen on living leaves of Thelypteris serrata from Brazil (morphology and key of genera of Parmulariaceae on fern see Guatimosim et al. 2014;
Wijayawardene et al. 2018, 2020). Rhagadolobiopsis is characterized by numerous, scattered, black, stromata forming
superficial, ellipsoid tar spot-like colonies on the abaxial
surfaces of leaves, multiloculate ascomata, opening by radiating fissures, cylindrical-clavate to clavate asci and fusiform, aseptate, hyaline ascospores (Guatimosim et al. 2014).
Rhagadolobium Henn. & Lindau, Bot. Jb. 23: 287 (1897).
Index Fungorum number: IF 4688; Facesoffungi number:
FoF 02335; – 11 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Rhagadolobium hemiteliae Henn. & Lindau [as ‘Rhagadolobium hemitheliae’], (1897).
Notes: Rhagadolobium is characterized by superficial, circular or elliptical, flattened, black ascostromata with upper
wall of stromata comprising a thin dark brown tissue (Guatimosim et al. 2014).
Rhipidocarpon (Theiss.) Theiss. & Syd., Annls mycol.
13(3/4): 197 (1915).
≡ Parmularia subgen. Rhipidocarpon Theiss., Annls
mycol. 11(5): 456 (1913).
Index Fungorum number: IF 4703; Facesoffungi number:
FoF 02337; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Rhipidocarpon javanicum (Pat.) Theiss. &
Syd., Annls mycol. 13(3/4): 197 (1915).
≡ Schneepia javanica Pat., Ann. Jard. Bot. Buitenzorg,
suppl. 1: 122 (1897).
Notes: This genus is only known from type location
Indonesia. It was reported as pathogenic on living leaves
of palm (morphology see Inácio and Cannon 2008; Wijayawardene et al. 2018, 2020). Rhipidocarpon is characterized
by solitary to gregarious, superficial, black, irregular, carbonaceous, flat ascostromata with ridges irregularly radiating
from the centre to the outer rim (Inácio and Cannon 2008).
127
Symphaeophyma Speg., Anal. Mus. nac. Hist. nat. B. Aires
23: 97 (1912).
Index Fungorum number: IF 5316; Facesoffungi number:
FoF 02339; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Symphaeophyma subtropicale Speg., Anal.
Mus. nac. Hist. nat. B. Aires 23: 97 (1912).
Notes: This monotypic genus was introduced by
Spegazzini (1912) to accommodate S. subtropicale, collected from a tropical region. This species was reported as
pathogenic on living leaves of Pouteria salicifolia (morphology see Inácio and Cannon 2008; Wijayawardene et al. 2018,
2020). Symphaeophyma is characterized by superficial, circular or elliptical, pulvinate, sinuate, black ascostromata
with thick internal part (Inácio and Cannon 2008).
Syrropeltis Bat., J.L. Bezerra & Matta, in Batista & Bezerra,
Portug. acta biol., Sér. B 7(4): 376 (1964).
Index Fungorum number: IF 5344; Facesoffungi number:
FoF 08122; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Syrropeltis xylopia Bat., J.L. Bezerra
& Matta, in Batista & Bezerra, Portug. acta biol., Sér. B
7(4):377 (1964).
Notes: The monotypic genus was introduced by Batista
and Bezerra (1964), and was treated as Dothideomycetes
genera incertae sedis by Wijayawardene et al. (2014a, 2018).
Doilom et al. (2018) re-examined the holotype of S. xylopia
and transferred Syrropeltis to Parmulariaceae based on its
morphologic similarity to this family. Syrropeltis is characterized by large and black ascostromata on host substrate and
1-septate ascospores that are ovoid with upper cell broader
and longer, slightly constricted and pale brown to brown at
the septum (Doilom et al. 2018).
Thallomyces H.J. Swart, Trans. Br. mycol. Soc. 65(1): 84
(1975).
Index Fungorum number: IF 5395; Facesoffungi number:
FoF 02341; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Thallomyces oritis (Hansf.) H.J. Swart,
Trans. Br. mycol. Soc. 65(1): 85 (1975).
≡ Stigmatea oritis Hansf. [as ‘oritidis’], Proc. Linn. Soc.
N.S.W. 79(3–4): 102 (1954).
Notes: This genus is only known from the type collection,
which was pathogenic on living leaves of Orites lancifolius
(morphology see Inácio and Cannon 2008; Wijayawardene
et al. 2018, 2020). Thallomyces is characterized by superficial, dark brown to black, coriaceous, flat ascomata forming
on branching, brown hyphae, which produce round to irregular, brown to dark brown appressoria (Inácio and Cannon
2008).
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128
Fungal Diversity (2020) 105:17–318
Fig. 62 Phylogram generated
from maximum likelihood
analysis (RAxML) of Patellariales based on ITS, LSU and
SSU sequence data. Maximum
likelihood bootstrap values
equal or above 70%, Bayesian
posterior probabilities equal
or above 0.90 (MLBS/PP) are
given at the nodes. Isolate/
specimen number is noted after
the species name. The tree is
rooted to Lichenothelia calcarea (L1324) and L. convexa
(L1609). The ex-type strains are
indicated in bold. Hyphen (-)
represents support values below
70% MLBS and 0.90 PP
Patellaria chromolaenae MFLUCC 17-1479
Patellaria chromolaenae MFLUCC 17-1482
100/1.00
Patellaria chromolaenae MFLU 16-0578
Patellaria microspora MFLU 16-0567
100/Patellaria quercus CPC 27232
89/Patellaria atrata CBS 958.97
Patellaria atrata BCC 28877
-/0.99
Patellaria atrata BCC 28876
Hysteropatella elliptica G.M. 2013-05-06
100/1.0 Hysteropatella prostii G.M. 2014-05-20
Hysteropatella clavispora CBS 247.34
100/1.0 Glyphium elatum EB 0388
Glyphium elatum EB 0342
Patellariaceae
Yuccamyces pilosus CBS 579.92
99/1.0
Yuccamayces citri CBS 143161
Coniosporium apollinis CBS 100213
100/1.0
-/1.0
Coniosporium apollinis CBS 352.97
Phaeococcomyces rothmanniae CPC 22668
Pseudoarthrographis phlogis CPC 32759
80/1.0 Holmiella junipericola TASM 6136
99/1.0
Holmiella sabina G.M. 2015-04-29 2
Holmiella juniper-semiglobosae MFLUCC 17-1955
Spissiomyces ramosus CGMCC 3.17075
Cryomyces minteri CCFEE 5187
Incertae sedis
97/1.0
Cryomyces antarcticus CCFEE 536
100/1.0 Abrothallus buellianus SPO303
100/1.0
Abrothallus pareliarum AB36
Abrothallus cladoniae AB53
Lichenothelia calcarea L1324
Outgroup
Lichenothelia convexa L1609
100/1.00
Patellariales
100/1.0
Abrothallales
100/1.0
0.04
Viegasella Inácio & P.F. Cannon, Mycol. Res. 107(1): 82
(2003).
Index Fungorum number: IF 28709; Facesoffungi number: FoF 02343; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Viegasella pulchella (Speg.) Inácio & P.F.
Cannon, Mycol. Res. 107(1): 83 (2003).
≡ Schneepia pulchella Speg., Anal. Soc. cient. argent.
26(1): 55 (1888).
Notes: Viegasella was revised by Inácio and Cannon
(2003) with description and illustration. It was reported as
pathogen on living leaves in Brazil (morphology see Inácio
and Cannon 2003,2008; Tian et al. 2015; Wijayawardene
et al. 2018, 2020). Viegasella is characterized by gregarious
to solitary, black ascostromata with multi-ascomata gregarious and star-shaped, olive-cream, clavate to cylindrical asci
and brown, ellipsoid, 1-septate ascospores (Inácio and Cannon 2003,2008; Tian et al. 2015).
Economic and ecological significance
Parmulariaceae is a large family, usually distributed in
tropical regions. Taxa in this family are usually epiphytic or
pathogenic on fern fronds or plant leaves, occasionally on
thallus of lichens (Inácio and Cannon 2008; Guatimosim
13
et al. 2015). Some genera, such as Aulacostroma are reported
as plant pathogens which can cause leaf spots.
Patellariales D. Hawksw. & O.E. Erikss.
Index Fungorum number: IF 90491; Facesoffungi number: FoF 08123.
Species of the Patellariales are saprobes on bark or wood,
although some are lichenized (Jones et al. 2015). Ascomata
are apothecial. Pseudoparaphyses are initially attached to
both the base and apex, but later become free at the apex.
Asci are cylindric-clavate and ascospores are mostly fusoid
(Kutorga and Hawksworth 1997). Some species have aposphaeria-like and diplodia-like asexual morphs (Yacharoen et al. 2015), but these are rarely observed. Species are
frequently found in terrestrial habitats and occasionally in
aquatic habitats (Kirk et al. 2008; Jones et al. 2015). Multigene phylogenetic study on this group was provided by Pem
et al. (2018). Phylogenetic placement of genera in this order
are shown in Fig. 62. The divergence time for Patellariales
is estimated as 276 MYA (stem age, Hongsanan et al. 2020).
Accepted families: Patellariaceae.
Patellariaceae Corda, Icon. fung. (Prague) 2: 37 (1838).
Fungal Diversity (2020) 105:17–318
129
Fig. 63 Patellaria atrata
(MFLU 18-1818). a Substrate.
b Apothecia on wood. c Cross
section of an apothecium, d
Filiform paraphyses. e–g, h
Cylindrical asci. i Clavate
ascospores. Scale bars: c = 100
µm, d = 50 µm, e–h = 75 µm, i
= 10 µm
Index Fungorum number: IF 81111; Facesoffungi number: FoF 00342, 143 species.
Sexual morph: Apothecia sessile, black, circular, humidity sensitive and turn into hysterothecia when dry. Ectal
excipulum composed of carbonized pseudoparenchymatous
tissue. Hamathecium comprising paraphysoids or paraphyses, their apices are branched, pigmented and covered by
dark exudates. Asci cylindrical, fissitunicate and arising from
croziers. Ascospores overlapping 2–3-seriate, cylindrical,
ellipsoid, fusiform, or clavate, hyaline or brown, with septa
(Kutorga and Hawksworth 1997; Yacharoen et al. 2015).
Asexual morph: Coelomycetous (Yacharoen et al. 2015).
Type: Patellaria Fr.
Notes: Patellariaceae was introduced by Corda (1838).
The family includes 21 genera and around 52 species (Jaklitsch et al 2016; Wijayawardene et al. 2017a). This is a
widely distributed family mostly found on bark or wood as
saprobes (Kutorga and Hawksworth 1997; Yacharoen et al.
2015; Jaklitsch et al 2016).
Patellaria Fr., Syst. mycol. (Lundae) 2(1): 158 (1822).
Index Fungorum number: IF 3765; Facesoffungi number:
FoF 00343; 48 morphological species (Species Fungorum
2020; this study), 4 species with molecular data
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Fungal Diversity (2020) 105:17–318
Fig. 64 Patellaria chromolaenae (MFLU 16-0578). a Substrate. b
Apothecia on wood. c Apothecium on wood, d: Cross section of an
apothecium (in lacto glycerine). e Vertical section of the apothecium
at margin (in lacto glycerine). f Apically enlarged paraphyses. g–j
Cylindrical asci, k–n allantoid ascospores. o germinating spore. Scale
bars: b = 500 µm, c = 200 µm, d = 300 µm, e = 70 µm, f = 40 µm,
g–j = 20 µm, k–n = 15 µm, o = 100 µm
Type species: Patellaria atrata (Hedw.) Fr.
Notes: This genus was introduced by Fries (1822)
and is characterized by black or dark coloured, sessile or
sub-pedicellate apothecia which appear to be hysterothecia
when dry, filiform paraphyses with branched, pigmented apices, cylindrical, fissitunicate asci which arise from croziers,
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and hyaline, brown or greenish ascospores (Kutorga and
Hawksworth 1997; Yacharoen et al. 2015).
Patellaria atrata (Hedw.) Fr., Syst. mycol. (Lundae) 2(1):
158 (1822).
≡ Lichen atratus Hedw., Descr. micr.-anal. musc. frond.
(Lipsiae) 2: 61 (1789).
Index Fungorum number: IF 398671; Facesoffungi number: FoF 00344; Fig. 63
Saprobic on dead stems. Sexual morph: Apothecia
600–1000 × 250–350 µm ( x̄ = 700 × 270 µm, n = 10) arising singly or in small groups, sessile, erumpent from the
substrate, black when fresh. Receptacle pulvinate, disc convex, disc and the margins are black in when fresh. Ectal
excipulum 20–25 µm ( x̄ = 22.7 µm, n = 10) a narrow layer,
3–4 cells deep, thick, black cells of subglobose to subangularis. Medullary excipulum 10–15 µm ( x̄ = 13.5 µm, n = 10)
composed of narrow, long, thin-walled, hyaline to brownish
cells of textura porrecta. Hymenium brownish to greenish.
Hamathecium comprising 2–3 µm wide ( x̄ = 2.7 µm, n =
20) at the apex, numerous, filiform, septate, branched and
swollen at the apex, apices glued together to form epithecium, paraphyses. Epithecium dark green. Asci 75–120 ×
12–20 µm ( x̄ = 101.5 × 17.3 µm, n = 30) 8-spored, short
to medium pedicellate, cylindric to clavate, rounded at the
apex, J-, croziers abscent at the asci base. Ascospore 25–30
× 6–10 µm ( x̄ = 28.1 × 6.8 µm, n = 40), partially bi-seriate,
upper and lower spores are uni-seriate, sometimes multiseriate, hyaline, smooth-walled, ellipsoid, 6–7-septate, distinctly more tapered towards the distal end. Asexual morph:
Undetermined.
Material examined: UK, New Forest, on Ulex wood, 9
June 2015, E. B. G. Jones, GJ183 (MFLU 18-1818).
Notes: Our collection failed in ascospore germination and
DNA extraction. However, characteristics of our collection
are similar to the description of Patellaria atrata provided
by Yacharoen et al. (2015).
Patellaria chromolaenae Mapook & K.D. Hyde, in Mapook
et al., Fungal Diversity 101: 127 (2020).
Index Fungorum number: IF 557350; Facesoffungi number: 07836; Fig. 64
Saprobic on dead stems. Sexual morph: Apothecia
622–693 × 108–182 µm ( x̄ = 652.2 × 150.4 µm, n = 10)
arising singly or in small groups, sessile, erumpent from
the substrate, black when fresh. Receptacle pulvinate, disc
convex, disc and the margins are greenish black when fresh,
in lacto glycerin colour change in to magenta. Excipulum
18–32 µm ( x̄ = 25.1 µm, n = 10) ectal excipulum and medullary excipulum are not clearly separable, ectal excipulum
a narrow layer, 3–4 cells deep, thick, greenish black colour
cells of subglobose to subangularis, medullary excipulum
composed of narrow, long, thin-walled, hyaline to greenish
131
cells of textura porrecta. Hymenium hyaline or greenish.
Hamathecium comprising 1.2–1.8 µm wide ( x̄ = 1.5 µm,
n = 20), numerous, obtuse, exceed asci in length, apically
enlarged and pigmented, apices glued together to form
pseudo epithecium, paraphyses. Asci 60–80 × 15–20 µm ( x̄
= 69.3 × 17.1 µm, n = 30) 8-spored, short pedicellate, cylindric–globose, rounded at the apex, J –at the ascus apex, croziers abscent at the base of asci. Ascospore 27–34 × 5.2–6.0
µm ( x̄ = 31.5 × 5.8 µm, n = 40), multi-seriate, hyaline to
greenish, smooth-walled, amerosporae, allantoid, regularly
seven septate. Asexual morph: Undetermined
Material examined: Thailand, Chiang Rai Province, Mae
Fah Luang University, 21 March 2015, A. H. Ekanayaka, HD
016, (MFLU 16-0578).
GenBank number: ITS: MW136695, LSU: MW142387,
SSU: MW127179
Notes: Patellaria chromolaenae is originally identified
from dead stems of Chromolaenae odorata. The dead stem
of the present collection was not identified. In the phylogenetic analysis (Fig. 62), our collection grouped with the holotype of Patellaria chromolaenae with strong bootstrap support. In our collection of P. chromolaenae (MFLU 16-0578),
we observed slightly larger apothecia and the ascospores
with 7 septa which was not observed in the holotype collection (Mapook et al. 2020).
Patellaria microspora Ekanayaka & K.D. Hyde, sp. nov.
Index Fungorum number: IF 557819; Facesoffungi number: FoF 08125; Fig. 65
Etymology: The specific epithet microspora refers small
ascospores.
Holotype: MFLU 16-0567.
Saprobic on dead stems. Sexual morph: Apothecia
396–402 × 165–170 µm ( x̄ = 395.4 × 167.9 µm, n = 10)
arising singly or in small groups, sessile, erumpent from
the substrate, black when fresh. Receptacle turbinate, disc
slightly convex, disc and the margins are black when fresh.
Ectal excipulum 15–21 µm ( x̄ = 16.7 µm, n = 10) a narrow
layer, 3–4 cells deep, thick, black colour cells of subglobose to subangularis. Medullary excipulum 22–30 µm ( x̄ =
27.1 µm, n = 10) composed of narrow, long, thin-walled,
hyaline to brownish cells of textura porrecta. Hymenium
brownish. Hamathecium 1.3 – 1.8 µm wide ( x̄ = 1.5 µm, n =
20), numerous, filiform towards the apex, branched, septate,
paraphyses. Asci 27–38 × 6.2–7.0 µm ( x̄ = 30.6 × 6.8 µm,
n = 30), 8-spored, short to medium pedicellate, cylindric to
clavate, rounded at the apex, asci become orange to brown
with the presence of Meltzer’s reagent, croziers absent at
the base of asci. Ascospores 19.2–20.3 × 8.7–9.5 µm ( x̄ =
19.88 × 9.21 µm, n = 40), partially bi-seriate, lower spores
are uni-seriate, hyaline, smooth-walled, ellipsoid, regularly
eight septate, distinctly more tapered towards the distal end.
Asexual morph: Undetermined
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132
Fig. 65 Patellaria microspora (MFLU 16-0567, holotype). a Substrate. b Apothecia on wood. c Apothecium on wood. d Cross section
of an apothecium. e Vertical section of the apothecium at margin. f
Ectal excipular hypha. g–i Cylindrical asci. j Reaction of the ascus
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in the Meltzer’s reagent. k Septate and branched paraphyses. l Germinated spore. m–p clavate ascospores. Scale bars: b = 500 µm, c =
200 µm, d = 100 µm, e, i, j = 50 µm, f = 40 µm, g, h, l = 30 µm, k,
m–p = 20 µm
Fungal Diversity (2020) 105:17–318
Material examined: UK, Lulworth Cove, Dorset, 5 February 2015, E. B. G. Jones, GJ95082 (MFLU 16-0567,
holotype).
GenBank number: ITS: MW136696, LSU: MW142388.
Note: Patellaria microspora is phylogenetically related to
P. atrata. However, P. atrata has larger ascospores (Yacharoen et al. 2015).
Other genera included
Baggea Auersw., Hedwigia 5: 1 (1866).
Index Fungorum number: IF 491; Facesoffungi number:
FoF 00345; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Baggea pachyascus Auersw., Hedwigia 5:
1 (1866).
Note: Baggea is characterized by apothecial or hysterothecia-like ascomata, polysporous asci and allantoid
ascospores (Yacharoen et al. 2015).
Banhegyia L. Zeller & Tóth, Sydowia 14: 326 (1960).
Index Fungorum number: IF 509; Facesoffungi number:
FoF 00601; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Banhegyia setispora L. Zeller & Tóth,
Sydowia 14: 327 (1960).
Note: Banhegyia is characterized by apothecia-like ascomata, bitunicate asci and epithecium formed from pseudoparaphyses, and 8-spored, bitunicate, subglobose to obovoid
asci (Yacharoen et al. 2015).
Colensoniella Hafellner, Beihefte zur Nova Hedwigia 62:
160 (1979).
Index Fungorum number: IF 1172; Facesoffungi number:
FoF 08126; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Colensoniella torulispora (W. Phillips)
Hafellner, Beihefte zur Nova Hedwigia 62: 160 (1979).
≡ Patellaria torulispora W. Phillips, Grevillea 15(no. 73):
16 (1886).
Note: Colensoniella is characterized by superficial
apothecia with raised margins, 8-spored asci, and oblong
ascospores which break into two part-spores at maturity
(Tian et al. 2014).
Endotryblidium Petr., Sydowia 13 (1–6): 244 (1959).
Index Fungorum number: IF 1812; Facesoffungi number:
FoF 00347; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Endotryblidium insculptum (Cooke) Petr.,
Sydowia 13 (1–6): 245 (1959).
≡ Triblidium insculptum Cooke [as ‘Tryblidium’], Grevillea 4(no. 32): 182 (1876)
133
Note: Endotryblidium is characterized by apothecia
aggregated on the host, a pseudoparenchymatous peridium, 4–6-, rarely 8-spored, short pedicellate, clavate asci,
numerous filamentous paraphyses and oblong to ellipsoidal
ascospores (Yacharoen et al. 2015).
Glyphium Nitschke ex F. Lehm., Nova Acta Acad. Caes.
Leop.-Carol. German. Nat. Cur.: 139 (1886).
Index Fungorum number: IF 2095 Facesoffungi number:
FoF 08127; – 6 morphological species (Species Fungorum
2020), 2 species with molecular data.
Type species: Glyphium dolabriforme (Wallr.) F. Lehm.:
139 (1886).
≡ Lophium dolabriforme Wallr., Fl. crypt. Germ. (Norimbergae) 2: 433 (1833).
Note: Glyphium is characterized by erect, carbonaceous
ligulate to dolabrate ascomata that are strongly laterally
compressed and dehisce along a longitudinal slit (Boehm
et al. 2015).
Haematomyxa Sacc., Botanisches Centralblatt 18: 250
(1884).
Index Fungorum number: IF 2198; Facesoffungi number:
FoF 08128; – 4 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Haematomyxa vinosa (Cooke & Ellis)
Sacc., Sylloge Fungorum 8: 646 (1889).
≡ Haematomyces vinosus Cooke & Ellis, Grevillea 4(no.
32): 179 (1876).
Note: Haematomyxa is characterized by subglobose,
gelatinous apothecia, broadly clavate, 8-spored asci, and
muriform, brown ascospores (Seaver 1951).
Holmiella Petrini, Samuels & E. Müll., Berichte der Schweizerischen Botanischen Gesellschaft 89: 83 (1979).
Index Fungorum number: IF 2344; Facesoffungi number:
FoF 00349; – 4 morphological species (Species Fungorum
2020), 3 species with molecular data.
Type species: Holmiella sabina (De Not.) Petrini, Samuels & E. Müll., Berichte der Schweizerischen Botanischen
Gesellschaft 89: 84 (1979).
≡ Triblidium sabinum De Not. [as ‘Tryblidium’], Comm.
Soc. crittog. Ital. 2(fasc. 3): 491 (1867).
Note: Holmiella is characterized by globose, black apothecia, filamentous, septate pseudoparaphyses, bitunicate,
clavate asci and clavate to ellipsoidal, 1-septate ascospores
(Yacharoen et al. 2015).
Hysteropatella Rehm, Rabenhorst’s Kryptogamen-Flora,
Pilze - Ascomyceten 1(3): 367 (1890).
Index Fungorum number: IF 2469; Facesoffungi number:
FoF 00351; – 7 morphological species (Species Fungorum
2020), 3 species with molecular data.
13
134
Type species: Hysteropatella prostii (Duby) Rehm,
Rabenhorst’s Kryptogamen-Flora, Pilze - Ascomyceten 1(3):
367 (1890).
≡ Hysterium prostii Duby, Mém. Soc. Phys. Hist. nat.
Genève 16(1): 38 (1861) [1862].
Note: Hysteropatella is characterized by hysterothecial
ascomata, carbonaceous excipulum, bitunicate asci and reniform to ellipsoid or fusoid, 3-septate ascospores (Yacharoen
et al. 2015).
Hysteropeltella Petr., Annales Mycologici 21 (1–2): 9
(1923).
Index Fungorum number: IF 2470; Facesoffungi number:
FoF 00352; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Hysteropeltella moravica Petr., Annales
Mycologici 21 (1–2): 10 (1923).
Note: Hysteropeltella is characterized by apothecial ascomata, bitunicate asci and hyaline ascospores with a sheath
(Yacharoen et al. 2015).
Lahmiomyces Cif. & Tomas., Atti Ist. bot. Univ. Lab. crittog. Pavia, sér. 5 10: 39, 66 (1953).
Index Fungorum number: IF 2632; Facesoffungi number:
FoF 08129; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Lahmiomyces piceae Cif. & Tomas., Atti
Ist. bot. Univ. Lab. crittog. Pavia, sér. 5 10(2–3): 270 (1953).
Note: Jaklitsch et al. (2016) treated Lahmiomyces as a
genus in Patellariaceae.
Lecanidiella Sherwood, Sydowia 38: 272 (1986).
Index Fungorum number: IF 25053; Facesoffungi number: FoF 00353; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Lecanidiella contortae Sherwood, Sydowia
38: 274 (1986).
Note: Lecanidiella is characterized by apothecia,
erumpent, black, serrated at the rim, a brown and powdery epithecium, bitunicate asci, and 3-septate ascospores
(Yacharoen et al. 2015).
Lirellodisca Aptroot, Nova Hedwigia 67: 485 (1998).
Index Fungorum number: IF 27901; Facesoffungi number: FoF 00355; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Lirellodisca pyrenulispora Aptroot, Nova
Hedwigia 67: 485 (1998).
Note: Lirellodisca is characterized by subglobose, deeply
cup-shaped apothecia, filamentous, individual, apically
branched, hyaline pseudoparaphyses and thick-walled distoseptate ascospores (Yacharoen et al. 2015).
13
Fungal Diversity (2020) 105:17–318
Murangium Seaver, The North American Cup-fungi
(Inoperculates) (3): 367 (1951).
Index Fungorum number: IF 3289; Facesoffungi number:
FoF 00357; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Murangium sequoiae (Plowr.) Seaver, The
North American Cup-fungi (Inoperculates) (3): 368 (1951).
≡ Cenangium sequoiae Plowr., Grevillea 7(no. 41): 23
(1878).
Note: Murangium is characterized by cup-shaped apothecia, bitunicate asci and muriform ascospores (Yacharoen
et al. 2015).
Poetschia Körb., Parerga lichenologica. Ergänzungen zum
Systema lichenum Germaniae: 280 (1861).
Index Fungorum number: IF 4289; Facesoffungi number:
FoF 00359; – 4 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Poetschia buellioides Körb., Parerga
lichenologica. Ergänzungen zum Systema lichenum Germaniae: 280 (1861).
Note: Poetschia is characterized by globose, black apothecial ascomata, pseudoparenchymatous excipulum, obovoid,
clavate asci and ellipsoidal to obovoid, 1-septate ascospores
(Yacharoen et al. 2015).
Pseudoparodia Theiss. & Syd., Annls mycol. 15 (1–2): 138
(1917).
Index Fungorum number: IF 4466; Facesoffungi number:
FoF 00377; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Pseudoparodia pseudopeziza (Pat.) Theiss.
& Syd., Sydowia 1 (4–6): 169 (1947).
≡ Parodiella pseudopeziza Pat., in Patouillard & Lagerheim, Bull. Herb. Boissier 3(1): 67 (1895).
Note: Pseudoparodia is characterized by apothecia which
are superficial and swollen, with dense pseudoparaphyses
forming epithecium above the asci (Yacharoen et al. 2015).
Rhizodiscina Hafellner, Beihefte zur Nova Hedwigia 62:
195 (1979).
Index Fungorum number: IF 4709; Facesoffungi number:
FoF 00361; – 2 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Rhizodiscina lignyota (Fr.) Hafellner, Beihefte zur Nova Hedwigia 62: 195 (1979).
≡ Peziza lignyota Fr., Syst. mycol. (Lundae) 2(1): 150
(1822).
Note: Rhizodiscina is characterized by apothecial ascomata, massive exciple and bitunicate asci (Yacharoen et al.
2015).
Fungal Diversity (2020) 105:17–318
-/1.0
Phaeotrichum benjaminii CBS 541.72
Phaeotrichum setosum CBS 151.77
100/1.0
Trichodelitschia bisporula CBS 262.69
Phaeotrichaceae
100/1.0
100/1.0
Trichodelitschia munkii Kruys 201 (UPS)
100/1.0
Zeloasperisporium hyphopodioides CBS 218.95
Zeloasperisporium wrightiae MFLUCC 15-0225
Zeloasperisporiaceae
Zeloasperisporium siamense IFRDCC 2194
Zeloasperisporiales
100/1.0
Phaeotrichales
Fig. 66 Phylogram generated
from maximum likelihood
analysis (RAxML) of Phaeotrichales based on LSU and
SSU sequence data. Maximum
likelihood bootstrap values
equal or above 70%, Bayesian
posterior probabilities equal
or above 0.90 (MLBS/PP) are
given at the nodes. Isolate/specimen number is noted after the
species name. The tree is rooted
to Venturia inaequalis (CBS
815.69 and CBS 594.70). The
ex-type strains are indicated in
bold. Hyphen (-) represents support values below 70% MLBS
and 0.90 PP
135
Venturia inaequalis CBS 815.69
Venturiales (Outgroup)
1001.0
Venturia inaequalis CBS 594.70
0.02
Rimula Velen., Monographia Discomycetum Bohemiae: 38
(1934).
Index Fungorum number: IF 4756; Facesoffungi number:
FoF 08130; – 28 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Rimula faginea Velen., Monogr. Discom.
Bohem. (Prague): 38 (1934).
Note: Jaklitsch et al. (2016) treated Rimula as a genus in
Patellariaceae.
Schrakia Hafellner, Nova Hedwigia, Beihefte zur 62:
204(1979).
Index Fungorum number: IF 4917; Facesoffungi number:
FoF 00363; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Schrakia crassula (Starbäck) Hafellner,
Nova Hedwigia, Beihefte zur. 62: 204 (1979).
≡ Karschia crassula Starbäck, Bih. K. svenska VetenskAkad. Handl., Afd. 3 25(no. 1): 10 (1899).
Note: The characters of this genus are typical of Patellariaceae and characterised by carbonaceous apothecia and
exciple with thick-walled and pseudoparenchymatous cells
and bitunicate asci (Yacharoen et al. 2015).
Stratisporella Hafellner, Beihefte zur Nova Hedwigia 62:
207 (1979).
Index Fungorum number: IF 20613; Facesoffungi number: FoF 00365; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Stratisporella episemoides (Nyl.) Hafellner, Beih. Nova Hedwigia 62: 207 (1979).
≡ Lecidea episemoides Nyl., Bull. Soc. linn. Normandie,
sér. 2 2: 515 (1868).
Note: Taxa form apothecioid ascomata and are characterized by carbonaceous excipulum, unitunicate asci and 1-septate ascospores with thick exospores (Yacharoen et al. 2015).
Tryblidaria (Sacc.) Rehm, Hedwigia 42(Beibl.): (172)
(1903).
≡ Blitridium subgen. Tryblidaria Sacc., Syll. fung. (Abellini) 8: 805 (1889).
Index Fungorum number: IF 5620; Facesoffungi number:
FoF 00367; – 27 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Tryblidaria fenestrata (Cooke & Peck)
M.E. Barr, Bulletin of the New York State Museum 459:
22 (1986).
≡ Patellaria fenestrata Cooke & Peck, Ann. Rep. N.Y. St.
Mus. nat. Hist. 28: 68 (1876) [1875].
Note: Tryblidaria taxa are saprobic and mostly found in
terrestrial, subtropical environments. Sexual morphs are
apothecial cupulate and carbonaceous (Yacharoen et al.
2015). Asexual morphs are undetermined (Yacharoen et al.
2015).
Economic and ecological significance
Patellariaceae is a widely distributed family. Most Patellariaceae taxa are saprobes and weak parasites on a wide variety
of plants. Few members are recorded on lichen thalli. Rarely
reported endophytic and fungicolous life styles are also
known. For example, Rhizodiscina lignyota was recorded
from Basidiomycetes and Holmiella sabina was recorded
from living needles and wood of Juniperus spp. (Kutorga
and Hawksworth 1997).
Phaeotrichales Ariyaw., Jian K. Liu & K.D. Hyde, in Hyde
et al.
13
136
Fungal Diversity (2020) 105:17–318
or arranged in little groups or bundles and evanescent at
maturity. Ascospores overlapping, dark brown to reddish
brown, sometimes fragmenting, composed of terminal germ
pores, with or without a sheath. Asexual morph: Unknown
(adapted from Hyde et al. 2013).
Type: Phaeotrichum Cain & M.E. Barr.
Notes: Phaeotrichaceae was introduced by Cain (1956)
to accommodate a single genus Phaeotrichum, characterized
by dark brown, septate ascospores with terminal germ pores.
Two coprophilous species were described, P. hystricinum
(type species) and P. circinatum from porcupine dung and
lemming dung, respectively. Cain (1956) pointed out some
morphological similarities of Phaeotrichum with Trichodelitschia species based on features of the ascospores and setose
ascomata. Lundqvist (1964) assigned Trichodelitschia to
Phaeotrichaceae. The family comprises three genera, Echinoascotheca, Phaeotrichum and Trichodelitschia (Zhang
et al. 2012; Hyde et al. 2013).
Fig. 67 Phaeotrichum hystricinum (redrawn from Cain 1956).
a Ascomata with black appendages. b, c Appendages. d Black and
opaque basal appendages. e, f Asci. g Ascospores
Index Fungorum number: IF 805301; Facesoffungi number: FoF 08131.
Phaeotrichales was introduced by Hyde et al. (2013)
to accommodate Phaeotrichum as the type genus. Phaeotrichales is considered an order of coprophilous fungi
within the class Dothideomycetes. Species in this order are
characterized by unilocular, stromatic, globose ascomata
with a thin, carbonaceous peridium and dark brown to reddish brown ascospores with terminal germ pores (Hyde et al.
2013). According to the recent outline of Wijayawardene
et al. (2018), only a single family (Phaeotrichaceae) is
placed in Phaeotrichales. Phylogenetic placement of genera
in this order are shown in Fig. 66. The divergence time for
Phaeotrichales is estimated as 229 MYA (stem age, Hongsanan et al. 2020).
Accepted families: Phaeotrichaceae.
Phaeotrichaceae Cain, Can. J. Bot. 34: 676 (1956).
Index Fungorum number: IF 81144; Facesoffungi numbers: FoF 08131, 11 species.
Saprobic (coprophilous) in terrestrial habitats. Sexual
morph: Ascomata superficial, solitary, or in small groups,
dark brown to black, globose, stromatic, perithecial or cleistothecial setose. Peridium thin-walled, carbonaceous,
membraneous. Hamathecium comprising tissue absent or
of evanescent cellular pseudoparaphyses. Asci 8-spored,
bitunicate, fissitunicate, clavate, pedicellate, in irregular
13
Phaeotrichum Cain & M.E. Barr, Can. J. Bot. 34: 676
(1956).
Index Fungorum number: IF 3967; Facesoffungi number:
FoF 08132; 5 morphological species (Species Fungorum
2020), 4 species with molecular data.
Type species: Phaeotrichum hystricinum Cain & M.E.
Barr.
Notes: Phaeotrichum was introduced by Cain (1956) to
accommodate two coprophilous species, P. hystricinum and
P. circinatum. Phaeotrichum species are characterized by
dark brown, septate ascospores with terminal germ pores,
superficial cleistothecial ascomata covered by long hairy
appendages and a coprophilous habitat (Cain 1956). Phaeotrichum contains only five species (Index Fungorum 2020).
Four species occur in dung and one from soil: P. benjaminii (on dung of Rodentia), P. circinatum (on dung of Lemmini), P. cylindrosporum (on dung of Erythizon dorsatus),
P. hystricinum (on dung of porcupine) and P. setosum (from
soil) (Fig. 67).
Other genera included:
Echinoascotheca Matsush., Matsush. Mycol. Mem. 8: 19
(1995).
Index Fungorum number: IF 27584; Facesoffungi number: FoF 08133; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Echinoascotheca duplooformis Matsush.,
Matsush. Mycol. Mem. 8: 19 (1995).
Notes: Echinoascotheca is a monotypic genus (Species
Fungorum 2020). Trichodelitschia species differ from Echinoascotheca in having an apical ring in asci (Hyde et al.
2013). For more details on morphology of Echinoascotheca
species see Matsushima (1995).
Fungal Diversity (2020) 105:17–318
100/1.0
73/-
Stigmatodiscus tamaricis L113
Stigmatodiscus tamaricis L114
Stigmatodiscus oculatus AP161116
100/1.0
Stigmatodiscus enigmaticus L69
82/1.0
100/1.0
Stigmatodiscus pruni L167
Stigmatodiscales
97/1.0
Muyocopron garethjonesii MFLU 16-2664a
100/0.98
Muyocopron dipterocarpi MFLUCC 14-1103
Acrospermum compressum M151
99/1.0
Acrospermum gramineum M152
Muyoco- Acrosperpronales matales
Fig. 68 Phylogram generated
from maximum likelihood
analysis (RAxML) of Stigmatodiscales based on ITS, LSU,
SSU and tef1 sequence data.
Maximum likelihood bootstrap
values equal or above 70%,
Bayesian posterior probabilities
equal or above 0.90 (MLBS/PP)
are given at the nodes. Isolate/
specimen number is noted after
the species name. The tree is
rooted to Jahnula aquatica
(R68-1) and J. sangamonensis
(F81-1). The ex-type strains are
indicated in bold. Hyphen (-)
represents support values below
70% MLBS and 0.90 PP
137
Jahnula sangamonensis F81-1
100/1.0
Jahnulales (Outgroup)
Jahnula aquatica R68-1
0.06
Trichodelitschia Munk, Dansk bot. Ark. 15(no. 2): 109
(1953).
Index Fungorum number: IF 5552; Facesoffungi number:
FoF 08134; – 5 morphological species (Species Fungorum
2020), 2 species molecular data.
Type species: Trichodelitschia bisporula (P. Crouan &
H. Crouan) Munk, Dansk bot. Ark. 15(no. 2): 109 (1953).
≡ Hormospora bisporula P. Crouan & H. Crouan, Florule
Finistère (Paris): 21 (1867).
Notes: Trichodelitschia was erected by Munk (1953) to
accommodate Trichodelitschia bisporula, which was previously known as Hormospora bisporula. Initially, Trichodelitschia was placed in Sporormiaceae by Munk (1957), but
later Lundqvist (1964) suggested it be transferred to Phaeotrichaceae. Trichodelitschia species are characterized by
having dark thick-walled spiny appendages, dark two-celled
ascospores with hyaline to subhyaline tenninal apophyses
and circular germ pores at the spore apices (Ebersohn and
Eicker 1992). Five Trichodelitschia species are accepted in
Species Fungorum (2020), i.e. T. adelphica, T. bisporula,
T. lundqvistii, T. microspora, and T. munkii.
Economic and ecological significance
Most Phaeotrichaceae species have been recorded on dung
of various animals as saprobes.
Stigmatodiscales Voglmayr & Jaklitsch, in Voglmayr, Gardiennet & Jaklitsch.
Index Fungorum number: IF 815325; Facesoffungi number: FoF 08771.
This order was established by Voglmayr et al. (2016)
to accommodate a single family, Stigmatodiscaceae. Two
genera (Asterodiscus and Stigmatodiscus) were included
in the family based on differences in ascomatal shape and
hyaline versus brown ascospores. However, Voglmayr and
Amengual (2018) synonymized Asterodiscus under Stigmatodiscus. Our phylogenetic tree (Fig. 68) indicates the same
result as in Voglmayr and Amengual (2018) which supports
Stigmatodiscales as a distinct order in Dothideomycetes. The
divergence time for Stigmatodiscales is estimated as 114
MYA (stem age, Hongsanan et al. 2020).
Accepted families: Stigmatodiscaceae.
Stigmatodiscaceae Voglmayr & Jaklitsch, in Voglmayr
et al., Fungal Diversity 80: 275 (2016): 77 (1903).
Index Fungorum number: IF 815326; Facesoffungi numbers: FoF 08772, 6 species species.
On dead twigs. Sexual morph: Ascomata apothecioid,
embedded in cortex of dead twigs, dark brown to black,
without distinct margin. Hamathecium comprising septate,
unbranched or rarely branched and anastomosing above,
with swollen free apical ends, paraphyses, embedded in a
rubber-like gel matrix and covered by an epithecium. Hymenial gel I-. Asci 8-spored, bitunicate, fissitunicate, J-, variable in shape from fusoid to saccate or clavate to broadly
fusiform broadly, with thin ecto- and thick endotunica, with
wide ocular chamber, without a ring. Ascospores 2–3-seriate,
13
138
Fungal Diversity (2020) 105:17–318
Fig. 69 Stigmatodiscus enigmaticus (redrawn from Voglmayr et al.
2016). a Ascomata erumpent from bark in face view. b Paraphyses.
c Asci, d–f Ascospores. g Phialides producing conidia. h Falcate
conidia. Notes: a, b, d–f from WU 35914 (holotype), c from WU
35925, g from WU 35925, h fromWU 35922
slightly to strongly asymmetric, broadly fusiform, upper cell
broader, with broadly rounded, hyaline to brown, 1-euseptate
when immature, with 2 distosepta at maturity, slightly to
strongly constricted at the septum, particularly the middle
septum, surrounded by a separate gelatinous sheath. Asexual
morph: Coelomycetous. Conidiomata in nature immersed,
peridermal, pycnidial (adapted from Voglmayr et al. 2016).
Type: Stigmatodiscus Voglmayr & Jaklitsch.
Notes: Voglmayr et al. (2016) established Stigmatodiscaceae to accommodate two genera, Asterodiscus and Stigmatodiscus, with two new species, Asterodiscus tamaricis
and Stigmatodiscus enigmaticus based on phylogenetic
placement and morphological characters. However further
phylogenetic analyses in Voglmayr and Amengual (2018)
with additional strains indicated that Asterodiscus should
be synonymized under Stigmatodiscus. Thus, Stigmatodiscaceae currently comprises only Stigmatodiscus.
13
Stigmatodiscus Voglmayr & Jaklitsch, in Voglmayr et al.,
Fungal Diversity 80: 278 (2016).
Index Fungorum number: IF 815327; Facesoffungi number: FoF 01654; 6 morphological species (Species Fungorum 2020), 6 species with molecular data.
Fungal Diversity (2020) 105:17–318
= Asterodiscus Voglmayr, Gardiennet & Jaklitsch, Fungal
Diversity 80: 275 (2016)
Type species: Stigmatodiscus enigmaticus Voglmayr &
Jaklitsch.
Notes: Stigmatodiscus was placed in Stigmatodiscaceae
based on morphology and phylogeny. Four new species were
added to this genus by Voglmayr et al. (2017) and Voglmayr
and Amengual (2018). Asterodiscus was also synonymized
under Stigmatodiscus based on sequence data of A. tamaricis
that formed among other Sigmatodiscus species (Voglmayr
and Amengual 2018). Six species are accepted in this genus
based on both morphology and phylogeny. Sequence data
are available for all Sigmatodiscus species.
Stigmatodiscus enigmaticus Voglmayr & Jaklitsch, in Voglmayr et al., Fungal Diversity 80: 278 (2016).
Index Fungorum number: IF 815328; Facesoffungi number: FoF08773; Fig. 69
Description: see Voglmayr et al. (2016).
Economic and ecological significance
Species in this genus were found on dead twigs and we
assume that they are saprobic.
Strigulales Lücking, M.P. Nelsen & K.D. Hyde.
Index Fungorum number: IF 805302; Facesoffungi number: FoF 08774.
The order was established by Hyde et al. (2013) to accommodate lichenized fungi in Dothideomycetes. Members of
Strigulales differ from most Trypetheliales in having euseptate ascospores with few septa and the lack of secondary
chemistry (Hyde et al. 2013). It also differs from Monoblastiales in having unbranched physes and the macroconidia
producing gelatinous appendages (Hyde et al. 2013). Two
families are accepted in this order. The divergence time for
Strigulales is estimated as 201 MYA (stem age, Hongsanan
et al. 2020).
Accepted families: Strigulaceae, Tenuitholiascaceae.
Strigulaceae Zahlbr. in Engler, Syllabus, Edn 2 (Berlin):
46 (1898).
MycoBank number: MB 81870; Index Fungorum number:
IF 81870; Facesoffungi number: FoF 08853; approximately
80 species (Lücking et al. 2017).
= Phyllobatheliaceae Bitter & F. Schill., Hedwigia,
Beibl. 67: 272 (1927). Type: Phyllobathelium (Müll. Arg.)
Müll. Arg., Flora, Regensburg 73: 195 (1890).
Lichenized on leaves, bark or rocks in terrestrial, chiefly
lowland to montane tropical to subtropical habitats, with
few species extending into temperate regions. Thallus
reduced and ecorticate, often white, to (pseudo-)corticate, grey-green to bright green, sometimes with metallic
appearance. Photobiont Trentepohlia sensu lato (including
139
Cephaleuros and Phycopeltis). Sexual morph: Ascomata
perithecia, scattered, clustered, or aggregated in pseudostromata, immersed to prominent (to rarely sessile), mostly
brown or black, rarely pale, often at least partly covered by
thallus, globose to pear-shaped or conical, coriaceous to carbonaceous, ostiolate, ostiole round. Involucrellum present in
some genera, usually carbonized. Excipulum dense, consisting of compressed hyphae, appearing prosoplectenchymatous in thin sections, but structure may be difficult to observe
due to carbonization, hyaline to brown or brown-black.
Hamathecium comprising 0.5–0.7(–1.5) µm wide paraphyses, hyaline, flexuous, unbranched or branched to sparsely
anastomosing. Asci (1–)8-spored, bitunicate, fissitunicate,
obclavate to mostly cylindrical, shortly pedicellate, with
narrow to somewhat broader, ocular chamber, non-amyloid.
Ascospores irregularly arranged to uni- or biseriate, fusiform
to ellipsoid, hyaline, septate to muriform, with eusepta and
rectangular lumina, smooth-walled, often constricted at the
septa. Asexual morph: Pycnidia common, immersed to
erumpent, often visible as black dots, rarely pale and inconspicuous, sometimes in specific, pseudostromatic areas, or
whole thalli only producing pycnidia. Conidia acrogenous,
either macro- or microconidia; macroconidia 1-septate to
rarely muriform, ellipsoid to bacillar, sometimes becoming
rather large, with variously shaped gelatinous appendages,
hyaline; microconidia usually aseptate, fusiform-ellipsoid to
bacillar, small, hyaline.
Chemistry: Most species do not contain secondary substances. Crystalline anthraquinone pigments are found in
Flavobathelium.
Type: Strigula Fr., Syst. Mycol. 2(2): 535 (1823).
Notes: For a general discussion of the family see Lücking
& Nelsen in Hyde et al. (2013; Strigulaceae). Recently, a
new family, Tenuitholiascaceae, with the single genus, Tenuitholiascus, was established to accomodate a novel lineage
sister to Strigulaceae, but with a different ascus type (Jiang
et al. 2020a). Simultaneously, molecular phylogenetic analysis focusing on a broad set of foliicolous species hitherto
assigned to the genus Strigula demonstrated that the latter is
heterogeneous and can be divided into six genera, all characterized by a combination of morphological and anatomical features (Jiang et al. 2020b). Given that S. jamesii, the
only non-foliicolous representative of Strigula sensu lato
sequenced so far, is unrelated to the six foliicolous clades
and positioned far from Strigula sensu stricto (Fig. 70),
here we consequently also reassess the generic classification of non-foliicolous representatives so far placed in this
genus. Based on characters of the ascospores and macroand microconidia, three further genera are recognized:
Phyllocharis Fée, for the anatomically unique foliicolous
taxon, Strigula orbicularis Fr.; Dichoporis Clem., for nonfoliicolous species with 1-septate ascospores, 1-septate macroconidia and chiefly fusiform microconidia; and the newly
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Fungal Diversity (2020) 105:17–318
Fig. 70 Best scoring maximum-likelihood tree (also including Bayesian posterior probabilities) of Strigulaceae showing the currently recognized and sequenced genera (adapted from Jiang et al. 2020b; see that reference for further details)
established genus Swinscowia S.H. Jiang, Lücking & Sérus.,
for non-foliicolous species with multi-septate to muriform
ascospores, multi-septate to muriform macroconidia and
chiefly bacillar microconidia. Dichoporis is closely related
to Phylloporis Clem., but we keep the two groups separate
largely based on the different substrate and photobiont, as
molecular data for foliicolous lineages indicate photobiont
preference, to be interpreted as a mycological character of
mycobiont-photobiont compatibility, be consistent within a
13
lineage. The same applies to Swinscowia in relation to the
foliicolous genus Phyllocratera Sérus. & Aptroot.
Both Index Fungorum and MycoBank list Strigulaceae
A.B. Frank (Frank in Leunis 1877) as valid name for the
family, antedating Strigulaceae Zahlbr. However, this is not
correct. Frank did not publish a new name but cited Strigulini Fr., which was originally established as the name of a
suborder (Fries 1825). Names cannot be used unchanged
at different ranks unless they are descriptive and not
Fungal Diversity (2020) 105:17–318
automatically typified (ICN Art. 6, Note 3, and Art. 16.1),
and the name Strigulini is derived from the genus name
Strigula and hence automatically typified. Also, the use of
names at different ranks, yet unchanged, does not apply to
the level of family. Therefore, Frank’s use of the name Strigulini Fr. as “Familie” (Frank in Leunis 1877: 1699) does not
constitute valid publication of a name at family level that
is to be corrected with unchanged authorship (ICN 18.4);
instead it is to be considered a misapplication of an existing
name at a different rank.
Strigula Fr., Syst. Mycol. 2(2): 535 (1823) [nom. sanct.];
Fries, Kongl. Vet.-Akad. Handl. 1821: 323 (1821) [nom.
nud.].
MycoBank number: MB 5285; Index Fungorum number:
IF 5285; Facesoffungi number: FoF 07706; 30 morphologically delimited species (Jiang et al. 2020b); molecular data
available for approximately 15 species (Jiang et al. 2016,
2017a, b, 2020b, Ford et al. 2019; Oh et al. 2019; Woo et al.
2020).
= Nemathora Fée, Essai Crypt Écorc. 1: LVIII, XCIV,
XCIX (1825); Santesson, Symb. Bot. Upsal. 12(1): 138
(1952). Type species: Nemathora viridissima Fée [= Strigula
smaragdula Fr.].
= Craspedon Fée, Essai Crypt Écorc. 1: LIX, XCIV, C
(1825); Santesson, Symb. Bot. Upsal. 12(1): 138 (1952).
Type species: Craspedon concretum Fée [≡ Strigula concreta (Fée) R. Sant.].
= Melanophthalmum Fée, Essai Crypt. Écorc. 1: LX,
XCIV, C (1825); Santesson, Symb. Bot. Upsal. 12(1): 138
(1952). Type species: Melanopthalmum antillarum Fée [≡
Strigula antillarum (Fée) Müll. Arg.].
= Haploblastia Trevis., Conspectus Verrucarinarum: 16
(1860); Santesson, Symb. Bot. Upsal. 12(1): 138 (1952).
Type species: Haploblastia nitidula (Mont.) Trevis. [≡ Strigula nitidula Mont.].
= Discosiella Syd. in Sydow & Sydow, Leafl. Philipp.
Bot. 5: 1546 (1912); Eriksson & Hawksworth, Syst. Ascomycetum 11: 56 (1992). Type species: Discosiella cylindrospora Syd. [= Strigula sp.].
= Strigulomyces [as Strigulomyces] Cif. & Tomas., Atti
Ist. Bot. Univ. Lab. Crittogam. Pavia, Ser. 5, 10: 61 (1953)
[nom. illeg.]; Harris, More Florida Lichens: 152 (1995);
Lücking & Hawksworth, Taxon 56: 1274 (2007). Type species: Strigula elegans (Fée) Müll. Arg. [= Strigula smaragdula Fr.].
= Catenata Bat. in Batista & Bezerra, Publ. Inst. Micol.
Univ. Recife 321: 15 (1961) [nom. inval.]; Lücking et al.,
Lichenologist 30: 161. (1998). Original species: Catenata
antillarum (Fée) Bat. [≡ Strigula antillarum (Fée) Müll.
Arg.].
= Kilikiostroma Bat. & J. L. Bezerra, Publ. Inst. Micol.
Univ. Recife 321: 13 (1961); Lücking et al., Lichenologist
141
30: 144 (1998). Type species: Kilikiostroma peresii Bat. &
J. L. Bezerra [= Strigula prasina Müll. Arg.].
= Pycnociliospora Bat. in Batista et al., Publ. Inst. Micol.
Univ. Recife 251: 6 (1962); Lücking et al., Lichenologist 30:
154 (1998). Type species: Pycnociliospora belluciae Bat. &
J. A. Lima [= Strigula antillarum (Fée) Müll. Arg.].
= Porina sect. Sagediastrella Vain., Ann. Acad. Sci.
Fenn., Ser. A, 19: 10 (1923); Santesson, Symb. Bot. Upsal.
12(1): 138. 1952. Type species: Porina nitens Vain. [= Strigula nitidula Mont.].
= Shanoria Anahosur in Harris, More Florida Lichens:
144 (1995) [nom. inval., non Shanoria Subram. & K. Ramakr., non Shanoria sensu Anahosur (1967)]. Original species: Shanoria indica Anahosur [= Strigula cf. nitidula
Mont.].
Lichenized on leaves in terrestrial, lowland to montane, tropical to subtropical, and extending into temperate,
habitats. Thallus (pseudo-)corticate, grey-green to bright
green, subcuticular and sometimes damaging the supporting leaf. Photobiont Cephaleuros. Sexual morph: Ascomata perithecia, erumpent to prominent, black or covered
by thallus layer, lens-shaped to wart-shaped or conical,
carbonaceous, ostiolate. Involucrellum present, carbonized.
Excipulum prosoplectenchymatous, dark brown to brownblack. Hamathecium comprising 0.5–0.7 µm wide paraphyses, hyaline, flexuose, unbranched or very rarely branched
and anastomosing. Asci usually 8-spored but sometimes
appearing 9–16-spored due to ascospores breaking in halves
within the asci, bitunicate, fissitunicate, cylindrical to narrowly obclavate, shortly pedicellate, with narrow ocular
chamber, non-amyloid. Ascospores irregularly arranged to
uni- or biseriate, fusiform to ellipsoid or bacillar, hyaline,
1-septate, rarely 3-septate, with thin eusepta and rectangular
lumina, smooth-walled, often constricted (and sometimes
breaking apart) at the septa. Asexual morph: Pycnidia common, immersed to erumpent, visible as black dots, rarely in
pseudostromatic clusters, often whole thalli only producing
pycnidia. Conidia acrogenous, either macro- or microconidia; macroconidia mostly 1-septate, ellipsoid to bacillar,
with gelatinous appendages, hyaline; microconidia usually
aseptate, ellipsoid to fusiform, small, hyaline.
Chemistry: No secondary substances known.
Type species: Strigula smaragdula Fr., Linnaea 5: 550
(1830).
Notes: Strigula as here defined includes the bulk of foliicolous species centered around S. smaragdula, after separation of the genera Dichoporis, Phyllocharis, Phylloporis,
Puiggariella, Raciborskiella, and Racoplaca, and introduction of the new genera Serusiauxiella and Swinscowia (see
below). As shown by Jiang et al. (2020b), a much more narrowly defined Strigula is still heterogeneous: whereas the
species with a morphology similar to S. smaragdula form a
supported clade, three basal lineages differ in morphology
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142
and/or anatomy, including the S. nitidula and S. prasina
groups and S. macaronesica. While Strigula sensu stricto
forms thickened, usually bright green thalli, those of the
S. nitidula group are very thin and more similar to those
of Racoplaca, and S. prasina and relatives have hypophyllous thalli with bluish tinge and branched and anastomosing
paraphyses. Should these be confirmed as separate clades
in a broader sampling of taxa, the names Haploblastia (S.
nitidula group) and Kilikiostroma (S. prasina group) would
be available. Currently, about 30 exclusively foliicolous species are to be assigned to the genus Strigula sensu stricto as
here defined.
Harris (1995) listed the name Shanoria Anahosur as a
synonym of Strigula, given that the only species described
by that author, S. indica Anahosur, represents the pycnidial
stage of Strigula cf. nitidula. While the latter is correct (see
also Raj 1981), Harris (1995) misinterpreted its protologue,
since Anahosur (1967) did not establish a new genus but
placed his new species in the already existing genus, Shanoria Subram. & K. Ramakr., typified by S. bambusarum
Subram. & K. Ramakr. (Subramanian and Ramakrishnan
1956). The latter has been established as a synonym of Placonema (Sacc. & D. Sacc.) Petr. (asexual morph; Sutton
1977) and Kiehlia Viégas (sexual morph; Farr 1968), with
Placonema having priority. The name Shanoria Anahosur
does therefore not exist with that place and date of publication (Anahosur 1967), but was inadvertently, yet effectively created by Harris (1995), who ascribed the name to
Anahosur. Unfortunately, this is a rather common lapsus,
frequently seen in early mycological literature, which has
perpetuated many presumed homonyms that were in fact
never established as such by their ascribed authors. The
Code (Shenzen Code; Turland et al. 2018) does not include
a specific provision for names created through erroneous
citation, and so various rules must be consulted to ascertain
whether such a name is actually valid (and consequently
illegitimate). In the case of Shanoria Anahosur in Harris
(1995), Harris (1995) provided a full and direct reference
to the Latin description of S. indica and cited the latter as
type of the genus, so the provisions of ICN Art. 38.1, 38.13,
39.1 and 40.1 are theoretically fulfilled. Since Harris (1995)
did not use an explicit rank designation, such as ‘gen. nov.’,
because he did not intend to describe a new genus, one could
cite ICN Art. 37.1 to render the name invalid; however, that
article does not include a specific provision how it should
be applied to genus names, and the genus rank of the name
is clear from context. Further, ICN Art. 36.1(b) may suggest the name as not validly published, since it is merely
cited in synonymy. However, that article specifically refers
to the intention of the author of the name, not the author
of the publication. As Harris (1995) ascribed the name to
Anahosur and not to himself [ICN Art. 46.2], his placement
of Shanoria as synonym of Strigula appears irrelevant with
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Fungal Diversity (2020) 105:17–318
respect to ICN Art. 36.1(b). ICN Art. 38.5 provides another
potential avenue, as for a simultaneously established new
genus and species, validation through reference to an earlier description is not possible. That article does not apply,
though, as Harris (1995) only established a new genus name,
whereas the name Shanoria indica Anahosur had already
been validly described earlier (Anahosur 1967). Fortunately,
this case finds its solution in ICN Art. 38.11(b), which specifies that “…for a name of a genus or subdivision of a genus,
the earlier description or diagnosis must be that of a genus
or subdivision of a genus”. Hence, the name Shanoria Anahosur in Harris (1995) is not validated by reference to the
Latin description of Shanoria indica Anahosur (1967), and
it its therefore invalid. It appears ineffective that one has to
fully explore the Code to eventuelly render a name invalid
that was inadvertently created through erroneous citation.
ICN Art. 48.1 may seemingly reflect such cases, but in reality it does not apply, as it specifically refers to names intentionally adopted by an author but in a sense different from
the protologue, i.e. deliberately excluding the type of the
name. Harris (1995) did not intentionally adopt the name
Shanoria in a different sense, he simply cited it incorrectly,
due to a misinterpretation of Anahosur’s (1967) paper. He
also did not explicitly exclude its original type but did so
by implication, through citing another species erroneously
as type of a name that did not actually exist, and through
indicating Shanoria Anahosur as illegitimate, which means
Harris (1995) was aware of the existence of Shanoria Subram. & K. Ramakr. but did not realize that Anahosur (1967)
was using precisely that name.
Molecular data show that the type species, Strigula
smaragdula, in its current definition is a collective species
(Jiang et al. 2016, 2017a, b; Oh et al. 2019; Woo et al. 2020).
Given that the type is from Nepal, the name is possibly to
be applied to a taxon restricted to eastern Asia, whereas the
bulk of tropical species will have to be redispositioned using
in part available names currently in synonymy of S. smaragdula. This may include reinstatement of the epithet elegans
for a common and possibly widespread tropical taxon, with
the type being from the Caribbean region.
Other genera:
Dichoporis Clem., Genera of Fungi: 40, 173 (1909).
MycoBank number: MB 1517; Index Fungorum number:
IF 1517; Facesoffungi number: FoF 08854; 18 morphologically delimited species (this paper); molecular data not
available.
= Diporina Clem., Genera of Fungi: 40, 173 (1909). Type
species: Diporina subsimplicans (Nyl.) Clem. [= Dichoporis
subsimplicans (Nyl.) S.H. Jiang, Lücking & Sérus.].
Lichenized on bark and rocks in terrestrial, (sub-)tropical
to rarely temperate habitats. Thallus usually ecorticate, whitish to brownish. Photobiont Trentepohlia. Sexual morph:
Fungal Diversity (2020) 105:17–318
Ascomata perithecia, usually dispersed but often dense, very
rarely fused, immersed-erumpent to prominent, usually black
or covered by thallus layer, lens-shaped to wart-shaped, usually carbonaceous, ostiolate. Involucrellum usually present
and carbonized, very rarely reduced or absent. Excipulum
prosoplectenchymatous, pale to brown. Hamathecium comprising 0.5–0.7 µm wide paraphyses, hyaline, flexuose,
typically branched and sometimes somewhat anastomosing.
Asci usually 8-spored, bitunicate, fissitunicate, cylindrical to
narrowly obclavate, shortly pedicellate, with narrow ocular
chamber, non-amyloid. Ascospores irregularly arranged to
uni- or biseriate, fusiform to ellipsoid or bacillar, hyaline,
1-septate, very rarely with two obscure, additional septa,
with thin eusepta and rectangular lumina, smooth-walled,
often constricted (and sometimes breaking apart) at the
septa. Asexual morph: Pycnidia common, immersed to
erumpent, usually visible as black dots. Conidia typically
acrogenous, either macro- or microconidia; macroconidia
1-septate, oblong-bacillar, typically with narrow gelatinous
appendages 1–2 µm broad, about 3–6 times as long as broad,
and about one fifth to half as long as the conidia, hyaline;
microconidia aseptate, typically fusiform, small, hyaline.
Chemistry: No secondary substances known.
Type species: Dichoporis schizospora (Vain.) Clem., [as
‘schizopora’], Gen. Fung.: 173 (1909) [= Dichoporis ziziphi
(A. Massal.) S.H. Jiang, Lücking & Sérus. (see below)].
Notes: Dichoporis is here adopted for a group of nonfoliicolous species that was already recognized as separate
group (Strigula taylorii group) by Roux and Sérusiaux
(2004), though not exactly in the same sense. Two names
are available for this group, besides Dichoporis also Diporina Clem., established in the same work (Clements 1909)
and based on D. subsimplicans (Nyl.) Clem. We prefer to
use Dichoporis, as it better expresses the potential relationships with the genus Phylloporis and is less confusing than
Diporina, since these lichen fungi are entirely unrelated to
Porina and its allies. The implied differences between the
two genera as given by Clements (1909), namely ascospores
remaining intact (Diporina) vs. breaking into two part spores
(Dichoporis) are not considered taxonomically relevant at
the genus level, as molecular data show foliicolous species
with ascospores remaining intact or breaking into part spores
to be closely related (Jiang et al. 2020b).
Dichoporis is one of two genera here recognized for the
bulk of non-foliicolous species thus far placed in Strigula
sensu lato, the other one being Swinscowia, established
below. The difference between these two groups was also
recognized by Roux and Sérusiaux (2004) and can be
emended as follows, including their Strigula affinis, S.
calcarea, and S. tagananae groups within Swinscowia:
ascospores and macroconidia 1-septate in Dichoporis vs.
3-septate to muriform in Swinscowia; macroconidial appendages relatively long and narrow, 1–2 µm broad, 3–6 times as
143
long as wide, about 20–50% of conidial length in Dichoporis
vs. relatively short and broad, (1–)2–4 µm, 1–2(–5) times as
long as narrow, 15–30% of conidial length in Swinscowia;
and microconidia typically fusiform(-oblong) in Dichoporis
vs. (oblong-)bacillar in Swinscowia. Thus far, only the type
of Swinscowia, S. jamesii, has been sequenced and shown
to be outside the clade containing the foliicolous species of
Strigula sensu stricto and sensu lato, being more closely
related to Flavobathelium and Phyllobathelium (Fig. 70).
The latter two genera, although predominantly foliicolous,
share the multi-septate to muriform ascospores and macroconidia with Swinscowia. We therefore predict Dichoporis to
form part of the clade containing Strigula sensu stricto and
relatives, as it shares ascospore and conidial features with
the latter (see also Roux and Sérusiaux 2004).
Whereas the separation of Dichoporis from Swinscowia
is straightforward, it is currently unclear how both genera
relate to the foliicolous Phylloporis and Phyllocratera,
respectively. Apart from the substrate, the principal difference lies in the photobiont, which in the chiefly foliicolous
taxa represents the Phycopeltis morphodeme, with cells
forming specific, plate-like arrangements (Aptroot et al.
1997; Lücking 2008). This is even the case in occasional
corticolous occurrences of e.g. Phylloporis (see below).
The nature of the photobiont is generally considered a nonmycological character of no taxonomic value (e.g. Harris
1995), but this is an oversimplification. Photobiont choice
is clearly a mycological character, as it is determined by
genetically fixed preferences of the mycobiont for specific
photobiont types, amply documented in numerous cases.
This does not mean that closely related mycobionts cannot
associate with disparate photobiont lineages, but it implies
that each case has to be evaluated in a phylogenetic context.
Whereas such comparative data are currently unavailable
for the target taxa, i.e. Dichoporis vs. Phylloporis (only
the latter sequenced) and Swinscowia vs. Phyllocratera
(only the former sequenced), data from foliicolous lineages
show that each genus-level mycobiont lineage consistently
associates with a specific photobiont lineage representing
either Cephaleuros, Phycopeltis or Trentepohlia (Jiang et al.
2020b), although the generic delimitation within Trentepohlia sensu lato is not settled (Lopez-Bautista et al. 2006;
Rindi et al. 2009; Nelsen et al. 2011b; Hametner et al. 2014;
Zhu et al. 2017). We therefore maintain this concept to keep
Dichoporis and Swinscowia separate from Phylloporis and
Phyllocratera, respectively. This appears also supported by
the unbranched paraphyses in the latter two genera vs. the
generally branched and sometimes anastomosing paraphyses
in Dichoporis and Swinscowia.
While the bulk of the 18 species here included in Dichoporis appears rather homogeneous, one taxon is placed in
this genus with great hesitation, namely D. dichosporidii.
The latter was described in Strigula due to its 1-septate
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144
macroconidia with gelatinous appendages, but differs from
all other species by its lichenicolous habit [on thalli of
Dichosporidium nigrocinctum (Ehrenb.) G. Thor], the urnshaped perithecia, and the lack of an involucrellum (Etayo
2002). It is unclear whether this species actually belongs in
Strigulaceae, but in lieu of a more convincing alternative
placement, it is here provisionally assigned to Dichoporis.
Among Strigulaceae, Dichoporis is the genus most similar and most easily confused with Anisomeridium, as the latter typically features a similar thallus morphology and photobiont, black perithecia, and mostly 1-septate ascospores
(Harris 1995). In the presence of pycnidia producing macroconidia, the distinction is straightforward, as Anisomeridium
produces aseptate macroconidia lacking gelatinous appendages but typically embedded in a gelatinous mass extruded
as drops from the pycnidia, very different from Strigulaceae.
In the absence of macroconidia, the distinction may pose
difficulties, as the differences in ascus structure are rather
subtle, although the asci of Anisomeridium are usually cylindrical and those of Strigula obclavate-fusiform. The paraphyses, usually stated to be unbranched in Strigula sensu
lato vs. anastomosing in Anisomeridium, can be branched
and slightly anastomosing in the former as well. Still, the
paraphyses in Anisomeridium typically appear net-like with
numerous anastomoses, whereas those of Strigula are usually lax and not net-like. Also, many species of Anisomeridium have lateral ostioles and/or larger ascospores, often
with granular ornamentation. In addition, the ascospores
in Anisomeridium are usually relatively broader, with a
length/width ratio in the range of 2–3(–3.5), whereas those
of Dichoporis are in the range of (2.5–)3–5.5. This somewhat narrows down the number of species that could potentially confused, to about seven taxa in Dichoporis with relatively broader ascospores (2.5–3.5 times as long as broad)
and those species of Anisomeridium with apical ostiole and
smooth ascospores shorter than 30 µm and relatively narrow
(about 2.5–3.5 times as long as broad). Unfortunately, that
still involves about 20 taxa in Anisomeridium, many of them
frequent and widespread. These taxa are largely summarized
in the world key to Anisomeridium by Harris (1995: 126),
between couplets 12 and 19 (for species growing on bark
or wood) and couplets 67 and 72 (for species growing on
rock). Thus, when in doubt whether a specimen at hand is
indeed a Dichoporis, species of Anisomeridium with similar
ascospore dimensions should be checked.
New combinations in Dichoporis:
Dichoporis bermudana (Tuck.) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836370; Index Fungorum number: IF 836370; Facesoffungi number: FoF 08855.
Bas.: Verrucaria bermudana Tuck. in Nylander,
Sert. Lich. Trop. Labuan Singapore: 43 (1891); Porina
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Fungal Diversity (2020) 105:17–318
bermudana (Tuck.) Zahlbr., Cat. Lich. Univers. 1: 367
(1922); Strigula bermudana (Tuck.) R.C. Harris, More
Florida Lichens: 155 (1995).
Notes: The species is usually cited as “Tuck ex Nyl.” but
Nylander (1891) ascribed the name to Tuckerman (ICN Art.
46.2), so the ‘ex’ citation format does not apply.
Dichoporis brevis (Bricaud & Cl. Roux) S.H. Jiang, Lücking
& Sérus. comb. nov.
MycoBank number: MB 836371; Index Fungorum number: IF 836371; Facesoffungi number: FoF 08856.
Bas.: Strigula brevis Bricaud & Cl. Roux in Roux &
Sérusiaux, Biblioth. Lichenol. 90: 53 (2004); Strigula brevis Bricaud & Cl. Roux in Roux & Bricaud, Bull. Soc. Linn.
Provence 44: 107 (1993) [nom. inval., ICN Art. 36.1(a)].
Dichoporis connivens (R.C. Harris) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836372; Index Fungorum number: IF 836372; Facesoffungi number: FoF 08857.
Bas.: Strigula connivens R.C. Harris, More Florida
Lichens: 155 (1995).
Dichoporis dichosporidii (Etayo) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836373; Index Fungorum number: IF 836373; Facesoffungi number: FoF 08858.
Bas.: Strigula dichosporidii Etayo, Biblioth. Lichenol.
84: 127 (2002).
Dichoporis elixii (P.M. McCarthy) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836374; Index Fungorum number: IF 836374; Facesoffungi number: FoF 08859.
Bas.: Strigula elixii P.M. McCarthy, Biblioth. Lichenol.
78: 279 (2001).
Dichoporis fractans (P.M. McCarthy) S.H. Jiang, Lücking
& Sérus. comb. nov.
MycoBank number: MB 836375; Index Fungorum number: IF 836375; Facesoffungi number: FoF 08860.
Bas.: Strigula fractans P.M. McCarthy, Lichenologist 29:
516 (1997).
Dichoporis maritima (H. Harada) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836376; Index Fungorum number: IF 836376; Facesoffungi number: FoF 08861.
Bas.: Strigula maritima H. Harada, Bryologist 101: 605
(1998).
Dichoporis minutula (P.M. McCarthy) S.H. Jiang, Lücking
& Sérus. comb. nov.
Fungal Diversity (2020) 105:17–318
MycoBank number: MB 836377; Index Fungorum number: IF 836377; Facesoffungi number: FoF 08862.
Bas.: Strigula minutula P.M. McCarthy, Muelleria 8: 327
(1995).
Dichoporis natalis (P.M. McCarthy) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836378; Index Fungorum number: IF 836378; Facesoffungi number: FoF 08863.
Bas.: Strigula natalis P.M. McCarthy, Biblioth. Lichenol.
78: 281 (2001).
Dichoporis nipponica (H. Harada) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836379; Index Fungorum number: IF 836379; Facesoffungi number: FoF 08864.
Bas.: Strigula nipponica H. Harada, Nova Hedwigia 60:
488 (1995).
Dichoporis occulta (P.M. McCarthy & Malcolm) S.H. Jiang,
Lücking & Sérus. comb. nov.
MycoBank number: MB 836380; Index Fungorum number: IF 836380; Facesoffungi number: FoF 08865.
Bas.: Strigula occulta P.M. McCarthy & Malcolm, Mycotaxon 60: 323 (1996).
Dichoporis phaea (Ach.) S.H. Jiang, Lücking & Sérus.
comb. nov.
MycoBank number: MB 836381; Index Fungorum number: IF 836381; Facesoffungi number: FoF 08866.
Bas.: Verrucaria phaea Ach., Syn. Meth. Lich.: 88
(1814); Porina phaea (Ach.) Müll. Arg., Flora 68: 261
(1885); Strigula phaea (Ach.) R.C. Harris in Tucker & Harris, Bryologist 83: 18 (1980).
= Porina mundula Müll. Arg., Bot. Jahrb. 6: 402 (1885).
= Strigula diederichiana Etayo, Cl. Roux & Sérus. in
Roux & Bricaud, Bull. Soc. Linn. Provence 44: 129 (1993)
[nom. inval., ICN Art. 36.1].
Dichoporis subprospersella (Vain.) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836382; Index Fungorum number: IF 836382; Facesoffungi number: FoF 08867.
Bas.: Porina subprospersella Vain., Mycologia 21: 39
(1929); Strigula subprospersella (Vain.) P.M. McCarthy,
Biblioth. Lichenol. 78: 281 (2001).
Dichoporis subsimplicans (Nyl.) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836383; Index Fungorum number: IF 836383; Facesoffungi number: FoF 08868.
Bas.: Verrucaria subsimplicans Nyl., Lich. Nov. Zeland.:
130 (1888); Diporina subsimplicans (Nyl.) Clem., Gen.
145
Fung.: 40, 173 (1909); Porina subsimplicans (Nyl.) Müll.
Arg., Bull. Herb. Boissier 2(1): 91 (1894); Segestria subsimplicans (Nyl.) Hellb., Bih. K. Svenska Vetensk. Akad.
Handl., Afd. 3, 21(13): 132 (1896); Strigula subsimplicans
(Nyl.) R.C. Harris, Biblioth. Lichenol. 57: 175 (1995).
Dichoporis taylorii (Carroll) S.H. Jiang, Lücking & Sérus.
comb. nov.
MycoBank number: MB 836384; Index Fungorum number: IF 836384; Facesoffungi number: FoF 08869.
Bas.: Verrucaria taylorii Carroll [as ‘taylori’] in
Nylander, Expos. Synopt. Pyrenocarp.: 82 (1858); Arthopyrenia taylorii (Carroll) Mudd, Man. Brit. Lich.: 302 (1861);
Porina taylorii (Carroll) Swinscow, Lichenologist 2: 169
(1962); Spermatodium taylorii (Carroll) Trevis. Conspect.
Verruc.: 11 (1860); Strigula taylorii (Carroll) R.C. Harris in
Hawksworth et al., Lichenologist 12: 107 (1980).
Notes: The species is usually cited as “Carroll ex Nyl.”
but Nylander (1858) ascribed the name to Carroll (ICN Art.
46.2), so the ‘ex’ citation format does not apply.
Dichoporis tenuis (R.C. Harris) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836385; Index Fungorum number: IF 836385; Facesoffungi number: FoF 08870.
Bas.: Arthopyrenia tenuis R.C. Harris, Michigan Bot. 12:
16 (1973); Strigula americana R.C. Harris in Tucker & Harris, Bryologist 83: 18 (1980).
Notes: Unfortunately, in the new generic disposition, the
original epithet tenuis has priority over the rather well-established replacement name Strigula americana.
Dichoporis viridiseda (Nyl.) S.H. Jiang, Lücking & Sérus.
comb. nov.
MycoBank number: MB 836386; Index Fungorum number: IF 836386; Facesoffungi number: FoF 08871.
Bas.: Verrucaria viridiseda Nyl., Expos. Synopt. Pyrenocarp.: 55 (1858); Strigula viridiseda (Nyl.) R.C. Harris in
Tucker & Harris, Bryologist 83: 18 (1980).
Dichoporis wilsonii (Riddle) S.H. Jiang, Lücking & Sérus.
comb. nov.
MycoBank number: MB 836387; Index Fungorum number: IF 836387; Facesoffungi number: FoF 08872.
Bas.: Porina wilsonii Riddle in Britton & Millspaugh,
The Bahama Flora: 523 (1920); Porina wilsonii Riddle,
Mycologia 15: 73 (1923) [nom. inval., ICN Art. 6.23, Note
2]; Strigula wilsonii (Riddle) R.C. Harris in Egan, Bryologist 90: 164 (1987).
Notes: The publication of Porina wilsonii by Riddle
(1923) is an exact copy of the earlier publication by the
same author in the chapter “Lichenes” in Britton & Millspaugh’s (1920) Bahama Flora. The later publication is
13
146
usually considered a superfluous validation, but likely there
was some confusion about publishing the same content
twice in different outlets. Riddle had possibly overlooked
that he had already published the species, because in the
treatment on the Bahamas he included several species that
he simultaneously described from other areas, such as in
this case the Isle of Pines (Cuba), and in 1923 he focused
on taxa from the latter area, without referring back to the
Bahama Flora.
Fungal Diversity (2020) 105:17–318
5.
5.
6.
6.
Dichoporis ziziphi (A. Massal.) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836388; Index Fungorum number: IF 836388; Facesoffungi number: FoF 08873.
Bas.: Sagedia ziziphi A. Massal., Miscell. Lichenol.:
30 (1856); Porina ziziphi (A. Massal.) Zahlbr., Cat. Lich.
Univers. 1: 410 (1922); Spermatodium ziziphi (A. Massal.)
Trevis., Conspect. Verruc.: 12 (1860); Strigula ziziphi (A.
Massal.) Cl. Roux & Sérus. in Roux & Sérusiaux, Biblioth.
Lichenol. 90: 55 (2004).
= Strigula mediterranea Etayo, Lichenologist 25: 258
(1993); Porina schizospora Vain., Term. Füz. 22: 340 (1899)
[non Strigula schizospora R. Sant., Symb. Bot. Upsal. 12(1):
175 (1952)].
Notes: The epithet schizospora, based on Porina schizospora Vain. (Vainio 1899), was blocked for use in Strigula
by S. schizospora R. Sant. (Santesson 1952), which required
the replacement name S. mediterranea (Etayo 1993). The
epithet schizospora is theoretically available in Dichoporis
but is predated by Sagedia ziziphi (Massalongo 1856). The
identity of both taxa was established by Roux and Sérusiaux
(2004). The species was recently reported also from Brazil
(Cavalcante and Cáceres 2014).
7.
7.
8.
8.
9.
9.
10.
10.
11.
11.
12.
Key to species of Dichoporis:
12.
1.
Lichenicolous (on Dichosporidium); involucrellum
absent………………………Dichoporis dichosporidii
1. Lichenized (on bark, wood or rock); involucrellum present
………………………………………………………2
2. On bark………………………………………………3
2. On rock………………………………………………10
3. Ascomata aggregate-fused; ascospores 7–12 µm long;
southeastern North America……Dichoporis connivens
3. Ascomata solitary (but sometimes dense); ascospores
(9–)12–28 µm long……………………………………4
3. Thallus rather dark olive-brown to olive-green, with the
black perithecia barely contrasting in colour; perithecia
0.1–0.3(–0.35) mm diam.……………………………5
4. Thallus white-grey, with the black perithecia
strongly contrasting; perithecia (0.3–)0.4–0.6 mm
diam.…………………………………………………8
13.
13.
14.
14.
15.
15.
13
Ascospores 9–14 µm long; macroconidia 6–10 × 1.5–
2.5 µm; macroconidiogenous cells rather long (mostly
15–25 µm); pantropical……………Dichoporis phaea
Ascospores 13–27 µm long; macroconidia 8–20 ×
(2–)2.5–3.5 µm; macroconidiogenous cells rather short
(mostly 5–10 µm)……………………………………6
Ascospores 13–20 µm long, partly obscurely 3-septate,
remaining intact; perithecia 0.1–0.15 mm diam.; New
Zealand……………………Dichoporis subsimplicans
Ascospores 17–27 µm long, consistently 1-septate,
partly breaking into part spores outside the asci; perithecia 0.2–0.3 mm diam.; Europe……………………7
Macroconidia 8–14 µm long………Dichoporis brevis
Macroconidia 13–20 µm long……Dichoporis taylorii
Ascospores 12–17 µm long; macroconidia 7–10 ×
2–2.5 µm; North America……Dichoporis viridiseda
Ascospores 15–28 µm long; macroconidia 8–19 × 3–4.5
µm……………………………………………………9
Macroconidia 8–12 µm long; mediterranean Europe…
……………………………………Dichoporis ziziphi
Macroconidia 14–19 µm long; eastern North America……… Dichoporis tenuis (= Strigula americana)
On calcareous rock (limestone)……………………11
On siliceous rock (granite)…………………………15
Ascospores 8–12 × 2.5–3.5 µm; perithecia 0.15–0.2
mm diam.; strongly prominent to sessile; Australasia
(Christmas Island)…………………Dichoporis natalis
Ascospores 11–22 × (3–)3.5–6(–7.5) µm; perithecia 0.2–0.5(–0.9) mm diam.; immersed-erumpent to
prominent (if smaller than 0.2 mm then immersed-eru
mpent)………………………………………………12
Perithecia 0.4–0.9 mm diam.; ascospores 5–7.5 µm
broad; chiefly Caribbean including southeasternmost North America, also reported from Papua New
Guinea……………………………Dichoporis wilsonii
Perithecia (0.15–)0.2–0.5 mm diam.; ascospores 3.5–
5(–6) µm broad………………………………………13
Ascospores 11–15 µm long; macroconidia 10–15 µm
long, about the same length as the ascospores; Caribbean and Australasia (Christmas Island)………………
………………………………Dichoporis bermudana
Ascospores 13–21 µm long; macroconidia (known
from D. elixii) 8–12 µm long, much shorter than the asc
ospores………………………………………………14
Perithecia 0.15–0.2 mm diam., immersed-erumpent;
ascospores 16–25 µm long; Caribbean………………
…………………………Dichoporis subprospersella
Perithecia 0.25–0.5 mm diam., erumpent to prominent;
ascospores 13–20 µm long; Australasia (Christmas
Island)………………………………Dichoporis elixii
Ascospores 6–10 × 2–3 µm; Australasia………………
…………………………………Dichoporis minutula
Ascospores 11–32 × 3.5–6.5 µm……………………16
Fungal Diversity (2020) 105:17–318
16. Ascospores breaking into part spores outside the
asci; perithecia 0.2–0.4 mm diam.; macroconidia
(known from D. fractans) 6–8 × 2.5–3 µm; Australasia……………………………………………………17
16. Ascospores remaining intact; perithecia 0.3–0.8 mm
diam.; macroconidia 12–19 × 2–4 µm; Japan………18
17. Ascospores 22–32 × 4–6.5 µm; New Zealand…………
……………………………………Dichoporis occulta
17. Ascospores 11–15 × 3.5–5 µm; Australia (Lord Howe
Island)……………………………Dichoporis fractans
18. Ascospores 15–26 µm long; perithecia partly covered
by thallus; montane……………Dichoporis nipponica
18. Ascospores 13–17 µm long; perithecia more or less
exposed; maritime rocks……… Dichoporis maritima
Flagellostrigula Lücking, S.H. Jiang & Sérus. gen. nov.
MycoBank number: MB 836389; Index Fungorum number: IF 836389; Facesoffungi number: FoF 08874; one morphologically delimited species (this paper); molecular data
not available.
Lichenized on bark in terrestrial, (sub-)tropical habitats.
Thallus corticate, greenish. Photobiont Trentepohlia. Sexual
morph: Ascomata unknown. Asexual morph: Pycnidia
common, prominent, hemisphaerical to wart-shaped, rather
large (up to 0.8 mm diam.), covered by rather thick thalline layer. Conidia typically acrogenous, only macroconidia
known; macroconidia 1-septate, broadly bacillar, with long
and thin (15–35 × 1 µm) gelatinous appendage at one end
only, hyaline.
Chemistry: No secondary substances known.
Type species: Flagellostrigula laureriformis (Aptroot &
Lücking) Lücking, S.H. Jiang & Sérus. (see below).
Notes: This new genus is established for an oddball that
agrees with Dichoporis in its 1-septate macroconidia. Flagellostrigula laureriformis is only known through pycnidia
producing macroconidia, which differ from Dichoporis and
other genera in having only a single, very long, name-giving
flagelliform appendage at the proximal end, about 3–4 times
as long as the conidia (Aptroot et al. 2008; Jiang et al. 2016;
Cáceres et al. 2017). In addition, the pycnidia are unusually
large and covered by a rather thick thalline layer. The single
species is with certainty known from the Neotropics only
(Costa Rica, Brazil; Aptroot et al. 2008; Cáceres et al. 2017).
The report from China (Jiang et al. 2016) may represent an
additional species, given the slightly deviating morphology
of the pycnidia.
Flagellostrigula laureriformis (Aptroot & Lücking) Lücking, S.H. Jiang & Sérus. comb. nov.
MycoBank number: MB 836390; Index Fungorum number: IF 836390; Facesoffungi number: FoF 08875.
147
Bas.: Strigula laureriformis Aptroot & Lücking in Aptroot et al., Biblioth Lichenol. 97: 131 (2008).
Flavobathelium Lücking, Aptroot & Thor, Lichenologist
29: 221 (1997).
MycoBank number: MB 27825; Index Fungorum number:
IF 27825; Facesoffungi number: FoF 08876; one morphologically and phylogenetically delimited species (Nelsen
et al. 2011a).
Lichenized on leaves in terrestrial, lowland to lower montane tropical forests. Thallus thinly corticate, grey-green with
characteristic metallic appearance. Photobiont Trentepohlia.
Sexual morph: Ascomata perithecia, immersed in groups in
slightly prominent pseudostromata covered by thallus layer
and with a medulla containing ochraceous-yellow, K+ dark
red, powdery crystals; individual perithecia ostiolate. Involucrellum absent. Excipulum prosoplectenchymatous, brown.
Hamathecium comprising 1.5–2 µm wide paraphyses, hyaline, flexuose, unbranched to sparsely branched at the base.
Asci 8-spored, bitunicate, fissitunicate, narrowly clavate to
oblong-fusiform, shortly pedicellate, with narrow ocular
chamber, non-amyloid. Ascospores irregularly arranged to
biseriate, fusiform, hyaline, 3(–5)-septate, with thin eusepta
and rectangular lumina, smooth-walled, not or slightly constricted at the septa. Asexual morph: Pycnidia common,
immersed in groups in slightly prominent pseudostromata
covered by thallus layer and with a medulla containing
ochraceous-yellow, K+ dark red, powdery crystals; often
whole thalli producing pycnidia only. Conidia acrogenous,
only macroconidia known; macroconidia 3(–5)-septate, narrowly bacillar to acicular, distal end with a very narrow,
ciliate appendage about one third as long as the conidia,
proximal end with a short, gelatinous cap, hyaline.
Chemistry: Unidentified ochraceous-yellow anthraquinone crystals in ascomatal and pycnidial pseudostromata.
Type species: Flavobathelium epiphyllum Lücking et al.,
Lichenologist 29: 221 (1997).
Notes: Flavobathelium is thus far a monospecific, tropical
genus growing on living leaves (Lücking et al. 1997; Lücking 2008). Given the inclusion of non-foliicolous species
with multi-septate to muriform ascospores and macroconidia in a single genus, Swinscowia (see below), one could
theoretically consider Flavobathelium and Phyllobathelium
conspecific. However, the two genera are not directly related
phylogenetically, with Flavobathelium sister to Swinscowia
and Phyllobathelium sister to the clade including the two
other genera, with strong support (Fig. 70). This also supports the separation of Swinscowia from Phyllocratera and
may indicate that Swinscowia as here circumscribed is still
heterogeneous, requiring assessment of some of the genuinely muriform-spored species currently included in the latter genus.
13
148
Phyllobathelium (Müll. Arg.) Müll. Arg., Flora 73: 195
(1890).
MycoBank number: MB 4057; Index Fungorum number:
IF 4057; Facesoffungi number: FoF 08877; five morphologically delimited species (Lücking et al. 2017); molecular data
available for one species (Nelsen et al. 2011a).
Lichenized on leaves or rarely on bark in terrestrial, lowland to lower montane tropical to subtropical forests. Thallus
corticate, grey-green with characteristic metallic appearance,
often minutely verrucose. Photobiont Trentepohlia. Sexual
morph: Ascomata perithecia, scattered, prominent to sessile, wart-shaped but often irregularly bumpy, immersed in
pseudostromata covered by thallus layer and usually filled
with a mass of black, powdery crystals, ostiolate. Involucrellum reduced. Excipulum prosoplectenchymatous, colourless. Hamathecium comprising 1.5–2 µm wide paraphyses, hyaline, flexuose, unbranched to sparsely branched.
Asci 8-spored, bitunicate, fissitunicate, broadly cylindrical,
shortly pedicellate, with broad ocular chamber, non-amyloid. Ascospores irregularly arranged to uni- or 2-seriate,
fusiform to ellipsoid, hyaline, muriform, with thin eusepta
and rectangular lumina, smooth-walled, often constricted
at the septa. Asexual morph: Pycnidia common, immersed
in slightly raised, applanate to conical pseudostromata filled
with black, powdery crystals, visible as brown to black dots,
often whole thalli only producing pycnidia. Conidia acrogenous, only macroconidia known; macroconidia septate to
muriform, ellipsoid to bacillar, with more or less ciliate but
rather thick gelatinous appendages, hyaline.
Chemistry: Unidentified black, powdery crystals in ascomata warts and pycnidial pseudostromata.
Type species: Phyllobathelium epiphyllum (Müll. Arg.)
Müll. Arg. [= Phyllobathelium firmum (Stirt.) Vězda in
Lücking & Kalb, Bot. Jb. 122: 44 (2000)].
Notes: For a detailed discussion of this genus, see Lücking (2008) and Lücking & Nelsen (in Hyde et al. 2013;
Strigulaceae). Over time, 13 taxa, including 11 at species
level, have been assigned to Phyllobathelium. Of these, two
were subsequently transferred to Strigula (Harris 1995) and
are currently treated in Swinscowia (see below), namely P.
albolinitum (Nyl.) H. Mayrhofer and P. obtectum (Vain.) H.
Mayrhofer. Phyllobathelium nudum Zahlbr. was recognized
as a second species of Phyllocratera (Lücking & Sérusiaux
2013). Of the remaining ten names, P. epiphyllum (Müll.
Arg.) Müll. Arg. and P. epiphyllum var. majus F. Schill.
are synonyms of P. firmum (Stirt.) Vězda, and P. thaxteri
(Vain.) Zahlbr., P. thaxteri var. heterogena (Vain.) Zahlbr.,
and P. megapotamicum (Malme) R. Sant. are synonymous
with P. chlorogastricum (Müll. Arg.) Aptroot & Lücking
(Lücking 2008; Aptroot and Lücking 2016). This leaves five
species currently accepted in the genus, including also P.
anomalum Lücking, P. leguminosae (Cavalc. & A.A. Silva)
Lücking & Sérus., and P. nigrum R. Sant. & Tibell. The
13
Fungal Diversity (2020) 105:17–318
pseudostromatic perithecia with black, crystalline medulla
are particularly well-developed in P. firmum and P. chlorogastricum, whereas in the other three species, the perithecia
appear naked, with only remnants of a black medullary mass
as irregular outer cover on the perithecia.
The macroconidia of Phyllobathelium chlorogastricum
are perhaps the largest known among all fungi, reaching 200
× 35 µm in size (Lücking 2008, as P. thaxteri).
Phyllocharis Fée, Essai Crypt Écorc. 1: LIX, XCIV, XCIX
(1825).
MycoBank number: MB 4062; Index Fungorum number:
IF 4062; Facesoffungi number: FoF 08878; one morphologically delimited species (this paper); molecular data not
available.
Lichenized on leaves in terrestrial, lowland to montane,
tropical to subtropical habitats. Thallus (pseudo-)corticate, pale grey-green, usually subcuticular and sometimes
damaging the supporting leaf. Photobiont Cephaleuros.
Sexual morph: Ascomata perithecia, prominent, black
but largely covered by thallus layer, wart-shaped, carbonaceous, ostiolate. Involucrellum present, weakly carbonized.
Excipulum prosoplectenchymatous, brownish. Hamathecium
comprising 0.5–0.7 µm wide paraphyses, hyaline, flexuose, unbranched. Asci 8-spored, bitunicate, fissitunicate,
obclavate to fusiform, shortly pedicellate, with narrow ocular chamber, non-amyloid. Ascospores irregularly arranged
to biseriate, oblong, hyaline, 3-septate, with thin eusepta
and rectangular lumina, smooth-walled. Asexual morph:
Pycnidia common, immersed to erumpent, visible as black
dots. Conidia acrogenous, either macro- or microconidia;
macroconidia 3–9-septate, long-filiform, with gelatinous
appendages, hyaline; microconidia aseptate, fusiform, small,
hyaline.
Chemistry: No secondary substances known.
Type species: Phyllocharis complanata Fée [= Phyllocharis orbicularis (Fr.) S.H. Jiang, Lücking & Sérus. (see
below)].
Notes: Although no genuine molecular data are available
for Strigula orbicularis, the 3-septate ascospores set it apart
from all other foliicolous, subcuticularly growing genera
(Jiang et al. 2020b), and the peculiar, filiform macroconidia
are unique among the entire family. The genus Phyllocharis
is therefore resurrected here to reflect this. The name Phyllocharis Fée has a later homonym, Phyllocharis Diels, a
plant genus in the family Lobeliaceae. The latter had been
proposed for conservation (van den Brink et al. 1961), but
the proposal was not accepted (Rickett 1963). Fée’s name is
therefore available.
Based on ascospore and macroconidial features, we predict S. orbicularis to cluster in the clade with Flavobathelium, Phyllobathelium, and Swinscowia. Incidentally, an ITS
sequence for S. orbicularis (GenBank accession KU509981)
Fungal Diversity (2020) 105:17–318
was published by Krishnamurthy and Subramanya (2016),
based on material from India, but unfortunately that
sequence represents a contaminant in the genus Aspergillus.
New combination in Phyllocharis:
Phyllocharis orbicularis (Fr.) S.H. Jiang, Lücking & Sérus.
comb. nov.
MycoBank number: MB 836391; Index Fungorum number: IF 836391; Facesoffungi number: FoF 08879.
Bas.: Strigula orbicularis Fr., Linnaea 5: 549 (1830).
Phyllocraterina Sérus. & Aptroot nom. nov.
MycoBank number: MB 836392; Index Fungorum number: IF 836392; Facesoffungi number: FoF 08880; two
morphologically delimited species (Lücking and Sérusiaux
2013); molecular data not available.
Replaced syn.: Phyllocratera Sérus. & Aptroot in Aptroot
et al., Biblioth. Lichenol. 64: 132 (1997) [nom. illeg., ICN
Art. 53.2; non Phyllocrater Wernham].
Lichenized on leaves in terrestrial, lowland to montane,
tropical habitats. Thallus thinly corticate, grey-green, supracuticular. Photobiont Phycopeltis. Sexual morph: Ascomata
perithecia, prominent, black, wart-shaped with spreading
base, carbonaceous, ostiolate. Involucrellum present, carbonized. Excipulum prosoplectenchymatous, brownish to blackish. Hamathecium comprising 1.5–2 µm wide paraphyses,
hyaline, unbranched. Asci 8-spored, bitunicate, fissitunicate,
clavate, shortly pedicellate, with rather broad ocular chamber, non-amyloid. Ascospores irregularly arranged, fusiformellipsoid, hyaline, muriform, with thin eusepta and rectangular lumina, smooth-walled, slightly constricted at the median
septum. Asexual morph: unknown.
Chemistry: No secondary substances known.
Type species: Phyllocraterina papuana (Sérus. & Aptroot.) Sérus. & Aptroot (see below).
Notes: As response to an anonymous request, the Nomenclatural Committee for Fungi considered the name Phyllocratera and Phyllocrater, a monospecific plant genus in
the family Rubiaceae known from Indonesia (Wernham in
Gibbs 1914), sufficiently alike to be confused and recommended to consider them as homonyms (so-called parahomonyms; Norvell 2011). We therefore introduce here a slightly
altered replacement name ascribed to the original authors, in
order to limit nomenclatural disruption and to honor original
authorship of this enigmatic taxon.
This small, exclusively foliicolous genus, currently containing two species (Lücking and Sérusiaux 2013), largely
corresponds to non-foliicolous species of Swinscowia with
muriform ascospores. Phyllocraterina is here maintained
as a separate genus because of the different substrate and
photobiont and because of the unbranched paraphyses (see
below Dichoporis for further discussion). However, its
149
relationships to some of the muriform-spored species placed
in Swinscowia need to be investigated further.
New combinations in Phyllocraterina:
Phyllocraterina papuana (Sérus. & Aptroot.) Sérus. & Aptroot. comb. nov.
MycoBank number: MB 836393; Index Fungorum number: IF 836393; Facesoffungi number: FoF 08881.
Bas.: Phyllocratera papuana Sérus. & Aptroot in Aptroot
et al., Biblioth. Lichenol. 64: 132 (1997).
Phyllocraterina nuda (Zahlbr.) Lücking & Sérus. comb.
nov.
MycoBank number: MB 836394; Index Fungorum number: IF 836394; Facesoffungi number: FoF 09210.
Bas.: Phyllobathelium nudum Zahlbr., Annals Cryptog.
Exot. 1: 115 (1928); Phyllocratera nuda (Zahlbr.) Lücking
& Sérus., Lichenologist 45: 691 (2013).
Phylloporis Clem., Gen. Fungi: 41, 173 (1909); Vězda,
Folia Geobot. Phytotax. 19: 185 (1984); Jiang et al., Fung.
Div. 102: 287.
MycoBank number: MB 4070; Index Fungorum number:
IF 4070; Facesoffungi number: FoF 08882; nine morphologically delimited species (Jiang et al. 2020b; this paper);
molecular data available for two species (Jiang et al. 2020b).
= Porinomyces Bat. in Bezerra et al., Atas Inst. Micol.
Univ. Pernambuco 5: 410 (1967) [nom. inval.]; Lücking
et al., Lichenologist 30: 168 (1998). Original species: Porinomyces phyllogenus (Müll. Arg.) Bat. [≡ Strigula phyllogena
(Müll. Arg.) R. C. Harris].
= Didymaster Bat. & H. Maia, Atas Inst. Micol. Univ.
Pernambuco 5: 58 (1967); Lücking et al., Lichenologist 30:
140 (1998). Type species: Didymaster myrtaciicola Bat., H.
Maia & Castro [= Strigula platypoda (Müll. Arg.) R. C.
Harris].
= Manaustrum Cavalc. & A. A. Silva in Cavalcante et al.,
Publ. Inst. Micol. Univ. Fed. Pernambuco 647: 13 (1972);
Lücking et al., Lichenologist 30: 146 (1998). Type species:
Manaustrum palmae Cavalc. & A. A. Silva [= Strigula multipunctata (R. Sant.) R. C. Harris].
= Porina sect. Sagediastrum Müll. Arg., Flora 66: 335
(1883); Vainio, Ann. Acad. Sci. Fenn., Ser. A, 15 (1921);
Santesson, Symb. Bot. Upsal. 12(1): 202 (1952). Sectional
type. Porina phyllogena Müll. Arg. [≡ Strigula phyllogena
(Müll. Arg.) R. C. Harris].
Lichenized on leaves or more rarely on bark and rock
in terrestrial, lowland to montane, tropical (to subtropical)
habitats. Thallus ecorticate, grey-green to grey, supracuticular. Photobiont Phycopeltis. Sexual morph: Ascomata perithecia, erumpent to prominent, black or covered
by thallus layer, wart-shaped to conical, carbonaceous,
ostiolate. Involucrellum present, carbonized. Excipulum
13
150
prosoplectenchymatous, dark brown to brown-black.
Hamathecium comprising 0.5–0.7 µm wide paraphyses,
hyaline, flexuose, unbranched. Asci 8-spored, bitunicate, fissitunicate, cylindrical, shortly pedicellate, with narrow ocular chamber, non-amyloid. Ascospores irregularly arranged
to biseriate, fusiform to ellipsoid, hyaline, 1-septate, with
thin eusepta and rectangular lumina, smooth-walled, slightly
constricted at the septa. Asexual morph: Pycnidia common, erumpent, visible as black dots, often whole thalli only
producing pycnidia. Conidia acrogenous, either macro- or
microconidia; macroconidia 1-septate, ellipsoid to bacillar,
with gelatinous appendages, hyaline; microconidia aseptate,
ellipsoid to fusiform, small, hyaline.
Chemistry: No secondary substances known.
Type. Phylloporis phyllogena (Müll. Arg.) Clem., Gen.
Fung.: 173 (1909).
Notes: This genus has long been recognized as a separate
group within Strigula sensu lato (Clements 1909; Vězda
1984) and has even been treated under Porina instead of
Strigula (Santesson 1952). Its inclusion in Strigula based
on internal anatomy and ascus type (e.g. Harris 1995; Lücking 2008) expresses close phylogenetic relationship, but a
schematic broad genus concept, based on shared anatomical features, is not warranted, as shown in numerous other
fungal families where genera agree in ascoma anatomy but
differ in thallus morphology (e.g. Parmeliaceae). Phylloporis is a basally diverging clade among the foliicolous clades
of Strigula sensu lato (Jiang et al. 2020b), indicating that its
supracuticular growth may be a plesiomorphy shared with
Flavobathelium and Phyllobathelium and the subcuticular
growth of all other clades evolved as a single synapomorphy.
Phylloporis currently includes seven foliicolous species
(Jiang et al. 2020b). Here, we also accept two at least partly
non-foliicolous species in this genus (see new combinations
proposed below). One of these, P. cinefaciens, is chiefly
foliicolous but also rarely found on smooth bark, whereas
the other, P. hypothallina, is thus far only known from the
corticolous type material. Both may be distinguished from
Dichoporis by the very thin thallus featuring a Phycopeltislike photobiont (see also above).
New combinations in Phylloporis:
Phylloporis cinefaciens (Nyl.) S.H. Jiang, Lücking & Sérus.
comb. nov.
MycoBank number: MB 836708; Index Fungorum number: IF 836708; Facesoffungi number: FoF 08883.
Bas.: Verrucaria cinefaciens Nyl., Ann. Sci. Nat., Bot.,
Sér. 5(7): 180 (1873); Arthopyrenia cinefaciens (Nyl.)
Zahlbr., Cat. Lich. Univers. 1: 320 (1921); Strigula cinefaciens (Nyl.) R.C. Harris, More Florida Lichens: 155 (1995).
= Porina multipunctata G. Merr. ex R. Sant., Symb. Bot.
Upsal. 12(1): 216 (1952); Phylloporina multipunctata (G.
Merr. ex R. Sant.) Szatala, Ann. Mus. Natn. Hung., N.S. 7:
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Fungal Diversity (2020) 105:17–318
21 (1956); Phylloporis multipunctata (G. Merr. ex R. Sant.)
Vězda, Folia Geobot. Phytotax. 19: 183 (1984); Strigula
multipunctata (G. Merr. ex R. Sant.) R.C. Harris, More
Florida Lichens: 158 (1995).
Notes: This species agrees with the foliicolous Phylloporis austropunctata (P.M. McCarthy) S.H. Jiang, Lücking &
J.C. Wei and P. multipunctata (G. Merr. ex R. Sant.) Vězda
in the thallus featuring numerous black dots. Apart from
the corticolous growth, it is in fact identical to the latter,
with ascospores 14–15 × 4–5 µm large and breaking into
part spores outside the asci; the ascospore size range for P.
multipunctata was given as 12–16 × 3.5–4.5 µm (Lücking
2008; McCarthy 2009). We therefore synonymize the latter
with P. cinefaciens.
Phylloporis hypothallina (R.C. Harris) S.H. Jiang, Lücking
& Sérus. comb. nov.
MycoBank number: MB 836709; Index Fungorum number: IF 836709; Facesoffungi number: FoF 08884.
Bas.: Strigula hypothallina R.C. Harris, More Florida
Lichens: 149 (1995).
Notes: This corticolous species is also placed in Phylloporis, rather than Dichoporis, based on its thallus anatomy.
Puiggariella Speg., Anal. Soc. Cient. Argentina 12: 99
(1881); Jiang et al., Fung. Div. 102: 280.
MycoBank number: MB 4546; Index Fungorum number:
IF 4546; Facesoffungi number: FoF 08885; four morphologically delimited species (Jiang et al. 2020b); molecular data
available for two species (Jiang et al. 2020b).
= Amoebomyces Bat. & H. Maia, Atas Inst. Micol. Univ.
Pernambuco 2: 353 (1965); Lücking et al., Lichenologist 30:
135 (1998). Type species: Amoebomyces pseudolmediae Bat.
& H. Maia [= Strigula nemathora Mont.].
Lichenized on leaves in terrestrial, lowland to montane,
tropical habitats. Thallus (pseudo-)corticate, light greygreen to whitish green, subcuticular, typically rugoseplicate and with numerous small, white papillae. Photobiont Cephaleuros. Sexual morph: Ascomata perithecia,
erumpent to prominent, pale, usually covered by thallus
layer, wart-shaped, non-carbonaceous, ostiolate. Involucrellum present, pale to brownish. Excipulum prosoplectenchymatous, pale to brownish. Hamathecium comprising
0.5–0.7 µm wide paraphyses, hyaline, flexuose, unbranched
or very rarely branched and anastomosing. Asci 8-spored,
bitunicate, fissitunicate, narrowly obclavate, shortly pedicellate, with narrow ocular chamber, non-amyloid. Ascospores
irregularly arranged to biseriate, fusiform, hyaline, 1-septate,
with thin eusepta and rectangular lumina, smooth-walled,
constricted at the septa. Asexual morph: Pycnidia infrequent, immersed, visible as dark dots. Conidia acrogenous,
either macro- or microconidia; macroconidia 1-septate,
Fungal Diversity (2020) 105:17–318
bacillar, with gelatinous appendages, hyaline; microconidia
usually aseptate, ellipsoid to fusiform, small, hyaline.
Chemistry: No secondary substances known.
Type species: Puiggariella apiahyna Speg. [= Puiggariella nemathora (Mont.) S.H. Jiang, Lücking & J.C.Wei in;
Jiang et al., Fungal Div. 102: 282.
Notes: The only species previously recognized in this
group by Santesson (1952) and subsequent workers, Strigula nemathora, was long considered unique due to its pale,
non-carbonized perithecia, in combination with its thallus
appearing folded and featuring numerous small, white papillae. Phylogenetically, this taxon not only forms a separate
clade, but the available data also indicate that Puiggariella
nemathora is a collective species, including several related
lineages differing largely in their morphology (Jiang et al.
2020b). Currently, three species are accepted in the genus
(Jiang et al. 2020b).
New combinations in Puiggariella:
Puiggariella hypothelia (P.M. McCarthy) S.H. Jiang, Lücking & Sérus. comb. nov.
MycoBank number: MB 836710; Index Fungorum number: IF 836710; Facesoffungi number: FoF 08886.
Bas.: Strigula hypothelia Nyl., Bull. Soc. Linn. Normandie, Sér. 2(2): 520 (1868); S. nemathora var. hypothelia
(Nyl.) R. Sant., Symb. Bot. Upsal. 12(1): 156 (1952); S.
nemathora f. hypothelia (Nyl.) Lücking, Fl. Neotrop., Monogr. 103: 248 (2008).
Raciborskiella Höhnel, Sitz.ber. Math.-Nat. Kl. Akad. Wiss.
Wien, Ser. 1, 118: 1176 (1909); Santesson, Symb. Bot.
Upsal. 12(1): 194 (1952); Jiang et al., Fungal Div. 102: 278.
MycoBank number: MB 4636; Index Fungorum number:
IF 4636; Facesoffungi number: FoF 08887; two morphologically and phylogenetically delimited species (Jiang et al.
2020b).
Lichenized on the underside of leaves in terrestrial,
lowland to montane, tropical habitats. Thallus (pseudo-)
corticate, greenish to bluish grey, subcuticular. Photobiont
Cephaleuros. Sexual morph: Ascomata perithecia, prominent, black, conical, carbonaceous, ostiolate. Involucrellum
present, carbonized. Excipulum prosoplectenchymatous,
dark brown to brown-black. Hamathecium comprising
0.5–0.7 µm wide paraphyses, hyaline, flexuose, unbranched
or very rarely branched and anastomosing. Asci 8-spored
but appearing 9–16-spored due to ascospores breaking in
halves within the asci, bitunicate, fissitunicate, cylindrical to
narrowly obclavate, shortly pedicellate, with narrow ocular
chamber, non-amyloid. Ascospores irregularly arranged to
biseriate, fusiform, hyaline, 1-septate, with thin eusepta and
rectangular lumina, smooth-walled, strongly constricted and
breaking apart at the septa, comparatively large (over 30 µm
and up to 70 µm long). Asexual morph: Pycnidia common,
151
erumpent, visible as black dots. Conidia acrogenous, either
macro- or microconidia; macroconidia 1-septate, bacillar,
with gelatinous appendages, hyaline; microconidia usually
aseptate, ellipsoid to fusiform, small, hyaline.
Chemistry: No secondary substances known.
Type species: Raciborskiella talaumae (Racib.) Höhn.,
Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 118:
1176 (1909).
Notes: This genus was kept separate by Santesson (1952),
on account of the hypophyllous growth and the branched
and anastomosing paraphyses particularly in Raciborskiella
janeirensis. However, molecular data demonstrate that R.
prasina, which differs considerably from R. janeirensis
in morphology and anatomy, is unrelated to the latter and
belongs in the genus currently recognized as Strigula sensu
stricto above. As a result, the most distinctive character of
Raciborskiella are the unusually large ascospores, up to
three to four times longer that in species of Strigula sensu
stricto.
Santesson (1952) included Clypeolum talaumae Racib.
and Melanopsamma areolatum Rehm in the synonymy of
Raciborskiella janeirensis, in spite of the reportedly smaller
ascospores of the former. Revision of type material and
specimens with molecular data showed that indeed two species with differently sized ascospores are involved, the one
with smaller ascospores represented by the eastern paleotropical C. talaumae, which was therefore reinstated (Jiang
et al. 2020b). However, the Brazilian type of M. areolatum
has mostly immature and in part larger ascospores, indicating its conspecificity with R. janeirensis, which appears to
be a neotropical species. Thus, currently two species are
recognized in this genus.
The name ‘Raciborskiella Speg.’ (MycoBank: MB 4637),
a presumed illegitimate later homonym of Raciborskiella
Höhn., does not exist. When Spegazzini (1919) made the
combination R. montana (Racib.) Speg., he did not describe
a new genus, so his use of the name must refer to von Höhnel’s (1909c) genus. This non-lichenized fungus is currently
treated as Trichopeltella montana (Racib.) Höhn. (Hyde
et al. in Hyde et al. 2013; Brefeldiellaceae; Wu et al. 2014).
von Höhnel (1909a) also recombined Micropeltis orbicularis Cooke into Raciborskiella, as R. orbicularis (Cooke)
Höhn.; this name is a synonym of Strigula nitidula (Santesson 1952). Raciborskiella parva L. Xavier is a synonym of
Racoplaca subtilissima (see below), R. zollerniae Bat. &
J.A. Lima [nom. inval.] is the same as S. microspora Lücking (Lücking et al. 1998), and R. minor Vězda is currently
treated in the genus Strigula (Roux and Sérusiaux 2004).
Racoplaca Fée, Essai Crypt Écorc. 1: LVIII, XCIV, XCIX
(1825); Jiang et al., Fung. Div. 102: 285.
MycoBank number: MB 4640; Index Fungorum number: IF 4640; Facesoffungi number: FoF 08888; five
13
152
morphologically delimited species (Jiang et al. 2020b);
molecular data available for two species (Jiang et al. 2020b).
= Heterodothis Syd. in Sydow & Sydow, Philip. J. Sci.
Bot. 9: 170 (1914). Type species: Heterodothis leptotheca
Syd. [= Strigula maculata (Cooke & Massee) R. Sant.].
Lichenized on leaves in terrestrial, lowland to montane,
tropical habitats. Thallus (pseudo-)corticate, olive-brown
to dark (olive-)green, usually with metallic glance, subcuticular and sometimes damaging the supporting leaf,
laciniate, with a thin black, continuous or interrupted line
along the margins of the laciniae. Photobiont Cephaleuros.
Sexual morph: Ascomata perithecia, prominent, black or
covered by thin thallus layer, wart-shaped or mostly conical, carbonaceous, ostiolate. Involucrellum present, carbonized. Excipulum prosoplectenchymatous, dark brown
to brown-black. Hamathecium comprising 0.5–0.7 µm wide
paraphyses, hyaline, flexuose, unbranched or very rarely
branched and anastomosing. Asci 8-spored, bitunicate,
fissitunicate, cylindrical to narrowly obclavate, shortly
pedicellate, with narrow ocular chamber, non-amyloid.
Ascospores irregularly arranged to biseriate, ellipsoid to
fusiform, hyaline, 1-septate, with thin eusepta and rectangular lumina, smooth-walled, at least slightly constricted at
the septa. Asexual morph: Pycnidia common, erumpent,
visible as black dots. Conidia acrogenous, either macro- or
microconidia; macroconidia aseptate or 1-septate, bacillar,
with gelatinous appendages, hyaline; microconidia aseptate, ellipsoid to fusiform, small, hyaline.
Chemistry: No secondary substances known.
Type species: Racoplaca subtilissima Fée, Essai Crypt.
Exot. (Paris): xciv, xcix (1825).
Notes: This newly recognized genus includes some of
the prettiest foliicolous lichens, characterized by narrowly
laciniate thalli bordered by a thin black line. It appears to be
an early diverging clade within Strigula sensu lato but has
no supported relationship with any of the other newly recognized genera (Jiang et al. 2020b). Five species are currently
assigned to this genus (Jiang et al. 2020b).
New combinations in Racoplaca:
Racoplaca melanobapha (Kremp.) S.H. Jiang, Lücking &
J.C. Wei comb. nov.
MycoBank number: MB 836395; Index Fungorum number: IF 836395; Facesoffungi number: FoF 09211.
Bas.: Verrucaria melanobapha Kremp., Lich. Foliic.
Leg. Beccari: 18 (1874); Strigula melanobapha (Kremp.)
R. Sant., Symb. Bot. Upsal. 12(1): 188 (1952).
Notes: This combination was published invalidly in Jiang
et al. (2020b) as the basionym was not indicated.
Serusiauxiella S.H. Jiang, Lücking & J.C.Wei in Jiang et al.,
Fung. Div. 102: 274.
13
Fungal Diversity (2020) 105:17–318
MycoBank number: MB 833565; Index Fungorum number: IF 833565; Facesoffungi number: FoF 08889; three
morphologically and phylogenetically delimited species
(Jiang et al. 2020b).
Lichenized on leaves in terrestrial, lowland to montane,
tropical to subtropical habitats. Thallus (pseudo-)corticate,
light grey-green, subcuticular. Photobiont Trentepohlia.
Sexual morph: Ascomata perithecia, immersed-erumpent,
black but mostly covered by thallus layer, lens-shaped to
wart-shaped, carbonaceous, ostiolate. Involucrellum present, carbonized. Excipulum prosoplectenchymatous, pale.
Hamathecium comprising 0.5–0.7 µm wide paraphyses,
hyaline, flexuose, unbranched or very rarely branched and
anastomosing. Asci 8-spored, bitunicate, fissitunicate, cylindrical to narrowly obclavate, shortly pedicellate, with narrow ocular chamber, non-amyloid. Ascospores irregularly
arranged to biseriate, fusiform, hyaline, 1-septate, with thin
eusepta and rectangular lumina, smooth-walled, constricted
at the septa. Asexual morph: Pycnidia common, immersederumpent, visible as black dots. Conidia acrogenous, either
macro- or microconidia; macroconidia 1-septate, bacillar, with gelatinous appendages becoming extremely long
in squash mounts (as if suspended on gelatinous threads),
hyaline; microconidia aseptate, ellipsoid to fusiform, small,
hyaline.
Chemistry: No secondary substances known.
Type species: Serusiauxiella filifera S.H. Jiang, Lücking
& J.C. Wei in Jiang et al., Fung. Div. 102: 275.
Notes: This entirely novel genus was descovered in eastern Asia, with three species known thus far from China
(Jiang et al. 2020b). It most closely resembles Strigula
sensu stricto and differs mainly in the highly unusual macroconidial appendages. The known species otherwise have
a morphology akin towards S. microspora.
Swinscowia S.H. Jiang, Lücking & Sérus. gen. nov.
MycoBank number: MB 836396; Index Fungorum number: IF 836396; Facesoffungi number: FoF 08890; 34 morphologically delimited species (this paper); molecular data
available for one species (Nelsen et al. 2011a).
Lichenized on bark and rocks in terrestrial, temperatesubalpine to tropical habitats. Thallus usually ecorticate,
whitish to brownish. Photobiont Trentepohlia. Sexual
morph: Ascomata perithecia, usually dispersed but often
dense, immersed-erumpent to prominent, usually black or
covered by thallus layer, rarely pale, lens-shaped to wartshaped, usually carbonaceous, rarely with pale to brownish
walls, ostiolate. Involucrellum usually present and carbonized, rarely reduced or pale. Excipulum prosoplectenchymatous, pale to brown or blackish. Hamathecium comprising
0.5–0.7 µm wide paraphyses, hyaline, flexuose, typically
Fungal Diversity (2020) 105:17–318
branched and sometimes somewhat anastomosing. Asci
usually 8-spored, rarely as little as 2-spored, bitunicate, fissitunicate, cylindrical to narrowly obclavate, shortly pedicellate, with narrow, non–amyloid ocular chamber. Ascospores
irregularly arranged to uni- or biseriate, fusiform to ellipsoid or bacillar, hyaline, 1-septate to mostly 3–7-septate to
(sub-)muriform, with thin eusepta and rectangular lumina,
smooth-walled, often constricted (and sometimes breaking
apart) at the septa. Asexual morph: Pycnidia common,
immersed to erumpent, usually visible as black dots. Conidia
typically acrogenous, either macro- or microconidia; macroconidia (1–)3–7(–11)-septate to more rarely submuriform,
oblong-ellipsoid to bacillar, with variably shaped gelatinous
appendages, hyaline; microconidia aseptate, ellipsoid to
fusiform, small, hyaline.
Chemistry: No secondary substances known.
Type species: Swinscowia jamesii (Swinscow) S.H. Jiang,
Lücking & Sérus. (see below).
Notes: This new genus, named after the British lichenologist Thomas Douglas Victor Swinscow, who first described
the type species, Geissleria jamesii (Swinscow 1967),
is here introduced for non-foliicolous species centered
around Strigula jamesii. Since the latter is the only species
sequenced so far, it is unclear whether the taxa included here
all form a monophyletic group. Given the phylogenetic and
morpho-anatomical differentiation in the foliicolous lineages, likely more than one genus may be recognized for the
non-foliicolous species here placed in Swinscowia. In their
treatment of European taxa, Roux and Sérusiaux (2004)
already arranged the non-foliicolous species into two major
groups, their Strigula affinis-tagananae-calcarea group
(“ensemble 2”) corresponding to Swinscowia.
New combinations in Swinscowia:
Swinscowia affinis (A. Massal.) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836397; Index Fungorum number: IF 836397; Facesoffungi number: FoF 08891.
Bas.: Sagedia affinis A. Massal., Memor. Lich.: 138
(1853); Spermatodium affine (A. Massal.) Trevis., Conspect. Verruc.: 10 (1860); Segestria affinis (A. Massal.)
Zwackh, Flora 45: 550 (1862); Verrucaria affinis (A. Massal.) Cromb., J. Bot., Lond. 14: 360 (1876); Porina affinis
(A. Massal.) Zahlbr., Öst. Bot. Z. 51: 277 (1901); Arthopyrenia affinis (A. Massal.) R.C. Harris, Michigan Bot. 12: 10
(1973); Strigula affinis (A. Massal.) R.C. Harris in Hawksworth et al., Lichenologist 12: 107 (1980).
= Pyrenula minuta Nägeli, in Hepp, Flecht. Europ.: no.
458 (1857); Sagedia affinis var. sporophorum Wartm. & B.
Schenk, Schweiz. Kryptogam.: no. 673.
Notes: The name Sagedia affinis var. sporophorum was
published in Wartmann & Schenk’s Exsiccate Schweizerische Kryptogamen, listing Pyrenula minuta based on
153
Hepp’s Flechten Europas no. 458 as synonym. The variety
name is thus validly published, as it formally constitutes a
replacement name at species level, based on the same type.
Swinscowia albicascens (Nyl.) S.H. Jiang, Lücking & Sérus.
comb. nov.
MycoBank number: MB 836398; Index Fungorum number: IF 836398; Facesoffungi number: FoF 08892.
Bas.: Verrucaria albicascens Nyl., Lich. Nov. Zeland.
(Paris): 129 (1888); Porina albicascens (Nyl.) Müll. Arg.,
Bull. Soc. R. Bot. Belg. 31(2): 38 (1892); Segestria albicascens (Nyl.) Hellb., Bih. K. Svenska Vetensk Akad. Handl.,
Afd. 3, 21(13): 132 (1896); Strigula albicascens (Nyl.) R.C.
Harris, More Florida Lichens: 154 (1995).
= Verrucaria indutula Nyl., Lich. Nov. Zeland.: 129
(1888); Porina indutula (Nyl.) Müll. Arg., Bull. Herb. Boissier 2(1): 92 (1894); Segestria indutula (Nyl.) Hellb., Bih. K.
Svenska Vetensk. Akad. Handl., Afd. 3, 21(13): 133 (1896);
Strigula indutula (Nyl.) R.C. Harris, More Florida Lichens:
157 (1995).
Notes: See discussion under Swincsowia glabra.
Swinscowia albolinita (Nyl.) S.H. Jiang, Lücking & Sérus.
comb. nov.
MycoBank number: MB 836399; Index Fungorum number: IF 836399; Facesoffungi number: FoF 08893.
Bas.: Verrucaria albolinita Nyl., Lich. Ins. Guin.: 38
(1889); Microglaena albolinita (Nyl.) Zahlbr., Cat. Lich.
Univers. 1: 188 (1921); Phyllobathelium albolinitum (Nyl.)
H. Mayrhofer, Biblioth. Lichenol. 26: 79 (1987); Strigula
albolinita (Nyl.) R.C. Harris [as ‘albolinitum’], More Florida Lichens: 155 (1995).
Swinscowia alpestris (Vězda) S.H. Jiang, Lücking & Sérus.
comb. nov.
MycoBank number: MB 836400; Index Fungorum number: IF 836400; Facesoffungi number: FoF 08894.
Bas.: Porina faginea var. alpestris Vězda, Acta Mus.
Silesiae, Ser. A, 10: 5 (1961); Arthopyrenia faginea var.
alpestris (Vězda) Swinscow, Lichenologist 3: 73 (1965);
Strigula stigmatella var. alpestris (Vězda) Coppins in Hawksworth et al., Lichenologist 12: 107 (1980); Strigula alpestris (Vězda) Hafellner, Mitt. Naturw. Ver. Steierm. 132:
131 (2002).
Notes: We follow here Hafellner (2002) in separating
this taxon from Swinscowia stigmatella (see also discussion
under S. glabra), on account of the different habitat and substrate ecology and the longer asci. McCarthy & Malcolm
(1996) reported Strigula stigmatella var. alpestris as silicicolous from Australia, but later this was apparently reidentified as Strigula decipiens.
13
154
Swinscowia amphora (Aptroot & Lücking) S.H. Jiang,
Lücking & Sérus. comb. nov.
MycoBank number: MB 836401; Index Fungorum number: IF 836401; Facesoffungi number: FoF 08895.
Bas.: Strigula amphora Aptroot & Lücking in Aptroot
et al., Biblioth. Lichenol. 97: 130 (2008).
Swinscowia aquatica (H. Harada) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836402; Index Fungorum number: IF 836402; Facesoffungi number: FoF 08896.
Bas.: Strigula aquatica H. Harada, Nova Hedwigia 66:
420 (1998).
Swinscowia australiensis (P.M. McCarthy) S.H. Jiang,
Lücking & Sérus. comb. nov.
MycoBank number: MB 836403; Index Fungorum number: IF 836403; Facesoffungi number: FoF 08897.
Bas.: Strigula australiensis P.M. McCarthy, Muelleria 8:
323 (1995).
Swinscowia bahamensis (Riddle) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836404; Index Fungorum number: IF 836404; Facesoffungi number: FoF 08898.
Bas.: Lithothelium bahamense Riddle in Britton & Millspaugh, The Bahama Flora: 532 (1920); Strigula bahamensis
(Riddle) R.C. Harris, More Florida Lichens: 155 (1995)
Swinscowia bispora (Aptroot & K.H. Moon) S.H. Jiang,
Lücking & Sérus. comb. nov.
MycoBank number: MB 836405; Index Fungorum number: IF 836405; Facesoffungi number: FoF 08899.
Bas.: Strigula bispora Aptroot & K.H. Moon, Herzogia
27: 354 (2014).
Fungal Diversity (2020) 105:17–318
MycoBank number: MB 836408; Index Fungorum number: IF 836408; Facesoffungi number: FoF 08902.
Bas.: Strigula confusa Fryday, Coppins & Common in
Fryday & Coppins, Lichenologist 36: 92 (2004).
Swinscowia decipiens (Malme) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836409; Index Fungorum number: IF 836409; Facesoffungi number: FoF 08903.
Bas.: Porina decipiens Malme, Ark. Bot. 23A(1):
26 (1929); Strigula decipiens (Malme) P.M. McCarthy,
Lichenologist 29: 513 (1997).
Swinscowia divisa (P.M. McCarthy) S.H. Jiang, Lücking &
Sérus. comb. et stat. nov.
MycoBank number: MB 836410; Index Fungorum number: IF 836410; Facesoffungi number: FoF 08904.
Bas.: Strigula decipiens var. divisa P.M. McCarthy,
Lichenologist 32: 34 (2000).
Notes: McCarthy (2000) considered this taxon a subspecies of S. decipiens, stating that the only difference
were occasional longitudinal septa in otherwise similarly
sized ascospores. However, not only is ascospore septation an important taxonomic criterion in this genus, but
the ascospores of S. decipiens are also distinctly longer and
narrower (see key below). As the few other known species
with submuriform ascospores are different in ascospore size,
morphology and/or substrate ecology, the recognition of S.
divisa as a separate species is justified.
Swinscowia calcarea (Bricaud & Cl. Roux) S.H. Jiang,
Lücking & Sérus. comb. nov.
MycoBank number: MB 836406; Index Fungorum number: IF 836406; Facesoffungi number: FoF 08900.
Bas.: Strigula calcarea Bricaud & Cl. Roux, Bull. Soc.
Linn. Provence 42: 131 (1991).
Swinscowia endolithea (Cl. Roux & Bricaud) S.H. Jiang,
Lücking & Sérus. comb. nov.
MycoBank number: MB 836411; Index Fungorum number: IF 836411; Facesoffungi number: FoF 08905.
Bas.: Strigula endolithea Cl. Roux & Bricaud in Roux
& Sérusiaux, Biblioth. Lichenol. 90: 88 (2004); S. endolithea Cl. Roux & Bricaud, Bull. Soc. Linn. Provence 44:
129 (1993) [nom. inval.; ICN Art. 36.1].
Notes: Tretiach (2006) provided a detailed description of
the perithecia, asci, and ascospores of this species, which
originally was described based on pycnidia and macroconidia only (Roux and Sérusiaux 2004).
Swinscowia cavicola (Cl. Roux & Bricaud) S.H. Jiang,
Lücking & Sérus. comb. nov.
MycoBank number: MB 836407; Index Fungorum number: IF 836407; Facesoffungi number: FoF 08901.
Bas.: Strigula cavicola Cl. Roux & Bricaud in Roux &
Sérusiaux, Biblioth. Lichenol. 90: 87 (2004).
Swinscowia fracticonidia (R.C. Harris) S.H. Jiang, Lücking
& Sérus. comb. nov.
MycoBank number: MB 836412; Index Fungorum number: IF 836412; Facesoffungi number: FoF 08906.
Bas.: Strigula fracticonidia R.C. Harris, More Florida
Lichens: 156 (1995).
Swinscowia confusa (A. Massal.) S.H. Jiang, Lücking &
Sérus. comb. nov.
Swinscowia glabra (A. Massal.) S.H. Jiang, Lücking &
Sérus. comb. nov.
13
Fungal Diversity (2020) 105:17–318
MycoBank number: MB 836413; Index Fungorum number: IF 836413; Facesoffungi number: FoF 08907.
Bas.: Sagedia glabra A. Massal., Ric. Auton. Lich. Crost.:
161 (1852); Spermatodium glabrum (A. Massal.) Trevis.,
Conspect. Verruc.: 11 (1860); Porina glabra (A. Massal.)
Zahlbr., Cat. Lich. Univers. 1: 382 (1922); Arthopyrenia
glabra (A. Massal.) J. Nowak & Tobol., Flora Polska: 1117
(1975); Strigula glabra (A. Massal.) V. Wirth, Flechtenflora:
531 (1980).
= Pyrenula netrospora Nägeli, Flecht. Europ.: no. 461
(1857); Spermatodium netrosporum (Nägeli) Trevis., Conspect. Verruc.: 11 (1860); Sagedia netrospora (Nägeli) Anzi,
Comm. Soc. Crittog. Ital. 1(3): 162 (1862); Arthopyrenia
netrospora (Nägeli) Müll. Arg., Mém. Soc. Phys. Hist. Nat.
Genève 16(2): 429 (1862); Verrucaria netrospora (Nägeli)
Nyl., Jahresber. St. Gall. Naturw. Ges.: 504 (1882); Porina
netrospora (Nägeli) Lettau, Hedwigia 52: 104 (1912).
= Sagedia candida Anzi, Comm. Soc. Crittog. Ital. 1(3):
162 (1862).
Notes: Swinscowia glabra forms a species complex with
S. albicascens and S. stigmatella, all three growing on bark
and producing chiefly 7-septate ascospores with overlapping
size ranges. There seem to be no discernable differences
between the Australasian S. albicascens and the European S.
glabra. Roux and Sérusiaux (2004) do not discuss S. albicascens in the context of S. glabra, and McCarthy (2009) does
not mention S. glabra in the discussion of S. albicascens,
although he did synonymize Verrucaria indutula with the
latter. We keep S. glabra and S. albicascens separate for
the time being, as macroconidia have not yet been reported
for the latter and might turn out to provide a diagnostic difference, although we consider this unlikely. Roux & Sérusiaux (2004) discuss the similarity between S. glabra and
S. stigmatella and keep both separate on account of slightly
different ascospore length (22–26 µm in S. glabra vs. 26–36
µm in S. stigmatella) and differences in substrate (smooth
bark in S. glabra and rough bark in S. stigmatella). Harris
(1995) gives ascospore length for North American material
of S. stigmatella as 24–42 µm, whereas McCarthy (2009)
cites 16–32 µm for Australasian S. albicascens. McCarthy
(1993) reported S. stigmatella as saxicolous from Australia
but later corrected these records to S. decipiens (McCarthy
2009). Thus, the taxonomic treatments for corticolous taxa
with 7-septate ascospores are inconsistent between North
America, Europe, and Australasia, using one name in North
America (stigmatella), two in Europe (glabra, stigmatella),
and one but a different name in Australasia (albicascens).
This problem is aggravated by the notion that the type of
Lichen stigmatellus Ach. grows on smooth bark and apparently fits the concept of S. glabra by Roux & Sérusiaux
(2004). This requires a critical revision of the available
material and a reassessment of the types to properly name
the taxa distinguished in this complex.
155
Swinscowia griseonitens (R.C. Harris) S.H. Jiang, Lücking
& Sérus. comb. nov.
MycoBank number: MB 836414; Index Fungorum number: IF 836414; Facesoffungi number: FoF 08908.
Bas.: Strigula griseonitens R.C. Harris, More Florida
Lichens: 156 (1995).
Swinscowia jamesii (Swinscow) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836415; Index Fungorum number: IF 836415; Facesoffungi number: FoF 08909.
Bas.: Geisleria jamesii Swinscow, Lichenologist 3: 420
(1967); Strigula jamesii (Swinscow) R.C. Harris in Hawksworth et al., Lichenologist 12(1): 107 (1980).
Swinscowia johnsonii (P.M. McCarthy) S.H. Jiang, Lücking
& Sérus. comb. nov.
MycoBank number: MB 836416; Index Fungorum number: IF 836416; Facesoffungi number: FoF 08910.
Bas.: Strigula johnsonii P.M. McCarthy, Muelleria 8: 324
(1995).
Swinscowia laceribracae (R.C. Harris) S.H. Jiang, Lücking
& Sérus. comb. nov.
MycoBank number: MB 836417; Index Fungorum number: IF 836417; Facesoffungi number: FoF 08911.
Bas.: Strigula laceribracae R.C. Harris, More Florida
Lichens: 158 (1995).
Swinscowia muriconidiata (Aptroot, L.I. Ferraro & M.
Cáceres) S.H. Jiang, Lücking & Sérus. comb. nov.
MycoBank number: MB 836418; Index Fungorum number: IF 836418; Facesoffungi number: FoF 08912.
Bas.: Strigula muriconidiata Aptroot, L.I. Ferraro & M.
Cáceres, Lichenologist 46: 99 (2014).
Notes: This taxon may be the macroconidial stage of S.
obtecta.
Swinscowia muriformis (Aptroot & Diederich) S.H. Jiang,
Lücking & Sérus. comb. nov.
MycoBank number: MB 836419; Index Fungorum number: IF 836419; Facesoffungi number: FoF 08913.
Bas.: Strigula muriformis Aptroot & Diederich in Aptroot
et al., Biblioth. Lichenol. 64: 188 (1997).
Swinscowia muscicola (F. Berger, Coppins, Cl. Roux &
Sérus.) S.H. Jiang, Lücking & Sérus. comb. nov.
MycoBank number: MB 836420; Index Fungorum number: IF 836420; Facesoffungi number: FoF 08914.
Bas.: Strigula muscicola F. Berger, Coppins, Cl. Roux
& Sérus. in Sérusiaux et al., Lichenologist 37: 481 (2005).
13
156
Swinscowia obtecta (Vain.) S.H. Jiang, Lücking & Sérus.
comb. nov.
MycoBank number: MB 836421; Index Fungorum number: IF 836421; Facesoffungi number: FoF 08915.
Bas.: Thelenella obtecta Vain., Acta Soc. Fauna Flora
Fenn. 7(2): 218 (1890); Polyblastiopsis obtecta (Vain.)
Zahlbr., Cat. Lich. Univers. 1: 351 (1922); Phyllobathelium
obtectum (Vain.) H. Mayrhofer, Biblioth. Lichenol. 26: 82
(1987); Strigula obtecta (Vain.) R.C. Harris, More Florida
Lichens: 158 (1995).
Swinscowia pallida (Aptroot & K.H. Moon) S.H. Jiang,
Lücking & Sérus. comb. nov.
MycoBank number: MB 836422; Index Fungorum number: IF 836422; Facesoffungi number: FoF 08916.
Bas.: Strigula pallida Aptroot & K.H. Moon in Moon &
Aptroot, Biblioth. Lichenol. 99: 308 (2009); Strigula pallidocarpa Aptroot & K.H. Moon, Herzogia 27: 354 (2014)
[nom. illeg., ICN Art. 52.1] [non Strigula pallida Kalchbr.
in Stizenberger, Ber. Tät. St Gall. Naturw. Ges.: 218 (1891);
nom. inval., ICN Art. 38.1–3].
Notes: Aptroot and Moon (2014) provided a replacement
name for their Strigula pallida, described five years prior
(Moon and Aptroot 2009), under the impression that the
name S. pallida had already been used for another species,
namely S. pallida Kalchbr. in Stizenberger (1891). However,
as already pointed out by Santesson (1952), that name is
invalid, as it does not have a validating description [ICN Art.
38.3] and hence has no standing under the Code [ICN Art.
12.1]. Thus, the name S. pallida Aptroot & K.H. Moon is
legitimate, although it does not follow the recommendation
of the Code to avoid giving a previously invalidly published
name to a different taxon [ICN Rec. 38C.1].
Swinscowia porinoides (Canals, Boqueras & Gómez-Bolea)
S.H. Jiang, Lücking & Sérus. comb. nov.
MycoBank number: MB 836423; Index Fungorum number: IF 836423; Facesoffungi number: FoF 08917.
Bas.: Strigula porinoides Canals, Boqueras & GómezBolea, Mycotaxon 55: 391 (1995).
Swinscowia rhodinula (Zahlbr.) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836424; Index Fungorum number: IF 836424; Facesoffungi number: FoF 08918.
Bas.: Porina rhodinula Zahlbr., Denkschr. Akad. Wiss.
Wien Math.-Naturwiss. Kl. 104: 254 (1941).; Strigula rhodinula (Zahlbr.) P.M. McCarthy, Australas. Lichenol. 78: 3
(2016).
Swinscowia rostrata (R.C. Harris & Aptroot) S.H. Jiang,
Lücking & Sérus. comb. nov.
13
Fungal Diversity (2020) 105:17–318
MycoBank number: MB 836425; Index Fungorum number: IF 836425; Facesoffungi number: FoF 08919.
Bas.: Strigula rostrata R.C. Harris & Aptroot in Harris,
More Florida Lichens: 159 (1995).
Swinscowia rupestris (P.M. McCarthy) S.H. Jiang, Lücking
& Sérus. comb. nov.
MycoBank number: MB 836426; Index Fungorum number: IF 836426; Facesoffungi number: FoF 08920.
Bas.: Strigula rupestris P.M. McCarthy, Lichenologist
29: 518 (1997).
Swinscowia stigmatella (Ach.) S.H. Jiang, Lücking & Sérus.
comb. nov.
MycoBank number: MB 836427; Index Fungorum number: IF 836427; Facesoffungi number: FoF 08921.
Bas.: Lichen stigmatellus Ach., Lich. Suec. Prodr.: 15
(1799); Verrucaria stigmatella (Ach.) Ach., Methodus, Sectio Prior: 117 (1803): Verrucaria punctiformis var. stigmatella (Ach.) Ficinus & C. Schub., Fl. Dresden 2: 147 (1823);
Arthopyrenia stigmatella (Ach.) A. Massal., Symmict. Lich.:
119 (1855); Leiophloea punctiformis var. stigmatella (Ach.)
Trevis., Conspect. Verruc.: 9 (1860); Verrucaria epidermidis var. stigmatella (Ach.) Garov., Tentam. Dispos. Lich.
Langob.: 85 (1865); Verrucaria epidermidis f. stigmatella
(Ach.) Malbr., Bull. Soc. Amis Sci. Nat. Rouen, Sér. II, 5:
315 (1869); Arthopyrenia cinereopruinosa var. stigmatella
(Ach.) Jatta, Syll. Lich. Ital.: 530 (1900); Didymella stigmatella (Ach.) Szatala in Degen, Flora Velebitica 3: 309
(1938); Strigula stigmatella (Ach.) R.C. Harris in Hawksworth et al., Lichenologist 12: 107 (1980).
= Porina faginea var. alpestris Vězda, Acta Mus. Silesiae, Ser. A 10: 5 (1961); Arthopyrenia faginea var. alpestris (Vězda) Swinscow, Lichenologist 3: 73 (1965); Strigula
stigmatella var. alpestris (Vězda) Coppins in Hawksworth
et al., Lichenologist 12: 107 (1980); Strigula alpestris
(Vězda) Hafellner, Mitt. Naturw. Ver. Steierm. 132: 131
(2002).
MycoBank: MB 351488
Notes: See discussion under Swincsowia glabra.
Swinscowia submuriformis (R.C. Harris) S.H. Jiang, Lücking & Sérus. comb. nov.
MycoBank number: MB 836428; Index Fungorum number: IF 836428; Facesoffungi number: FoF 08922.
Bas.: Arthopyrenia submuriformis R.C. Harris, Michigan
Bot. 12: 15 (1973); Strigula submuriformis (R.C. Harris)
R.C. Harris in Egan, Bryologist 90: 164 (1987).
Swinscowia tagananae (Harm.) S.H. Jiang, Lücking &
Sérus. comb. nov.
MycoBank number: MB 836429; Index Fungorum number: IF 836429; Facesoffungi number: FoF 08923.
Fungal Diversity (2020) 105:17–318
Bas.: Verrucaria tagananae Harm. in Pitard & Harmand,
Bull. Soc. Bot. Fr. 58(22): 68 (1911); Polyblastiopsis tagananae (Harm.) Zahlbr., Cat. Lich. Univers. 1: 352 (1922);
Strigula tagananae (Harm.) R.C. Harris, More Florida
Lichens: 160 (1995).
= Strigula lateralis Aptroot & van den Boom, Mycotaxon
56: 3 (1995).
MycoBank: MB 414225
157
8.
9.
9.
10.
10.
Swinscowia thelopsidoides (Coppins, Cl. Roux & Sérus.)
S.H. Jiang, Lücking & Sérus. comb. nov.
MycoBank number: MB 836430; Index Fungorum number: IF 836430; Facesoffungi number: FoF 08924.
Bas.: Strigula thelopsidoides Coppins, Cl. Roux & Sérus.
in Roux & Sérusiaux, Biblioth. Lichenol. 90: 90 (2004).
Key to species of Swinscowia:
1.
1.
2.
2.
3.
3.
4.
4.
5.
5.
6.
6.
7.
7.
8.
Ascospores transversely 3–7(–9)-septate, very rarely
few ascospores with some longitudinal septa, mostly
less than 7 µm broad; macroconidia transversely septate……………………………………………………2
Ascospores regularly submuriform to muriform (i.e.
most or all ascospores with at least a few longitudinal septa); macroconidia transversely septate to muriform…………………………………………………17
On bark………………………………………………3
On rock (or rarely mosses over rock)…………………9
Ascospores regularly 3-septate; macroconidia (where
known) 13–18 µm long………………………………4
Ascospores irregularly 3–7(–9)-septate to regularly
7-septate, very rarely few ascospores with some longitudinal septa; macroconidia (where known) 18–30
µm long………………………………………………7
Perithecial with carbonized involucrellum, black; pycnidia with macroconidia frequent; macroconidia 13–18
× 2.5–4 µm; chiefly Europe…………………………5
Perithecial lacking or with non-carbonized involucrellum, pale to orange-brown; pycnidia with macroconidia unknown………………………………………6
Ascospores 15–24 × 4.5–6 µm……Swinscowia affinis
Ascospores 13–17 × 3.5–5.5 µm…Swinscowia jamesii
Ascospores 17–22 × 6–8 µm; perithecia pale; Korea
……………………………………Swinscowia pallida
Ascospores 10–14 × 2.5–3.5 µm; perithecia orangebrown; Europe…………Swinscowia thelopsidioides
Ascospores 25–42 µm long, largely over 30 µm; typically on rough bark; subcosmopolitan………………
………………………………Swinscowia stigmatella
Ascospores 16–30 µm long; typically on smooth
bark…………………………………………………8
Collected in Europe; macroconidia 18–28 × 3.5–4
µm…………………………………Swinscowia glabra
11.
11.
12.
12.
13.
13.
14.
14.
15.
15.
16.
16.
17.
17.
18.
18.
19.
20.
Collected in Australasia, including New Zealand; macroconidia unknown………… Swinscowia albicascens
Notes: See discussion for the three species keyed out
under couplets 7 and 8 under Swincsowia glabra
On calcareous rock (limestone)……………………10
On siliceous rock (granite) or over mosses…………14
Perithecial with carbonized involucrellum, black; extraEuropean……………………………………………11
Perithecial lacking involucrellum, pale; Europe
………………………………………………………12
Perithecia with lateral ostioles, fused; ascospores irregularly 3–7-septate; Caribbean…Swinscowia bahamensis
Perithecia with apical ostioles, solitary; ascospores regularly 3-septate; New Zealand…Swinscowia rhodinula
Ascospores (5–)7-septate; macroconidia (3)4–7-septate, 20–30 µm long……… Swinscowia endolithea
Ascospores (where known) 3-septate; macroconidia
3(–4)-septate, 10–20 µm long………………………13
Macroconidia 15–20 µm long…Swinscowia calcarea
Macroconidia 10–15 µm long…Swinscowia cavicola
Notes: Strigula cavicola is here keyed out together
with S. calcarea following Roux and Sérusiaux (2004),
who hypothesized a close relationship between these
taxa. However, perithecia and ascospores are thus far
unknown from S. cavicola
Ascospores 3-septate, 18–21 µm long; over mosses;
Europe………………………Swinscowia muscicola
Ascospores 7(–9)-septate, 25–42 µm long; on rock,
rarely over mosses on rock…………………………15
Over mosses on rock; involucrellum carbonized but
developed only around ostiole…Swinscowia alpestris
Directly on rock; involucrellum carbonized and reaching down to substrate, or involucrellum pale to brownish…………………………………………………16
Involucrellum carbonized, reaching down to substrate;
pantropical……………………Swinscowia decipiens
Involucrellum pale to brownish, developed only around
ostiole; Japan……… …………Swinscowia aquatica
Notes: Swinscowia aquatica was considered by
McCarthy (2001) to possibly represent a pale form of
S. decipiens, but the structure of the involucrellum suggests a closer relationship with S. alpestris
On bark………………………………………………18
On rock (or rarely mosses over rock)………………27
Ascospores 20–33 × 6–10 µm long, submuriform with
few longitudinal septa; macroconidia transversely septate to submuriform…………………………………19
Ascospores 35–135 × 8–48 µm long, regularly
muriform; macroconidia submuriform to muriform…………………………………………………23
Perithecial wall pale; North America………………
………………………………Swinscowia griseonitens
Perithecial wall carbonized…………………………20
13
158
20. Pycnidia producing macroconidia rostrate; macroconidia submuriform; Madagascar………………………
…………………………………Swinscowia rostrata
21. Pycnidia producing macroconidia not rostrate; macroconidia transversely septate…………………………21
21. Ostiole lateral; macroconidia 7–11-septate,
35–45 × 4–5 µm; Europe, Australia………
………………………………Swinscowia tagananae
22. Ostiole apical; macroconidia 3–7-septate, 20–25 × 3–10
µm…………………………………………………22
22. Macroconidia 3-septate, 15–20 × 3–4 µm, about 5–6
times as long as broad, fragmenting into part spores;
New Zealand………………Swinscowia fracticonidia
23. Macroconidia 7-septate, 20–25 × 8–10 µm, about 2–3
times as long as broad, remaining intact; North America………………………Swinscowia submuriformis
23. Ascomata unknown; macroconidia regularly muriform,
90–105 × 30–35 µm; South America…………………
……………………………Swinscowia muriconidiata
24. Ascomata present; macroconidia unknown………24
24. Ostiole lateral; ascospores 42–50 × 14–17 µm, about
2.5–3.5 times as long as broad; Neotropics (Costa
Rica)……………………………Swinscowia amphora
25. Ostiole apical; ascospores variously sized but relatively narrower, about 3.5–4.5 times as long as
broad………………………………………………25
25. Ascospores 115–135 × 38–48 µm, 2/ascus;
Korea……………………………Swinscowia bispora
26. Ascospores 35–90 × 8–26 µm, 8/ascus………………26
26. Ascospores 35–45 × 8–10 µm; perithecial wall pale;
North America…………… Swinscowia laceribracae
27. Ascospores 60–90 × 18–26 µm; perithecial wall carbonized but covered by thallus; Neotropics (Brazil)…
……………………………………Swinscowia obtecta
27. On calcareous rock (limestone) or mosses over limestone; Europe………………………………………28
28. On siliceous rock (granite); Africa, Australasia……………………………………………………29
28. Directly on rock; ascospores submuriform, 20–33 ×
5–10 µm; macroconidia submuriform, 14–27 × 4–9
µm……………………………Swinscowia porinoides
29. On mosses over rock; ascospores muriform, 35–60 ×
7–12 µm; macroconidia unknown……………………
…………………………………Swinscowia confusa
29. Ascospores 16–36 × 5.5–11.5 µm…………………30
30. Ascospores 37–63 × 10–27 µm……………………32
30. Macroconidia muriform, 19–30 × 6–9 µm; ascospores
muriform, 23–36 × 7–11.5 µm; continental Australia…
……………………………Swinscowia australiensis
30. Macroconidia 3-septate, 11–20 × 3–4.5 µm, or macroconidia unknown; ascospores submuriform, 16–30 ×
5.5–9 µm……………………………………………31
13
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31. Perithecia 0.2–0.3 mm diam.; involucrellum 30–60 µm
thick; thallus thin; Cook Islands………………………
……………………………………Swinscowia divisa
31. Perithecia 0.3–0.6 mm diam.; involucrellum 50–120
µm thick; thallus thick; Lord Howe Island……………
…………………………………Swinscowia rupestris
32. Perithecial wall pale; ascospores 2–2.5 times as
long as broad; African Paleotropics (São Tomé and
Príncipe)………………………Swinscowia albolinita
32. Perithecial wall carbonized; ascospores 2.5–4 times as
long as broad………………………………………33
33. Perithecia 0.2–0.3 mm diam.; Papua New Guinea……
………………………………Swinscowia muriformis
33. Perithecia 0.4–0.8 mm diam.; New Zealand…………
…………………………………Swinscowia johnsonii
Excluded genera:
Oletheriostrigula Huhndorf & R.C. Harris, Brittonia 48:
551 (1996).
MycoBank number: MB 27751; Index Fungorum number:
IF 27751; Facesoffungi number: FoF 08925.
Type species: Oletheriostrigula papulosa (Durieu &
Mont.) Huhndorf & R.C. Harris, Brittonia 48: 551 (1996);
Sphaeria papulosa Durieu & Mont., Expl. Sci. Alg., Fl.
Algér. 1(14): 536 (1848) [1846–49]; Leptosphaeria papulosa (Durieu & Mont.) Sacc., Nuovo G. Bot. Ital. 7: 313
(1875); Metasphaeria papulosa (Durieu & Mont.) Sacc.,
Syll. Fung. 2: 168 (1883); Massarina papulosa (Durieu &
Mont.) S.K. Bose, Phytopath. Z. 41: 176 (1961).
Notes: Huhndorf and Harris (1996) suggested placement of this unique, non-lichenized, apparently saprotrophic fungus in Strigulaceae, based on hamathecium and
ascus structure. Close relationship of non-lichenized and
lichenized fungi has been demonstrated for other lineages,
such as Heleiosa in the Monoblastiaceae (Nelsen et al.
2011a; Lücking et al. in Hongsanan et al. 2020, this volume;
Monoblastiaceae). The paraphyses, asci and ascospores of
Oletheriostrigula indeed closely resemble those of e.g. Flavobathelium. However, based on Bose (1961), Huhndorf and
Harris (1996) gave the asexual morph of Oletheriostrigula
as Diplodia stage, which is described as forming rather
large (around 20–40 × 10–15 µm), aseptate, oval-ellipsoid,
rather thick-walled conidia that remain mostly hyaline but
partially become brown and septate with age and never
produce gelatinous appendages (Burgess et al. 2003; Alves
et al. 2004; Damm et al. 2007; Phillips et al. 2007). Bose
(1961) described the conidia of Massarina populosa as 7–16
× 2.5–5 µm in size, gradually (not abruptly in age) turning brown, and irregularly pyriform-clavate to elliptical,
thus different from Diplodia sensu stricto. Even so, while
the conidia of Oletheriostrigula do not seem to represent a
Fungal Diversity (2020) 105:17–318
genuine Diplodia asexual morph, they are also very different from macroconidia in Strigulaceae, which are septate to
muriform, persistently hyaline, and with gelatinous appendages; hence Oletheriostrigula must be excluded from that
family.
On a side note, the genus Diplodia was established in
Montagne (1834: 302), ascribed to Fries, based on Sphaeria mutila Fr., with the description “Asci elliptico-oblongi,
didymi, sporidiis binis referti”. Also, Fries (1823) had originally described the species in Sphaeria, which he explicitly
characterized as producing asci. In spite of this, the name
Diplodia has become firmly established for a frequently
encountered, economically important asexual morph of
various ascomycetous fungi (Kirk et al. 2008). This interpretation, which is in conflict with the protologue, goes
back to Stevens (1933), who reportedly saw only pycnidia
with conidia in the material identified in Montagne (1834;
cited as no. 498) and therefore established the name Diplodia mutila (Fr.) Mont. [sic] as having priority over the
previously used name Sphaeropsis malorum (Berk.) Berk.
for said asexual morph. The ascospores and conidia of Botryosphaeria stevensii, the current name covering Sphaeria
mutila and Diplodia mutila (Shoemaker 1964; Alves et al.
2004), are remarkably similar in size, shape, and development, remaining aseptate and hyaline for a long time and
eventually becoming brown and septate in part. Thus, in
superficial observation, it is conceivable that pycnidia with
conidia might be mistaken for perithecia with asci, and viceversa. However, if that was the case and the original material
seen by Fries and Montagne only contained pycnidia, one
would still have to proof that neither author actually saw at
least one perithecium with asci, in order to superseed the
explicit description of asci in the protologue of Diplodia
(Montagne 1834). That seems impossible, given that both
stages may occur together, both perithecia and pycnidia are
formed in black stromata, and perithecia possibly present in
the original material might have been used up by previous
examinations. Fries (1849) himself later elaborated on the
formation of ascospores and conidia in otherwise identical fruiting bodies of Sphaeria sensu lato species, including Diplodia. According to Sutton (1980) and Alves et al.
(2004), original material of Sphaeria mutila seen by Fries
(Scleromycetes Suecici 164, 385) contains only empty fruiting bodies, which at least in one case (164) were interpreted
by Sutton (1980) as ascomata, who therefore suggested to
adopt the name for a sexual morph. Alves et al. (2004) also
argued that Montagne (1834) did not ascribe to Fries a new
combination for Sphaeria mutila Fr., as Diplodia mutila (Fr.)
Fr., but instead ascribed to that author a new species, Diplodia mutila Fr. This is incorrect, as Montagne (1834, p. 302)
clearly states that “… this species …”, namely Sphaeria
mutila Fr., will be the type of Diplodia Fr., providing a valid
description for the latter, ascribed to Fries. Hence, the genus
159
Diplodia is to be based on Sphaeria mutila, with its type to
be cited as D. mutila (Fr.) Fr., and cannot be interpreted otherwise. This also means that the relevant elements to assess
the status of Diplodia as representing a sexual or asexual
morph genus are the material used by Fries (Scleromycetes
Suecici 164, 385), with only empty fruiting bodies at least
partly interpreted as perithecia, plus Fries’s description in
Montagne (1834) explicitly describing asci. The assessment
of Stevens (1933) and Alves et al. (2004) of Montagne’s
no. 498 is insofar irrelevant as it does not proof that Fries’s
original material contained only pycnidia and that therefore
Fries’s placement of the species in a genus characterized by
the formation of asci was in error; neither does it proof that
Fries did not see asci in Montagne’s material either. Consequently, if Sphaeria mutila is accepted as name of a sexual
morph and placed in the genus Botryosphaeria Ces. & De
Not., Diplodia Fr. has priority over the latter name.
On top of this, there is also confusion with the correct
name applying to the species originally described as Sphaeria mutila and currently known as Botryosphaeria stevensii Shoemaker (Shoemaker 1964; Alves et al. 2004). Stevens (1936) claimed to have discovered the sexual morph
of Sphaeria mutila (Fr.) for the first time and proposed
the combination into Physalospora, as P. mutila (Fr.) N.E.
Stevens. At the time when Shoemaker (1964) treated this
taxon, the Montreal Code (Lanjouw et al. 1961) was in
effect, which included Art. 59 on dual nomenclature and
the phrase: “The author who first describes a perfect state
may adopt the specific epithet applied to the corresponding
imperfect state, but his binomial for the perfect state is to be
attributed to him alone, and is not to be regarded as a new
combination.” Thus, assuming that Stevens’s (1936) description was the first discovery of the sexual morph and applying the Montreal Code, Shoemaker (1964) changed Steven’s
name to Physalospora mutila N.E. Stevens, i.e. a new taxon
with a type different from Sphaeria mutila Fr. This is problematic for two reasons: (1) the Cambridge Rules (Briquet
1935), which were in effect in 1936, did not include such a
provision, and hence Stevens’s combination was legitimate,
and (2) said provision hinges on the correct assessment of
what constitutes the first description of a sexual morph. For
the reasons outlined above, the name Sphaeria mutila Fr.
must be associated with the sexual morph, even if Fries
(1823) did not explicitly describe asci and ascospores for
that species, because by placing the species in Sphaeria,
Fries (1823) implicitly adopted the name for a taxon producing asci. Therefore, Stevens (1936) cannot be claimed
as the first description of the sexual morph, even if he provided essential details beyond Fries’s (1823) morphological diagnosis. Shoemaker (1964) also erroneously thought
that the combination of Physalospora mutila N.E. Stevens,
when interpreted according to the Montreal Code, ínto
Botryosphaeria was blocked by the name Botryosphaeria
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Fungal Diversity (2020) 105:17–318
mutila Cooke [sic]. As so often in mycological literature,
that name does not exist. Cooke (1885) explicitly referred to
Sphaeria mutila as treated in Schweinitz’s (1832) work on
North American fungi, who in turn explicitly used the name
Sphaeria mutila Fr. for his North American material. Therefore, Cooke’s name is a combination of Sphaeria mutila Fr.
into Botryosphaeria, as B. mutila (Fr.) Cooke, accepting that
the fungus in question was a widespread species conspecific
with Fries’s original material. Incidentally, Cooke (1885)
gave a description of the asci and ascospores of the taxon
five decades prior to Stevens (1936), a fact overlooked by
Shoemaker (1964), hence also clearly associating the name
Sphaeria mutila Fr. with a sexual morph. This further underlines the complications from the provisions in Art. 59 of the
Montreal Code, since the description of the asexual morph
for S. mutila by Stevens (1936) was arguably not the first,
and hence that rule did not apply. In subsequent version of
the Code (starting with the Edinburgh Code and ending with
the Vienna Code; Lanjouw et al. 1966; McNeill et al. 2006),
this provision was amended in a way that only combinations
accompanied by elements required for valid publication of a
new taxon, e.g. a Latin description or diagnosis, were to be
corrected to names of new taxa and else were to be treated
as new combinations. Thus, under the rules applicable at
the time, under the amendment in the Edinburgh Code and
thereafter, and particularly starting with the Melbourne
Code (Barrie et al. 2012), not only is the name Physalospora
mutila (Fr.) N.E. Stevens legitimate, but the correct name
for the species to be adopted in the genus Botryosphaeria
is B. mutila (Fr.) Cooke. The name B. stevensii Shoemaker
is thus a superfluous, illegitimate replacement name and all
names considered here, namely Sphaeria mutila Fr., Diplodia mutila (Fr.) Fr., Botryosphaeria mutila (Fr.) Cooke,
Physalospora mutila (Fr.) N.E. Stevens, and Botryosphaeria
stevensii Shoemaker, are based on the same type and are to
be attributed to a sexual morph or holomorph. Given that
Fries’s original material is sterile, it is an entirely different
matter what species Botryosphaeria mutila exactly represents, and this can only be settled by epitypification.
Key to genera of Strigulaceae:
1.
1.
2.
2.
3.
Growing on living leaves (foliicolous)………………2
Growing on bark or rock or sometimes over bryophytes
or rarely lichenicolous on other lichens (corticolous,
saxicolous, muscicolous, hepaticolous, lichenicolous)…………………………………………………11
Ascospores 1-septate, very rarely with two obscure
additional septa………………………………………3
Ascospores 3-septate to muriform ……………………8
Growing supracuticularly, easily removable from the
leaf surface; photobiont representing the Phycopeltis
13
3.
4.
4.
5.
5.
6.
6.
7.
7.
8.
8.
9.
9.
morphodeme, with more or less rectangular cells forming net-like plates (Fig. 71a, b)……………Phylloporis
Growing subcuticularly, difficult to remove from the
leaf surface; photobiont representing the Cephaleuros
or Trentepohlia morphodeme, with rounded to oblong
cells forming irregular groups or plates often in more
than one layer…………………………………………4
Perithecial wall not carbonized; thallus often conspicuously folded or laciniate, with numerous white papillae
(Fig. 71c)…………………………………Puiggariella
Perithecial wall carbonized; thallus smooth to uneven,
rarely with white papillae……………………………5
Ascospores rather large, over 30 µm long; thallus
thin, typically dark bluish grey, always hypophyllous
(Fig. 71d)………………………………Raciborskiella
Ascospores small to medium-sized, up to 25 µm; thallus
thin to thickened, usually bright to dark green to greygreen or olive-brown, usually epiphyllous, very rarely
bluish grey or hypophyllous…………………………6
Thallus olive-brown to dark green with often metallic glance, composed of dichotomously branched but
sometimes confluent lacinia bordered by a continuous
or interrupted black line; perithecia covered by thin
thallus layer up to ostiole, therefore not pure black
(Fig. 71e, f)…………………………………Racoplaca
Thallus bright green to grey-green or rarely with bluish
tinge, with entire to crenulate or lobed margin but rarely
laciniate, not bordered by a thin black line except in the
S. nitidula aggregate and then thallus green and perithecia exposed, black…………………………………7
Macroconidia with appendages becoming very long
after release from pycnidia (up to 70–100 μm), appearing as if suspended along gelatinous strands; photobiont representing the Trentepohlia morphodeme, forming a single layer (Fig. 71g, h)…………Serusiauxiella
Macroconidia with appendages remaining rather
short after release from pycnidia (below 35 μm or
typically much shorter); photobiont representing the
Cephaleuros morphodeme, forming more than one
layer (Fig. 71i–o)………………Strigula sensu stricto
Ascospores 3(–5)-septate; macroconidia acicular to
filiform, multi-septate………………………………9
Ascospores muriform; macroconidia bacillar to
oblong-ellipsoid, multi-septate to muriform………10
Thallus supracuticular; perithecia aggregate in
slightly raised pseudostromata filled with ochraceousyellow, K+ dark red crystals; macroconidia acicular,
3(–5)-septate (Fig. 72a, b)……………Flavobathelium
Thallus subcuticular; perithecia solitary, strongly
prominent and laterally covered by a thin thallus layer;
macroconidia filiform, 3–9-septate (Fig. 72c)…………
…………………………………………Phyllocharis
Fungal Diversity (2020) 105:17–318
161
Fig. 71 Habit of genera and species in Strigulaceae. a Phylloporis
phyllogena. b P. obducta. c Puiggariella nemathora. d Raciborskiella
janeirensis. e Racoplaca subtilissima. f R. melanobapha. g Seru-
siauxiella sinensis. h S. filifera. i Strigula smaragdula. j S. macrocarpa. k S. schizospora. l S. antillarum. m S. bella. n S. prasina. o S.
nitidula
10. Perithecia immersed in thalline verrucae filled with
black, pulveraceous crystals or exposed and then with
somewhat roughened surface and wart-shaped to subglobose; pycnidia usually aggregate in pseudostromata,
rarely solitary in thalline verrucae (Fig. 72d–h)………
………………………………………Phyllobathelium
10. Perithecia exposed or covered by very thin thallus
layer, not immersed in thalline verrucae, with smooth
13
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Fig. 72 Habit of genera and species in Strigulaceae. a, b Flavobathelium epiphyllum. c Phyllocharis orbicularis. d Phyllobathelium
firmum. e, f P. thaxteri (in f macroconidia). g P. leguminosae. h P.
anomalum. i Phyllocraterina papuana. j Swinscowia glabra. k Flagellostrigula laureriformis. l Dichoporis ziziphii. m D. phaea. n D.
viridiseda. o D. wilsonii
surface, lens-shaped to hemisphaerical; pycnidia not
known (Fig. 72i)……………………Phyllocraterina
11. Ascospores and macroconidia 3-septate to muriform;
macroconidial appendages relatively short and broad,
(1–)2–4 µm, 1–2(–5) times as long as narrow, 15–30%
of conidial length in Swinscowia; microconidia typically (oblong-)bacillar (Fig. 72j)…………Swinscowia
11. Ascospores and macroconidia 1-septate; macroconidial
appendages relatively long and narrow, 1–2 µm broad,
3–6 times as long as wide, about 20–50% of conidial
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12.
12.
13.
13.
length (or rarely much longer); microconidia typically
fusiform(-oblong)…………………………………12
Perithecia and ascospores unknown; macroconidia
with a single, flagelliform, very long appendage at the
proximal end, about 3–4 times as long as the conidia;
pycnidia producing macroconidia very large (0.4–
0.8 mm diam.), covered by rather thick thallus layer
(Fig. 72k)……………………………Flagellostrigula
Perithecia and ascospores frequently present; macroconidia with similar appendages at each end, up to half of
the length of the conidia; pycnidia small (usually less
than 0.2 mm diam.), visible as dark dots……………13
Photobiont representing the Phycopeltis morphodeme,
with more or less rectangular cells forming net-like
plates; paraphyses unbranched………………………
…………………Phylloporis (rare instances on bark)
Photobiont representing the Trentepohlia morphodeme,
with rounded to oblong cells forming irregular groups
or plates; paraphyses generally branched and sometimes slightly anastomosing (Fig. 72l–o)……………
……………………………………………Dichoporis
Checklist of names published in the genus Strigula:
Strigula abietina Fr. [nom. inval.] = Lophodermium abietis
Rostr. (Rhytismataceae)
Strigula actinoplaca Nyl. = Actinoplaca strigulacea Müll.
Arg. (Gomphillaceae)
Strigula actinoplacoides Vain. = Puiggariella nemathora
(Mont.) S.H. Jiang, Lücking & J.C. Wei
Strigula acuticonidiarum S.H. Jiang, X.L. Wei & J.C. Wei
Strigula affinis (A. Massal.) R.C. Harris ≡ Swinscowia
affinis (A. Massal.) S.H. Jiang, Lücking & Sérus.
Strigula africana Vain. = Strigula smaragdula Fr.
Strigula africana var. africana Vain. = Strigula smaragdula
Fr.
Strigula africana var. crenulata Vain. = Strigula smaragdula Fr.
Strigula africana var. natalensis Vain. ex Van der Byl =
Strigula smaragdula Fr.
Strigula africana var. vegetior Vain. = Strigula subelegans
Vain.
Strigula albicascens (Nyl.) R.C. Harris ≡ Swinscowia albicascens (Nyl.) S.H. Jiang, Lücking & Sérus.
Strigula albolinita (Nyl.) R.C. Harris ≡ Swinscowia albolinita (Nyl.) S.H. Jiang, Lücking & Sérus.
Strigula albomaculata Sérus.
Strigula alpestris (Vezda) Hafellner ≡ Swinscowia alpestris
(Vězda) S.H. Jiang, Lücking & Sérus.
Strigula amblyoloba Müll. Arg. = Racoplaca subtilissima
(Fée) S.H. Jiang, Lücking & J.C. Wei
Strigula americana R.C. Harris ≡ Dichoporis tenuis (R.C.
Harris) S.H. Jiang, Lücking & Sérus.
163
Strigula amphora Aptroot & Lücking ≡ Swinscowia
amphora (Aptroot & Lücking) S.H. Jiang, Lücking &
Sérus.
Strigula angustata Cl. Roux & Sérus.
Strigula angustata Sérus. & Cl. Roux [nom. inval.] ≡ Strigula angustata Cl. Roux & Sérus.
Strigula angustissima Vain. = Phyllocharis orbicularis (Fr.)
S.H. Jiang, Lücking & Sérus.
Strigula antillarum (Fée) Müll. Arg.
Strigula aquatica H. Harada ≡ Swinscowia aquatica (H.
Harada) S.H. Jiang, Lücking & Sérus.
Strigula argentea (Fée) Trevis. = Strigula sp.
Strigula argyronema Müll. Arg. = Puiggariella nemathora
(Mont.) S.H. Jiang, Lücking & J.C. Wei
Strigula argyronema var. argyronema Müll. Arg. = Puiggariella nemathora (Mont.) S.H. Jiang, Lücking & J.C. Wei
Strigula argyronema var. confluens Müll. Arg. ≡ Puiggariella confluens (Müll. Arg.) S.H. Jiang, Lücking & J.C.
Wei
Strigula astroidea Vain. = Strigula smaragdula Fr.
Strigula astroidiza Vain. = Puiggariella nemathora (Mont.)
S.H. Jiang, Lücking & J.C. Wei
Strigula atrocarpa Vain. = Strigula concreta (Fée) R. Sant.
Strigula atrocarpa var. atrocarpa Vain. = Strigula concreta
(Fée) R. Sant.
Strigula atrocarpa var. fumigata Vain. [in Zahlbruckner
(1931)] ≡ Strigula atrocarpoides var. fumingata Vain. =
Strigula nitidula Mont.
Strigula atrocarpoides Vain. = Strigula nitidula Mont.
Strigula atrocarpoides var. atrocarpoides Vain. = Strigula
nitidula Mont.
Strigula atrocarpoides var. fumigata Vain. [in Zahlbruckner
(1931)] ≡ Strigula atrocarpoides var. fumingata Vain. =
Strigula nitidula Mont.
Strigula atrocarpoides var. fumingata Vain. = Strigula
nitidula Mont.
Strigula australiensis P.M. McCarthy ≡ Swinscowia australiensis (P.M. McCarthy) S.H. Jiang, Lücking & Sérus.
Strigula austropunctata P.M. McCarthy ≡ Phylloporis
austropunctata (P.M. McCarthy) S.H. Jiang, Lücking &
J.C.Wei
Strigula babingtonii Berk. ≡ Dennisiella babingtonii
(Berk.) Bat. & Cif. (Coccodiniaceae)
Strigula bahamensis (Riddle) R.C. Harris ≡ Swinscowia
bahamensis (Riddle) S.H. Jiang, Lücking & Sérus.
Strigula banksii Schwein. [nom. inval.] = [unidentified
fungus]
Strigula bella G. Thor, Lücking & Tat. Matsumoto
Strigula bermudana (Tuck. ex Nyl.) R.C. Harris ≡ Dichoporis bermudana (Tuck.) S.H. Jiang, Lücking & Sérus.
Strigula bispora Aptroot & K.H. Moon ≡ Swinscowia bispora
(Aptroot & K.H. Moon) S.H. Jiang, Lücking & Sérus.
13
164
Strigula brevis Bricaud & Cl. Roux ≡ Dichoporis brevis
(Bricaud & Cl. Roux) S.H. Jiang, Lücking & Sérus.
Strigula brevis Bricaud & Cl. Roux [nom. inval.] ≡ Dichoporis brevis (Bricaud & Cl. Roux) S.H. Jiang, Lücking &
Sérus.
Strigula buxi Chodat
Strigula caerulensis P.M. McCarthy
Strigula calcarea Bricaud & Cl. Roux ≡ Swinscowia calcarea (Bricaud & Cl. Roux) S.H. Jiang, Lücking & Sérus.
Strigula cavicola Cl. Roux & Bricaud ≡ Swinscowia cavicola (Cl. Roux & Bricaud) S.H. Jiang, Lücking & Sérus.
Strigula ciliata Mont. = Cephaleuros sp. (Trentepohliaceae)
Strigula cinefaciens (Nyl.) R.C. Harris ≡ Phylloporis cinefaciens (Nyl.) S.H. Jiang, Lücking & J.C.Wei
Strigula complanata (Fée) Nyl. [IF 370567] ≡ Strigula complanata (Fée) Mont. = Phyllocharis orbicularis (Fr.) S.H.
Jiang, Lücking & Sérus.
Strigula complanata Mont. [IF 547404] ≡ Strigula complanata (Fée) Mont. = Phyllocharis orbicularis (Fr.) S.H.
Jiang, Lücking & Sérus.
Strigula complanata f. complanata Mont. = Phyllocharis
orbicularis (Fr.) S.H. Jiang, Lücking & Sérus.
Strigula complanata f. major Nyl. = Puiggariella
nemathora f. hypothelia (Nyl.) S.H. Jiang, Lücking &
Sérus.
Strigula complanata var. ciliata (Mont.) Müll. Arg. =
Cephaleuros sp. (Trentepohliaceae)
Strigula complanata var. complanata Mont. = Phyllocharis
orbicularis (Fr.) S.H. Jiang, Lücking & Sérus.
Strigula complanata var. diplomorpha Müll. Arg. = Phyllocharis orbicularis (Fr.) S.H. Jiang, Lücking & Sérus.
Strigula complanata var. genuina Müll. Arg. [nom. inval.]
≡ Strigula complanata var. complanata (Fée) Mont. = Phyllocharis orbicularis (Fr.) S.H. Jiang, Lücking & Sérus.
Strigula complanata var. major Nyl. = Puiggariella
nemathora f. hypothelia (Nyl.) S.H. Jiang, Lücking &
Sérus.
Strigula complanata var. mesotropa Müll. Arg. =
Cephaleuros sp. (Trentepohliaceae)
Strigula complanata var. stellata Nyl. & Cromb. ex H.M.
Ward ≡ Strigula smaragdula var. stellata (Nyl. & Cromb.
ex H.M. Ward) Farkas
Strigula complanata var. subtilis Müll. Arg. = Phycopeltis
sp. (Trentepohliaceae)
Strigula complanata var. virescens Nyl. [IF 604058] ≡
Strigula complanata var. virescens (Kunze ex Fr.) Nyl. ≡
Cephaleuros virescens Kunze ex Fr. (Trentepohliaceae)
Strigula concentrica Müll. Arg. = Strigula smaragdula Fr.
Strigula concreta (Fée) R. Sant.
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Strigula confusa Fryday, Coppins & Common ≡ Swinscowia
confusa (A. Massal.) S.H. Jiang, Lücking & Sérus.
Strigula connivens R.C. Harris ≡ Dichoporis connivens
(R.C. Harris) S.H. Jiang, Lücking & Sérus.
Strigula contristans Zahlbr. = Racoplaca maculata (Cooke
& Massee) S.H. Jiang, Lücking & J.C. Wei
Strigula dacrydiospora Zahlbr. = Strigula subelegans Vain.
Strigula decipiens (Malme) P.M. McCarthy ≡ Swinscowia
decipiens (Malme) S.H. Jiang, Lücking & Sérus.
Strigula decipiens var. decipiens (Malme) P.M. McCarthy
≡ Swinscowia decipiens (Malme) S.H. Jiang, Lücking &
Sérus.
Strigula decipiens var. divisa P.M. McCarthy ≡ Swinscowia
divisa (P.M. McCarthy) S.H. Jiang, Lücking & Sérus.
Strigula delicata Sérus. ≡ Strigula delicata Sérus.
Strigula deplanata Müll. Arg. = Puiggariella nemathora
f. hypothelia (Nyl.) (Nyl.) S.H. Jiang, Lücking & Sérus.
Strigula dichosporidii Etayo ≡ Dichoporis dichosporidii
(Etayo) S.H. Jiang, Lücking & Sérus.
Strigula diederichiana Etayo, Cl. Roux & Sérus. = Dichoporis phaea (Ach.) S.H. Jiang, Lücking & Sérus.
Strigula difformis Vain. = Puiggariella nemathora (Mont.)
S.H. Jiang, Lücking & J.C. Wei
Strigula difformis var. arimensis Vain. = Puiggariella
nemathora (Mont.) S.H. Jiang, Lücking & J.C. Wei
Strigula difformis var. difformis Vain. = Puiggariella
nemathora (Mont.) S.H. Jiang, Lücking & J.C. Wei
Strigula dilatata Vain. [IF 454751] ≡ Porina dilatata Vain.
= Strigula nitidula Mont.
Strigula dispersa Vain. = Strigula prasina Müll. Arg.
Strigula donacis Vain. = Puiggariella nemathora f.
hypothelia (Nyl.) (Nyl.) S.H. Jiang, Lücking & Sérus.
Strigula effusa Fr. = Echinoplaca sp. (Gomphillaceae)
Strigula elatior Stirt. = Strigula smaragdula Fr.
Strigula elegans (Fée) Müll. Arg. = Strigula smaragdula Fr.
Strigula elegans f. elegans (Fée) Müll. Arg. = Strigula
smaragdula Fr.
Strigula elegans f. feei (Mont.) Müll. Arg. = Strigula smaragdula Fr.
Strigula elegans f. fuscata Müll. Arg. [as var. genuina f. fuscata; nom. inval.] ≡ Strigula elegans var. elegans f. fuscata
Müll. Arg. = Strigula smaragdula Fr.
Strigula elegans f. hirtella (Fée) Nyl. [IF 477956] ≡ Strigula
elegans var. elegans f. hirtella Müll. Arg. = Cephaleuros
sp. (Trentepohliaceae)
Strigula elegans f. hirtella Müll. Arg. [as var. genuina f.
hirtella; nom. inval.] ≡ Strigula elegans var. elegans f.
hirtella Müll. Arg. = Cephaleuros sp. (Trentepohliaceae)
Fungal Diversity (2020) 105:17–318
Strigula elegans subsp. elegans (Fée) Müll. Arg. = Strigula
smaragdula Fr.
Strigula elegans subsp. intermedia Müll. Arg. [IF 451609]
≡ Strigula elegans var. intermedia Müll. Arg. = Strigula
concreta (Fée) R. Sant.
Strigula elegans var. antillarum (Fée) R. Sant. ≡ Strigula
antillarum (Fée) Müll. Arg.
Strigula elegans var. concreta (Fée) Zahlbr. ≡ Strigula concreta (Fée) R. Sant.
Strigula elegans var. elatior (Stirt.) Zahlbr. = Strigula smaragdula Fr.
Strigula elegans var. elegans (Fée) Müll. Arg. = Strigula
smaragdula Fr.
Strigula elegans var. eumorpha Müll. Arg. = Strigula
microspora Lücking
Strigula elegans var. feei (Mont.) Müll. Arg. = Strigula
smaragdula Fr.
Strigula elegans var. genuina Müll. Arg. [nom. inval.] ≡
Strigula elegans var. elegans (Fée) Müll. Arg. = Strigula
smaragdula Fr.
Strigula elegans var. hirtella Fée [IF 451584] ≡ Strigula
elegans var. elegans f. hirtella Müll. Arg. = Cephaleuros
sp. (Trentepohliaceae)
Strigula elegans var. intermedia Müll. Arg. = Strigula concreta (Fée) R. Sant.
Strigula elegans var. nematora (Nyl.) Müll. Arg. [IF 588134]
≡ Strigula elegans var. nematora (Mont.) Müll. Arg. =
Puiggariella nemathora (Mont.) S.H. Jiang, Lücking &
J.C. Wei
Strigula elegans var. oxyloba Müll. Arg. = Strigula smaragdula Fr.
Strigula elegans var. pertenuis Müll. Arg. = Cephaleuros
sp. (Trentepohliaceae)
Strigula elegans var. stellata (Nyl. & Cromb. ex H.M. Ward)
R. Sant. ≡ Strigula smaragdula var. stellata (Nyl. &
Cromb. ex H.M. Ward) Farkas
Strigula elegans var. subciliata Müll. Arg. = Strigula smaragdula Fr.
Strigula elegans var. tremula Müll. Arg. = Strigula smaragdula Fr.
Strigula elegans var. viridissima Müll. Arg. = Strigula
smaragdula Fr.
Strigula elegantior Vain. = Strigula subelegans Vain.
Strigula elixii P.M. McCarthy ≡ Dichoporis elixii (P.M.
McCarthy) S.H. Jiang, Lücking & Sérus.
Strigula endolithea Cl. Roux & Bricaud ≡ Swinscowia
endolithea (Cl. Roux & Bricaud) S.H. Jiang, Lücking &
Sérus.
165
Strigula endolithea Cl. Roux & Bricaud [nom. inval.] ≡
Swinscowia endolithea (Cl. Roux & Bricaud) S.H. Jiang,
Lücking & Sérus.
Strigula epiphylla Eschw. [IF 454754] ≡ Verrucaria epiphylla Eschw. [nom. illeg.] = Strigula smaragdula Fr.
Strigula feei Mont. = Strigula smaragdula Fr.
Strigula fibrillosa Zahlbr. = Racoplaca melanobapha
(Kremp.) S.H. Jiang, Lücking & J.C. Wei
Strigula fossulicola P.M. McCarthy, Streimann & Elix
Strigula fossulicoloides Sérus.
Strigula fractans P.M. McCarthy ≡ Dichoporis fractans
(P.M. McCarthy) S.H. Jiang, Lücking & Sérus.
Strigula fracticonidia R.C. Harris ≡ Swinscowia fracticonidia (R.C. Harris) S.H. Jiang, Lücking & Sérus.
Strigula gibberosa Müll. Arg. = Strigula schizospora R.
Sant. [nom. cons.]
Strigula gibbosa Vain. = Strigula smaragdula Fr.
Strigula glabra (A. Massal.) V. Wirth ≡ Swinscowia glabra
(A. Massal.) S.H. Jiang, Lücking & Sérus.
Strigula glaziovii Müll. Arg. = Cephaleuros sp.
(Trentepohliaceae)
Strigula graminicola R. Sant.
Strigula griseonitens R.C. Harris ≡ Swinscowia griseonitens (R.C. Harris) S.H. Jiang, Lücking & Sérus.
Strigula guangxiensis S.H. Jiang, X.L. Wei & J.C. Wei
Strigula hymenaeicola Bat. & J.L. Bezerra [nom. inval.] =
Strigula sp.
Strigula hypothallina R.C. Harris ≡ Phylloporis hypothallina (R.C. Harris) S.H. Jiang, Lücking & Sérus.
Strigula hypothelia Nyl. ≡ Puiggariella nemathora f.
hypothelia (Nyl.) S.H. Jiang, Lücking & Sérus.
Strigula indutula (Nyl.) R.C. Harris = Swinscowia albicascens (Nyl.) S.H. Jiang, Lücking & Sérus.
Strigula insignis Jatta = Racoplaca melanobapha (Kremp.)
S.H. Jiang, Lücking & J.C. Wei
Strigula jamesii (Swinscow) R.C. Harris ≡ Swinscowia
jamesii (Swinscow) S.H. Jiang, Lücking & Sérus.
Strigula janeirensis (Müll. Arg.) Lücking ≡ Raciborskiella
janeirensis (Müll. Arg.) R. Sant.
Strigula johnsonii P.M. McCarthy ≡ Swinscowia johnsonii
(P.M. McCarthy) S.H. Jiang, Lücking & Sérus.
Strigula kaitokensis Sérus. & Polly
Strigula laceribracae R.C. Harris ≡ Swinscowia laceribracae (R.C. Harris) S.H. Jiang, Lücking & Sérus.
Strigula lacericola P.M. McCarthy
Strigula lacinulata Vain. = Phyllocharis orbicularis (Fr.)
S.H. Jiang, Lücking & Sérus.
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166
Strigula lacinulata var. compactior Zahlbr. = Phyllocharis
orbicularis (Fr.) S.H. Jiang, Lücking & Sérus.
Strigula lacinulata var. lacinulata Vain. = Phyllocharis
orbicularis (Fr.) S.H. Jiang, Lücking & Sérus.
Strigula lamellosa Zahlbr. = Strigula subelegans Vain.
Strigula lamprocarpa Müll. Arg. [IF 454755] ≡ Porina lamprocarpa Müll. Arg. = Strigula nitidula Mont.
Strigula lateralis Aptroot & van den Boom = Swinscowia
tagananae (Harm.) S.H. Jiang, Lücking & Sérus.
Strigula laureriformis Aptroot & Lücking ≡ Flagellostrigula laureriformis (Aptroot & Lücking) Lücking, S.H.
Jiang & J.C. Wei
Strigula linearis Vain. = Racoplaca melanobapha (Kremp.)
S.H. Jiang, Lücking & J.C. Wei
Strigula lobulosa Kunze ex Fr. = Strigula sp.
Strigula mabae Zahlbr. = Strigula smaragdula Fr.
Strigula macaronesica Sérus.
Strigula macrocarpa Vain.
Strigula maculata (Cooke & Massee) R. Sant. ≡ Racoplaca
maculata (Cooke & Massee) S.H. Jiang, Lücking & J.C.
Wei
Strigula maritima H. Harada ≡ Dichoporis maritima (H.
Harada) S.H. Jiang, Lücking & Sérus.
Strigula mediterranea Etayo = Dichoporis ziziphi (A. Massal.) S.H. Jiang, Lücking & Sérus.
Strigula melanobapha (Kremp.) R. Sant. ≡ Racoplaca melanobapha (Kremp.) S.H. Jiang, Lücking & J.C. Wei
Strigula melanophthalma Mont. = Strigula antillarum
(Fée) Müll. Arg.
Strigula microspora Lücking
Strigula microthyrium Mont. = [unidentified fungus]
Strigula minor (Vezda) Cl. Roux & Sérus.
Strigula minuta Lücking
Strigula minutula P.M. McCarthy ≡ Dichoporis minutula
(P.M. McCarthy) S.H. Jiang, Lücking & Sérus.
Strigula mori Schulzer ex Keissl. = [unidentified fungus]
Strigula multipunctata (G. Merr. ex R. Sant.) R.C. Harris
= Phylloporis cinefaciens (Nyl.) S.H. Jiang, Lücking &
J.C.Wei
Strigula muriconidiata Aptroot, L.I. Ferraro & M. Cáceres
≡ Swinscowia muriconidiata (Aptroot, L.I. Ferraro & M.
Cáceres) S.H. Jiang, Lücking & Sérus.
Strigula muriformis Aptroot & Diederich ≡ Swinscowia
muriformis (Aptroot & Diederich) S.H. Jiang, Lücking
& Sérus.
Strigula muscicola F. Berger, Coppins, Cl. Roux & Sérus. ≡
Swinscowia muscicola (F. Berger, Coppins, Cl. Roux &
Sérus.) S.H. Jiang, Lücking & Sérus.
Strigula natalis P.M. McCarthy ≡ Dichoporis natalis (P.M.
McCarthy) S.H. Jiang, Lücking & Sérus.
Strigula nemathora Mont. ≡ Puiggariella nemathora
(Mont.) S.H. Jiang, Lücking & J.C. Wei
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Fungal Diversity (2020) 105:17–318
Strigula nemathora f. hypothelia (Nyl.) Lücking ≡ Puiggariella nemathora f. hypothelia (Nyl.) S.H. Jiang, Lücking
& Sérus.
Strigula nemathora f. nemathora Mont. ≡ Puiggariella
nemathora (Mont.) S.H. Jiang, Lücking & J.C. Wei
Strigula nemathora var. hypothelia (Nyl.) R. Sant. ≡ Puiggariella nemathora f. hypothelia (Nyl.) S.H. Jiang, Lücking & Sérus.
Strigula nemathora var. nemathora Mont. ≡ Puiggariella
nemathora (Mont.) S.H. Jiang, Lücking & J.C. Wei
Strigula nemathora var. pulchella (Müll. Arg.) R. Sant. =
Puiggariella nemathora (Mont.) S.H. Jiang, Lücking &
J.C. Wei
Strigula nemathora var. pulchella Mont. [IF 451376] ≡
Strigula nemathora var. pulchella (Müll. Arg.) R. Sant.
= Puiggariella nemathora (Mont.) S.H. Jiang, Lücking
& J.C. Wei
Strigula nematora Nyl. [in Zahlbruckner 1922)] ≡ Puiggariella nemathora (Mont.) S.H. Jiang, Lücking & J.C. Wei
Strigula nigrocarpa Lücking
Strigula nigrocincta Müll. Arg. ≡ Puiggariella nigrocincta
(Müll. Arg.) S.H. Jiang, Lücking & J.C. Wei
Strigula nigrocincta var. nigrocincta Müll. Arg. ≡ Puiggariella nigrocincta (Müll. Arg.) S.H. Jiang, Lücking &
J.C. Wei
Strigula nigrocincta var. soluta Müll. Arg. = Puiggariella
nigrocincta (Müll. Arg.) S.H. Jiang, Lücking & J.C. Wei
Strigula nipponica H. Harada ≡ Dichoporis nipponica (H.
Harada) S.H. Jiang, Lücking & Sérus.
Strigula nitidula Mont.
Strigula novae-zelandiae (Nag Raj) Sérus.
Strigula nylanderi Vain. = Strigula smaragdula Fr.
Strigula nylanderiana F. Schill. = Puiggariella nemathora
f. hypothelia (Nyl.) S.H. Jiang, Lücking & Sérus.
Strigula obducta (Müll. Arg.) R.C. Harris ≡ Phylloporis
obducta (Müll. Arg.) R. Sant. & Tibell
Strigula obtecta (Vain.) R.C. Harris ≡ Swinscowia obtecta
(Vain.) S.H. Jiang, Lücking & Sérus.
Strigula obvelata Vain. = Strigula macrocarpa Vain.
Strigula occulta P.M. McCarthy & Malcolm ≡ Dichoporis
occulta (P.M. McCarthy & Malcolm) S.H. Jiang, Lücking
& Sérus.
Strigula oceanica P.M. McCarthy, Streimann & Elix
Strigula orbicularis Fr. ≡ Phyllocharis orbicularis (Fr.)
S.H. Jiang, Lücking & Sérus.
Strigula pachyneura Müll. Arg. = Phyllocharis orbicularis
(Fr.) S.H. Jiang, Lücking & Sérus.
Strigula pallida Aptroot & K.H. Moon ≡ Swinscowia pallida (Aptroot & K.H. Moon) S.H. Jiang, Lücking & Sérus.
Strigula pallida Kalchbr. & Stizenb.[IF 479700; nom. inval.]
≡ Strigula pallida Kalchbr. = Strigula sp.
Fungal Diversity (2020) 105:17–318
Strigula pallidocarpa Aptroot & K.H. Moon ≡ Swinscowia
pallida (Aptroot & K.H. Moon) S.H. Jiang, Lücking &
Sérus.
Strigula parksii Räsänen = Strigula subelegans Vain.
Strigula phaea (Ach.) R.C. Harris ≡ Dichoporis phaea
(Ach.) S.H. Jiang, Lücking & Sérus.
Strigula philippina Vain. = Strigula smaragdula Fr.
Strigula phyllogena (Müll. Arg.) R.C. Harris ≡ Phylloporis
phyllogena (Müll. Arg.) Clem.
Strigula plana Müll. Arg. = Strigula smaragdula Fr.
Strigula platypoda (Müll. Arg.) R.C. Harris ≡ Phylloporis
platypoda (Müll. Arg.) Vězda
Strigula porinoides Canals, Boqueras & Gómez-Bolea ≡
Swinscowia porinoides (Canals, Boqueras & GómezBolea) S.H. Jiang, Lücking & Sérus.
Strigula prasina Müll. Arg.
Strigula pulchella Müll. Arg. = Puiggariella nemathora
(Mont.) S.H. Jiang, Lücking & J.C. Wei
Strigula puncticulata Müll. Arg. = Puiggariella nemathora
f. hypothelia (Nyl.) S.H. Jiang, Lücking & Sérus.
Strigula racoplaca Mont. = Racoplaca subtilissima (Fée)
S.H. Jiang, Lücking & J.C. Wei
Strigula radiata Lücking ≡ Phylloporis radiata (Lücking)
S.H. Jiang, Lücking & J.C. Wei
Strigula rhodinula (Zahlbr.) P.M. McCarthy ≡ Swinscowia
rhodinula (Zahlbr.) S.H. Jiang, Lücking & Sérus.
Strigula rostrata R.C. Harris & Aptroot ≡ Swinscowia
rostrata (R.C. Harris & Aptroot) S.H. Jiang, Lücking &
Sérus.
Strigula rotula Mont. ≡ Mazosia rotula (Mont.) A. Massal.
(Roccellaceae)
Strigula rugulosa Müll. Arg. = Strigula concreta (Fée) R.
Sant.
Strigula rugulosa var. irregularis Müll. Arg. = Strigula
microspora Lücking
Strigula rugulosa var. rugulosa Müll. Arg. = Strigula concreta (Fée) R. Sant.
Strigula rupestris P.M. McCarthy ≡ Swinscowia rupestris
(P.M. McCarthy) S.H. Jiang, Lücking & Sérus.
Strigula schizospora R. Sant. [nom. cons.]
Strigula setacea Müll. Arg. = Cephaleuros sp.
(Trentepohliaceae)
Strigula sinoaustralis S.H. Jiang, X.L. Wei & J.C. Wei
Strigula smaragdula Fr.
Strigula smaragdula var. smaragdula Fr. ≡ Strigula smaragdula Fr.
Strigula smaragdula var. stellata (Nyl. & Cromb. ex H.M.
Ward) Farkas ≡ Strigula smaragdula var. stellata (Nyl.
& Cromb. ex H.M. Ward) Farkas
Strigula stigmatella (Ach.) R.C. Harris ≡ Swinscowia stigmatella (Ach.) S.H. Jiang, Lücking & Sérus.
167
Strigula stigmatella var. alpestris (Vĕzda) Coppins = Swinscowia stigmatella (Ach.) S.H. Jiang, Lücking & Sérus.
Strigula stigmatella var. stigmatella (Ach.) R.C. Harris ≡
Swinscowia stigmatella (Ach.) S.H. Jiang, Lücking &
Sérus.
Strigula stilboideum Bat. = Strigula sp.
Strigula subelegans Vain.
Strigula submuriformis (R.C. Harris) R.C. Harris ≡ Swinscowia submuriformis (R.C. Harris) S.H. Jiang, Lücking
& Sérus.
Strigula subprospersella (Vain.) P.M. McCarthy ≡ Dichoporis subprospersella (Vain.) S.H. Jiang, Lücking & Sérus.
Strigula subsimplicans (Nyl.) R.C. Harris ≡ Dichoporis subsimplicans (Nyl.) S.H. Jiang, Lücking & Sérus.
Strigula subtilissima (Fée) Müll. Arg. ≡ Racoplaca subtilissima (Fée) S.H. Jiang, Lücking & J.C. Wei
Strigula sulcata Vain. = Strigula concreta (Fée) R. Sant.
Strigula sychnogonoides (Nitschke) R.C. Harris ≡ Geisleria
sychnogonoides Nitschke (Stictidaceae)
Strigula tagananae (Harm.) R.C. Harris ≡ Swinscowia tagananae (Harm.) S.H. Jiang, Lücking & Sérus.
Strigula taylorii (Carroll ex Nyl.) R.C. Harris ≡ Dichoporis
taylorii (Carroll) S.H. Jiang, Lücking & Sérus.
Strigula tenuis Müll. Arg. = Strigula smaragdula Fr.
Strigula thelopsidoides Coppins, Cl. Roux & Sérus. ≡ Swinscowia thelopsidoides (Coppins, Cl. Roux & Sérus.) S.H.
Jiang, Lücking & Sérus.
Strigula transversoundulata Sipman ≡ Racoplaca transversoundulata (Sipman) S.H. Jiang, Lücking & J.C. Wei
Strigula tremens Müll. Arg. ≡ Racoplaca tremens (Müll.
Arg.) S.H. Jiang, Lücking & J.C. Wei
Strigula umbilicata Müll. Arg. ≡ Asterothyrium umbilicatum (Müll. Arg.) Müll. Arg. (Gomphillaceae)
Strigula undulata Müll. Arg. = Strigula smaragdula Fr.
Strigula urticae Bonord. = [unidentified fungus]
Strigula vincentina Vain. [IF 454756] ≡ Porina vincentina
Vain. = Racoplaca subtilissima (Fée) S.H. Jiang, Lücking
& J.C. Wei
Strigula virescens (Kunze ex Fr.) Trevis. ≡ Cephaleuros
virescens Kunze ex Fr. (Trentepohliaceae)
Strigula viridis (Lücking) R.C. Harris ≡ Phylloporis viridis
Lücking
Strigula viridiseda (Nyl.) R.C. Harris ≡ Dichoporis viridiseda (Nyl.) S.H. Jiang, Lücking & Sérus.
Strigula viridissima (Fée) Trevis. = Strigula smaragdula Fr.
Strigula vulgaris (Müll. Arg.) Lücking ≡ Phylloporis vulgaris (Müll. Arg.) S.H. Jiang, Lücking & J.C. Wei
Strigula wandae M. Cáceres & Lücking
Strigula wilsonii (Riddle) R.C. Harris ≡ Dichoporis wilsonii
(Riddle) S.H. Jiang, Lücking & Sérus.
Strigula xylopiae Bat. & Cavalc. = Strigula schizospora R.
Sant. [nom. cons.]
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Strigula ziziphi (A. Massal.) Cl. Roux & Sérus. ≡ Dichoporis ziziphi (A. Massal.) S.H. Jiang, Lücking & Sérus.
thereby refers to the remarkable similarity with Porina both
in morphological and anatomical features.
Economic and ecological significance
For foliicolous species of Strigulaceae, indicator values
have been published that allow applications in monitoring
the conservation status of forest ecosystems (Lücking 1997).
Particularly species of Strigula sensu lato now dispositioned
in the genera Phylloporis, Puiggariella, Raciborskiella,
Racoplaca, and Strigula sensu stricto allow for a rather precise monitoring, as their scores are highly specific to individual genera and species (Lücking 1997). In a recent study,
the population genetics of a new species segregated from
Strigula smaragdula, S. multiformis, and its correlation with
modeled habitat preferences was studied (Oh et al. 2019).
Tenuitholiascus S.H. Jiang, Lücking & J.C. Wei, IMA Fungus 11: 1 (2020).
Fungal Names: FN570581; Facesoffungi number: FoF
08776, 1 morphological species (Jiang et al. 2020a), 1 species with molecular data.
Type species: Tenuitholiascus porinoides S.H. Jiang,
Lücking & J.C. Wei.
Notes: Besides ascus type, the only genus in the family is distinguished from genera in the sister family Strigulaceae as follows: from Phylloporis in the 3-septate, oblong
ascospores, from Phyllocratera in the small, 3-septate
ascospores, and from Flavobathelium in the external habit
with exposed perithecia. Tenuitholiascus closely resembles
Porina in external morphology and ascospore type, as well
as the thin-walled asci and unbranched paraphyses. Yet, it is
entirely unrelated to the latter, which belongs in Gyalectales
in class Lecanoromycetes (Kraichak et al. 2018; Lücking
2019).
Tenuitholiascaceae S.H. Jiang, Lücking & J.C. Wei, in Jiang
et al., IMA Fungus 11: 1 (2020).
Fungal Names: FN570578; Facesoffungi number: FoF
08775, 1 species.
Lichenized on leaves chiefly in lowland to montane tropical to subtropical habitats. Thallus supracuticular, easily
separated from the leaf surface, smooth, pale green. Photobiont Phycopeltis, cells rectangular, composed of anastomosing filaments in one layer forming regular, radiating plates.
Ascomata perithecia, wart-shaped, scattered or somewhat
clustered, ostiolate, exposed but covered by thin thallus layer
up to the ostiole, with the basal part spreading to form a horizontal plate. Involucrellum carbonized, black. Exciple dense,
prosoplectenchymatous, colourless to brown. Hamathecium
comprising unbranched paraphyses. Asci bitunicate in structure but lacking a distinct tholus, apex rounded, I-, appearing
nearly unitunicate in some stages, clavate to almost cylindrical, I-, KI-, 8-spored. Ascospores irregularly arranged to
2-seriate, fusiform, 3-septate, hyaline. Asexual state: Pycnidia common, wart-shaped, immersed to erumpent, visible as black dots. Conidia acrogenous, only microconidia
known; microconidia fusiform, hyaline, aseptate. Chemistry:
No secondary substances detected in TLC.
Type: Tenuitholiascus S.H. Jiang, Lücking & J.C. Wei.
Notes: The family was introduced by Jiang et al. (2020a)
to accomodate a novel lineage sister to Strigulaceae, but
with a different ascus type. To assess placement of the new
lineage within Dothideomycetes, a dataset consisting of
three loci (SSU, LSU, and tef1-α) was constructed and analysed. Even if superficially similar to Porina in Lecanoromycetes, it was evident that the new lineage was a member
of class Dothideomycetes, forming a clade supported sister
to Strigulaceae. Since the asci of the novel lineage differed
from those of Strigulaceae in lacking a thickened tholus and
ocular chamber, it was considered prudent to introduce the
monospecific family Tenuitholiascaceae and genus Tenuitholiascus for this taxon. The specific epithet, T. porinoides,
13
Tenuitholiascus porinoides S.H. Jiang, Lücking & J.C. Wei,
IMA Fungus 11: 1 (2020).
Fungal Names: FN570580; Fig. 73
Thallus supracuticular, easily separated from the leaf
surface, continuous, smooth, pale green, 3–12 mm diam,
30–52.5 μm thick. Algal partner: Phycopeltis, cells rectangular, 8–14 × 3–5 μm, composed of anastomosing filaments
lying in one layer and forming regular radial plates or irregular nets. Ascomata perithecia, globose, scattered or clustered,
exposed but covered by thin thallus layer up to the ostiole,
central part wart-shaped, sometimes basal part broadly
spreading to form horizontal plate, 0.25–0.5 mm diam and
80–150 μm high, greyish black. Involucrellum carbonized,
black, 55–125 μm thick. Exciple dense, prosoplectenchymatous, 10–12.5 μm thick, colourless to brown. Interascal filaments: unbranched or simply branched, thin. Asci bitunicate
in structure, apex with a I- rounded, sometimes appearing
almost unitunicate in some developmental stages, due to the
gradually thinner inner walls (Fig. 73f), clavate to cylindrical, 75–90 × 10–12.5 μm, I-, KI-, 8-spored. Ascospores
fusiform, 3-septate, colourless, 25–30 × 6–8 μm. Pycnidia
common, wart-shaped, immersed to erumpent, 0.05–0.1 mm
diam, black. Conidia (microconidia) fusiform, hyaline, aseptate, 4–5 × 1.5–2 μm.
Economic and ecological significance
Since this family is poorly known, it is difficult at the
moment to elaborate on this topic. As foliicolous lichen
apparently found in shady conditions, Tenuitholiascaceae
is a potential indicator of well-reserved forest ecosystems,
Fungal Diversity (2020) 105:17–318
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Fig. 73 Tenuitholiascus porinoides (HMAS–L0139638). a Thallus. b The Phycopeltis algal partner. c Perithecia in vertical section.
d Ascus (HMAS–L0139639). e Ascus (HMAS–L0139640). f Ascus
apex (HMAS–L0141346). g Ascus with iodine reaction (HMAS–
L0139638). h Ascus with iodine reaction (HMAS–L0141348). i
Ascospores (HMAS–L0139639). j Ascospores (HMAS–L0139638).
Scale bars: a = 300 μm, c = 20 μm, b, d, i, j = 10 μm, e–h = 5 μm
similar to other ecologically equivalent foliicolous lichens
(Lücking 1997).
Superstratomycetales was introduced to accommodate
previously unknown taxa in Pezizomycotina which were
recovered from oil-treated timber after outdoor exposure
in Australia and the Netherlands (Nieuwenhuijzen et al.
2016). The phylogenetic tree (Fig. 74) indicates that species of Superstratomyces form a clade with high support
(100% ML, 1.0 PP, Fig. 74) within Dothideomycetes. The
Superstratomycetales van Nieuwenh., Miądl., Houbraken,
Adan, Lutzoni & Samson.
Index Fungorum number: IF 819160; Facesoffungi number: FoF 08926.
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Fungal Diversity (2020) 105:17–318
Fig. 74 Phylogram generated
from maximum likelihood
analysis (RML) of genera in
Superstratomycetales based
on ITS, LSU, rpb-2, SSU and
tef1 sequence data. Maximum
likelihood bootstrap values
equal or above 70%, Bayesian
posterior probabilities equal
or above 0.90 (MLBS/PP) are
given at the nodes. An original
isolate number is noted after the
species name. The tree is rooted
to Venturia inaequalis (CBS
815.69). The ex-type strains are
indicated in bold. Hyphen (-)
represents support values below
70% MLBS and 0.90 PP
Fig. 75 Superstratomyces albomucosus (redrawn from holotype
H-22668 in van Nieuwenhuijzen et al. 2016). a Pycnidia. b Conidiogenous cells. c Conidia. Scale bar: c = 5 μm
divergence time for Superstratomycetales is estimated as 156
MYA (stem age, Hongsanan et al. 2020).
Accepted families: Superstratomycetaceae.
Superstratomycetaceae van Nieuwenh., Miądl., Houbraken,
Adan, Lutzoni & Samson, Stud. Mycol. 85: 115 (2016).
Index Fungorum number: IF 819161; Facesoffungi numbers: FoF 08777, 4 species.
Saprobic on oil treated wood or darkened surface of a
wall, further nature of the substrates varied from human and
environmental, therefore probably they showed a wide distribution. Sexual morph: Undetermined. Asexual morph:
Coelomycetous. Colonies olive to grey-green or light greygreen with thin white edge; reverse dark green or dark green
with thin white edge; knotted cone-shaped. Hyphae hyaline
to brown, smooth- and thin- to thick-walled, septate. Conidiomata pycnidial, superficial, solitary or confluent, brown
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to black, glabrous, globose, filled by a white mass of slimy
conidia; pycnidial wall pseudoparenchymatous, of textura
angularis cells, composed of several layers of pale brown to
brown, flattened. Setae erect to recurved, hyaline to subhyaline at apex and turning brown towards the base, 1–2-septate,
strongly verrucose to tuberculate. Conidiophores branched,
hyaline, smooth-walled, bearing lateral conidiogenous cells.
Conidiogenous cells phialidic, hyaline, cylindrical to barrelshaped or ampulliform, smooth-walled, solitary or laterally
disposed on the conidiophores. Conidia cylindrical to navicular, hyaline, aseptate, smooth- and thin-walled, guttulate.
Type: Superstratomyces van Nieuwenh., Miądl. &
Samson.
Notes: Superstratomycetaceae comprises one genus
named Superstratomyces, and it was typified by Superstratomyces albomucosus. Multi-locus phylogenies are necessary for the identification of species in Superstratomyces.
Superstratomycetaceae lacks unique characteristics and,
therefore, cannot be identified based only on morphology.
With the recent introduction of Superstratomyces tardicrescens, the number of species in Superstratomyces, family
Superstratomycetaceae has been increased up to four (Crous
et al. 2018a).
Superstratomyces van Nieuwenh., Miądl. & Samson, in van
Nieuwenhuijzen et al., Stud. Mycol. 85: 115 (2016).
Index Fungorum number: IF 819162; Facesoffungi number: FoF 08778; 4 morphological species (Species Fungorum 2020), 4 species with molecular data.
Type species: Superstratomyces albomucosus van Nieuwenh. & Samson.
Notes: Superstratomyces was introduced by van Nieuwenhuijzen et al. (2016) to accommodate the species that
was recovered from outdoor wood in the Netherlands. Currently there are four Superstratomyces species including
Fungal Diversity (2020) 105:17–318
S. albomucosus, S. flavomucosus, S. atroviridis and S.
tardicrescens.
Superstratomyces albomucosus van Nieuwenh., Miądl. &
Samson, in van Nieuwenhuijzen et al., Stud. Mycol. 85: 115
(2016).
Index Fungorum: IF 819163; Facesoffungi number: FoF
08779, Fig. 234
Description: see van Nieuwenhuijzen et al. (2016)
(Fig. 75).
Economic and ecological significance
Superstratomyces albomucosus and S. atroviridis were isolated from exposed, oil-treated wood and may decay wood.
Superstratomyces flavomucosus and S. tardicrescens were
recovered from leaves of Hakea multilinearis in Australia
and a human eye specimen in the USA (Crous et al. 2018a).
Trypetheliales Lücking, Aptroot & Sipman.
Index Fungorum number: IF 90793; Facesoffungi number: FoF 08780.
Aptroot et al. (2008) established Trypetheliales to accommodate a lichen-forming family Trypetheliaceae, which is
phylogenetically distinct from other orders and families in
Dothideomycetes (Del Prado et al. 2006). Based on molecular phylogeny and morphology, a second family Polycoccaceae, with lichenicolous fungi, was introduced by Ertz
et al. (2015). The order Trypetheliales accommodates primarily lichenized and less commonly non-lichenized taxa,
characterized by perithecioid ascomata appearing solitary
or aggregated in pseudostromata, branched and anastomosing paraphysoids forming a distinct network, and hyaline
or rarely brown ascospores, transversely septate to muriform
ascospores often with diamond-shaped lumina and angular wall thickenings (Aptroot et al. 2008; Hyde et al. 2013;
Nelsen et al. 2014; Ertz et al. 2015). In a case study of divergence time estimates in Dothideomycetes (Liu et al. 2017),
the order was diversified approximately 215 (140–298)
MYA (crown age) and 309 (231–388) MYA (stem age).
Polycoccaceae Ertz, Hafellner & Diederich, in Ertz et al.,
Fungal Diversity 74: 82 (2015).
Index Fungorum number: IF 814032; Facesoffungi numbers: FoF 08781, 50 species.
Lichenicolous. Sexual morph: Ascomata arising singly,
often becoming grouped, sometimes united by a clypeus or
enclosed in galls, immersed with only the ostiole visible to
erumpent and the upper half exposed when mature, perithecioid, subglobose to obpyriform, dark brown to black, ostiolate, neck not extended and scarcely distinguishable from
the ascomatal wall. Peridium thick or somewhat broader
near the ostiole, comprises 3–6 layers of polyhedral pseudoparenchymatous cells, radially compressed in vertical
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section and roughly isodiametric in surface view, forming a
textura angularis, thick, brown to dark brown and continuing
below the centrum in outer layers, less intensely pigmented
to hyaline with thin-walled cells at inner layers. Hamathecium comprising branched, anastomosing net of thin, septate to remotely septate narrow hyphal filaments, probably
trabeculate pseudoparaphyses (paraphysoids), periphyses
absent, hymenial gel I+ blue to violet or unchanged. Asci
2–8-spored, bitunicate, fissitunicate, broadly cylindrical to
subclavate, shortly stalked, wall thicker in the upper part of
mature asci, with a small internal apical beak. Ascospores
irregularly distichously arranged in the asci, ellipsoid, brown
to dark brown when mature, 1-septate (euseptate), somewhat
constricted at the septum, normally equal in size or the upper
cell larger and broader, generally rounded or rarely attenuated at apex, smooth or delicately verruculose, sometimes
with a gelatinous sheath (adapted from Ertz et al. 2015).
Type: Polycoccum Saut. ex Körb.
Notes: Ertz et al. (2015) included species of Clypeococcum and Polycoccum in phylogenetic analyses and found
that members of the two genera formed a new lineage
within Trypetheliales. Therefore, the sister family of Trypetheliaceae was introduced and named as Polycoccaceae
to accommodate Clypeococcum and Polycoccum based on
molecular and morphological data.
Polycoccum Saut. ex Körb., Parerga lichenol. (Breslau) 5:
470 (1865).
Index Fungorum number: IF 4309; Facesoffungi number:
FoF 08782; 41 morphological species (Species Fungorum
2020), 4 species with molecular data.
Type species: Polycoccum sauteri Körb.
Notes: Polycoccum is considered as polyphyletic, with
species being presented in two distantly related clades in
Trypetheliales and Pleosporales (Ertz et al. 2015). Polycoccum sensu stricto and species of Clypeococcum form a lineage in Trypetheliales, thus Ertz et al. (2015) established
Polycoccaceae to accommodate these genera. Polycoccum
sensu stricto (in Polycoccaceae) is characterized by ascomata formed on the host thallus, induce the formation of
galls, thick-walled with cells ± isodiametric in longitudinal
section, relatively thick paraphysoids, broadly cylindrical
to subclavate asci, generally with irregularly distichously or
irregularly uni-seriate arranged ascospores, often distinctly
ornamented ascospores (Ertz et al. 2015). The genus has 57
species accepted by Lawrey and Diederich (2015) and new
species are described almost every year (Ertz et al. 2015). It
has become one of the most species-rich genera of lichenicolous fungi. The most complete key to species of Polycoccum so far was provided by Hawksworth and Diederich
(1988). Other regional keys were established such as key for
13 species from Spain (Atienza et al. 2003), and key for 14
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Fig. 76 Morphology of genera in Polycoccaceae (a–d = Polycoccum pulvinatum, Belgium, Ertz 18114 (BR); e–h = Clypeococcum
cladonema, Belgium, Ertz 15260 (BR)). a, b Galls with immersed
ascomata on Physcia caesia. c Asci with ascospores in water. d
Ascospores in water, the one in the lower right corner showing the
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Fungal Diversity (2020) 105:17–318
verruculose surface. e, f ascomata (= black areas) on Xanthoparmelia (= brownish host thallus). g Section of ascomata in water. h,
ascospores in water. Scale bars: a, e = 1 mm, b = 500 µm, f = 250
µm, g = 50 µm, c, d, h = 10 µm
Fungal Diversity (2020) 105:17–318
species from Sweden (Ihlen and Wedin 2008), and a key for
13 species from France (Gardiennet 2012) (Fig. 76).
Other genera included:
Clypeococcum D. Hawksw., J. Linn. Soc., Bot. 75: 196
(1977).
Index Fungorum number: IF 1114; Facesoffungi number:
FoF 08783; – 9 morphological species (Species Fungorum
2020), 2 species with molecular data.
Type species: Clypeococcum cladonema (Wedd.) D.
Hawksw., Bot. J. Linn. Soc. 75(2): 197 (1977).
≡ Lecidea cladonema Wedd., Bull. Soc. bot. Fr. 21: 345
(1874).
Notes: According to Hawksworth and Diederich (1988),
this genus shares some characters with Polycoccum, but
differs by hyphal rather than pseudoparenchymatous ascomatal walls, thickened around the ostiole, and by the ascomata occurring in groups united by a common clypeus. In
phylogenetic tree of Ertz et al. (2015), Clypeococcum represented by C. placopsiiphilum and C. psoromatis is paraphyletic among Polycoccum. Ertz et al. (2015) noted that
the long branches within Polycoccaceae clade showed the
fast-evolving contrasting with low morphological variability,
which results in few phenotypic characters useful for the
recognition of different genera (Ertz et al. 2015). Clypeococcum is however included in Polycoccaceae but its placement will need to be confirmed by molecular data of type
species. Fourteen epithets and 9 epithets of Clypeococcum
are listed in Index Fungorum (2020) and Species Fungorum
(2020), respectively, while sequence data are available for
two species.
Economic and ecological significance
Species in this family are lichenicolous living on lichens as
parasites.
Trypetheliaceae Zenker in Goebel & Kunze, Pharmaceutische Waarenkunde: 123 (1827); Fée, Essai Crypt.
Écorc. 1: xxxvi (1824), nom. inval.; Eschweiler, Syst. Lich.:
17 (1824), nom. inval.
MycoBank number: MB 81884; Index Fungorum number:
IF 81884; Facesoffungi number: FoF 08784; approximately
440 species (Aptroot and Lücking 2016; Aptroot et al. 2016,
2019; Lücking et al. 2017; this paper).
Lichenized or more rarely saprobic on bark or rarely
on bryophytes over soil; in terrestrial, chiefly lowland to
lower montane tropical habitats, with few species extending into temperate regions. Thallus reduced and ecorticate,
white, to distinctly corticate, yellow-brown to olive-green,
sometimes partly or completely bright yellow, orange or
red due to superficial anthraquinone pigments. Photobiont
Trentepohlia. Ascomata scattered, clustered, aggregated in
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pseudostromata, or fused with common ostiole, immersed
to sessile, brown-black but usually covered by thallus,
globose to pear-shaped or conical, coriaceous to carbonaceous, ostiolate, ostiole round. Excipulum dense, consisting
of compressed hyphae, appearing prosoplectenchymatous
in thin, bleached sections but structure usually difficult to
observe due to heavy carbonization, generally dark brown
to brown-black. Hamathecium comprising 0.5–0.7(–1.5)
µm wide paraphysoids, hyaline, straight or rarely flexuose,
branched and usually anastomosing, usually embedded in a
thick, gelatinous matrix, occasionally with hyaline or yellow oil inspersion. Asci 1–8-spored, bitunicate, fissitunicate,
obclavate to cylindrical, short pedicellate, with refractive
ring and very wide, non-amyloid ocular chamber comprising
more than half of the width of the ascus. Ascospores irregularly arranged to biseriate, fusiform-ellipsoid to oblong,
hyaline to dark brown, septate to muriform, with distosepta
and sometimes additional eusepta and rectangular to diamond-shaped (generally six-angled) lumina, smooth-walled
or rarely ornamented, not or slightly constricted at the septa,
with evanescent mucilaginous material on the outside, either
as polar pads or irregular median pads or a complete sheath
enveloping the whole ascospore. Pycnidia known from a few
species, immersed, visible as black dots in specific, often
pseudostromatic areas on fertile thalli, or whole thalli only
producing pycnidia; rarely old ascospores within asci transforming into pycnidia. Conidia acrogenous, hyaline, aseptate, bacillar. Chemistry: lichexanthone sometimes produced
on thallus surface; yellow to orange or red anthraquinones
and perylenequinones often produced in the medulla of the
thalline layer covering the perithecia and sometimes in the
thallus medulla or superficial on various parts of the thallus.
Type: Trypethelium Spreng.
Notes: Trypetheliaceae (Zenker in Goebel and Kunze
1827) is one of the oldest described families of lichenized
Ascomycota. Its delimitation was obscure throughout the past
nearly 200 years, but in general the family included pyrenocarpous, epiphytic lichens with crustose thalli containing a
Trentepohlia photobiont, anastomosing paraphysoids forming a network embedded in a gelatinous matrix, bitunicate
asci, and hyaline, distoseptate ascospores with diamondshaped lumina (Aptroot 1991; Harris 1995; Aptroot et al.
2008; Sweetwood et al. 2012). Several genera that had been
included at some point have subsequently been placed in
other families, such as Megalotremis, Ornatopyrenis, and
Trypetheliopsis (Aptroot 1991; Harris 1995; Lumbsch and
Huhndorf 2010). Traditionally, the family included seven
core genera: Pseudopyrenula (ecorticate; ascospores astrothelioid), Polymeridium (ecorticate; ascospores euseptate),
Trypethelium (corticate; ascomata with apical ostiole;
ascospores astrothelioid, transversely septate), Laurera (corticate; ascomata with apical ostiole; ascospores astrothelioid,
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Fungal Diversity (2020) 105:17–318
Fig. 77 Best-scoring maximumlikelihood tree of Trypetheliaceae, showing all currently
sequenced and recognized
genera, based on the mtSSU
marker, with a length of 962
bases. The tree was reconstructed in RAxML 8, using
the universal GTR-Gamma
model without site partitioning,
and the final likelihood was
− 7368.672686. The currently
accepted genera are highlighted
muriform), Astrothelium (corticate; ascomata with lateral or
shared ostiole; ascospores astrothelioid, transversely septate),
Campylothelium (corticate; ascomata with lateral ostiole;
ascospores astrothelioid, muriform), and Cryptothelium (corticate; ascomata with shared ostiole; ascospores astrothelioid,
13
muriform). Four additional genera were included or recognized in more recent treatments (Aptroot 1991; Tucker and
Harris 1980; Harris 1986, 1995; Lücking et al. 2007): Exiliseptum (corticate; ascomata with shared ostiole; ascospores
euseptate), Bathelium (corticate; ascomata pseudostromatic,
Fungal Diversity (2020) 105:17–318
with walls composed of brown, jigsaw-puzzle-shaped hyphal
cells), Architrypethelium (corticate; ascomata with apical or
lateral ostiole; ascospores transversely septate, very large,
hyaline or brown), and Aptrootia (corticate; ascomata with
apical ostiole; ascospores muriform, dark brown).
Molecular phylogenetic data (Fig. 77) have profoundly
changed both the family circumscription of Trypetheliaceae
and the delimitation of its genera (Del Prado et al. 2006;
Nelsen et al. 2009, 2011a, 2014; Aptroot and Lücking 2016;
Lücking et al. 2016; Miranda-González et al. 2020). Apart
from the core Trypetheliaceae, the family now also contains
a large number of species previously placed in the genera
Arthopyrenia and Mycomicrothelia, for which the genera
Bogoriella, Constrictolumina and Novomicrothelia have
been established or reinstated (Aptroot and Lücking 2016;
Lücking et al. 2016). In addition, Polypyrenula has been
shown to be another member of Trypetheliaceae (MirandaGonzález et al. 2020). Other newly added elements are
Alloarthopyrenia (Hyde et al. 2016b) and Julella fallaciosa
(Nelsen et al. 2011a; Lücking et al. 2016). Based on molecular data, also the core Trypetheliaceae were reorganized,
with most species now in a single genus, Astrothelium, and
additional lineages with special characters allocated in the
genera Architrypethelium, Aptrootia, Bathelium, Dictyomeridium (segregated from Polymeridium), Marcelaria (Laurera
purpurina group), Nigrovothelium (Trypethelium tropicum
group), Polymeridium sensu stricto, Pseudopyrenula, and
Trypethelium sensu stricto (T. eluteriae group). The previously recognized genera Campylothelium, Cryptothelium,
and Laurera are now considered synonyms of Astrothelium (Aptroot and Lücking 2016). As a whole, Trypetheliaceae forms a strongly supported, somewhat isolated clade
within Dothideomycetes, and has therefore been assigned its
own order, Trypetheliales (Aptroot et al. 2008). Recently,
the mostly lichenicolous family Polycoccaceae was also
assigned to this order (Ertz et al. 2015).
The hamathecial hyphae (physes) have variously been
labeled paraphysoids or (trabeculate or trabecular) pseudoparaphyses (Henssen and Jahns 1973; Barr 1979a; Eriksson 1981; Liew et al. 2000). These terms were originally
coined for hamathecium physes of presumably ascolocular
fungi, to distinguish from true paraphyses. As per formal
definition, the various types are clearly distinguished (Barr
1979a; Henssen and Jahns 1973; Eriksson 1981): true paraphyses develop from below and remain mostly unbranched;
paraphysoids develop from stretching interascal plectenchyma; and pseudoparaphyses grow from top to bottom
during ontogeny, attaching below and eventually becoming
free above. However, the ontogeny of most Ascomycota
regarding these features has not been studied and there are
intermediate types. Furthermore, the terminology is confounded, e.g. by paraphysoids being named with ‘trabecular pseudoparaphyses’ (Eriksson 1981), thus combining
175
two supposedly different types into one. Also, according
to Henssen and Jahns (1973), paraphysoids can either be
precursors of true paraphyses or the only physes, which by
extension means that paraphysoids can occur both in ascohymenial and in ascolocular fungi. In addition, ascohymenial and ascolocular fungi do not form phylogenetic entities
and taxa thought to be closely related are now known to
be unrelated and viceversa (Liew et al. 2000; Schoch et al.
2009a, b). For instance, Henssen and Jahns (1973) included
Trypethelium in Pyrenulaceae, a family forming paraphyses (e.g. Pyrenula) or paraphysoids (e.g. Trypethelium),
but whose elements actually belong to two different classes
(Eurotiomycetes and Dothideomycetes). Eriksson (1981)
also hypothesized a close relationship between these two lineages, using the term paraphyses for Pyrenulaceae and paraphysoids for Trypetheliaceae. We consider this terminology
largely misleading and suggest to use only paraphysoids.
Trypetheliaceae now includes 18 genera plus some currently orphaned lineages (see key below) and well over 400
species, with about 800 predicted globally (Aptroot et al.
2016, 2019), mostly found in tropical lowland to lower montane, rain forest, dry forest, and savanna habitats.
Trypethelium Spreng., Anleit. Kennt. Gew. 3: 350 (1804).
MycoBank number: MB 5628; Index Fungorum number:
IF 5628; Facesoffungi number: FoF 08785; 16 morphologically defined species (Aptroot and Lücking 2016; Lücking
et al. 2017); molecular data available for six species (Lücking et al. 2016).
Lichenized on bark in terrestrial, lowland to lower montane tropical habitats. Thallus corticate, yellow-brown to
olive-green. Photobiont Trentepohlia. Ascomata aggregated in pseudostromata, sessile, brown-black or covered
by algal-free, sterile tissue filled with crystalline, yellow to
orange, KOH+ red to purple pigment, globose, coriaceous
to carbonaceous, ostiolate, ostiole round. Excipulum prosoplectenchymatous, dark brown to brown-black. Hamathecium comprising 0.5–0.7 µm wide paraphysoids, hyaline,
straight, branched and anastomosing, embedded in a thick,
gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate,
obclavate, short pedicellate, with refractive ring and nonamyloid ocular chamber. Ascospores irregularly arranged
to biseriate, fusiform, hyaline, multi-septate, with thin distosepta and more or less rectangular lumina, smooth-walled,
not constricted at the septa, surrounded by an evanescent
mucilaginous sheath. No asexual state known. Chemistry:
Yellow to orange or red anthraquinones and perylenequinones produced in the pseudostromata.
Type species: Trypethelium eluteriae Spreng., Anleit.
Kennt. Gew. 3: 350 (1804).
Notes: A tropical, lichenized genus found on bark of
branches and trunks of trees, usually in (semi-)exposed
situations. Traditionally, Trypethelium included all
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Trypetheliaceae with thallus-dominated perithecia with
separate, apical ostioles and transversely septate, astrothelioid ascospores. In its current circumscription, the genus
is limited to species with pseudostromatic perithecia, the
pseudostromata typically containing yellow to orange, crystalline pigments, and multi-septate ascospores with more or
less thin septa and walls and rectangular lumina (Aptroot
and Lücking 2016).
Other genera included
Alloarthopyrenia Phukhams., Lücking & K.D. Hyde in
Hyde et al., Fungal Diversity 80: 122, 131 (2016).
MycoBank number: MB 552236; Index Fungorum number: IF 552236; Facesoffungi number: FoF 02379; one phylogenetically and morphologically defined species (Hyde
et al. 2016b).
Saprobic on tree branches in terrestrial, temperate forest habitats. Thallus and photobiont absent. Ascomata
solitary, erumpent, carbonaceous, ostiolate, ostiole apical,
periphysate. Involucrellum distinct, dark brown. Excipulum
prosoplectenchymatous, brownish above, colorless below.
Hamathecium comprising 0.5–1 µm wide paraphysoids,
hyaline, straight, branched and anastomosing, embedded in
a gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate,
clavate, short pedicellate, with a non-amyloid ocular chamber. Ascospores irregularly arranged to biseriate, fusiformellipsoid to oblong, hyaline, 1-septate, euseptate, with secondary constrictions in each cell, rough-walled, constricted
at the septum, with a gelatinous halo. Pycnidia unknown.
Chemistry: no substances detected by TLC.
Type species: Alloarthopyrenia italica Phukhams.,
Camporesi, Ariyaw. & K.D. Hyde in Hyde et al., Fungal
Diversity 80: 122, 135 (2016).
Notes: This monospecific genus was recently introduced
for a previously undescribed species growing saprobically
on tree branches in Italy. It represents one of now four separate clades in Trypetheliaceae that include members of the
collective genus Arthopyrenia, besides Constrictolumina,
Macroconstrictolumina (see below) and “Arthopyrenia”
aff. minor. Three of these lineages feature ascospores with
characteristic secondary constrictions in each of the cells.
The discovery of this novel lineage further challenges the
systematics of the families Arthopyreniaceae and Naetrocymbaceae the genera Arthopyrenia sensu stricto and Naetrocymbe. Lücking and Nelsen (in Hyde et al. 2013) argued
that the type of Arthopyrenia, A. cerasi (Schrad.) A. Massal.
differs from the lineages now included in Trypetheliaceae in
various anatomical characters, thus likely representing an
unrelated lineage; the same argument was made for Naetrocymbe, with the type species N. fuliginea Körb. (Dai et al. in
Hyde et al. 2013). However, almost every sequenced member of the collective genera Arthopyrenia and Julella has so
far been found to represent a lineage of Trypetheliaceae, and
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so the fate of Arthopyrenia sensu stricto and Naetrocymbe,
whose type species have not yet been sequenced, remains
uncertain.
Aptrootia Lücking & Sipman in Lücking et al., Lichenologist 39: 188 (2007).
MycoBank number: MB 29134; Index Fungorum number:
IF 29134; Facesoffungi number: FoF 08786; three morphologically and phylogenetically defined species (Aptroot and
Lücking 2016; Lücking et al. 2016, 2017).
Lichenized on bark or bryophytes or soil in terrestrial,
chiefly montane tropical forest habitats. Thallus distinctly
corticate, yellow-brown to mostly green, or ecorticate
and greyish. Photobiont Trentepohlia. Ascomata solitary,
immersed to erumpent, brown-black but usually covered
by thick thallus layer except for ostiolar area, globose to
pear-shaped or ovoid, coriaceous to carbonaceous, ostiolate,
ostiole apical. Involucrellum reduced. Excipulum prosoplectenchymatous, colorless to brownish. Hamathecium
comprising 0.5–1 µm wide paraphysoids, hyaline, straight,
branched and anastomosing, embedded in a gelatinous
matrix. Asci 1(–2)-spored, bitunicate, fissitunicate, oblongellipsoid, short pedicellate, with a non-amyloid ocular chamber. Ascospores oblong-ellipsoid, initially hyaline and I+
amyloid, but soon becoming dark brown, richly muriform,
weakly distoseptate, wall smooth to granular ornamented,
outer wall easily breaking under pressure, with gelatinous
sheath. Pycnidia unknown. Chemistry: no substances
detected by TLC.
Type species: Aptrootia terricola (Aptroot) Lücking,
Umaña & Chaves in Lücking et al., Lichenologist 39: 188
(2007).
Notes: For key and discussion see Aptroot and Lücking
(2016). The genus currently includes three species (Fig. 78).
Architrypethelium Aptroot, Biblioth. Lichenol. 44: 120
(1991).
MycoBank number: MB 26294; Index Fungorum number:
IF 26294; Facesoffungi number: FoF 08787; eight morphologically defined species (Aptroot and Lücking 2016; Lücking et al. 2017; Luangsuphabool et al. 2018); molecular data
available for four species (Lücking et al. 2016; Luangsuphabool et al. 2018).
Lichenized on bark in terrestrial, chiefly lowland to
lower montane tropical habitats. Thallus distinctly corticate, yellow-brown to olive-green. Photobiont Trentepohlia.
Ascomata solitary to aggregated, erumpent to prominent,
brown-black but usually covered by thallus layer except for
ostiolar area, coriaceous to carbonaceous, ostiolate, ostiole apical or lateral. Involucrellum reduced, brown-black.
Excipulum prosoplectenchymatous, brownish to colorless.
Hamathecium comprising 0.5–1 µm wide paraphysoids,
hyaline, straight, branched and anastomosing, embedded in
Fungal Diversity (2020) 105:17–318
177
Fig. 78 Aptrootia terricola (Sipman 35503; f, j: Sipman 51849). a
Herbarium package and specimen. b, c, f Thallus with ascomata. d
Vertical section through an ascoma. e Ostiolar region with periphyses. g–h Perithecial wall in thin section, and the outer layer become
reddish in 10% KOH. i Paraphysoids. j Trentepohlioid photobiont. k,
l Asci. m, n Ascospores. o Ascospore in Lugol’s iodine. Scale bars:
b, c = 1 mm, d = 200 µm, e, k–n = 50 µm, f = 500 µm, g–i = 20 µm,
j = 10 µm
a gelatinous matrix, in some species inspersed with oil droplets. Asci 2–8-spored, bitunicate, fissitunicate, oblong-ellipsoid, short pedicellate, with a non-amyloid ocular chamber.
Ascospores irregularly arranged, oblong-ellipsoid, hyaline to
mostly dark brown, 3–5-septate and very large (usually over
100 µm) or rarely muriform and small, somewhat distoseptate but not astrothelioid, smooth-walled or with longitudinal
wall folds or crystals, not or slightly constricted at the septa,
13
178
surrounded by a mucilaginous sheath. Pycnidia unknown.
Chemistry: lichexanthone sometimes produced on thallus
surface.
Type species: Architrypethelium seminudum (Mont.) Aptroot. [= Architrypethelium nitens (Fée) Aptroot in Aptroot
et al., Biblioth. Lichenol. 97: 38 (2008)].
Notes: For discussion see Aptroot and Lücking (2016).
The genus currently includes eight species. With the recent,
surprising discovery of a species with rather small, muriform ascospores falling into this clade (Luangsuphabool
et al. 2018), the generic boundaries towards Astrothelium
have become more diffuse.
Astrothelium Eschw., Syst. Lich.: 18, 26 (1824).
MycoBank number: MB 443; Index Fungorum number:
IF 443; Facesoffungi number: FoF 08788; Over 250 morphologically defined species (Aptroot and Lücking 2016;
Lücking et al. 2017); molecular data available for over 50
species (Lücking et al. 2016).
Lichenized on bark in terrestrial, chiefly lowland to lower
montane tropical habitats. Thallus distinctly corticate,
yellow-brown to mostly green. Photobiont Trentepohlia.
Ascomata scattered, clustered, aggregated in pseudostromata, or fused with common ostiole, immersed to sessile,
brown-black but usually covered by thick thallus layer
except for ostiolar area, globose to pear-shaped or conical,
coriaceous to carbonaceous, ostiolate, ostiole round. Excipulum prosoplectenchymatous, dark brown to brown-black.
Hamathecium comprising 0.5–0.7 µm wide paraphysoids,
hyaline, straight, branched and anastomosing, embedded in
a thick, gelatinous matrix. Asci 1–8-spored, bitunicate, fissitunicate, obclavate, short pedicellate, with a non-amyloid
ocular chamber. Ascospores irregularly arranged to uni- or
biseriate, fusiform-ellipsoid to oblong-cylindrical, hyaline,
septate to muriform, with distinct distosepta and sometimes
eusepta and diamond-shaped lumina (best visible in septate
ascospores), smooth-walled, not or slightly constricted at
the septa, often surrounded by an evanescent mucilaginous
material on the outside, either as polar pads, or irregular
median pads or a complete sheath enveloping the whole
ascospore. Pycnidia known from a few species, immersed
to erumpent, visible as black dots. Conidia acrogenous, rodshaped, hyaline. Chemistry: lichexanthone sometimes produced on thallus surface; yellow to orange or red anthraquinones and perylenequinones often produced in the medulla
of the thalline layer covering the perithecia and sometimes
in the thallus medulla or superficial on various parts of the
thallus.
Type species: Astrothelium conicum Eschw., Syst. Lich.:
26 (1824) [= Astrothelium cinnamomeum (Eschw.) Müll.
Arg., Flora 67: 670 (1884)].
Notes: Astrothelium traditionally included species with
thallus-dominated perithecia featuring lateral, fused ostioles
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Fungal Diversity (2020) 105:17–318
and transversely septate, astrothelioid ascospores. In its modern circumscription (Aptroot and Lücking 2016), practically
all species with thickly corticate thalli, thallus-dominated
perithecia, and astrothelioid ascospores are now included
in this genus, regardless of the disposition of the perithecia and their ostioles or ascospore septation, with a total
of over 250 taxa (Aptroot and Lücking 2016; Cáceres and
Aptroot 2017; Aptroot and Weerakoon 2018; Aptroot et al.
2019). Astrothelium is essentially a fusion of the previously
separated genera Astrothelium, Bathelium p.p., Campylothelium, Cryptothelium, Laurera, and Trypethelium p.p. Harris (1995) had already predicted the existence of this clade,
suggesting the use of the name Laurera. However, Laurera
postdates Astrothelium, a name Harris (1995) intended to
reserve for a small group of species that have been shown to
form part of a single, large clade including most species of
the family (Lücking et al. 2016).
Bathelium Ach., Method. Lich.: 111 (1803).
MycoBank number: MB 517; Index Fungorum number:
IF 517; Facesoffungi number: FoF 08789; 16 morphologically defined species (Aptroot and Lücking 2016; Lücking
et al. 2017); molecular data available for four species (Lücking et al. 2016).
Lichenized on bark in terrestrial, chiefly lowland to lower
montane tropical habitats, often in drier situations. Thallus
corticate, yellow-brown to olive-green. Photobiont Trentepohlia. Ascomata solitary to aggregated in distinct pseudostromata, prominent to sessile, brown-black and remaining
fully exposed, rarely pruinose, coriaceous to carbonaceous,
ostiolate, ostiole apical. Involucrellum reduced. Excipulum
brown-black. Hamathecium comprising 0.5–0.7 µm wide
paraphysoids, hyaline, straight, branched and anastomosing, embedded in a gelatinous matrix. Asci 1–8-spored,
bitunicate, fissitunicate, clavate to oblong, short pedicellate,
with a non-amyloid ocular chamber. Ascospores irregularly
arranged to biseriate, fusiform-ellipsoid, hyaline, septate to
mostly muriform, more or less euseptate, smooth-walled,
not or slightly constricted at the septa (often at the middle
septum only), surrounded by a gelatinous sheath. Pycnidia
unknown. Chemistry: lichexanthone sometimes produced
on thallus surface; yellow to orange anthraquinones often
produced in the medulla of the pseudostromata.
Type species: Bathelium mastoideum Afz. ex Ach.,
Method. Lich.: 111 (1803).
Notes: For key and discussion see Aptroot and Lücking
(2016).
Bogoriella Zahlbr., Annals Cryptog. Exot. 1(2): 111 (1928).
MycoBank number: MB 608; Index Fungorum number:
IF 608; Facesoffungi number: FoF 08790; 18 morphologically defined species (this paper); molecular data available
Fungal Diversity (2020) 105:17–318
179
Fig. 79 a–d Bogoriella complexoluminata (holotype). a Thallus with
ascomata. b–d Ascospores in various stages of development. e B.
isthmospora (Aptroot s.n.), ascospores in various stages of development. f Schummia angulata (Aptroot 14061), ascospores in various
stages of development. g–k Macroconstrictolumina megalateralis
(holotype). g Thallus with ascomata. h Hamathecium and ascospores.
i–k Ascospores in various stages of development. Scale bars: a, g = 1
mm, b–f, h–k = 10 µm. Photographs by André Aptroot except e and
f, by Felix Schumm
for four species (Lücking et al. 2016; Zhang et al. 2017b;
this paper).
= Ornatopyrenis Aptroot, Biblioth. Lichenol. 44: 127
(1991). Type species: Ornatopyrenis queenslandica (Müll.
Arg.) Aptroot [≡ Bogoriella queenslandica (Müll. Arg.)
Aptroot & Lücking].
= Distothelia Aptroot in Seaward & Aptroot, Bryologist
108: 284 (2005). Type species: Distothelia isthmospora
Aptroot in Seaward & Aptroot, Bryologist 108: 284 (2005).
= Novomicrothelia Aptroot, M.P. Nelsen & Lücking in
Lücking et al., Lichenologist 48: 757 (2016). Type species:
Novomicrothelia oleosa (Aptroot) Aptroot, M.P. Nelsen &
Lücking in Lücking et al., Lichenologist 48: 758 (2016).
Lichenized (sometimes barely so or apparently nonlichenized) on bark in terrestrial, chiefly lowland to lower
montane tropical habitats. Thallus ecorticate, mostly whitish to pale brownish. Photobiont Trentepohlia. Ascomata
solitary, erumpent to prominent, brown-black to carbonaceous, hemisphaerical to wart-shaped or conical, coriaceous
to carbonaceous, ostiolate, ostiole apical to rarely lateral.
Involucrellum usually well-developed, carbonaceous. Excipulum prosoplectenchymatous, usually brownish, in lateral
and apical parts typically fused with the involucrellum.
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Fungal Diversity (2020) 105:17–318
Fig. 80 Classification tree for non-sequenced species of Bogoriella
sensu lato and Constrictolumina sensu lato [QUERY] based on phenotype-based phylogenetic binning into a molecular reference tree of
sequenced species (in boldface). Numbers in parentheses behind each
query taxon represent bootstrap support for the corresponding placement. Queries with alternative placements supported with less than
70% are greyed out
Hamathecium comprising 1(–2) µm wide para-physoids,
hyaline, straight to somewhat wavy, branched and anastomosing, embedded in a thin, gelatinous matrix. Asci 8-spored,
bitunicate, fissitunicate, (ob-)clavate to fusiform, short pedicellate, with a non-amyloid ocular chamber. Ascospores
irregularly arranged to uni- or biseriate, fusiform-ellipsoid
to oblong, grey-brown, 1–5-septate to small muriform,
with eusepta and rectangular lumina or with thickened
distosepta making the lumina appear halter-shaped, wall
smooth to finely granular ornamented, not constricted at
the septa or rarely with constrictions. Pycnidia known from
a few species, immersed to erumpent, visible as black dots.
Conidia fusiform, aseptate, bacillar, hyaline, 4–5 × 0.8 µm.
Chemistry: no substances detected by TLC.
Type species: Bogoriella subpersicina Zahlbr., Annals
Cryptog. Exot. 1: 111 (1928) [= Bogoriella decipiens (Müll.
Arg.) Aptroot & Lücking].
Sequence data for Bogoriella decipiens: mtSSU:
MT968881.
Notes: Aptroot and Lücking (2016) adopted the name
Bogoriella for more or less lichenized, tropical species previously classified in the genus Mycomicrothelia (Hawksworth
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Fungal Diversity (2020) 105:17–318
1985). The expanded molecular phylogeny now available
shows that these taxa form several lineages (Fig. 77): two
in a paraphyletic grade formed by the most basally diverging lineages, and three in a distant clade, forming several
lineages on long branches. One of these had been named
Novomicrothelia (Aptroot and Lücking 2016; Lücking et al.
2016), whereas a second clade of previously unpublished
sequences includes the type species of Bogoriella, together
with a new taxon, both with small muriform ascospores,
one with eusepta and the other with peculiar distosepta
(Fig. 79a–d). The third clade is formed by the recently
described species Novomicrothelia pandanicola S.N. Zhang
& K.D. Hyde (Zhang et al. 2017b). In the original analysis,
nuLSU sequence data suggested a phylogenetic position
of the latter close to N. oleosa, prompting inclusion in that
genus. Our expanded phylogeny based on new mtSSU data
showed that the genus Bogoriella, for which no data were
originally available (Zhang et al. 2017b), is nested within
Novomicrothelia sensu lato. Novomicrothelia pandanicola
forms small, rather narrow (20–35 × 7–10 µm, about 3–3.5
times as long as broad), muriform ascospores with thin
walls and septa and constrictions at the septa, whereas in
the Bogoriella sensu stricto clade the ascospores are broadly
ellipsoid (20–30 × 12–16 µm, about 1.5–2 times as long
as broad) and muriform with either thin or conspicuously
thickened walls and septa. Novomicrothelia sensu stricto
has thin-walled, 1-septate ascospores without constrictions.
The species hitherto classified in Bogoriella sensu lato
(including Novomicrothelia) thus represent a polyphyletic
assembly. Given that most species of this morphodeme have
not been sequenced yet, we employed morphology-based
phylogenetic binning (Berger et al. 2011; Parnmen et al.
2012; Lücking and Kalb 2018) to assess the likely placement
of the remaining species. As a result, 13 species clustered
with support with the basal paraphyletic grade formed by
B. minutula, B. miculiformis, and B. hemisphaerica, seven
clustered with Novomicrothelia sensu stricto, none with N.
pandanicola, four outside but close to Novomicrothelia in an
unresolved position, and three with Bogoriella sensu stricto
(Fig. 80). Given this result, the underlying topology, and
the observation that taxa with different ascospore types (B.
complexoluminata and B. decipiens) are closely related, we
saw no alternative to merging Bogoriella and Novomicrothelia into a single genus, under the name Bogoriella. The
basal grade is here recognized conservatively as a separate,
single genus, under the name Pseudobogoriella (see below).
Based on the phylogenetic analysis and the binning
exercise, Bogoriella in its revised sense includes three
generic synonyms, since the type species of Distothelia (D.
isthmospora) and Ornatopyrenis (O. queenslandica) are also
assigned to this genus by the binning approach. Indeed, the
ascospores of the type species of the latter two genera are
181
similar to those of B. complexoluminata (Fig. 79b–e) in
development and endospore formation (Aptroot 1991; Harris
1995; Seaward and Aptroot 2005). One species previously
placed in Distothelia, D. angulata, is here transferred to the
new genus Schummia, as its ascospores are quite different
(Fig. 79f; see below).
New species and new combinations in Bogoriella
Bogoriella chiquitana (Flakus, Kukwa & Aptroot) Lücking,
R. Miranda & Aptroot comb. nov.
MycoBank number: MB 836785; Index Fungorum number: IF 836785; Facesoffungi number: FoF 08791
Bas.: Constrictolumina chiquitana Flakus, Kukwa & Aptroot, Lichenologist 48: 681 (2016).
Bogoriella complexoluminata Aptroot & Lücking sp. nov.
(Fig. 79a–d)
MycoBank number: MB 836786; Index Fungorum number: IF 836786; Facesoffungi number: FoF 08792
Etymology: Named for the complex lumina of the
ascospores.
Diagnosis: Corticolous Bogoriella with thallus UV-negative; ostioles apical, ascospores 27–37 × 10–15 µm, initially
hyaline and 1-septate with an euseptum, constricted at the
septum, wall thickened in two open distosepta two places in
each cell, so as to from three pseudoloculi per cell; mature
ascospores brown with 1 dark brown median euseptum and
a distoseptate wall that leaves six rows of 2–6 oval loculi of
c. 5 µm diam.
Holotype: A. Aptroot 78050 (CGMS; isotype: ABL).
GenBank accessions for type-based sequences: mtSSU:
MT968880.
Thallus inapparent, immersed in the soft, corky bark,
covering up to 10 cm diam., apparently not lichenized; no
algae observed. Ascomata erumpent to sessile, hemisphaerical with flattened top, black, 0.3–0.5 mm diam. and up to 0.4
mm high; ostiole black, minute, usually in a c. 0.1 mm wide
and 0.05 mm deep depression; clypeus covering the top and
the sides; wall c. 100 µm thick at the sides, only c. 20 µm
thick below the hamathecium. Hamathecium clear, hyaline,
of c. 1 µm wide filaments that are anastomosing above the
asci. Asci broadly cylindrical, c. 100 × 25 µm, wall c. 2
µm thick all around; without ocular chamber. Ascospores
8/ascus, 27–37 × 10–15 µm, initially hyaline and 1-septate
with an euseptum, constricted at the septum, wall thickened
in two open distosepta two places in each cell, so as to form
three pseudoloculi per cell; mature ascospores brown with
1 dark brown median euseptum and a distoseptate wall that
leaves six rows of 2–6 oval loculi of c. 5 µm diam. Pycnidia
not observed.
Chemistry: Thallus UV–, C–, P–, K–. TLC: nil.
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Ecology and distribution: On tree bark in rural and urban
areas; thus far only known from Brazil.
Material examined: Brazil. Mato Grosso do Sul, Campo
Grande, UFMS campus; 20°30’ S, 54°37’ W, 550 m; on soft,
corky bark of Aspidosperma tomentosa (Apocynaceae) tree
(identified by Dr Angela Sartori); 22 November 2018, A.
Aptroot 78050 (holotype: CGMS; isotype: ABL); Bonito,
outskirts, near tower; 21°07′44″ S, 56°30′41″ W, 475 m; on
bark of tree; 9 November 2018, A. Aptroot 78013 (paratype:
CGMS).
Notes: This new species is phylogenetically closely
related to the type species, Bogoriella decipiens, but the
latter differs in having regularly muriform ascospores. The
endospore development of B. complexoluminata appears to
be unique, producing a peculiar shape of the lumina.
3.
Bogoriella isthmospora (Aptroot) Lücking, R. Miranda &
Aptroot comb. nov. (Fig. 79e)
MycoBank number: MB 836787; Index Fungorum number: IF 836787; Facesoffungi number: FoF 08793
Bas.: Distothelia isthmospora Aptroot in Seaward & Aptroot, Bryologist 108: 284 (2005).
6.
Bogoriella pandanicola (S.N. Zhang & K.D. Hyde) S.N.
Zhang, Lücking & Aptroot comb. nov.
MycoBank number: MB 836822; Index Fungorum number: IF 836822; Facesoffungi number: FoF 09232.
Bas.: Novomicrothelia pandanicola S.N. Zhang & K.D.
Hyde in Zhang et al., Phytotaxa 321: 258 (2017).
Sequence data: mtSSU: MT967501.
7.
Bogoriella rubrostoma (Aptroot) Lücking, R. Miranda &
Aptroot comb. nov.
MycoBank number: MB 836788; Index Fungorum number: IF 836788; Facesoffungi number: FoF 08794
Bas.: Mycomicrothelia rubrostoma Aptroot, Biblioth.
Lichenol. 44: 135 (1991); Distothelia rubrostoma (Aptroot)
Aptroot & Lücking, Lichenologist 48(6): 927 (2016).
Key to species of Bogoriella:
1.
1.
2.
2.
Ascospores becoming distinctly muriform, with thin
eusepta and rectangular lumina………………………2
Ascospores 1-septate to rarely 3-septate or submuriform, with thin eusepta or with irregularly thickened
endospore and distosepta……………………………4
Ascospores becoming thick-walled and with thickened
median septum, 28–45 µm long………………………
…………………………………Bogoriella chiquitana
Ascospores remaining thin-walled, 20–35 µm
long…………………………………………………3
13
3.
4.
4.
5.
5.
6.
7.
8.
8.
9.
10.
10.
10.
11.
11.
12.
12.
13.
13.
14.
14.
15.
15.
16.
16.
17.
17.
Ascospores 20–30 × 12–16 µm, about 1.5–2 times as
long as broad……………………Bogoriella decipiens
Ascospores 20–35 × 7–10 µm, about 3–3.5 times as
long as broad…………………Bogoriella pandanicola
Ascospores with irregularly thickened endospore and
distosepta, 1-septate or rarely 3-septate to submuriform; lumina irregular to halter-shaped………………5
Ascospores with thin eusepta, 1-septate or rarely 3-septate; lumina rectangular………………………………8
Ascospores (1–)3-septate to appearing submuriform,
27–45 × 10–17 µm……………………………………6
Ascospores 1-septate but cells with irregular endospore
invaginations, 15–29 × 6–14 µm……………………7
Ascospores mostly distinctly 3-septate, occasionally
1-septate or with additional longitudinal septa………
……………………………Bogoriella queenslandica
Ascospores with one distinct, rather dark central septum and the two cells with irregular endospore invaginations giving them the appearance of a (sub-)muriform division…………Bogoriella complexluminata
Ostiole with red pigment; ascospores 23–29 × 10–14
µm, secondary endospore invaginations developing
late……………………………Bogoriella isthmospora
Ostiole lacking pigment; ascospores 15–18 × 6–8 µm,
secondary endospore invaginations developing soon…
………………………………Bogoriella rubrostoma
Ostiole lateral…………………Bogoriella triangularis
Ostiole apical…………………………………………9
Hamathecium inspersed……………………………10
Hamathecium clear…………………………………12
Ascospores 24–27 × 8–11 µm…………………………
……………………………………Bogoriella oleosa
Ascospores 30–45 × 10–15 µm……………………11
Ascomata 0.5–0.7 mm diam.…………………………
………………………………Bogoriella xanthonica
Ascomata 0.7–1.1 mm diam.…………………………
………………………………Bogoriella macrocarpa
Ascospores becoming 3-septate………………………
……………………………………Bogoriella obovata
Ascospores remaining 1-septate……………………13
Ascospores 28–40 µm long……Bogoriella megaspora
Ascospores 17–27 µm long…………………………14
Black prothallus line present………………………15
Black prothallus line absent…………………………16
Asci 10–20 µm broad………Bogoriella conothelena
Asci 20–30 µm broad………Bogoriella thelena
Ascomata 0.8–1 mm diam., basally broadly
expanded………………………Bogoriella confluens
Ascomata 0.2–0.4 mm diam., basally barely or not
expanded……………………………………………17
Ascomata conical……………Bogoriella pachytheca
Ascomata flattened………………Bogoriella modesta
Fungal Diversity (2020) 105:17–318
Constrictolumina Aptroot, M.P. Nelsen & Lücking in Lücking et al., Lichenologist 48: 756 (2016).
MycoBank number: MB 816872; Index Fungorum number: IF 816872; Facesoffungi number: FoF 08795; five morphologically defined species (this paper); molecular data
available for two species (Lücking et al. 2016).
Lichenized on bark in terrestrial, chiefly lowland to lower
montane tropical habitats. Thallus ecorticate, usually whitish. Photobiont Trentepohlia. Ascomata solitary to rarely
aggregate, erumpent to prominent, carbonaceous, ostiolate,
ostiole apical to rarely lateral. Involucrellum carbonized.
Excipulum prosoplectenchymatous, brownish to colorless
below. Hamathecium comprising ca. 1 µm wide, hyaline,
straight, branched and anastomosing, pseudoparaphyses,
embedded in a gelatinous matrix. Asci 4–8-spored, bitunicate, fissitunicate, clavate, short pedicellate, with a nonamyloid ocular chamber. Ascospores irregularly arranged
to biseriate, fusiform-ellipsoid, hyaline, 1–3-septate, euseptate, with secondary constrictions in each cell and often with
slightly thickened walls, smooth-walled, slightly constricted
at the septum, with a gelatinous halo, rather small (15–30
× 5–11 µm). Pycnidia unknown. Chemistry: no substances
detected by TLC.
Type species: Constrictolumina cinchonae (Ach.) Lücking, M.P. Nelsen & Aptroot in Lücking et al., Lichenologist
48(6): 756 (2016).
Notes: This genus was recently introduced for lichenized
species of Arthopyrenia sensu lato with secondary cell wall
constrictions clustering within Trypetheliaceae (Aptroot
and Lücking 2016; Lücking et al. 2016). At the time, it
was already noted that the species formed two lineages, C.
malaccitula coming out separate. A phenotype-based phylogenetic binning approach of 11 species included in the
genus demonstrated that about half of them each cluster with
either with notable differences in the ascospores, those of
the C. malaccitula group being mostly larger [(20–)30–50 ×
(7–)10–16 µm)], with more distinct secondary constrictions,
and granular ornamented. We therefore introduce the genus
Macroconstrictolumina for this group below. Constrictolumina in a strict sense then retains five species, which are
keyed out below.
Key to species of Constrictolumina sensu stricto
1.
1.
2.
2.
3.
Ascospores 17–23 × 5–8 µm, 1–3-septate……………2
Ascospores (20–)22–46 × 7–24 µm, 1-septate…………3
Perithecia aggregate…Constrictolumina esenbeckiana
Perithecia solitary………Constrictolumina cinchonae
Ascospores 40–46 × 16–24 µm…………………………
……………………………Constrictolumina porospora
3. Ascospores 20–30 × 7–16 µm…………………………4
4. Perithecia aggregate; ascospores 3-septate……………
…………………………Constrictolumina leucostoma
183
4. Perithecia solitary; ascospores 1(–3)-septate…………
……………………………Constrictolumina planorbis
Dictyomeridium Aptroot, M.P. Nelsen & Lücking in Lücking et al., Lichenologist 48: 756 (2016).
MycoBank number: MB 816873; Index Fungorum number: IF 816873; Facesoffungi number: FoF 08796; seven
morphologically defined species (Aptroot and Lücking
2016); molecular data available for one species (Lücking
et al. 2016).
Lichenized on bark in terrestrial, chiefly lowland to lower
montane tropical habitats, typically in drier situations. Thallus ecorticate, usually whitish. Photobiont Trentepohlia.
Ascomata solitary to sometimes aggregated, erumpent to
prominent, somewhat pyriform when seen from above,
brown-black to grey-black, carbonaceous, ostiolate, ostiole lateral. Involucrellum carbonized. Excipulum prosoplectenchymatous, brownish. Hamathecium comprising
0.5–0.7 µm wide paraphysoids, hyaline, straight, branched
and anastomosing, embedded in a gelatinous matrix. Asci
2–8-spored, bitunicate, fissitunicate, clavate, short pedicellate, with a non-amyloid ocular chamber. Ascospores
irregularly arranged to biseriate, oblong-ellipsoid, hyaline,
often I+ amyloid, muriform, euseptate, smooth-walled, not
constricted at the septa, with a gelatinous sheath. Pycnidia
known from a few species, immersed to erumpent, visible as
black dots. Conidia aseptate, bacillar, hyaline. Chemistry:
lichexanthone sometimes produced on thallus surface.
Type species: Dictyomeridium proponens (Nyl.) Aptroot,
M.P. Nelsen & Lücking in Lücking et al., Lichenologist 48):
757 (2016)
Notes: For key and discussion see Aptroot and Lücking
(2016).
“Julella” fallaciosa (Stizenb. ex Arnold) R.C. Harris, in
Egan, Bryologist 90(2): 163 (1987)
MycoBank number: MB 132219; Index Fungorum number: IF 132219; Facesoffungi number: FoF 08797.
Notes: The genus Julella has usually been considered a
muriform-spored counterpart of Arthopyrenia sensu lato
(Barr 1985, 1987b; Harris 1995). Species assigned to this
genus are typically non-lichenized and possibly saprobic,
about some doubtful lichenization has been discussed (Aptroot 2002a). The few studies available about this genus seem
to disagree widely about species taxonomy. Thus, while
Harris (1995) distinguished at least 13 species, Aptroot
and van den Boom (1995) only accepted three. The latter
authors included all forms with 2–4(-6) ascospores in a single species, J. lactea (A. Massal.) M.E. Barr, whereas Harris (1995) keyed out six species in this group, separated by
ascospore size and numbers of ascospores per ascus. Among
taxa with 8-spored asci, Aptroot and van den Boom (1995)
13
184
distinguished two species: J. vitrispora (Cooke & Harkn.)
M.E. Barr with longer ascospores and larger ascomata and
J. sericea (A. Massal.) Coppins with shorter ascospores and
smaller ascomata. In contrast, Harris (1995) accepted at least
seven species in this group, distinguished by ascus shape
and ascospore arrangement, size, and color, among other
features. These differences are less the result of a disparate
species concept (the theoretical approach of both treatments
not being that much different) and more due to the fact that
these authors studied a very different set of names and material and hence came to such different conclusions. Given the
findings in other, similarly defined genera such as Arthopyrenia sensu lato, it is conceivable that Julella is also a highly
heterogeneous assemblage not only including many species
but also representing several, partly unrelated genera.
Only two species of Julella have been sequenced so
far, viz. J. avicenniae (Borse) K.D. Hyde and J. fallaciosa
(Nelsen et al. 2011a, 2014; Ariyawansa et al. 2013a; Lücking et al. 2016), thus not including the type species, J. buxi.
Julella avicenniae was shown to be different from the type
species in ascus type, the brownish ascospores, and the
habitat on bark of mangrove trees in the genus Avicennia, a
notion anticipated by Harris (1995), and consequently was
placed in a separate, monospecific genus and family, Halojulella and Halojulellaceae (Ariyawansa et al. 2013a). The
sequenced material of J. fallaciosa, which agrees well with
the key provided by Harris (1995) and Wirth et al. (2013)
clustered within Trypetheliaceae (Nelsen et al. 2011a, 2014;
Lücking et al. 2016; Fig. 77), which is in line with the ascus
featuring a broad ocular chamber and the hamathecial physes
being thin and anastomosing. Notably, most species currently placed in Julella agree with J. fallaciosa in ascus and
hamathecium type and are likely congeneric with J. fallaciosa, whereas the type, J. buxi, differs by the asci lacking a distinct tholus and ocular chamber and by the physes
being flexuose and only sparsely branched (Barr 1987b;
Ariyawansa et al. 2013a). This situation is thus comparable
to that of Arthopyrenia (Lücking and Nelsen in Hyde et al.
2013).
Barr (1987b) was the first to propose synonymy of the
genera Polyblastiopsis, based on P. naegelii (= Julella lacteal), and Peltosphaeria, based on P. vitrispora (≡ Julella
vitrispora), with Julella. Various workers pointed out the
asci with broad ocular chamber and the thin, anastomosing
physes of the type species of both genera (Riedl 1971; Barr
1987b; Mayrhofer 1987), which suggests that these taxa
are not actually congeneric with J. buxi but would likely
cover the clade falling in Trypetheliaceae. We have refrained
from taking up either name for this clade, given that most
species of this complex have not been sequenced and also
considering the complex situation emerging with the taxa
previously placed on Arthopyrenia sensu lato and Mycomicrothelia sensu lato, which now form several clades each
13
Fungal Diversity (2020) 105:17–318
within Trypetheliaceae. Indeed, the two types of Polyblastiopsis and Peltosphaeria differ in ascospore shape, being
(ob-)clavate in Polyblastiopsis naegelii (J. lactea group) and
cylindrical in Peltosphaeria vitrispora (J. vitrispora group).
Thus, if the two generic names indeed correspond to two
separate phylogenetic units, then Polyblastiopsis would be
the name applicable to the J. fallaciosa clade, as the latter
belongs in the J. lactea group (Harris 1995; Aptroot and van
den Boom 1995). However, if both groups are congeneric,
then the older name Peltosphaeria has priority.
Index Fungorum currently lists Polyblastiopsis sericea
(≡ Julella sericea) as type of Polyblastiopsis. However,
Clements and Shear (1931) had listed P. naegelii as lectotype, a selection adopted by Riedl (1971) and Barr (1987b).
Polyblastiopsis was established as replacement name for
Polyblastia (Müller 1882), an illegitimate later homonym
of Polyblastia [nom. cons.]. Since Zahlbruckner (1907) did
not cite a type, Polyblastiopsis is to be typified by the type
of Polyblastia, which means the lectotypification by Clements and Shear (1931) must be in accordance with Müller’s
(1882) protologue. This fortunately is the case, as P. naegelii
is explicitly listed in the protologue. While Aptroot and van
den Boom (1995) synonymized P. naegelii under J. lactea,
both Riedl (1971) and Barr (1987b) recognized the taxon
as variety of the latter, according to Riedl (1971) differing
in the complete (var. lactea) vs. cap-shaped (var. naegelii)
gelatinous sheath of the ascospores and in substrate preferences (Fraxinus vs. conifers). Given that the two taxa have
overlapping ranges, it seems appropriate to recognize them
as separate species, in which case the type of Polyblastiopsis
is to be cited as P. naegelii and not as P. lactea or P. sericea.
Macroconstrictolumina Lücking, R. Miranda & Aptroot
gen. nov.
MycoBank number: MB 836789; Index Fungorum number: IF 836789; Facesoffungi number: FoF 08798; Four
morphologically defined species (this paper); molecular data
available for two species (Lücking et al. 2016; this paper).
Diagnosis: Differing from Constrictolumina chiefly in the
larger ascospores with granular ornamentation.
Etymology: The name refers to the relatively large
ascospores.
Lichenized on bark in terrestrial, chiefly lowland to lower
montane tropical habitats. Thallus ecorticate, usually whitish. Photobiont Trentepohlia. Ascomata solitary to rarely
aggregate, erumpent to prominent, carbonaceous, ostiolate,
ostiole apical to rarely lateral. Involucrellum carbonized.
Excipulum prosoplectenchymatous, brownish to colourless
below. Hamathecium comprising ca. 1 µm wide pseudoparaphyses, hyaline, straight, branched and anastomosing,
embedded in a gelatinous matrix. Asci 2–8-spored, bitunicate, fissitunicate, clavate, short pedicellate, with a nonamyloid ocular chamber. Ascospores irregularly arranged
Fungal Diversity (2020) 105:17–318
to biseriate, fusiform-ellipsoid, hyaline, 1(–3)-septate,
euseptate, with secondary constrictions in each cell and
often with slightly thickened walls, typically granular ornamented, slightly constricted at the septum, with a gelatinous
halo, medium-sized [30–50 × 10–16(–24) µm], only in one
species (M. lyrata) somewhat smaller. Pycnidia unknown.
Chemistry: no substances detected by TLC.
Type species: Macroconstrictolumina malaccitula (Nyl.)
Lücking, R. Miranda & Aptroot (see below).
Notes: This new genus accommodates the lineage phylogenetically separate from the Constrictolumina cinchonae
group (Fig. 77), but which had so far been included in that
genus (Aptroot and Lücking 2016; Lücking et al. 2016).
The difference in ascospore size, in combination with the
granular ornamented ascospore wall, appears to be consistent, but further species of both genera need to be sequenced
to test this taxonomic hypothesis developed from the binning
exercise (Fig. 80).
New species and new combinations in Macroconstrictolu
mina:
Macroconstrictolumina lyrata (R.C. Harris) Lücking, R.
Miranda & Aptroot comb. nov.
MycoBank number: MB 836790; Index Fungorum number: IF 836790; Facesoffungi number: FoF 08799.
Bas.: Arthopyrenia lyrata R.C. Harris in Tucker & Harris,
Bryologist 83: 6 (1980); Constrictolumina lyrata (R.C. Harris) Lücking, M.P. Nelsen & Aptroot in Aptroot & Lücking,
Lichenologist 48: 921 (2016).
Sequence data: mtSSU: MT948052.
Macroconstrictolumina majuscula (Nyl.) Lücking, R.
Miranda & Aptroot comb. nov.
MycoBank number: MB 836791; Index Fungorum number: IF 836791; Facesoffungi number: FoF 08800.
Bas.: Verrucaria majuscula Nyl., Ann. Sci. Nat., Bot.,
Sér. 4 20: 253 (1863); Arthopyrenia majuscula (Nyl.)
Zahlbr., Cat. Lich. Univers. 1: 332 (1922); Ciferriolichen
majusculus (Nyl.) R.C. Harris, Moscosoa 6: 215 (1990);
Constrictolumina majuscula (Nyl.) Lücking, M.P. Nelsen
& Aptroot in Aptroot & Lücking, Lichenologist 48: 921
(2016).
Macroconstrictolumina malaccitula (Nyl.) Lücking, R.
Miranda & Aptroot comb. nov.
MycoBank number: MB 836792; Index Fungorum number: IF 836792; Facesoffungi number: FoF 08801.
Bas.: Verrucaria malaccitula Nyl. in Nylander & Crombie, J. Linn. Soc., Bot. 20: 61 (1883); Arthopyrenia malaccitula (Nyl.) Zahlbr., Cat. Lich. Univers. 1: 284 (1921);
Constrictolumina malaccitula (Nyl.) Lücking, M.P. Nelsen
& Aptroot in Aptroot & Lücking, Lichenologist 48: 921
(2016).
185
= Tomasellia dispora Müll. Arg., Flora, Regensburg 70:
427 (1887).
= Arthopyrenia bifera Zahlbr., Annls mycol. 33(1/2): 34
(1935).
= Megalotremis biocellata Aptroot, Biblthca Lichenol.
44: 125 (1991).
Notes: This species is being discussed in detail in the
Monoblastiaceae treatment of this issue (Lücking et al. in
this paper), based on the finding that Megalotremis biocellata, previously classified in a genus belonging to Monoblastiaceae, was found to be conspecific with M. malaccitula.
Macroconstrictolumina megalateralis Aptroot sp. nov.
(Fig. 79 g–k)
MycoBank number: MB 836793; Index Fungorum number: IF 836793; Facesoffungi number: FoF 08802.
Diagnosis: Corticolous Macroconstrictolumina with
thallus UV-negative; ostioles lateral, ascospores granular
ornamented, 34–37 × 11–14 µm, with gelatinous sheath of
c. 4 µm.
Etymology: The epithet refers to the large ascomata and
ascospores and the lateral ostioles.
Holotype: A. Aptroot 77751 (CGMS).
Thallus crustose, endoperidermal, whitish to pinkish,
opaque, not surrounded by a prothallus. Ascomata solitary,
pyriform, 0.6–0.8 mm diam., black. Ostioles lateral, black.
Hamathecium clear. Ascospores 8/ascus, hyaline, 1-septate,
septum median, 34–37 × 11–14 µm, sole-shaped with upper
cell wider than lower cell, with conspicuous granular wall
ornamentation, more or less distinctly constricted but not
thickened at the median septum, with secondary constriction
or invagination in each cell, surrounded by a c. 4 µm thick
gelatinous sheath. Pycnidia not observed.
Chemistry: Thallus UV–, C–, P–, K–; no substances
detected by TLC.
Ecology and distribution: On tree bark in Atlantic Forest;
thus far only known from Brazil.
Material examined: Brazil. Mato Grosso do Sul: Serra
da Bodoquena, Bonito, Fazenda Marambaia; 20°58’ S,
56°42’ W, 650 m; Atlantic Forest, on tree bark in pasture
close to forest; 30 October 2018, A. Aptroot 77751 (holotype: CGMS; isotype: ABL); Serra da Bodoquena, Fazenda
Monte Negro; 20°54’25″ S, 56°48’04″ W, 530 m; Atlantic
Forest, on tree bark in forest; 8 November 2018, A. Aptroot
77888, 77930, 77933, 77940, 77941, 77948, 77966 (paratypes: CGMS). Acre: Núcleo Cazumbá; 09˚07’ S, 68˚57’ W,
150 m; disturbed tropical rain forest; 2019, M. Cáceres & A.
Aptroot (ABL, ISE 50171).
Notes: Four species of Macroconstrictolumina are so far
recognized. The new species is unmistakable by the lateral
ostioles.
Key to species of Macroconstrictolumina:
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186
1. Ascospores 18–30 × 7–10(–12) µm……………………
…………………………Macroconstrictolumina lyrata
1. Ascospores 30–48 × 10–16(–24) µm…………………2
2. Ostiole lateral; ascospores 1-septate……………………
…………………Macroconstrictolumina megalateralis
2. Ostiole apical; ascospores 1(–3)-septate………………3
3. Ascospores 27–37 × 10–12 µm, often 3-septate………
……………………Macroconstrictolumina majuscula
3. Ascospores 40–50 × 15–20 µm, mostly 1-septate………
…………………Macroconstrictolumina malaccitula
Marcelaria Aptroot, M.P. Nelsen & Parnmen, Glalia 5(2):
3 (2013), nom. cons. prop.
MycoBank number: MB 805450; Index Fungorum number: IF 805450; Facesoffungi number: FoF 08803; three
morphologically defined species (Aptroot and Lücking
2016); molecular data available for two species (Lücking
et al. 2016).
= Buscalionia Sambo nom. cons. prop., Ann. Bot., Roma
22: 27 (1940). Type species: Buscalionia rubra Sambo, Ann.
Bot., Roma 22: 27 (1940) [= Marcelaria purpurina (Nyl.)
Aptroot, Nelsen & Parnmen].
Lichenized on bark in terrestrial, chiefly lowland tropical
wet forest habitats. Thallus distinctly corticate, yellow-brown
to olive-green. Photobiont Trentepohlia. Ascomata solitary
to often crowded or loosely aggregate, prominent to sessile,
brown-black but covered by thick thallus and a thick layer
of yellow-orange to red pigment crystals, hemispherical to
wart-shaped, coriaceous, ostiolate, ostiole apical. Excipulum prosoplecten-chymatous, dark brown to carbonized.
Hamathecium comprising 0.5–0.7 µm wide para-physoids,
hyaline, straight, branched and anastomosing, embedded in
a gelatinous matrix. Asci 2–8-spored, bitunicate, fissitunicate, oblong-clavate, short pedicellate, with a non-amyloid
ocular chamber. Ascospores irregularly arranged to biseriate,
fusiform-ellipsoid, hyaline, richly muriform, with somewhat
thickened distosepta and rounded lumina, smooth-walled,
not or slightly constricted at the septa, often with a gelatinous sheath. Pycnidia unknown. Chemistry: lichexanthone
sometimes produced on thallus surface; yellow to orange or
red anthraquinones present on the ascomata.
Type species: Marcelaria purpurina (Nyl.) Aptroot,
Nelsen & Parnmen, Glalia 5(2): 9 (2013) [= Buscalionia
rubra Sambo, Ann. Bot., Roma 22: 27 (1940)].
Notes: For key and discussion see Aptroot and Lücking (2016). The older name Buscalionia has recently been
discovered to be congeneric with Marcelaria and has been
proposed for conservation (Aptroot and Nepi 2017), with
the decision pending.
Nigrovothelium Lücking, M.P. Nelsen & Aptroot in Lücking
et al., Lichenologist 48: 757 (2016).
13
Fungal Diversity (2020) 105:17–318
MycoBank number: MB 816875; Index Fungorum number: IF 816875; Facesoffungi number: FoF 08804; three
morphologically and partly phylogenetically defined species (Aptroot and Lücking 2016); molecular data available
for two species (Lücking et al. 2016).
Lichenized on bark in terrestrial, chiefly lowland to lower
montane tropical habitats, often in drier situations. Thallus
distinctly corticate, olive-green to yellowish brown. Photobiont Trentepohlia. Ascomata solitary, prominent to sessile,
black, subglobose to egg-shaped, carbonaceous, ostiolate,
ostiole apical. Involucrellum carbonized. Excipulum prosoplectenchymatous, brownish to carbonized. Hamathecium
comprising 0.5–0.7 µm wide paraphysoids, hyaline, straight,
branched and anastomosing, embedded in a gelatinous
matrix. Asci 8-spored, bitunicate, fissitunicate, clavate, short
pedicellate, with a non-amyloid ocular chamber. Ascospores
irregularly arranged to biseriate, fusiform-ellipsoid, hyaline,
3-septate, astrothelioid with distinct distosepta and diamondshaped lumina, smooth-walled, not constricted at the septa,
often with a gelatinous sheath. Pycnidia rare, immersed to
erumpent, visible as black dots. Conidia aseptate, bacillar,
hyaline. Chemistry: no substances detected by TLC.
Type species: Nigrovothelium tropicum (Ach.) Lücking,
M.P. Nelsen & Aptroot in Lücking et al., Lichenologist 48:
757 (2016).
Notes: For key and discussion see Aptroot and Lücking
(2016).
Polymeridium (Müll. Arg.) R.C. Harris in Tucker & Harris,
Bryologist 83: 12 (1980).
MycoBank number: MB 4316; Index Fungorum number:
IF 4316; Facesoffungi number: FoF 08805; Approximately
50 morphologically defined species (Aptroot and Lücking
2016; Aptroot and Weerakoon 2018); molecular data available for three species (Lücking et al. 2016).
Lichenized on bark in terrestrial, chiefly lowland to lower
montane tropical habitats, often in drier situations. Thallus usually ecorticate, rarely indistinctly corticate, whitish.
Photobiont Trentepohlia. Ascomata solitary to aggregate,
erumpent to prominent, black, hemisphaerical to wartshaped or conical, carbonaceous, ostiolate, ostiole apical,
rarely lateral or fused. Involucrellum carbonized. Excipulum
prosoplectenchymatous, brownish to carbonized. Hamathecium comprising 0.5–0.7 µm wide paraphysoids, hyaline,
straight, branched and anastomosing, embedded in a gelatinous matrix. Asci 4–8-spored, bitunicate, fissitunicate,
clavate, short pedicellate, with a non-amyloid ocular chamber. Ascospores irregularly arranged to biseriate, fusiformellipsoid, hyaline, multi-septate to rarely (sub-)muriform,
with thin eusepta and rectangular lumina, smooth-walled,
not or slightly constricted at the septa, often with a gelatinous sheath. Pycnidia rare, immersed to erumpent, visible
as black dots. Conidia aseptate, bacillar, hyaline. Chemistry:
Fungal Diversity (2020) 105:17–318
lichexanthone sometimes produced on thallus surface; ostiolar area or hamathecium rarely with anthraquinones.
Type species: Polymeridium contendens (Nyl.) R.C. Harris in Tucker & Harris, Bryologist 83: 12 (1980).
Notes: For key and discussion see Aptroot and Lücking
(2016).
Polypyrenula D. Hawksw., Bull. Br. Mus. Nat. Hist., Bot.
14: 165 (1985).
MycoBank number: MB 25730; Index Fungorum number:
IF 25730; Facesoffungi number: FoF 08806; one morphologically and phylogenetically defined species (MirandaGonzález et al. 2020).
Lichenized (sometimes indistinctly or photobiont apparently absent) on bark in terrestrial, lowland tropical, seasonally dry forest habitats. Thallus ecorticate, whitish to pale
greyish or brownish. Photobiont (when present) Trentepohlia. Ascomata dispersed, solitary, erumpent to prominent, black but basally covered by thin thallus layer, hemisphaerical to wart-shaped, carbonaceous, ostiolate, ostiole
apical. Involucrellum well-developed, carbonaceous. Excipulum prosoplectenchymatous, basally reddish brown, laterally
and apically thin and fused with involucrellum. Hamathecium comprising 0.5–1 µm wide paraphysoids, hyaline,
straight, branched and anastomosing, embedded in a gelatinous matrix. Asci (6–)8-spored, bitunicate, clavate, shortly
pedicellate, with a non-amyloid ocular chamber. Ascospores
biseriate, fusiform-ellipsoid and tapering towards the distal
end, reddish brown but with basal cell paler, 5(-6)-septate,
with distinct distosepta and rounded lumina except for the
lowermost cell which is separated by a thin euseptum and
has a triangular lumen, smooth-walled, not constricted at
the septa, lacking a gelatinous sheath. Pycnidia unknown.
Chemistry: no substances detected by TLC.
Type species: Polypyrenula sexlocularis (Müll. Arg.) D.
Hawksw., Bull. Br. Mus. Nat. Hist., Bot. 14: 165 (1985) [=
Polypyrenula albissima Aptroot, Biblioth. Lichenol. 44: 102
(1991); nom. illeg., ICN Art. 52.1].
Notes: This monospecific genus is characterized by highly
unique ascospores, which coincides with its position on a
very long branch but with unresolved affinities among the
basally diverging clades of the family (Fig. 77). For a long
time, Polypyrenula was classified in the families Pyrenulaceae and Requienellaceae (Aptroot 1991; Lumbsch and
Huhndorf 2007, 2010; Jaklitsch et al. 2016). However, the
paraphysoids are more in line with those of Trypetheliaceae
and molecular data recently demonstrated placement of the
genus in that family (Miranda-González et al. 2020), a result
anticipated in two recent classification papers (Lücking et al.
2017; Wijayawardene et al. 2017a). Miranda-González et al.
(2020) also showed that P. sexlocularis is the correct name
for the only species, P. albissima Aptroot being an illegitimate, homotypic synonym. The species appears to be found
187
mostly in dry forests across the Neotropics (known from the
Caribbean, Mexico and Bolivia) and has likely been overlooked due to the nondescript thallus and ascomata.
Pseudobogoriella Lücking, R. Miranda & Aptroot gen. nov.
MycoBank number: MB 836794; Index Fungorum number: IF 836794; Facesoffungi number: FoF 08807; 15 morphologically defined species (this paper); molecular data
available for three species (Lücking et al. 2016; this paper).
Diagnosis: Differing from Bogoriella chiefly in the
smaller ascospores always lacking endospore thickenings.
Etymology: The name refers to the notion as a “false”
Bogoriella, i.e. similar but not closely related to the latter.
Lichenized (sometimes barely so or apparently nonlichenized) on bark in terrestrial, chiefly lowland to lower
montane tropical habitats. Thallus ecorticate, mostly whitish to pale brownish or pinkish. Photobiont Trentepohlia.
Ascomata solitary, erumpent to prominent, brown-black to
carbonaceous, hemisphaerical to wart-shaped, coriaceous to
carbonaceous, ostiolate, ostiole apical to rarely lateral. Involucrellum usually well-developed, carbonaceous. Excipulum
prosoplectenchymatous, usually brownish, in lateral and apical parts typically fused with the involucrellum. Hamathecium comprising 1 µm wide para-physoids, hyaline, straight
to somewhat wavy, branched and anastomosing, embedded
in a thin, gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate, clavate to fusiform, short pedicellate, with a nonamyloid ocular chamber. Ascospores irregularly arranged to
uni- or biseriate, fusiform-ellipsoid, grey-brown, 1-septate,
with eusepta and rectangular lumina, wall finely granular
ornamented, not or slightly constricted at the septa, typically rather small (mostly around 15–20 × 6–8 µm), only
in the type species somewhat larger. Pycnidia known from
a few species, immersed to erumpent, visible as black dots.
Conidia aseptate, bacillar, hyaline. Chemistry: no substances
detected by TLC.
Type species: Pseudobogoriella hemisphaerica (Müll.
Arg.) Lücking, R. Miranda & Aptroot (see below).
Notes: The establishment of this new genus follows the
topology of the phylogenetic tree, in which former members of Mycomicrothelia sensu lato emerge on three separate clades (Fig. 77). The systematics of these taxa is not
settled, given that only few species have been sequenced.
Pseudobogoriella currently represents a paraphyletic grade
of two clades, but we have adopted the conservative solution
of merging the two clades into a single genus for the time
being. This reflects the fact that these species cannot be classified in Bogoriella sensu stricto for topological reasons, but
at the same time too few data are available to further resolve
this basally emerging grade. The main difference when compared to Bogoriella are the small, 1-septate ascospores lacking endospore thickenings.
13
188
New combinations in Pseudobogoriella:
Pseudobogoriella alata (Groenh. ex Aptroot) Lücking, R.
Miranda & Aptroot comb. nov.
MycoBank number: MB 836795; Index Fungorum number: IF 836795; Facesoffungi number: FoF 08808.
Bas.: Mycomicrothelia alata Groenh. ex Aptroot, Biblioth. Lichenol. 44: 130 (1991); Bogoriella alata (Groenh. ex
Aptroot) Aptroot & Lücking, Lichenologist 48: 905 (2016).
Pseudobogoriella annonacea (Müll. Arg.) Lücking, R.
Miranda & Aptroot comb. nov.
MycoBank number: MB 836796; Index Fungorum number: IF 836796; Facesoffungi number: FoF 08809.
Bas.: Microthelia annonacea Müll. Arg., [as ‘anonacea’],
Hedwigia 34: 145 (1895); Mycomicrothelia annonacea
(Müll. Arg.) D. Hawksw., [as ‘anonacea’] Bull. Br. Mus.
Nat. Hist., Bot. 14(2): 67 (1985); Bogoriella annonacea
(Müll. Arg.) Aptroot & Lücking, Lichenologist 48: 907
(2016).
Pseudobogoriella captiosa (Kremp.) Lücking, R. Miranda
& Aptroot comb. nov.
MycoBank number: MB 836797; Index Fungorum number: IF 836797; Facesoffungi number: FoF 08810.
Bas.: Verrucaria captiosa Kremp., Flora 59: 524 (1876);
Microthelia captiosa (Kremp.) Müll. Arg., Flora 66: 272
(1883); Mycomicrothelia captiosa (Kremp.) D. Hawksw.,
Bull. Br. Mus. Nat. Hist., Bot. 14: 73 (1985); Bogoriella
captiosa (Kremp.) Aptroot & Lücking, Lichenologist 48:
908 (2016).
Pseudobogoriella exigua (Müll. Arg.) Lücking, R. Miranda
& Aptroot comb. nov.
MycoBank number: MB 836798; Index Fungorum number: IF 836798; Facesoffungi number: FoF 08811.
Bas.: Microthelia exigua Müll. Arg., Bot. Jahrb. 6: 416
(1885); Mycomicrothelia exigua (Müll. Arg.) D. Hawksw.,
Bull. Br. Mus. Nat. Hist., Bot. 14: 84 (1985); Bogoriella
exigua (Müll. Arg.) Aptroot & Lücking, Lichenologist 48:
910 (2016).
Pseudobogoriella fumosula (Zahlbr.) Lücking, R. Miranda
& Aptroot comb. nov.
MycoBank number: MB 836799; Index Fungorum number: IF 836799; Facesoffungi number: FoF 08812.
Bas.: Microthelia fumosula Zahlbr. in Handel-Mazzetti,
Symb. Sinic. 3: 18 (1930); Mycomicrothelia fumosula
(Zahlbr.) D. Hawksw., Bull. Br. Mus. Nat. Hist., Bot. 14: 86
(1985); Bogoriella fumosula (Zahlbr.) Aptroot & Lücking,
Lichenologist 48: 910 (2016).
Pseudobogoriella hemisphaerica (Müll. Arg.) Lücking, R.
Miranda & Aptroot comb. nov.
13
Fungal Diversity (2020) 105:17–318
MycoBank number: MB 836800; Index Fungorum number: IF 836800; Facesoffungi number: FoF 08813.
Bas.: Microthelia hemisphaerica Müll. Arg., Bot. Jahrb.
6: 417 (1885); Mycomicrothelia hemisphaerica (Müll. Arg.)
D. Hawksw., Bull. Br. Mus. Nat. Hist., Bot. 14: 86 (1985);
Bogoriella hemisphaerica (Müll. Arg.) Aptroot & Lücking,
Lichenologist 48: 910 (2016).
Pseudobogoriella lateralis (Sipman) Lücking, R. Miranda
& Aptroot comb. nov.
MycoBank number: MB 836801; Index Fungorum number: IF 836801; Facesoffungi number: FoF 08814.
Bas.: Mycomicrothelia lateralis Sipman, Lichenologist
37: 309 (2005); Bogoriella lateralis (Sipman) Aptroot &
Lücking, Lichenologist 48: 911 (2016).
Pseudobogoriella leuckertii (D. Hawksw. & J.C. David)
Lücking, R. Miranda & Aptroot comb. nov.
MycoBank number: MB 836802; Index Fungorum number: IF 836802; Facesoffungi number: FoF 08815.
Bas.: Mycomicrothelia leuckertii D. Hawksw. & J.C.
David in David & Hawksworth, Biblioth. Lichenol. 57: 98
(1995); Bogoriella leuckertii (D. Hawksw. & J.C. David)
Aptroot & Lücking, Lichenologist 48: 911 (2016).
Pseudobogoriella minutula (Zahlbr.) Lücking, R. Miranda
& Aptroot comb. nov.
MycoBank number: MB 836803; Index Fungorum number: IF 836803; Facesoffungi number: FoF 08816.
Bas.: Microthelia minutula Zahlbr. in Handel-Mazzetti,
Symb. Sinic. 3: 18 (1930); Mycomicrothelia minutula
(Zahlbr.) D. Hawksw., Bull. Br. Mus. Nat. Hist., Bot. 14(2):
102 (1985); Bogoriella minutula (Zahlbr.) Aptroot & Lücking, Lichenologist 48: 912 (2016).
Pseudobogoriella nonensis (Stirt.) Lücking, R. Miranda &
Aptroot comb. nov.
MycoBank number: MB 836804; Index Fungorum number: IF 836804; Facesoffungi number: FoF 08817.
Bas.: Verrucaria nonensis Stirt., Proc. Roy. Phil. Soc.
Glasgow 11: 320 (1879); Microthelia nonensis (Stirt.)
Zahlbr., Cat. Lich. Univers. 1: 264 (1921); Mycomicrothelia nonensis (Stirt.) D. Hawksw., Bull. Br. Mus. Nat. Hist.,
Bot. 14: 105 (1985); Bogoriella nonensis (Stirt.) Aptroot &
Lücking, Lichenologist 48: 913 (2016).
Pseudobogoriella pachnea (Körb.) Lücking, R. Miranda &
Aptroot comb. nov.
MycoBank number: MB 836805; Index Fungorum number: IF 836805; Facesoffungi number: FoF 08818.
Bas.: Microthelia pachnea Körb., Parerga Lichenol. 5:
398 (1865); Amphisphaeria pachnea (Körb.) Rehm, Ann.
Fungal Diversity (2020) 105:17–318
Mycol. 4: 264 (1906); Mycomicrothelia pachnea (Körb.) D.
Hawksw., Bull. Br. Mus. Nat. Hist., Bot. 14: 108 (1985).
Pseudobogoriella punctata (Aptroot) Lücking, R. Miranda
& Aptroot comb. nov.
MycoBank number: MB 836806; Index Fungorum number: IF 836806; Facesoffungi number: FoF 08819
Bas.: Mycomicrothelia punctata Aptroot, Biblioth.
Lichenol. 44: 134 (1991); Bogoriella punctata (Aptroot)
Aptroot & Lücking, Lichenologist 48: 914 (2016).
Pseudobogoriella socialis (Zahlbr.) Lücking, R. Miranda &
Aptroot comb. nov.
MycoBank number: MB 836807; Index Fungorum number: IF 836807; Facesoffungi number: FoF 08820.
Bas.: Microthelia socialis Zahlbr., Mycologia 22: 69
(1930); Mycomicrothelia socialis (Zahlbr.) D. Hawksw.,
Bull. Br. Mus. Nat. Hist., Bot. 14: 110 (1985); Bogoriella
socialis (Zahlbr.) Aptroot & Lücking, Lichenologist 48: 915
(2016).
189
1.
1.
2.
2.
3.
3.
4.
4.
5.
5.
6.
6.
7.
7.
Pseudobogoriella striguloides (Sérus. & Aptroot) Lücking,
R. Miranda & Aptroot comb. nov.
MycoBank number: MB 836808; Index Fungorum number: IF 836808; Facesoffungi number: FoF 08821.
Bas.: Mycomicrothelia striguloides Sérus. & Aptroot,
Bryologist 101: 145 (1998); Bogoriella striguloides (Sérus.
& Aptroot) Aptroot & Lücking, Lichenologist 48: 915
(2016).
Notes: This species is highly unusual among the genus
due to its foliicolous, Strigula-like, subcuticular thallus.
Sérusiaux and Aptroot (1998) already discussed the possibility of this being a lichenicolous fungus on a Strigula thallus, but found no support for this hypothesis. The anatomical characters of the ascomata and asci are reminiscent of
the lichenicolous genus Polycoccum Saut. ex Körb. (Matzer
1996; Diederich et al. 2018), which has recently been identified as a close relative of Trypetheliaceae (Ertz et al.
2015). We refrain from proposing any taxonomic changes
at this point but we predict that this taxon is not a genuine
Trypetheliaceae.
8.
8.
9.
9.
10.
10.
11.
11.
12.
12.
13.
13.
Pseudobogoriella subfallens (Müll. Arg.) Lücking, R.
Miranda & Aptroot comb. nov.
MycoBank number: MB 836809; Index Fungorum number: IF 836809; Facesoffungi number: FoF 08822
Bas.: Microthelia subfallens Müll. Arg., Bot. Jahrb. 6:
416 (1885); Mycomicrothelia subfallens (Müll. Arg.) D.
Hawksw., Bull. Br. Mus. Nat. Hist., Bot. 14: 111 (1985);
Bogoriella subfallens (Müll. Arg.) Aptroot & Lücking,
Lichenologist 48: 915 (2016).
Key to species of Pseudobogoriella:
14.
14.
Ostiole lateral……………Pseudobogoriella lateralis
Ostiole apical…………………………………………2
Ascospores 24–28 µm long……………………………
……………………Pseudobogoriella hemisphaerica
Ascospores 12–22 µm long……………………………3
Thallus foliicolous, subcuticular, distinctly lobed……
………………………Pseudobogoriella striguloides
Thallus corticicolous, not lobed………………………4
Excipulum with K+ red, dissolving pigment…………
…………………………Pseudobogoriella leuckertii
Excipulum lacking pigment…………………………5
Black prothallus line with numerous pycnidia…………
……………………………Pseudobogoriella punctata
Black prothallus line lacking pycnidia or
absent…………………………………………………6
Thallus besides ascomata with numerous black dots representing pycnidia……………………………………7
Thallus lacking pycnidia………………………………8
Ascomata aggregate; ascospores 15–20 × 6–9 µm……
……………………………Pseudobogoriella socialis
Ascomata solitary; ascospores 11–17 × 5–7 µm……
…………………………Pseudobogoriella fumosula
Involucrellum broadly expanding basally to over 0.5
mm diam.……………………Pseudobogoriella alata
Ascomata including basally slightly expanding involucrellum 0.2–0.5 mm diam.……………………………9
Ascospores 12–16 µm long…………………………10
Ascospores 16–22 µm long…………………………12
Ascomata 0.3–0.5 mm diam.…………………………
…………………………Pseudobogoriella subfallens
Ascomata 0.2–0.3 mm diam.………………………11
Ascospores 4–5.5 µm broad, 2.5–3 times as long as
broad; involucrellum basally not expanding…………
……………………………Pseudobogoriella exigua
Ascospores 5.5–7 µm broad, 2–2.5 times as long as
broad; involucrellum basally expanding………………
…………………………Pseudobogoriella minutula
Black prothallus line present………………………13
Black prothallus line absent…………………………14
Ascomata conical, basally not expanded……………
……………………………Pseudobogoriella captiosa
Ascomata flattened, basally expanded………………
………………………Pseudobogoriella miculiformis
Ascomata basally broadly expanded, fringe 50–100 µm
broad……… …………Pseudobogoriella annonacea
Ascomata basally narrowly expanded, fringe 20–50 µm
broad……… ……………Pseudobogoriella nonensis
Pseudopyrenula Müll. Arg., Flora 66: 247 (1883).
MycoBank number: MB 4481; Index Fungorum number:
IF 4481; Facesoffungi number: FoF 08823; Approximately
20 morphologically defined species (Aptroot and Lücking
13
190
2016: Aptroot et al. 2019); molecular data available for four
species (Lücking et al. 2016).
Lichenized on bark in terrestrial, chiefly lowland to lower
montane tropical habitats, often in drier situations. Thallus ecorticate, whitish. Photobiont Trentepohlia. Ascomata
solitary, erumpent to prominent, black, hemisphaerical to
wart-shaped or conical, carbonaceous, ostiolate, ostiole
apical, rarely lateral. Involucrellum carbonized. Excipulum prosoplectenchymatous, brownish to carbonized.
Hamathecium comprising 0.5–0.7 µm wide paraphysoids,
hyaline, straight, branched and anastomosing, embedded in
a gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate,
clavate, short pedicellate, with a non-amyloid ocular chamber. Ascospores irregularly arranged to biseriate, fusiformellipsoid, hyaline, 3(–5)-septate, astrothelioid with distinct
distosepta and diamond-shaped lumina, smooth-walled,
not constricted at the septa, often with a gelatinous sheath.
Pycnidia rare, immersed to erumpent, visible as black dots.
Conidia aseptate, bacillar, hyaline. Chemistry: lichexanthone
sometimes produced on thallus surface; hamathecium and/or
ascospores sometimes with yellow anthraquinones.
Type species: Pseudopyrenula diluta (Fée) Müll. Arg.,
Flora 66: 249 (1883)
Notes: For key and discussion see Aptroot and Lücking
(2016).
Schummia Lücking, R. Miranda & Aptroot gen. nov.
MycoBank number: MB 836810; Index Fungorum number: IF 836810; Facesoffungi number: FoF 08824; one morphologically defined species (this paper); molecular data
thus far unavailable.
Diagnosis: Differing from Bogoriella in the ascospores
with lens-shaped lumina and very thick terminal walls.
Etymology: Named after our colleague and friend, Felix
Schumm, for his excellent contributions to our knowledge
of the morphology and anatomy of lichen fungi.
Lichenized (weakly so) on bark in terrestrial, lowland to
lower montane subtropical habitats (Azores). Thallus ecorticate, pale brownish (to pinkish). Photobiont Trentepohlia.
Ascomata solitary, erumpent, brown-black to carbonaceous,
applanately wart-shaped to irregular in outline, ostiolate,
ostiole apical. Involucrellum weakly developed, with periderm layers, carbonaceous. Excipulum prosoplectenchymatous, brown-black, in apical parts fused with involucrellum. Hamathecium comprising 1.5 µm wide paraphysoids,
hyaline, straight to somewhat wavy, slightly branched. Asci
8-spored, bitunicate, fissitunicate, clavate, short pedicellate,
with a non-amyloid, rather long ocular chamber. Ascospores
irregularly arranged to biseriate, ellipsoid to drop-shaped,
grey-brown, 1-septate, with a central euseptum and rectangular lumina, the terminal walls much thickened and
inflated, not or slightly constricted at the septa. Pycnidia
unknown. Chemistry: no substances detected by TLC.
13
Fungal Diversity (2020) 105:17–318
Type species: Schummia angulata (Aptroot & Schumm)
Lücking, R. Miranda & Aptroot (see below).
Notes: This new genus is introduced for a single species
with highly unusual ascospores that in their mature stage
are reminiscent of Arthoniaceae and Graphidaceae rather
than Trypetheliaceae. The species was originally described
in the genus Distothelia (Schumm and Aptroot 2013), but
the ascospores differ clearly in shape and endospore development, resulting in two strongly thickened terminal walls.
Immature ascospores appear 3-septate with three eusepta,
somewhat similar to ascospores in Novomicrothelia and
Pseudobogoriella except for the more numerous septa.
However, the two terminal lumina eventually become
filled entirely with endospore material and simultaneously
inflated, so that the two-remaining central lumina appear
much smaller than the thickened terminal walls (Fig. 79f).
Given the current distribution of ascospore types among
basally diverging lineages in the family, we predict that this
taxon represents its own lineage, and placement in Bogoriella, together with the other two species previously placed
in Distothelia, would be misleading.
Schummia angulata (Aptroot & Schumm) Lücking, R.
Miranda & Aptroot comb. nov.
MycoBank number: MB 836811; Index Fungorum number: IF 836811; Facesoffungi number: FoF 08825.
Bas.: Distothelia angulata Aptroot & Schumm in
Schumm & Aptroot, Flechten Madeiras, der Kanaren und
Azoren 2: 200 (2013).
Viridothelium Lücking, M.P. Nelsen & Aptroot in Lücking
et al., Lichenologist 48: 758 (2016).
MycoBank number: MB 816877; Index Fungorum number: IF 816877; Facesoffungi number: FoF 08826; 11 morphologically defined species (Aptroot and Lücking 2016;
Cáceres and Aptroot 2017); molecular data available for
three species (Lücking et al. 2016).
= ?Exiliseptum R.C. Harris, Acta Amazonica 14(1–2,
Suppl.): 65 (1986) [1984]. Type species: Exiliseptum ocellatum (Müll. Arg.) R.C. Harris, Acta Amazonica 14(1–2,
Suppl.): 66 (1986) [1984].
Lichenized on bark in terrestrial, chiefly lowland to lower
montane tropical to temperate forest habitats. Thallus distinctly corticate, yellow-brown to olive-green. Photobiont
Trentepohlia. Ascomata solitary to aggregated or diffusely
pseudostromatic, immersed to erumpent, brown-black but
usually covered by thallus layer except for ostiolar area,
coriaceous to carbonaceous, ostiolate, ostiole apical to lateral, sometimes fused. Excipulum prosoplectenchymatous,
brownish. Hamathecium comprising 0.5–0.7 µm wide paraphysoids, hyaline, straight, branched and anastomosing,
embedded in a thick, gelatinous matrix. Asci 4–8-spored,
bitunicate, fissitunicate, clavate to oblong, short pedicellate,
Fungal Diversity (2020) 105:17–318
191
which case it would provide an earlier name for that genus.
We have refrained from formal implementation of this finding, as the phylogenetic placement of odd lineages in Trypetheliaceae is somewhat unpredictable and we anticipate
collecting authentic material for sequencing.
Key to genera of Trypetheliaceae:
1.
1.
2.
2.
3.
3.
4.
4.
5.
5.
6.
6.
Fig. 81 Exiliseptum ocellatum (holotype). a, b habitus and asci with
ascospores. Scale bars: a = 1 mm, b = 10 µm
7.
7.
with a non-amyloid ocular chamber. Ascospores irregularly
arranged to biseriate, oblong-fusiform, hyaline, septate to
rarely muriform, with slightly thickened distosepta and more
or less rectangular lumina, smooth-walled, not constricted
at the septa, often with a gelatinous sheath. Pycnidia rare,
immersed to erumpent, visible as black dots. Conidia aseptate, bacillar, hyaline. Chemistry: orange anthraquinone
known from one species
Type species: Viridothelium virens (Tuck. ex Michener)
Lücking, M.P. Nelsen & Aptroot in Lücking et al., Lichenologist 48: 759 (2016).
Notes: For key and discussion see Aptroot and Lücking
(2016). The genus currently includes 11 species (Cáceres
and Aptroot 2017). The genus Exiliseptum, described by
Harris (1986), was later considered a member of Polymeridium with a corticate thallus (Aptroot and Cáceres 2014),
and this view was followed in the recent synopsis on Trypetheliaceae (Aptroot and Lücking 2016). However, upon
re-examination of the type material (Fig. 81), we believe
that this taxon may be congeneric with Viridothelium, in
8.
8.
9.
9.
10.
10.
11.
11.
12.
12.
13.
13.
14.
Ascospores remaining hyaline………………………2
Ascospores becoming brown………………………23
Ascospores 1-septate…………………………………3
Ascospores 3-septate to muriform……………………6
Ascospores up to 15 × 5 µm, lacking secondary constrictions…………………“Arthopyrenia” (aff. minor)
Ascospores larger than 15 × 5 µm, usually with secondary constrictions………………………………………4
Non-lichenized; saprobic on branches in southern
Europe……………………Alloarthopyrenia (italica)
Lichenized (often barely so); (sub-)tropical…………5
Ascospores smooth-walled, up to 30 × 11 µm (in one
species up to 46 × 24 µm) ……Constrictolumina p.p.
Ascospores granular ornamented, mostly larger than 30
× 11 µm (in one species 20–30 × 7–12 µm)…………
…………………………Macroconstrictolumina p.p.
Thallus ecorticate or indistinctly corticate, usually
whitish or greyish, contrasting with the black ascomata…………………………………………………7
Thallus distinctly corticate, olive-green to yellowish
brown………………………………………………13
Hamathecium physes basally somewhat thickened,
somewhat flexuose, branched but usually not anastomosing; ascospores (1–)3-septate………………………8
Hamathecium physes thin, straight, branched and anastomosing to form a net-like structure…………………9
Ascospores up to 23 × 8 µm, smooth-walled…………
…………………………………Constrictolumina p.p.
Ascospores mostly larger than 30 × 11 µm (in one species 20–30 × 7–12 µm), granular ornamented…………
…………………………Macroconstrictolumina p.p.
Ascospores astrothelioid, with diamond-shaped
lumina………………………………Pseudopyrenula
Ascospores euseptate, with thin septa and rectangular
lumina………………………………………………10
Ascospores transversely septate…Polymeridium p.p.
Ascospores muriform………………………………11
Ostioles lateral………………………Dictyomeridium
Ostioles apical………………………………………12
Non-lichenized; temperate………“Julella” (fallaciosa)
Lichenized; (sub-)tropical…………Polymeridium p.p.
Ascospores transversely septate……………………14
Ascospores muriform………………………………19
Ascospores very large with few (3–5) septa, typically
over 100 × 30 µm and up to 190 × 60 µm; septa and
13
192
14.
15.
15.
16.
16.
17.
17.
18.
18.
19.
19.
20.
20.
21.
21.
22.
22.
23.
23.
24.
24.
Fungal Diversity (2020) 105:17–318
walls somewhat thickened but lumina not astrothelioid…………………………Architrypethelium p.p.
Ascospores when 3–5-septate very rarely exceeding
100 × 30 µm and if so, ascospores distinctly astrothelioid, with diamond-shaped lumina…………………15
Ascospores astrothelioid, with diamond-shaped
lumina………………………………………………16
Ascospores euseptate, with rather thin septa and more or
less rectangular lumina……………………………17
Perithecia strongly prominent to sessile, completely exposed, pure black, somewhat egg-shaped
………………………………………Nigrovothelium
Perithecia immersed to erumpent or aggregate in
erumpent to sessile pseudostromata…………………
………………………………………Astrothelium p.p.
Perithecia aggregate in prominent to sessile, brownblack pseudostromata; ascospores with thin septa and
walls…………………………………Bathelium p.p.
Perithecia solitary to pseudostromatic but pseudostromata not as above; ascospores with slightly thickened
septa…………………………………………………18
Perithecia immersed to erumpent or rarely
indistinctly pseudostromatic; pigments mostly
absent………………………………Viridothelium p.p.
Perithecia aggregate in distinct, prominent to sessile
pseudostromata; internal and/or external pigments usually present……………………………Trypethelium
Ascospores with thin (eu-)septa and thin walls……20
Ascospores with thickened (disto-)septa, in young
stages often astrothelioid……………………………21
Perithecia aggregate in prominent to sessile, brownblack pseudostromata…………………Bathelium p.p.
Perithecia dispersed and with whitish rim or in
erumpent, whitish pseudostromata strongly contrasting with the blackish perithecia and the olive-brown
thallus……………Viridothelium (incl. Exsiliseptum)
Perithecia in prominent to sessile warts covered by a
thick layer of yellow-orange or red pigment…………
……………………………………………Marcelaria
Perithecia immersed to erumpent, rarely prominent,
usually covered by thallus, rarely with a thin layer of
pigment pruina………………………………………22
Perithecia aggregate in irregular, blackish pseudostromata; ascospores about 50 × 15 µm…………………
……………………Architrypethelium (murisporum)
Perithecia and ascospores not in the above combination…………………………………Astrothelium p.p.
Ascospores transversely septate……………………24
Ascospores (sub-)muriform…………………………30
Ascospores with a basal euseptum and the basal cell
pale, otherwise distoseptate and brown………………
…………………………Polypyrenula (sexlocularis)
Ascospores with either eu- or distosepta…………25
13
25. Ascospores large, usually over 100 µm long………26
25. Ascospores small, under 50 µm long………………27
26. Ascospores 3-septate, 25–50 µm broad, about 3–4
times as long as broad………Architrypethelium p.p.
26. Ascospores 11–15-septate, 25–30 µm broad, about 4–5
times as long as broad…Astrothelium (fuscosporum)
27. Ascospores wit h strongly t hickened distosepta………………………………………………28
27. Ascospores with thin eusepta………………………29
28. Ascospores drop-shaped, macrocephalic, with strongly
thickened terminal walls and two narrow, rectangular
lumina near the center…………Schummia (angulata)
28. Ascospores ellipsoid, with irregular wall thickenings
and halter-shaped to irregular lumina…………………
…………………………………………Bogoriella p.p.
29. Ascospores up to 20 × 8 µm, only in one species larger
(up to 30 × 10 µm)…………………Pseudobogoriella
29. Ascospores 20–50 × 8–15 µm ………Bogoriella p.p.
30. Ascospores small, under 50 × 20 µm, often with irregularly thickened endospore and halter-shaped lumina…
…………………………………………Bogoriella p.p.
30. Ascospores very large, 150–400 × 50–140 µm, with
outer wall often breaking under pressure……………
………………………………………………Aptrootia
Corrections for errors detected in previous synopsis of
Trypetheliaceae
Given the large size of the previous synopsis of the Trypetheliaceae (Aptroot and Lücking 2016), unfortunate errors
and omissions were almost inevitable. Corrections to most
of these were already listed in Aptroot et al. (2019). A few
more are reported and corrected below. When using the key,
it should also be noted that additional species have been
described in the meanwhile, particularly in Astrothelium but
also in other genera.
Page 783, Table 1: for (Astrothelium) subinterjectum, replace
+/+ with −/+ (lichexanthone on ascomata only).
Page 802, key couplet 24, first alternative, add: ascospores
48–65 × 17–20 µm.
Page 802, key couplet 24, second alternative, add:
ascospores 62–80 × 20–25 µm.
Page 806, key couplet 21, first alternative: replace
“Hamathecium inspersed” with “Lichexanthone on thallus only”.
Page 806, key couplet 21, second alternative: replace
“Hamathecium clear” with “Lichexanthone on thallus and
ascomata or on ascomata only”.
Page 808, key couplet 39, first alternative: replace “Thallus cracked” with “Thallus covered with yellow to orange
pigment”.
Page 808, key couplet 39, second alternative: replace “Thallus not cracked” with “Thallus olive-green to brownish
Fungal Diversity (2020) 105:17–318
yellow, lacking pigment, only pseudostromata covered with
yellow to orange pigment”.
Page 857, left column, description of Astrothelium defossum: replace “8 per ascus” with “4 per ascus”.
Page 911, left column, description of Bogoriella lateralis
(Sipman) Aptroot & Lücking comb. nov.: replace “Ascomata with apical ostioles” with “Ascomata with lateral
ostioles”.
Page 916, right column, description of Bogoriella triangularis (Aptroot) Aptroot & Lücking comb. nov.: replace
“Ascomata with apical ostioles” with “Ascomata with lateral ostioles”.
Page 961, right column, description of Pseudopyrenula
superans: replace “Hamathecium inspersed” with
“Hamathecium clear.”, “UV–” by “UV+ yellow”, and “no
substances found” by “lichexanthone”.
Economic and ecological significance
Trypetheliaceae is the second most diverse family of
tropical crustose lichens, with over 400 species known and
about 800 predicted (Aptroot et al. 2016). It is an important component of lichen communities particularly in wet
and dry tropical forest and savanna ecosystems (Komposch
and Hafellner 2000, 2003; Rivas Plata et al. 2008; Aptroot
2013; Menezes et al. 2018). There are no currently known
uses of economic importance of Trypetheliaceae; the family is chemically comparatively simple, but with diverse
anthraquinone and other pigments (Mathey 1979; Mathey
et al. 1980; Aptroot and Lücking 2016). Some of these pigments appear to have potential pharmaceutical properties
(Manojlovic et al. 2010; Sun et al. 2010; Takenaka et al.
2013; Elsebai et al. 2014; Basnet et al. 2019; Srinivasan
et al. 2019). The copious red pigment of the ascomata of
Marcelaria purpurina, collected during the 1899 expedition
of Luigi Buscalioni (Sambo 1940; Daly and Millozza 2007),
has apparently been used as dye by Amazonian indigenous
tribes. Harris (1986) reported the use of the same species by
the Bolivian Chácobo people as a treatment against headaches. Prance (1972) described the use of a pyrenocarpous
lichen by the Denís tribe in Amazonian Brazil as ‘Badhu’,
sniffed through the nose; this lichen may well represent a
Trypetheliaceae but its exact affinity is unknown. Trypetheliaceae is one of four families that has been studied in detail
regarding the latitudinal diversity gradient in lichens and
the potential impact of climate change on lichen distribution
ranges (Menezes et al. 2018).
Tubeufiales Boonmee & K.D. Hyde.
Index Fungorum number: IF 550704; Facesoffungi number: FoF 00203.
Tubeufiales was established by Boonmee et al. (2014b)
based on phylogenetic analyses and morphology. Liu et al.
193
(2017) treated Bezerromycetaceae and Wiesneriomycetaceae
as families of Tubeufiales on the basis of divergence time
estimates. The latest comprehensive revision on Tubeufiales
was carried out by Lu et al. (2018b), who expanded the circumscription of the type family Tubeufiaceae, accepted 42
genera in this family, and revised the taxonomy of tubeufiaceous species. This revision, in particular, helped to highlight fungi which were misidentified and resolved which
morphs are important in classifying species. Tubeufiaceous
fungi have the potential to produce bioactive compounds
(Ohtsu et al. 2003; Lu et al. 2018b; Fan et al. 2019). Fan
et al. (2019) tested the antifungal and anticancer effects of 19
tubeufiaceous strains, and found that most of them showed
obvious bioactivities. We introduce a new morphological
record of Tubeufia longiseta. The divergence time for Tubeufiales is estimated as 234 MYA (stem age, Hongsanan et al.
2020).
Accepted families: Bezerromycetaceae, Tubeufiaceae, and
Wiesneriomycetaceae (Fig. 82).
Bezerromycetaceae J.D.P. Bezerra, Souza-Motta & Crous,
Mycol. Progr. 16: 301 (2017).
Index Fungorum: IF 817521; Facesoffungi number: FoF
06809, 4 species.
Saprobic on dead wood in terrestrial habitats, or as endophytic fungi associated with cactus species in tropical dry
forests. Sexual morph: Ascomata superficial or immersed
in culture media, pseudothecial, unilocular, globose to
subglobose, gregarious or solitary, pale brown to brown at
maturity, minutely papillate with ostiole, collapsing cupulate or laterally, smooth or hairy. Peridium comprising pale
brown cells of textura angularis, and small cells of textura
prismatica. Hamathecium comprising numerous, filiform,
septate, branched, hyaline pseudoparaphyses. Asci 8-spored,
bitunicate, fissitunicate, cylindrical to cylindric-clavate,
short pedicellate, with or without a minute ocular chamber.
Ascospores 1–2-seriate, ellipsoidal, hyaline when young,
becoming pale brown to brown at maturity, muriformly
septate, smooth or minutely verrucose. Chlamydospores
sometimes linked to ascomata by hyphae, multi-septate,
brown, dictyochlamydospore-like, globose to subglobose
or ellipsoid to cylindrical. Asexual morph: Undetermined.
Type: Bezerromyces J.D.P. Bezerra, Souza-Motta &
Crous
Notes: Bezerromycetaceae was introduced based on
multi-gene phylogenetic analyses and morphological characteristics to accommodate the genera Bezerromyces and
Xiliomyces. Liu et al. (2017) treated Bezerromycetaceae as a
family of Tubeufiales by divergence time estimates. Lu et al.
13
194
Fungal Diversity (2020) 105:17–318
Tubeufia longiseta MFLUCC 15-0188
Tubeufia longiseta MFLUCC 17–1810
Tubeufia sessilis MFLUCC 16-0021
88/1.0
Tubeufia bambusicola MFLUCC 17-1803
100/1.0
Tubeufia inaequalis MFLUCC 17-0053
96/100
Tubeufia fangchengensis MFLUCC 17-0047
97/1.0
74/1.0
Tubeufia hechiensis MFLUCC 17-0052
100/1.0
Pseudohelicomyces menglunicus MFLUCC14-0689
Pseudohelicomyces aquaticus MFLUCC 16-0234
58/0.93
Helicomyces chiayiensis BCRC FU30842
Helicomyces hyalosporus MFLUCC 17-0051
Dematiohelicoma pulchrum MUCL 39827
93/1.0
Helicotruncatum palmigenum NBRC 32663
100/1.0 Neohelicomyces hyalosporus GZCC 16-0086
Neohelicomyces submersus MFLUCC 16-1106
89/1.0 50/0.93
Muripulchra aquatica MFLUCC 15-0249
100/1.0 Boerlagiomyces macrospora MFLUCC 12-0388
Manoharachariella tectonae MFLUCC 12-0170
97/1.0 Neohelicosporium ovoideum GZCC 16-0064
98/1.0 Neohelicosporium laxisporum MFLUCC 17-2027
91/1.0 Helicohyalinum aquaticum MFLUCC 16-1131
56/1.0
Helicodochium aquaticum MFLUCC 17-2016
100/1.0
Dematiohelicomyces helicosporus MFLUCC 16-0213
60/0.98
Neochlamydotubeufia khunkornensis MFLUCC10-0118
82/1.0
Neochlamydotubeufia fusiformis MFLUCC 16-0016
96/1.0
Pseudohelicoon subglobosum NCYU K3 2 1 3
-/0.99
Pseudohelicoon gigantisporum BCC 355
Helicotubeufia hydei MFLUCC 17-1980
90/1.0
Helicotubeufia jonesii MFLUCC 17-0043
Kevinhydea brevistipitata MFLUCC 18-1269
58/Aquaphila albicans MFLUCC 16-0010
Chlamydotubeufia cylindrica MFLUCC 16-1130
93/1.0 Chlamydotubeufia krabiensis MFLUCC 16 1134
Pleurohelicosporium parvisporum MFLUCC 17-1982
100/1.0 Tamhinispora srinivasanii NFCCI 4231
Neotubeufia krabiensis MFLUCC 16-1125
Helicosporium setiferum MFLUCC 17-1994
73/0.99
Helicosporium vesicarium MFLUCC 17-1795
96/1.0
100/1.0
Helicosporium vegetum CBS 941.72
100/1.0
Acanthostigma patagonicum BBB MVB 573
83/Helicoma vaccinii CBS 216-90
Acanthostigma perpusillum UAMH 7237
97/1.0
Acanthostigma chiangmaiensis MFLUCC10-0125
Thaxteriellopsis lignicola MFLUCC 10-0124
Helicangiospora lignicola MFLUCC 11-0378
Acanthostigmina minutum ANM 880
Acanthostigmina multiseptatum ANM 665
Acanthostigma filiforme ANM 514
71/- Acanthohelicospora pinicola MFLUCC 10-0116
Acanthohelicospora aurea GZCC 16-0059
54/Dictyospora thailandica MFLUCC 16-0001
100/1.0 Helicoma septoconstrictum MFLUCC 17-2001
-/1.0
100/1.0
Helicoma brunneisporum MFLUCC 17-1983
53/1.0
Helicoma rubriappendiculatum MFLUCC 18-0491
57/1.0
81
Dematiotubeufia chiangraiensis MFLUCC10-0115
Neohelicoma fagacearum MFLUCC11-0379
Helicoarctatus aquaticus MFLUCC 17-1996
KAmalomyces bambusicola MFLU 11-0228
70/90/1.0
KAmalomyces thailandicus MFLUCC 11-0158
Berkleasmium thailandicum MFLUCC 15-1248
93/1.0
Berkleasmium fusiforme MFLUCC 17-1978
52
Neoacanthostigma fusiforme MFLUCC 11-0510
63/Dematiohelicosporum guttulatum MFLUCC 17-2011
Pseudogliophragma indicum MTCC 11985
85/1.0
56/Parawiesneriomyces syzygii CBS 141333
83
Wiesneriomyces conjunctosporus BCC18606
Wiesneriomycetaceae
100/1.0
Wiesneriomyces laurinus DAOM 250029
99/1.0 90/1.0
Wiesneriomyces laurinus MFLUCC 17-0076
Speiropsis pedatospora CBS 397.59
82/1.0
Speiropsis scopiformis LAMIC007206
100/1.0 Bezerromyces brasiliensis URM 7411
100/1.0 Bezerromyces pernambucoensis URM 7412
Bezerromycetaceae
Xiliomyces brasiliensis URM 7413
68/Neorhamphoria garethjonesii MFLUCC 16-0210
100/1.0 Botryosphaeria dothidea CBS 115476
Botryosphaeriales (Outgroup)
Botryosphaeria agaves MFLUCC 10-0051
74/-
100/1.0
100/1.0
0.1
13
Tu b e u f i a l e s
T
u
b
e
u
f
i
a
c
e
a
e
Fungal Diversity (2020) 105:17–318
◂Fig. 82 Phylogram generated from maximum likelihood analysis
(RAxML) of genera in Tubeufiales based on ITS, LSU, rpb-2 and tef1
sequence data. Maximum likelihood bootstrap values equal or above
70%, Bayesian posterior probabilities equal or above 0.90 (MLBS/
PP) are given at the nodes. An original isolate number is noted
after the species name. The tree is rooted to Botryosphaeria agaves
(MFLUCC 10-0051) and B. dothidea (CBS 115476). The ex-type
strains are indicated in bold. Hyphen (-) represents support values
below 70% MLBS and 0.90 PP
(2018b) transferred Neorhamphoria to Bezerromycetaceae
based on phylogenetic and morphological evidence. Four
species of Bezerromycetaceae are reported, Bezerromyces
brasiliensis, B. pernambucoensis, Neorhamphoria garethjonesii, and Xiliomyces rasiliensis, and all have both morphology and DNA molecular data (Boonmee et al. 2016;
Bezerra et al. 2017).
Bezerromyces J.D.P. Bezerra, Souza-Motta & Crous, Mycol.
Progr. 16: 301 (2017).
Index Fungorum number: IF 817522; Facesoffungi number: FoF 06810; 2 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Bezerromyces brasiliensis J.D.P. Bezerra,
Souza-Motta & Crous, Mycol. Progr. 16: 302 (2017).
Notes: Bezerromyces was established based on the type
species B. brasiliensis and a second species B. pernambucoensis. They are known only as endophytes associated with
cactus species in tropical dry forests (Bezerra et al. 2017).
Their ascomata are superficial or immersed in culture media,
and sometimes there have multi-septate, brown chlamydospores linked to ascomata by hyphae (Bezerra et al. 2017).
Bezerromyces brasiliensis J.D.P. Bezerra, Souza-Motta &
Crous, in Bezerra et al., Mycol. Progr. 16(4): 302 (2016)
[2017].
Index Fungorum number: IF 817523; Facesoffungi number: FoF 06810; Fig. 83
Description: see Bezerra et al. (2017).
Material examined: Brazil, Pernambuco, Catimbau
National Park (8°36′35″S, 37°14′40″W), as endophyte from
cactus Tacinga inamoena, September 2013, J.D.P. Bezerra
(URM 89943, holotype).
Other genera included
Neorhamphoria Boonmee, Hüseyın & Selçuk, Mycosphere
7(9): 1450 (2016).
Index Fungorum number: IF 552704; Facesoffungi number: FoF 02823; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Neorhamphoria garethjonesii Boonmee,
Hüseyın & Selçuk, Mycosphere 7(9): 1451 (2016).
Notes: Neorhamphoria was introduced as a genus incertae sedis in Tubeufiales. Lu et al. (2018b) accepted it as a
195
member of Bezerromycetaceae based on phylogenetic and
morphological evidence. Neorhamphoria is characterized by
dark apothecial ascomata, broad cellular pseudoparaphyses,
with bitunicate, broad-clavate asci, and hyaline, muriform
ascospores.
Xiliomyces J.D.P. Bezerra, Souza-Motta & Crous, Mycol.
Progr. 16: 304 (2017).
Index Fungorum number: IF 817525; Facesoffungi number: FoF 06811; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Xiliomyces brasiliensis J.D.P. Bezerra,
Souza-Motta & Crous, Mycol. Progr. 16: 304 (2017).
Notes: Xiliomyces was established mainly based on phylogenetic analyses. Xiliomyces brasiliensis was reported as
an endophyte from cactus Tacinga inamoena, and its morphology is incomplete because it lacks well-defined sexual
or asexual structures (Bezerra et al. 2017).
Economic and ecological significance
There is no report on their economic significance but they
play roles in recycling organic matter.
Tubeufiaceae M.E. Barr, Mycologia 71(5): 948 (1979).
Index Fungorum: IF 81599; Facesoffungi number: FoF
00204, 377 species.
Saprobic on decaying wood and leaves in terrestrial and/
or aquatic habitats. Sexual morph: Ascomata superficial,
seated on a subiculum, unilocular, globose-subglobose to
obovate, solitary to gregarious, cream-white, yellow, pale
brown, brown to black, ostiolate, with or without setae.
Peridium composed of cells of textura angularis, thickwalled cells, yellow, pale brown, dark brown to black externally, with thin layers of textura prismatica inwardly, hyaline, yellow, pale brown to brown. Hamathecium comprising
numerous filiform, septate, branched, sometimes anastomosing, hyaline pseudoparaphyses, embedded in a gelatinous
matrix. Asci 8-spored, bitunicate, fissitunicate, saccate,
cylincrical to clavate, sometimes broadly oblong-subclavate,
with or without an apically rounded, distinct ocular chamber, with or without pedicel, smooth-walled. Ascospores
2–3-seriate to fasciculate, elongate, cylindric, fusiform
to filiform, tapering towards narrow ends, hyaline to pale
brown, sometimes yellow, multi-septate, smooth-walled.
Asexual morph: Hyphomycetous, 1) helicosporous. Colonies on the substratum superficial, effuse, gregarious, white
to pink, pale yellow to yellow green, pale brown to brown.
Mycelium mostly immersed, partly superficial, composed of
branched, septate hyphae, hyaline, pale brown to brown, with
masses of crowded conidia. Conidiophores macronematous,
mononematous, erect, cylindrical, branched or unbranched,
hyaline, pale brown to dark brown, smooth-walled. Conidiogenous cells holoblastic, mono- to polyblastic, discrete
13
196
Fungal Diversity (2020) 105:17–318
Fig. 83 Bezerromyces brasiliensis (URM 89943, holotype). a
Ascomata. b Pseudoparaphyses. c Chlamydospores. d, e Asci and
ascospores. f Ascospore. Bezerromyces pernambucoensis. g Asco-
mata on culture medium. h Ascoma and ascospores. i–l Asci and
ascospores. m, n Germinating ascospores. Scale bars: a, h = 25 µm,
b–f, i–n = 10 µm
or integrated, determinate or sympodial, terminal or intercalary, denticulate, arising laterally from the lower parts
of conidiophores as tiny tooth-like protrusions or bladderlike protrusions, sometimes truncate at apex after conidial
secession, hyaline to pale brown, smooth-walled. Conidia
solitary, acrogenous, or pleurogenous, or acropleurogenous,
helicoid, rounded at tip, hyaline to yellow green, yellow, pale
brown to brown, multi-septate, guttulate, smooth-walled; 2)
chlamydosporous. Conidiophores lacking. Chlamydospores
holoblastic, broadly oval to ellipsoid, dictyoseptate, pale
brown when immature, darkened to black when matured,
with terminal cells round and subhyaline; 3) phragmosporous. Conidiophores macronematous, mononematous, erect,
arising as lateral branches from creeping hyphae, branched
or unbranched, thin-walled and smooth, indistinctly septate,
flexuous, hyaline. Conidiogenous cells holoblastic, mono- to
polyblastic, sympodial, terminal or intercalary, cylindrical,
with denticles, hyaline. Conidia solitary, acrogenous, fusiform to obclavate, slightly curved and acute at both ends,
13
Fungal Diversity (2020) 105:17–318
hyaline, multi-septate, slightly constricted at the septa, guttulate, smooth-walled.
Type: Tubeufia Penz. & Sacc., Malpighia 11(11–12): 517
(1898) [1897].
Notes: Tubeufiaceae was introduced based on the generic
type Tubeufia to accommodate bitunicate ascomycetes
occurring as saprobes on decaying wood. Barr (1979a)
accepted six genera in the family, while Rossman (1987)
accepted 12 genera, Kirk et al. (2001) accepted 21 genera,
Lumbsch and Huhndorf (2010) accepted 23 genera, and
Boonmee et al. (2014b) accepted 19 genera. Brahamanage
et al. (2017) introduced Dictyospora, Chaiwan et al. (2017)
introduced Neotubeufia, Luo et al. (2017) introduced Muripulchra and Neohelicomyces, Liu et al. (2018) introduced
Helicotubeufia and Neohelicosporium to this family based
on morphology and phylogeny (Lu et al. (2018b) reappraised
Tubeufiaceae and introduced 13 genera, and accepted 43
genera in this family based on phylogenetic analyses and
morphological evidence. Liu et al. (2019) introduced Kevinhydea and Jayasiri et al. (2019) introduced Discotubeufia
and Hyde et al. (2020b) introduced Camporesiomyces. Currently, Tubeufiaceae contains 46 genera (Lu et al. 2018b;
Jayasiri et al. 2019; Liu et al. 2019).
Tubeufiaceous fungi are widespread distributed in tropical and temperate regions (Rossman 1987; Kirk et al. 2001;
Lumbsch and Huhndorf 2010; Boonmee et al. 2014b; Luo
et al. 2017; Lu et al. 2018a, b). Most species in this family
are saprobic on terrestrial woody substrates and some are
found on aquatic habitats (Barr 1979a, 1980; Rossman 1987;
Kirk et al. 2001; Zhao et al. 2007; Lumbsch and Huhndorf
2010; Boonmee et al. 2011, 2014b; Hyde et al. 2016b; Doilom et al. 2017; Luo et al. 2017; Phookamsak et al. 2018).
Interestingly, most asexual morphs of Tubeufiaceae were
collected from freshwater habitats (Hyde et al. 2016b,
2017; Brahamanage et al. 2017; Chaiwan et al. 2017; Lu
et al. 2017a, b, c, 2018a, b; Luo et al. 2017; Liu et al. 2018).
Tubeufia Penz. & Sacc., Malpighia 11(11–12): 517 (1898)
[1897].
Index Fungorum number: IF 5635; Facesoffungi number:
FoF 00063; 57 morphological species (Species Fungorum
2020), 38 species with molecular data.
Notes: Tubeufia is the type genus of Tubeufiaceae, established by Penzig and Saccardo (1897) based on the type
species T. javanica. Boonmee et al. (2014b) designated the
epitype of T. javanica with phylogenetic analyses. Lu et al.
(2018b) reappraised the taxonomy of Tubeufia and introduced 18 species and six combinations in this genus based
on phylogeny and morphology.
Tubeufia longiseta D.Q. Dai & K.D. Hyde, Fungal Diversity
82: 42 (2016).
197
Index Fungorum: IF 552029; Facesoffungi number: FoF
01985; Fig. 84
Saprobic on dead culms of bamboo in a forest. Sexual
morph: Ascomata 230–350 μm high × 170–220 μm diam.
superficial, gregarious, ellipsoidal, subglobose, setose, coriaceous, with a central ostiole, dark brown to black. Ostiole
single, central. Setae 320–410 × 3.5–4.5 μm, dense, flexuous, covering the whole ascoma, unbranched, rounded at
apical end, septate, dark brown. Peridium 25–45 μm wide,
composed cells of textura angularis, dark, with innermost
part comprising thin layers of hyaline to pale brown cells
of textura subprismatica. Hamathecium comprising numerous filiform, septate, branched, cellular pseudoparaphyses.
Asci 130–160 × 13–18 μm ( x̄ = 150 × 15 μm, n = 20),
8-spored, bitunicate, cylindrical, rounded at apex, with an
ocular chamber, thick-walled, short-pedicellate. Ascospores
55–75 × 4.5–6 μm ( x̄ = 65 × 5.3 μm, n = 50), 2–3-seriate,
broad fusiform, cylindrical to long subfusiform, elongate,
slightly curved, tapering towards rounded ends, hyaline,
13–17-septate, guttulate. Asexual morph: undetermined.
Culture characteristics: Ascospores germinating on water
agar and germ tubes produced from ascospores within 12
hours. Colonies growing on PDA, circular, with flat surface, edge undulate, reaching 11 mm in two weeks at 28
°C, brown to dark brown. Mycelium superficial and partially
immersed, branched, septate, hyaline to pale brown, smooth.
Material examined: Thailand, Chiang Rai, Mae Fah
Luang University, on dead culms of bamboo, 19 February
2017, Yong-Zhong Lu, MFU01 (MFLU 17-1119, HKAS
100789); living culture, MFLUCC 17-1810.
GenBank numbers: ITS: MW136949, TEF1: MW147764,
RPB2:MW147765.
Notes: Morphologically, our new collection resembles
Tubeufia longiseta in ascomata, asci, and ascospores (Dai
et al. 2017). Although the ascospores of our new collection are longer than the holotype of T. longiseta (55–75 vs.
38.5–55.5 μm), we identify them as the same species as
there is only a single base pair difference in the ITS sequence
data. We provide LSU, rpb-2 and tef1-α sequence data in
this study, but we could not compare them to the holotype
of T. longiseta as it has only ITS sequence data available
(Dai et al. 2017).
Other genera included
Acanthohelicospora Boonmee & K.D. Hyde, Fungal Diversity 68(1): 251 (2014).
Index Fungorum number: IF 550572; Facesoffungi number: FoF 00206; – 4 morphological species (Species Fungorum 2020), 4 species with molecular data.
Type species: Acanthohelicospora pinicola Boonmee &
K.D. Hyde, Fungal Diversity 68(1): 251 (2014).
Notes: Acanthohelicospora was introduced based on
morphological and phylogenetic evidence. Its sexual morph
13
198
Fig. 84 Tubeufia longiseta (MFLU 17-1119). a Superficial ascomata on substrate. Note ascomata surrounded by black setae. b Setae.
c Ascoma. d Peridium. e–h Asci. i–k Ascospores. l, m Colonies on
13
Fungal Diversity (2020) 105:17–318
PDA from above and below. Scale bars: a, d, i–k = 200 µm, b, c =
100 µm, e–h = 50 µm
Fungal Diversity (2020) 105:17–318
is characterized by superficial ascomata covered by setae
which taper to an acute apex, 8-spored, bitunicate asci and
slightly curved, fusiform ascospores, which taper towards
rounded ends (Boonmee et al. 2014b; Lu et al. 2018b). Its
asexual morph is characterized by helicosporous hyphomycetous, which are similar to Helicosporium in conidiophores, conidiogenous cells and conidial morphology (Lu
et al. 2017a, 2018b). Four species are accommodated in this
genus, A. aurea, A. guianensis, A. pinicola, and A. scopula
(Lu et al. 2018b).
Acanthophiobolus Berl., Atti Congr. Bot. Intern. di Genova,
1892: 571 (1893).
Index Fungorum number: IF 13; Facesoffungi number:
FoF 06812; – 3 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Acanthophiobolus helicosporus (Berk.
& Broome) J. Walker, Trans. Br. mycol. Soc. 58(3): 445
(1972).
≡ Sphaeria helicospora Berk. & Broome, Ann. Mag. nat.
Hist., Ser. 2 9: 383 (1852).
Notes: Acanthophiobolus was introduced with A. helicosporus as the type species (Berlese 1893, as A. helminthosporus). Boonmee et al. (2011) reexamined the holotype of A. helicosporus and accepted Acanthophiobolus as a
member of Tubeufiaceae. The ascomata of Acanthophiobolus are globose to subglobose, reddish brown to dark brown,
with much longer red brown to dark brown setae, and asci
are elongate-cylindrical with long filiform ascospores
(Boonmee et al. 2011, 2014). Lu et al. (2018b) listed three
species, A anogeissi, A. helicosporus, and A. indicus.
Acanthostigma De Not., Hedwigia 4: 28 [1865].
Index Fungorum number: IF 16; Facesoffungi number:
FoF 06813; – 41 morphological species (Species Fungorum
2020), 2 species with molecular data.
Type species: Acanthostigma perpusillum De Not., Hedwigia 4: 28 [1865].
Notes: Réblová and Barr (2000) reviewed Acanthostigma
and six species were accepted. Boonmee et al. (2011) introduced A. chiangmaiensis based on phylogenetic analyses
and morphology. Boonmee et al. (2014b) accepted only
A. perpusillum, A. chiangmaiensis, and A. minutum within
Acanthostigma, but did not change the status of other Acanthostigma species. The ascomata of Acanthostigma are globose to subglobose, dark brown to black and covered by dark
setae, asci are clavate with a short pedicel, and ascospores
are broadly fusiform to clavate, with one of the middle cells
often broader than others (Boonmee et al. 2011, 2014). Lu
et al. (2018b) accepted seven species, A. chiangmaiensis,
A. ellisii, A. longisporum, A. minutum, A. patagonicum, A.
perpusillum, and A. revocatum.
199
Acanthostigmina Höhn., Sber. Akad. Wiss. Wien, Math.naturw. Kl., Abt. 1 118: 1499 (1909).
Index Fungorum number: IF 19; Facesoffungi number:
FoF 06814; – 2 morphological species (Species Fungorum
2020), 2 species with molecular data.
Type species: Acanthostigmina minuta (Fuckel) Clem. &
Shear, Gen. fung., Edn 2 (Minneapolis): 270 (1931).
≡ Lasiosphaeria minuta Fuckel, Jb. nassau. Ver. Naturk.
23–24: 148 (1870) [1869–70].
Notes: Boonmee et al. (2014b) reported that A. minuta
(ANM810, ANM818 and ANM238) is related to two strains
of A. multiseptatum (ANM475 and ANM665) which represented Acanthostigmina. The ascomata of Acanthostigmina
are globose to subglobose, dark brown, and covered by
sparse dark brown to black setae, and asci are broadly cylindric-subclavate with curved fusiform ascospores (Boonmee
et al. 2014b). Lu et al. (2018b) accepted two species, A.
minuta and A. multiseptatum.
Acanthotubeufia Y.Z. Lu & K.D. Hyde, Fungal Diversity
92: 146 (2018).
Index Fungorum number: IF 554812; Facesoffungi number: FoF 04696; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Acanthotubeufia filiforme (Promp. &
A.N. Mill.) Y.Z. Lu & K.D. Hyde, Fungal Diversity 92: 146
(2018).
≡ Acanthostigma filiforme Promp. & A.N. Mill., Mycologia 102(3): 575 (2010).
Notes: Acanthotubeufia was introduced based on morphological and phylogenetic analyses. Its morphology is similar
to Neoacanthostigma and Acanthostigma with one-celled
or rarely one-septate setae covering the globose to subglobose ascomata, but differs by its longer, narrower and symmetrical filiform ascospores (Promputtha and Miller 2010,
as Acanthostigma filiforme; Lu et al. 2018b).
Aquaphila Goh, K.D. Hyde & W.H. Ho, Mycol. Res. 102(5):
588 (1998).
Index Fungorum number: IF 27791; Facesoffungi number: FoF 02356; – 2 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Aquaphila albicans Goh, K.D. Hyde &
W.H. Ho, Mycol. Res. 102(5): 588 (1998).
Notes: Castañeda-Ruíz et al. (2000) introduced A. edentata based on morphology. Tsui et al. (2007) accepted Aquaphila as a member of Tubeufiaceae based on phylogenetic
analyses. Boonmee et al. (2014b) linked its sexual-asexual
morphs based on phylogenetic evidence. The sexual morph
of Aquaphila is characterized by dark globose to subglobose
ascomata covered by brown setae, and cylindrical-clavate
asci with fusiform ascospores (Boonmee et al. 2014b).
Its asexual morphs are hyaline flexuous to geniculate
13
200
conidiophores with hyaline fusoid to falcate or sigmoid
conidia (Goh et al. 1998a; Boonmee et al. 2014b; Hyde et al.
2016b). Hyde et al. (2016b) reported a new record of A. albicans associated with submerged wood in southern Thailand.
Artocarpomyces Subram., Kavaka 22/23: 52 (1996) [1994].
Index Fungorum number: IF 28736; Facesoffungi number: FoF 06815; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Artocarpomyces paradoxus Subram. [as
‘paradoxa’], Kavaka 22/23: 52 (1996) [1994].
Notes: Artocarpomyces is characterized by macronematous conidiophores arising from a brown pseudoparenchymatous stroma, and conidia are muriform, irregular and
resembling the fruit of Artocarpus (Subramanian 1994). Lu
et al. (2018b) accepted Artocarpomyces as a genus of Tubeufiaceae based on morphological evidence.
Berkleasmium Zobel, Icon. fung. (Prague) 6: 4 (1854).
Index Fungorum number: IF 7362; Facesoffungi number:
FoF 01879; – 44 morphological species (Species Fungorum
2020), 7 species with molecular data.
Type species: Berkleasmium concinnum (Berk.) S.
Hughes, Canad. J. Bot. 36: 740 (1958).
≡ Sporidesmium concinnum Berk., London J. Bot. 4: 309
(1845).
Notes: Tanney and Miller (2017) accepted Berkleasmium
within Tubeufiaceae, and reported that the type species B.
concinnum, is the asexual morph of Neoacanthostigma
septoconstrictum based on phylogenetic analyses. Lu et al.
(2018b) accepted seven Berkleasmium species within Tubeufiaceae based on phylogenetic and morphological evidence.
Berkleasmium is characterized by superficial ascomata with
dark brown to black multi-celled setae, and cylindrical asci
with slightly curved fusiform ascospores (Tanney and Miller
2017; Lu et al. 2018b). Its asexual morphs comprise dictyosporous and helicosporous hyphomycetes (Tanney and
Miller 2017; Lu et al. 2018b).
Bifrontia Norman, Bot. Notiser: 18 (1872).
Index Fungorum number: IF 569; Facesoffungi number:
FoF 00175; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Bifrontia compactior Norman, Bot.
Notiser: 19 (1872).
Notes: Boonmee et al. (2014b) reexamined the holotype of B. compactior and accepted Bifrontia as a genus
of Tubeufiaceae. The ascomata of Bifrontia are globose to
subglobose, collapsing when mature or dried, and asci are
fusiform with straight to slightly curved fusiform ascospores
(Boonmee et al. 2014b).
Boerlagiomyces Butzin, Willdenowia 8(1): 39 (1977).
13
Fungal Diversity (2020) 105:17–318
Index Fungorum number: IF 607; Facesoffungi number:
FoF 00176; – 8 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Boerlagiomyces velutinus (Penz. & Sacc.)
Butzin, Willdenowia 8(1): 39 (1977).
≡ Boerlagella velutina Penz. & Sacc., Malpighia
11(9–10): 404 (1897).
Notes: Crane et al. (1998) reviewed and accepted six
Boerlagiomyces species in Tubeufiaceae. The ascomata of
Boerlagiomyces are globose and covered with flexuous and
hairy black velvety setae, and asci are cylindric-clavate with
hyaline muriform ascospores (Boonmee et al. 2014b). Lu
et al. (2018b) listed eight species in this genus.
Camporesiomyces D.P. Wanas. & K.D. Hyde, Fungal Diversity 100: 139 (2020).
Index Fungorum number: IF 557070; Facesoffungi number: FoF 07074; – 3 morphological species (Species Fungorum 2020), 3 species with molecular data.
Type species: Camporesiomyces mali D.P. Wei & K.D.
Hyde, Fungal Diversity 100: 139 (2020).
Notes: Camporesiomyces was introduced based on morphological and phylogenetic analyses. The type species of
Camporesiomyces is distinguished from other members
in Tubeufiaceae by multi-loculate ascomata (Hyde et al.
2020b). Acanthostigma patagonicum and Helicoma vaccinii
were transferred to Camporesiomyces patagonicus and C.
vaccinii based on phylogenetic evidence, respectively (Hyde
et al. 2020b).
Chaetosphaerulina I. Hino, Bull. Miyazaki Coll. Agric.
Forest. 10: 62 (1938).
Index Fungorum number: IF 971; Facesoffungi number:
FoF 06816; – 6 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Chaetosphaerulina yasudae I. Hino,
Canad. J. Plant Sci. 10: 62 (1938).
Notes: Pirozynski (1972) transferred C. yasudae and C.
vermicularispora to Herpotrichia. Sivanesan (1984) transferred these two species to Tubeufia. Crane (1998) transferred Thaxteriellopsis bambusicola and T. lignicola to
Chaetosphaerulina. Boonmee et al. (2011) accepted Crane’s
evaluation for Thaxteriellopsis bambusicola but reinstated T.
lignicola as the type species of Thaxteriellopsis. Boonmee
et al. (2014b) excluded Chaetosphaerulina from Tubeufiaceae. Lu et al. (2018b) accepted Chaetosphaerulina as a
member of Tubeufiaceae based on morphological evidence.
The ascomata of Chaetosphaerulina are ovoid to ellipsoid
and covered with hairy setae, asci are cylindrical and slightly
truncate at apex, with curved hyaline fusiform ascospores
which are constricted at the median (Boonmee et al. 2011).
Fungal Diversity (2020) 105:17–318
Chlamydotubeufia Boonmee & K.D. Hyde, Fungal Diversity 51(1): 78 (2011).
Index Fungorum number: IF 563500; Facesoffungi number: FoF 02357; – 5 morphological species (Species Fungorum 2020), 4 species with molecular data.
Type species: Chlamydotubeufia huaikangplaensis
Boonmee & K.D. Hyde, Fungal Diversity 51(1): 78 (2020).
Notes: Lu et al. (2018b) accepted five species in Chlamydotubeufia and excluded three species from this genus
based on phylogenetic inference. The sexual morph of
Chlamydotubeufia is characterized by globose to subglobose ascomata covered with dark brown to black setae, and
cylindrical to clavate asci with straight to slightly curved
fusiform ascospores (Boonmee et al. 2011; Lu et al. 2018b).
Its asexual morphs are mostly chlamydosporous and rarely
helicosporous hyphomycetous (Lu et al. 2018b).
Dematiohelicoma Y.Z. Lu, J.C. Kang & K.D. Hyde, Fungal
Diversity 92: 157 (2018).
Index Fungorum number: IF 554821; Facesoffungi number: FoF 04700; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Dematiohelicoma pulchrum (R.F.
Castañeda & Guarro) Y.Z. Lu & K.D. Hyde, Fungal Diversity 92: 158 (2018).
≡ Helicoma pulchrum R.F. Castañeda & Guarro [as ‘pulchra’], in Castañeda Ruíz, Kendrick, Guarro & Mayayo,
Mycol. Res. 102(1): 58 (1998).
Notes: Dematiohelicoma was introduced based on morphological and phylogenetic analyses. The conidia of Dematiohelicoma are characterized by hyaline or subhyaline basal
cells, tapering gradually towards a truncated base, while the
remaining cells are reddish brown, and their conidial septa
are dark brown and obviously darker than the adjacent parts
(Lu et al. 2018b).
Dematiohelicomyces Y.Z. Lu, Boonmee & K.D. Hyde, Fungal Diversity 92: 159 (2018).
Index Fungorum number: IF 554824; Facesoffungi number: FoF 04701; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Dematiohelicomyces helicosporus (Boonmee, Y.Z. Lu & K.D. Hyde) Y.Z. Lu, Fungal Diversity 92:
159 (2018).
≡ Chlamydotubeufia helicospora Boonmee, Y.Z. Lu &
K.D. Hyde, in Hyde et al., Fungal Diversity 80:123 (2016).
Notes: Dematiohelicomyces was introduced based on
morphological and phylogenetic analyses. Dematiohelicomyces resembles Helicomyces in having short conidiophores
that are 0–3-septate, and helicoid conidia, having a spathulate basal end cell, but can be distinguished by its brown
conidiophores (Lu et al. 2018b).
201
Dematiohelicosporum Y.Z. Lu, J.K. Liu & K.D. Hyde, Fungal Diversity 92: 159 (2018).
Index Fungorum number: IF 554826; Facesoffungi number: FoF 04703; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Dematiohelicosporum guttulatum Y.Z. Lu,
J.K. Liu & K.D. Hyde, Fungal Diversity 92: 160 (2018).
Notes: Dematiohelicosporum was introduced based on
morphological and phylogenetic analyses. Conidia of Dematiohelicosporum are slightly circinate in three dimensions,
clearly verrucose and guttulate, olivaceous to brown (Lu
et al. 2018b).
Dematiotubeufia Y.Z. Lu, Boonmee & K.D. Hyde, Fungal
Diversity 92: 160 (2018).
Index Fungorum number: IF 554828; Facesoffungi number: FoF 04705; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Dematiotubeufia chiangraiensis (Boonmee
& K.D. Hyde) Y.Z. Lu, Fungal Diversity 92: 162 (2018).
≡ Helicoma chiangraiense Boonmee & K.D. Hyde, in
Boonmee, Rossman, Liu, Crous, Bhat, Chukeatirote, Jones
& Hyde, Fungal Diversity 68: 271 (2014).
Notes: Dematiotubeufia was introduced based on morphology and phylogenetic analysis. It is characterized by
dark brown to black, shiny ascomata with distinct ostioles,
and cylindrical asci with straight to slightly curved fusiform ascospores (Boonmee et al. 2014b, as Helicoma chiangraiense; Lu et al. 2018b).
Dictyospora Brahaman., Y.Z. Lu, Boonmee & K.D. Hyde,
Mycosphere 8(7): 924 (2017).
Index Fungorum number: IF 553178; Facesoffungi number: FoF 03262; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Dictyospora thailandica Brahaman., Y.Z.
Lu, Boonmee & K.D. Hyde, Mycosphere 8(7): 924 (2017).
Notes: Lu et al. (2018b) described a new record of D.
thailandica bearing some peculiar morphological characteristics, and pointed out that Dictyospora resembles Chlamydotubeufia in both asexual and sexual morphs, but phylogenetic analyses indicated that they are distinct genera.
Discotubeufia Jayasiri, E.B.G. Jones & K.D. Hyde, Mycosphere 10(1): 156 (2019).
Index Fungorum number: IF 555585; Facesoffungi number: FoF 05300; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Discotubeufia browneae Jayasiri, E.B.G.
Jones & K.D. Hyde, Mycosphere 10(1): 158 (2019).
Notes: Discotubeufia is characterized by cup-shaped,
erumpent to superficial, light brown to dark, setiferous
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202
Fungal Diversity (2020) 105:17–318
ascomata and cylindrical to sub cylindrical asci (Jayasiri
et al. 2019).
terminal conidiogenous cells tapering towards apex, having
an inverted funnel shape at apex (Lu et al. 2018b).
Helicangiospora Boonmee, Bhat & K.D. Hyde, Fungal
Diversity 68(1): 259 (2014).
Index Fungorum number: IF 550574; Facesoffungi number: FoF 00210; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Helicangiospora lignicola Boonmee, Bhat
& K.D. Hyde, Fungal Diversity 68(1): 262 (2014).
Notes: Helicangiospora helicoma-like asexual morph was
found from the culture of sexual morph specimen (Boonmee
et al. 2014b).
Helicoma Corda, Icon. fung. (Prague) 1: 15 (1837).
Index Fungorum number: IF 8473; Facesoffungi number:
FoF 00211; – 64 morphological species (Species Fungorum
2020), 22 species with molecular data.
Type species: Helicoma muelleri Corda, Icon. fung.
(Prague) 1: 15 (1837).
Notes: Goos (1986) reviewed the status of Helicoma species. Lu et al. (2018b) introduced ten new Helicoma species,
and redefined the morphological concept of Helicoma to
include two additional asexual morphs, one is the typicalhelicoma morphology with conidia that are acrogenous or
acropleurogenous, helicoid, circinate, dry, tapering towards
apex, truncate at the base, coiled 1¼–1¾ times, not becoming loose in water. Another one is characterized by conidiogenous cells that are intercalary, cylindrical, with denticles,
arising laterally from the lower portion of conidiophores as
tooth-like protrusions, and conidia are pleurogenous, helicoid, hygroscopic, tapering towards apex and rounded at tip,
coiled 1½–5 times, becoming loosely coiled in water (Lu
et al. 2018b). Lu et al. (2018b) accepted 57 species within
Helicoma, and excluded twelve species from this genus. Liu
et al. (2019) reported a new species, H. hydei, from decaying
wood in Thailand.
Helicoarctatus Y.Z. Lu, J.C. Kang & K.D. Hyde, Fungal
Diversity 92: 165 (2018).
Index Fungorum number: IF 554830; Facesoffungi number: FoF 04707; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Helicoarctatus aquaticus Y.Z. Lu, J.C.
Kang & K.D. Hyde, Fungal Diversity 92: 166 (2018).
Notes: Helicoarctatus shares similar morphology with
Helicosporium by setiform conidiophores, and discrete,
tooth-like protrusions conidiogenous cells arising laterally
from lower portion of the conidiophores, with each bearing
1–2 tiny sporogenous conidiogenous loci (Lu et al. 2018b).
Helicodochium J.S. Monteiro, R.F. Castañeda, A.C. Cruz &
Gusmão, Mycotaxon 127: 6 (2014).
Index Fungorum number: IF 804995; Facesoffungi number: FoF 04709; – 2 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Helicodochium amazonicum J.S. Monteiro, R.F. Castañeda, A.C. Cruz & Gusmão, Mycotaxon
127: 6 (2014).
Notes: Lu et al. (2018b) accepted Helicodochium as a
genus of Tubeufiaceae based on morphological and phylogenetic evidence. Helicodochium is characterized by
sporodochial conidiomata, macronematous branched conidiophores, and conidiogenous cells that produce helicoids,
multi-septate, smooth and hyaline conidia (Monteiro et al.
2014; Lu et al. 2018b).
Helicohyalinum Y.Z. Lu, J.K. Liu & K.D. Hyde, Fungal
Diversity 92: 168 (2018).
Index Fungorum number: IF 554833; Facesoffungi number: FoF 04710; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Helicohyalinum infundibulum Y.Z. Lu,
J.K. Liu & K.D. Hyde, Fungal Diversity 92: 170 (2018).
Notes: Helicohyalinum is characterized by flexuous,
cylindrical, 0–3-septate, hyaline conidiophores with the
13
Helicomyces Link, Mag. Gesell. naturf. Freunde, Berlin
3(1–2): 21 (1809).
Index Fungorum number: IF 8476; Facesoffungi number:
FoF 00212; – 17 morphological species (Species Fungorum
2020), 4 species with molecular data.
Type species: Helicomyces roseus Link, Mag. Gesell.
naturf. Freunde, Berlin. 3: 21 (1809).
Notes: Linder (1929) redefined this genus, and many of
the species originally assigned to Helicomyces were reassigned to Helicosporium or Helicoma. Moore (1955) provided a key for all known Helicomyces species. Goos (1985)
reviewed the taxonomic status of Helicomyces, provided
updated descriptions and a key to accepted species. Lu et al.
(2018b) pointed out the confusion of Helicomyces and reappraised its taxonomy and phylogeny. Twelve species were
accepted in Helicomyces and four species were excluded (Lu
et al. 2018b). Helicomyces is characterized by conidiophores
that are 0–3-septate, arising as lateral branches from creeping hyphae, conidiogenous cells are integrated, sympodial
and truncate at apex after conidial secession, and conidia
are acrogenous or acropleurogenous, helicoid and rounded
at tip (Link 1809; Linder 1929; Moore 1955; Goos 1985;
Lu et al. 2018b).
Helicosporium Nees, Syst. Pilze (Würzburg): 68 (1817) –
13 species (Lu et al. 2018b).
Fungal Diversity (2020) 105:17–318
Index Fungorum number: IF 8484; Facesoffungi number:
FoF 00213; – 20 morphological species (Species Fungorum
2020), 8 species with molecular data.
Type species: Helicosporium vegetum Nees, Syst. Pilze
(Würzburg): 68 (1817).
Notes: Linder (1929) redefined the generic concept
of Helicosporium. Goos (1989) reviewed the status of
Helicosporium species and redefined the generic concept
including species with hyaline conidia whose filaments
do not exceed 3 μm diam. Lu et al. (2018b) reviewed all
Helicosporium species, redefined the generic concept, and
accepted 13 species and excluded 25 species from this genus.
Helicotruncatum Y.Z. Lu, J.C. Kang & K.D. Hyde, Fungal
Diversity 92: 220 (2018).
Index Fungorum number: IF 554859; Facesoffungi number: FoF 04730; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Helicotruncatum palmigenum (Penz. &
Sacc.) Y.Z. Lu & K.D. Hyde, Fungal Diversity 92: 220
(2018).
≡ Helicosporium intermedium var. palmigenum Penz. &
Sacc., Malpighia 15(7–9): 249 (1902) [1901].
Notes: Helicotruncatum was established based on morphological and phylogenetic evidence. Helicotruncatum is
characterized by the lateral cell wall of the conidiophore
and by the basal cells of conidia distinctively thickened (Lu
et al. 2018b).
203
Notes: Phookamsak et al. (2018) reported the phylogenetic relationship of Kamalomyces and first described its
asexual morph. Kamalomyces is characterized by subglobose to lemoniform ascomata with short stalks on a subiculum of black hyphae, and broadly cylindrical to clavate asci
with crowded, hyaline vermiform and septate ascospores
(Verma et al. 2008; Phookamsak et al. 2018). Six Kamalomyces species are accepted, K. bambusicola, K. indicus, K.
mahabaleshwarensis, K. mangrovei, K. polyseptatus and K.
thailandicus (Lu et al. 2018b, Hyde et al. 2019).
Kevinhydea N.G. Liu, Y.Z. Lu & J.K. Liu, Mycol. Prog. 18:
675 (2019).
Index Fungorum number: IF 555352; Facesoffungi number: FoF 04875; – 1 morphological species (Liu et al. 2019),
1 species with molecular data.
Type species: Kevinhydea brevistipitata N.G. Liu, Y.Z.
Lu & J.K. Liu, Mycol. Prog. 18: 677 (2019).
Notes: Kevinhydea was introduced based on phylogenetic
and morphological evidence. Kevinhydea is characterized by
dictyosporous asexual morph and has short conidiophores
which differentiate it from Chlamydotubeufia, Dictyospora,
Tamhinispora, and Tubeufia, genera that lack conidiophores
(Liu et al. 2019).
Helicotubeufia Y.Z. Lu & J.K. Liu, Mycosphere 9(3): 500
(2018).
Index Fungorum number: IF 554759; Facesoffungi number: FoF 04385; – 3 morphological species (Species Fungorum 2020), 3 species with molecular data.
Type species: Helicotubeufia guangxiensis Y.Z. Lu &
J.K. Liu, Mycosphere 9(3): 500 (2018).
Notes: Helicotubeufia was introduced based on phylogenetic and morphological evidence to accommodate three
species, the type species H. guangxiensis, H. hydei, and H.
jonesii. Helicotubeufia is characterized by subglobose to
ellipsoidal-ovate, dark brown to black ascomata, and cylindrical asci with hyaline fusiform, slightly curved ascospores,
and hyphomycetous helicosporous asexual morphs (Liu
et al. 2018).
Manoharachariella Bagyan., N.K. Rao & Kunwar, Mycotaxon 109: 301 (2009).
Index Fungorum number: IF 512919; Facesoffungi number: FoF 07530; – 4 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Manoharachariella lignicola Bagyan.,
N.K. Rao & Kunwar, Mycotaxon 109: 302 (2009).
Notes: Rajeshkumar and Singh (2012) introduced Manoharachariella indica and Selcuk and Ekici (2014) introduced M. elsadii based on morphological evidence. Doilom
et al. (2017) introduced the fourth species, M. tectonae, and
accepted this genus as a member of Tubeufiaceae based on
phylogenetic analysis and morphological evidence. Manoharachariella is characterized by macronematous, mononematous conidiophores with integrated monoblastic conidiogenous cells producing solitary, doliiform, obpyriform
dictyoseptate and apiculate conidia (Bagyanarayana et al.
2009; Rajeshkumar and Singh 2012; Selcuk and Ekici 2014;
Doilom et al. 2017).
Kamalomyces R.K. Verma, N. Sharma & Soni, Forest Fungi
of Central India: 196 (2008).
Index Fungorum number: IF 512509; Facesoffungi number: FoF 06817; – 6 morphological species (Species Fungorum 2020), 3 species with molecular data.
Type species: Kamalomyces indicus R.K. Verma, N.
Sharma & Soni, Forest Fungi of Central India: 196 (2008).
Muripulchra Z.L. Luo, Hong Y. Su & K.D. Hyde, Cryptog.
Mycol. 38(1): 36 (2017).
Index Fungorum number: IF 818825; Facesoffungi number: FoF 02647; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Muripulchra aquatica Z.L. Luo, Hong Y.
Su & K.D. Hyde, Cryptog. Mycol. 38(1): 39 (2017).
13
204
Notes: Muripulchra was introduced based on phylogenetic and morphological analyses. Muripulchra is characterized by its obpyriform conidia often carrying part of conidiogenous cells at the base as a broken frill (Luo et al. 2017).
Neoacanthostigma Boonmee, Bhat & K.D. Hyde, Fungal
Diversity 68(1): 278 (2014).
Index Fungorum number: IF 550576; Facesoffungi number: FoF 00214; – 3 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Neoacanthostigma fusiforme Boonmee,
Bhat & K.D. Hyde, Fungal Diversity 68(1): 279 (2014).
Notes: Neoacanthostigma was introduced based on phylogenetic and morphological evidence. Lu et al. (2017c)
introduced four new Neoacanthostigma species. Lu et al.
(2018b) accepted only the type species N. fusiforme based
on phylogenetic and morphological analyses. The ascomata
of Neoacanthostigma are globose to subglobose, reddish
brown to dark brown, to black, and are covered by black
setae. Asci are cylindric-clavate with a short rounded pedicel, ascospores are narrowly fusiform and tapering towards
the rounded ends. The asexual morph of Neoacanthostigma
is characterized by helicosporous hyphomycetous, which
lack conidiophores, and conidia are acrogenous with loosely
coiled conidial filaments (Boonmee et al. 2014b; Lu et al.
2018b).
Neochlamydotubeufia Y.Z. Lu, Boonmee & K.D. Hyde,
Fungal Diversity 92: 221 (2018).
Index Fungorum number: IF 554861; Facesoffungi number: FoF 04731; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Neochlamydotubeufia fusiformis Y.Z. Lu,
Boonmee & K.D. Hyde, Fungal Diversity 92: 221 (2018).
Notes: Neochlamydotubeufia was introduced based on
phylogenetic evidence. Neochlamydotubeufia shares similar sexual and asexual morphs with Chlamydotubeufia and
Dictyospora, but phylogenetic analyses support Neochlamydotubeufia as a distinct genus (Lu et al. 2018b).
Neohelicoma Y.Z. Lu, Boonmee & K.D. Hyde, Fungal
Diversity 92: 224 (2018).
Index Fungorum number: IF 554864; Facesoffungi number: FoF 04734; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Neohelicoma fagacearum (Boonmee &
K.D. Hyde) Y.Z. Lu, Fungal Diversity 92: 224 (2018).
≡ Helicoma fagacearum Boonmee & K.D. Hyde, in
Boonmee et al., Fungal Diversity 68: 271 (2014).
Notes: Neohelicoma was introduced based on phylogenetic evidence. Neohelicoma shares similar sexual morph
with Helicoma in ascomata, asci and ascospores, but phylogenetic analyses supported it as a distinct genus (Lu et al.
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Fungal Diversity (2020) 105:17–318
2018b). The type species Neohelicoma fagacearum is the
only accepted species within this genus.
Neohelicomyces Z.L. Luo, Bhat & K.D. Hyde, Cryptog.
Mycol. 38(1): 39 (2017).
Index Fungorum number: IF 818820; Facesoffungi number: FoF 02643; – 8 morphological species (Species Fungorum 2020), 8 species with molecular data.
Type species: Neohelicomyces aquaticus Z.L. Luo, Bhat
& K.D. Hyde, Cryptog. Mycol. 38(1): 40 (2017).
Notes: Neohelicomyces was introduced to accommodate N. aquaticus, N. grandisporus, and N. submersus. Lu
et al. (2018b) introduced Neohelicomyces hyalosporus, and
synonymized Helicosporium pallidum under Neohelicomyces based on phylogenetic and morphological evidence.
Tibpromma et al. (2018b) reported Neohelicomyces pandanicola from dead roots of Pandanus. Crous et al. (2019a)
introduced N. deschampsiae from the culm base of dead leaf
sheath of Deschampsia cespitosa. Neohelicomyces can be
recognized from other helicosporous hyphomycetous by its
hyaline conidiophores and conidia (Luo et al. 2017; Lu et al.
2018b; Crous et al. 2019a).
Neohelicosporium Y.Z. Lu, J.C. Kang & K.D. Hyde, Mycol.
Progr. 17 (5): 637 (2017).
Index Fungorum number: IF 822045; Facesoffungi number: FoF 03570; – 22 morphological species (Species Fungorum 2020), 18 species with molecular data.
Type species: Neohelicosporium parvisporum Y.Z. Lu,
J.C. Kang & K.D. Hyde, Mycol. Progr. 17 (5): 637 (2017).
Notes: Neohelicosporium was established based on phylogenetic and morphological evidence. Lu et al. (2018b)
accepted 22 species in this genus. Neohelicosporium differs from Helicosporium in having acrogenous and/or
acropleurogenous conidia developing from an integrated,
sympodial conidiogenous cells while Helicosporium species have pleurogenous conidia and discrete, determinate
conidiogenous cells (Lu et al. 2018a, b).
Neotubeufia Chaiwan, Boonmee, Y.Z. Lu & K.D. Hyde,
Mycosphere 8(9): 1149 (2017).
Index Fungorum number: IF 553871; Facesoffungi number: FoF 03777; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Neotubeufia krabiensis Chaiwan, Boonmee, Y.Z. Lu & K.D. Hyde, Mycosphere 8(9): 1149 (2017).
Notes: Neotubeufia was introduced based on phylogenetic
and morphological evidence. The type species N. krabiensis
is the only accepted species in this genus. The ascomata
of Neotubeufia are dark brown to black subglobose without
setae, and asci are broadly cylindrical with slightly curved
cylindric-fusiform ascospores (Chaiwan et al. 2017).
Fungal Diversity (2020) 105:17–318
Pleurohelicosporium Y.Z. Lu, J.C. Kang & K.D. Hyde,
Fungal Diversity 92: 247 (2018).
Index Fungorum number: IF 554884; Facesoffungi number: FoF 04743; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Pleurohelicosporium parvisporum Y.Z.
Lu, J.C. Kang & K.D. Hyde, Fungal Diversity 92: 248
(2018).
Notes: Pleurohelicosporium was introduced based on phylogenetic and morphological evidence. Pleurohelicosporium
shares similar asexual morphs with Neohelicosporium but
can be distinguished by its pleurogenous conidia, while the
conidia of Neohelicosporium are acrogenous or acropleurogenous (Lu et al. 2018b). The type species P. parvisporum is
the only accepted species in this genus.
Podonectria Petch, Trans. Br. mycol. Soc. 7(3): 146 (1921).
Index Fungorum number: IF 4280; Facesoffungi number:
FoF 06818; – 9 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Podonectria coccicola (Ellis & Everh.)
Petch, Trans. Br. mycol. Soc. 7(3): 146 (1921).
≡ Nectria coccicola Ellis & Everh., J. Mycol. 2(4): 39
(1886).
Notes: Rossman (1978) reviewed this genus and accepted
eight species. The ascomata of Podonectria are subgloboseglobose, light yellow reddish and covered with light brown
mycelium, asci are cylindric-clavate with short pedicele,
and ascospores are clavate to fusiform. Its asexual morph
is characterized by phragmosporous hyphomycetous, and
conidia are produced on sympodial conidiophores (Petch
1921; Rossman 1978; Boonmee et al. 2011). Boonmee
et al. (2014b) accepted Podonectria in Tubeufiaceae and
pointed out that its sexual morphs were compatible with
Tubeufiaceae.
Pseudohelicomyces Y.Z. Lu, J.K. Liu & K.D. Hyde, Fungal
Diversity 92: 248 (2018), nom. illegit., non Garnica & E.
Valenz. (2000).
Index Fungorum number: IF 554886; Facesoffungi number: FoF 04745; – 8 morphological species (Species Fungorum 2020), 7 species with molecular data.
Type species: Pseudohelicomyces talbotii (Goos) Y.Z. Lu
& K.D. Hyde, Fungal Diversity 92: 252 (2018).
≡ Helicosporium talbotii Goos, Mycologia 81(3): 368
(1989).
Notes: Pseudohelicomyces was introduced to accommodate five species, P. aquaticus, P. hyalosporus, P. indicus, P.
paludosus, and P. talbotii based on phylogenetic and morphological evidence (Lu et al. 2018b). Phookamsak et al.
(2019) reported Pseudohelicomyces menglunicus from an
unidentified seed in China. Jayasiri et al. (2019) introduced a
new Pseudohelicomyces species and two new records. Hyde
205
et al. (2019) described a new member, Pseudohelicomyces
menglunicus, from an unidentified seed coat. Recently, Lu
et al. (2020) submitted a proposal to conserve Pseudohelicomyces (Tubeufiaceae) against Pseudohelicomyces (Hymenogastraceae). Pseudohelicomyces shares similar conidial
characters with Helicomyces but can be distinguished by
its erect and branched cylindrical conidiophores (Lu et al.
2018b; Jayasiri et al. 2019).
Pseudohelicoon Y.Z. Lu & K.D. Hyde, Fungal Diversity
92: 254 (2018).
Index Fungorum number: IF 554919; Facesoffungi number: FoF 04748; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Pseudohelicoon subglobosum (Goh & C.H.
Kuo) Y.Z Lu & Hyde, Fungal Diversity 92: 255 (2018).
≡ Helicoon subglobosum Goh & C.H. Kuo [as ‘Helicoön’], Phytotaxa 346(2): 145 (2018).
Notes: Pseudohelicoon shares similar asexual morph with
Helicoön in non-proliferating and ellipsoidal to doliiform
conidia but phylogenetic analyses indicated that they are different genera (Lu et al. 2018b).
Tamhinispora Rajeshk. & Rah. Sharma, Mycosphere 4(2):
166 (2013).
Index Fungorum number: IF 803105; Facesoffungi number: FoF 00215; – 3 morphological species (Species Fungorum 2020), 3 species with molecular data.
Type species: Tamhinispora indica Rajeshk. & Rah.
Sharma, Mycosphere 4(2): 167 (2013).
Notes: Tamhinispora was established based on phylogenetic and morphological evidence. Rajeshkumar et al.
(2018) introduced Tamhinispora srinivasanii and Sommai
et al. (2019) introduced T. saraburiensis. The conidia of
Tamhinispora are irregular, dictyoseptate, dark brown and
adorned with brown to blackish apical appendages, which
are unique in Tubeufiaceae (Rajeshkumar and Sharma 2013;
Rajeshkumar et al. 2018; Sommai et al. 2019).
Thaxteriella Petr., Annls mycol. 22(1/2): 63 (1924).
Index Fungorum number: IF 5408; Facesoffungi number:
FoF 00189; – 10 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Thaxteriella corticola Petr., Annls mycol.
22(1/2): 63 (1924).
Notes: Boonmee et al. (2011) reexamined the type material of T. corticola and accepted Thaxteriella as a genus of
Tubeufiaceae. The ascomata of Thaxteriella are globose or
subglobose, shiny, and collapsing in the middle when dried,
asci are broadly clavate or subclavate with a short pedicel,
and ascospores are cylindrical to long fusiform (Boonmee
et al. 2011, 2014).
13
206
Thaxteriellopsis Sivan., Panwar & S.J. Kaur, Kavaka 4: 39
(1977) [1976].
Index Fungorum number: IF 5409; Facesoffungi number:
FoF 01866; – 1 morphological species (Lu et al. 2018b), 1
species with molecular data.
Type species: Thaxteriellopsis lignicola Sivan., Panwar
& S.J. Kaur, Kavaka 4: 39 (1977) [1976].
Notes: Crane et al. (1998) transferred T. lignicola to
Chaetosphaerulina. Boonmee et al. (2011) reexamined the
holotype of T. lignicola (IMI 197065), designated an epitype
(MFLU 10-0057), and retained Thaxteriellopsis as a distinct
genus within Tubeufiaceae. Doilom et al. (2017) introduced
an asexual record of T. lignicola based on phylogenetic evidence. Lu et al. (2018b) reported new records of T. lignicola
with new morphological characteristics. Thaxteriellopsis is
characterized by globose to subglobose ascomata are reddish brown to dark brown, collapsed when dry, and covered
with brown to black setae, cylindrical asci with a long pedicel, and straight or slightly curved ascospores are fusiform
to clavate with 5 septa. The conidia of Thaxteriellopsis are
typical helicoma-like but conidiophores are distinct from
Helicoma species (Boonmee et al. 2011; Doilom et al. 2017;
Lu et al. 2018b).
Economic and ecological significance
Tubeufiaceae fungi have the potential to produce a number of
bioactive compounds. Hanada et al. (1996) reported a novel
protein produced by a species of Helicosporium, which had
effects on neurite outgrowth via cultured cortical neurons
and NGF-treated PC12 cells. Ohtsu et al. (2003) obtained
five compounds from Helicomyces sp. No. 19353, and found
that one compound had a significant anti-diabetic activity
(Ohtsu et al. 2003; Yoshimura et al. 2003; Zenkoh et al.
2003). Dong et al. (2004) reported that mobility of nematodes was inhibited by 98.95% by the mycelial extracts from
Helicomyces roseus. Jiao et al. (2006) reported that seven
compounds were isolated from Tubeufiaceae sp. A-00471
and exhibited antibiotic activity against Gram-positive bacteria (Jiao et al. 2006). Hu et al. (2006) obtained one Diepoxin and five Decaspirones compounds from Helicoma viridis. Jung et al. (2012) reported that 2-methylresorcinol from
Helicosporium sp. KCTC 0635BP exhibited antimicrobial
activity against various types of bacteria and fungi, and also
exhibited considerable cytotoxicity activity against human
cancer cells. Lee et al. (2013) found that Helicosporium sp.
0635BP was effective against Fusarium oxysporum, Phytophthora drechsleri, and Rhizoctonia solani.
Wiesneriomycetaceae Suetrong, Rungjindamai, Somrithipol. & E.B.G. Jones, Phytotaxa 176: 288 (2014).
Index Fungorum: IF 550306; Facesoffungi number: FoF
06311, 20 species.
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Fungal Diversity (2020) 105:17–318
Saprobic on different substrates, colonies effuse, consisting of scattered conidiomata. Mycelium immersed. Asexual
morph: Conidiomata sporodochial or synnematal, solitary
to gregarious, with or without setae. Setae subulate, septate,
pigmented, thick-walled, erect, flexuous, arising from basal
pseudoparenchymatous stalk or mycelium immersed in the
substrate. Conidiophores macronematous, mononematous,
penicillate, septate, branched, straight or flexuous, solitary
or gregarious. Conidiogenous cells holoblastic, monoblastic,
polyblastic, discrete, determinate, terminate, clavate, cylindrical, slightly ampulliform. Conidia solitary to gregarious,
catenate, connected by narrow isthmi, fusiform to falcate,
cylindrical, subcylindrical, cuneiform, branched, hyaline to
slightly pigmented, aseptate. Sexual morph: undetermined.
Type: Wiesneriomyces Koord.
Notes: Suetrong et al. (2014) introduced Wiesneriomycetaceae as order incertae sedis, based on morphology and
molecular phylogenetic inference. Pratibha et al. (2015)
placed Wiesneriomycetaceae in Tubeufiales, and included
Pseudogliophragma in this family. Santos (2015) included
Speiropsis in this family and suggested Pseudogliophragma
as a synonym of Wiesneriomyces. Parawiesneriomyces was
included in this family by Crous et al. (2016b). Bezerra et al.
(2017) introduced Wiesneriomycetales to accommodate
Wiesneriomycetaceae based on morphological characteristics and phylogenetic analyses. Liu et al. (2017) assigned
Wiesneriomycetales as a synonym of Tubeufiales based on
phylogenetic inference and divergence times estimates which
estimated as 40 MYA (crown age).
Wiesneriomyces Koord., Verh. K. Akad. Wet., tweede
sect. 13(4): 246 (1907).
Index Fungorum number: IF 10438; Facesoffungi number: FoF 06312; 3 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Wiesneriomyces laurinus (Tassi) P.M.
Kirk, Trans. Br. mycol. Soc. 82(4): 748 (1984).
≡ Volutellaria laurina Tassi, Atti R. Accad. Fisiocrit.
Siena, Sér. 4 8: 5 (1897).
Notes: Kirk (1984) proposed the new combination,
Wiesneriomyces laurinus, based on the holotype of Volutellaria laurina and showed that it was an earlier name
for Wiesneriomyces javanicus. No sexual morph has been
reported for this genus (Suetrong et al. 2014; Bezerra et al.
2017; Lu et al. 2018b).
Wiesneriomyces laurinus (Tassi) P.M. Kirk, Trans. Br.
mycol. Soc. 82(4): 748 (1984).
Index Fungorum number: IF 107371; Facesoffungi number: FoF 06313; Fig. 85
Saprobe, on bark of Hevea brasiliensis; Sexual morph:
Undetermined. Asexual morph: Colonies 385–440 μm
diam., on the substrate inconspicuous. Conidiomata
Fungal Diversity (2020) 105:17–318
207
Fig. 85 Wiesneriomyces laurinus (MFLU 18-1384, new host
record). a, b Habit on substrate. c Conidioma. d Seta. e Base of seta.
f Tip of seta. g, h Conidiophores. i Germinated spores. j Conidia. k
Culture characteristic on PDA after 21 days (k = colony from above, l
= colony from below). Scale bars: a–c = 100 µm, d–j = 30 µm
sporodochial, superficial, solitary to gregarious, scattered.
Setae 130–430 × 3–12 µm, arising from basal pseudostromatal stalk, straight or ± bent, abundant, dark brown to
black, verruculose, 5–10-septate, tapering to subulate apex.
Conidiophores 18–38 × 2–3 µm, macronematous, hyaline to
subhyaline, smooth-walled, branched. Conidiogenous cells
terminal, clavate to slightly inflated, hyaline, smooth-walled,
5.5–8.5 × 1.5–2.5 µm. Conidia [(4)5–7 cells], 46–67 × 2–4
µm, in chains, straight, long cylindrical, hyaline to subhyaline, aseptate, smooth-walled.
Culture characteristics: Conidia germinated on MEA
within 12 hours and germ tube produced from one or both
end or several cell of the conidia. Colonies on PDA reaching
4.5–5 cm diam. after 21 days at room temperature (20–25
13
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Fungal Diversity (2020) 105:17–318
Wiesneriomyces laurinus YMF1.04044
Wiesneriomyces laurinus DAOM.250029
W
Wiesneriomyces laurinus BCC9453
92/1.00
i
Wiesneriomyces laurinus BCC2922
-/0.96
e
Wiesneriomyces laurinus BCC40684
-/0.99
Wiesneriomyces laurinus BCC3922
s
78/0.99
Wiesneriomyces laurinus MFLUCC 17-0076
n
Wiesneriomyces laurinus BCC18609
e
Wiesneriomyces laurinus BCC40614
-/0.98
r
Phalangispora sinensis YMF1.03656
100/i
90/Phalangispora sinensis YMF1.02689
o
100/0.99
Phalangispora constricta MUCL 40953
m
Phalangispora nawawii LAMIC041712
y
Setosynnema
yunnanense
YMF1.03964
98/1.00
Setosynnema yunnanense YMF1.02680
-/1.00
c
Wiesneriomyces conjunctosporus BCC18608
e
96/1.00
Wiesneriomyces conjunctosporus BCC18606
98/1.00
t
Wiesneriomyces conjunctosporus BCC4027
96/0.99
a
74/Wiesneriomyces conjunctosporus BCC20803
99/1.00
c
Parawiesneriomyces syzygii CBS 141333
100/1.00
e
Pseudogliophragma indicum MTCC 11985
a
Speiropsis pedatospora GUFCC 18008
100/1.00
Speiropsis pedatospora CBS 397.59
e
99/1.00
Speiropsis scopiformis LAMIC005111
100/1.00
Speiropsis scopiformis LAMIC007206
100/1.00 Tubeufia chiangmaiensis MFLUCC 17-1801
Tubeufia chiangmaiensis MFLUCC 11-0514
Helicomyces hyalosporus MFLUCC 17-0051
100/1.00
Neohelicomyces hyalosporus GZUCC 16-0086
100/1.00
Tubeufiaceae
Helicomyces roseus CBS 283.51
98/1.00
Acanthostigma chiangmaiense MFLUCC 10-0125
Bezerromyces pernambucoensis URM 7411
100/1.00
Bezerromycetaceae
Bezerromyces brasiliensis URM 7411
Botryosphaeria agaves MFLUCC 10-0051
Botryosphaeriales (Outgroup)
Botryosphaeria dothidea CBS 115476
-/0.98
87/1.00
TubeufIales
100/1.00
100/1.00
0.04
Fig. 86 Phylogram generated from maximum likelihood analysis
(RAxML) of genera in Wiesneriomycetaceae based on ITS, LSU,
rpb-2, SSU and tef1 sequence data. Maximum likelihood bootstrap
values equal or above 70%, Bayesian posterior probabilities equal
or above 0.90 (MLBS/PP) are given at the nodes. An original iso-
late number is noted after the species name. The tree is rooted to
Botryosphaeria agaves (MFLUCC 10-0051) and B. dothidea (CBS
115476). The ex-type strains are indicated in bold. The new sequence
is in blue. Hyphen (-) represents support values below 70% MLBS
and 0.90 PP
°C), circular, effuse, velvety to hairy, initially white, becoming slightly radiating with greyish-brown concentric ring,
white at the margin from above, dark grey to blackish at the
center from below, with white circular edge.
Material examined: Thailand, Phayao Province, Muang,
inner bark of Hevea brasiliensis (Euphorbiaceae), 5 December 2016, Senwanna C., RBPY019 (MFLU 18-1384; living
culture MFLUCC 17-0076)
Sequence data: ITS: MN168764, LSU: MN168761, rpb2: MN174863, SSU: MN168759, tef1: MT050455.
Notes: During our study of microfungi on Para rubber in
Thailand, a fresh specimen was obtained from inner bark
of Hevea brasiliensis. The morphological characters of our
fresh collection fit well with the description of W. laurinus
(Suetrong et al. 2014, Chen and Tzean 2015). Based on our
phylogenetic analysis of combined LSU and SSU sequence
data, our strain groups with other W. laurinus strains with
moderate bootstrap support (Fig. 86). Thus, this fresh collection is identified as W. laurinus.
13
Other genera included
Parawiesneriomyces Crous & M.J. Wingf., in Crous et al.,
Persoonia 36: 389 (2016).
Index Fungorum number: IF 817060; Facesoffungi number: FoF 06556; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Parawiesneriomyces syzygii Crous & M.J.
Wingf., in Crous et al., Persoonia 36: 389 (2016).
Notes: Parawiesneriomyces was introduced based
on phylogeny and morphology (Crous et al. 2016b).
Fungal Diversity (2020) 105:17–318
100/1.0
100/1.0
100/1.0
Valsaria insitiva CBS 127882
Valsaria spartii CBS 139070
Valsaria lopadostomoides CBS 139062
Bambusaria bambusae DDQ00253
100/1.0
Bambusaria bambusae DDQ00254
Valsariales
Fig. 87 Phylogram generated
from maximum likelihood
analysis (RAxML) of genera in
Valsariales based on ITS, LSU,
SSU and tef1 sequence data.
Maximum likelihood bootstrap
values equal or above 70%,
Bayesian posterior probabilities
equal or above 0.90 (MLBS/
PP) are given at the nodes. An
original isolate number is noted
after the species name. The tree
is rooted to Capnodium coffeae
(OSC 100414) and Leptoxyphium fumago (CBS 123.26). The
ex-type strains are indicated in
bold. Hyphen (-) represents support values below 70% MLBS
and 0.90 PP
209
Myrmaecium rubrum CBS 345.86
71/100/1.0
100/1.0
100/1.0
Myrmaecium fulvopruinatum CBS 139058
Lasiodiplodia crassispora CBS 118741
Botryosphaeria dothidea AFTOL-ID 946
Cladoriella xanthorrhoeae culture CBS 143398
Cladoriella kinglakensis culture CPC 32730
Cladoriellales
100/1.0
Botryosphaeriales
Capnodium coffeae OSC 100414
100/1.0
Outgroup
Leptoxyphium fumago CBS 123.26
0.02
Parawiesneriomyces is similar to Wiesneriomyces in its
conidial characters, however they differ in conidiomata and
setae characters (Crous et al. 2016b). Thus, further collections and sequence data are necessary to resolve and to confirm the validity of the new genus.
Phalangispora Nawawi & J. Webster, Trans. Br. mycol. Soc.
79(1): 65 (1982).
Index Fungorum number: IF 9328; Facesoffungi number:
FoF 07568; – 4 morphological species (Species Fungorum
2020), 3 species with molecular data.
Type species: Phalangispora constricta Nawawi & J.
Webster, Trans. Br. mycol. Soc. 79(1): 65 (1982).
Notes: Phalangispora is characterized by setose, sporodochial conidiomata and branched, tetraradiate conidia
(Nawawi and Webster 1982). A combined dataset of ITS,
LSU, tub1 and tub2 sequence data placeed Phalangispora
within Wiesneriomyces (Santos 2015). Recent molecular
phylogenetic data showed that Phalangispora is a distinct
group within Wiesneriomycetaceae (Guo et al. 2019).
Pseudogliophragma Phadke & V.G. Rao, Norw. J Bot. 27:
127 (1980).
Index Fungorum number: IF 9579; Facesoffungi number:
FoF 06557; – 1 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Pseudogliophragma indica Phadke & V.G.
Rao, Norw. Jl Bot. 27(2): 127 (1980).
Notes: Pseudogliophragma was introduced to accommodate a single species characterized by synnematous conidiomata, 4–8-celled, fusiform to falcate conidia. Recent molecular phylogenetic data showed that Pseudogliophragma is a
distinct genus within Wiesneriomycetaceae (Pratibha et al.
2015).
Setosynnema D.E. Shaw & B. Sutton, J. Linn. Soc., Bot.
91: 33 (1985).
Index Fungorum number: IF 9328; Facesoffungi number:
FoF 07569; – 3 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Setosynnema isthmosporum D.E. Shaw &
B. Sutton, Bot. J. Linn. Soc. 91(1–2): 34 (1985).
Notes: Setosynnema is charactered by synnematous conidiomata, filiform conidia with an isthmus at the central septum (Shaw and Sutton 1985). The genus was introduced as a
genus incertae sedis and reported as aquatic fungi (Bai et al.
2013; Rajeshkumar 2014; Guo et al. 2019). Guo et al. (2019)
accepted Setosynnema as a member of Wiesneriomycetaceae
based on phylogenetic and morphological evidence.
Speiropsis Tubaki, J. Hattoribot. Lab. 20: 171 (1958).
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210
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Fungal Diversity (2020) 105:17–318
◂Fig. 88 Valsaria ostryae (MFLU 17-0837, holotype). a, b Habit
and appearance of ectostromata on host surface. c Vertical section
of stroma. d Peridium wall. e Apically free pseudoparaphyses. f, g
Immature asci h Mature asci. i–k Immature ascospores l, m Mature
ascospore. q Hyphae with denticles and conidia (partly budding). n–p
Phialides. r–w Conidia. Scale bars: a = 500 μm, b, c = 200 μm, f–h
= 50 μm, d, n–q = 10 μm, e, i–m, r–w = 5 μm
Index Fungorum number: IF 9965; Facesoffungi number:
FoF 06558; – 8 morphological species (Species Fungorum
2020), 2 species with molecular data.
Type species: Speiropsis pedatospora Tubaki, J. Hattori
bot. Lab 20: 171 (1958).
Notes: Speiropsis is characterized by erect, simple,
straight, mostly mononematous conidiophores, polyblastic
conidiogenous cells and catenate conidia in branched or
unbranched chains connected by narrow isthmi (Barbosa and
Gusmao 2005; Pratibha et al. 2016). Based on morphology
and molecular analyses, Pratibha et al. (2016) referred Speiropsis to Wiesneriomycetaceae. There are 28 DNA sequence
data available in GenBank, comprising S. pedatospora and
S. scopiformis.
Economic and ecological significance
Genera included in Wiesneriomycetaceae play a role as saprobes, occurring on different substrates with a worldwide
distribution (Suetrong et al. 2014; Pratibha et al. 2015;
2016; Crous et al. 2016b). In addition, Speiropsis has been
reported as an aquatic hyphomycete (Barbosa and Gusmao
2005; Santos 2015).
Valsariales Jaklitsch, K.D. Hyde & Voglmayr.
Index Fungorum number: IF 811900; Facesoffungi number: FoF 08827.
Valsariales was introduced by Jaklitsch et al. (2015) based
on multi-gene phylogeny analyses to accommodate the single family Valsariaceae. Valsariales species have perithecioid ascomata, immersed in eu- or pseudostromata, bitunicate asci, without obvious fissitunicate dehiscence and dark
brown, bicelled ascospores. Member of Valsariales have a
worldwide distribution on various substrates as saprobes,
plant pathogens or necrotrophs (Ju et al. 1996; Jaklitsch et al.
2015). There are only three genera with molecular data. Our
phylogenetic analyses generated from ITS, LSU, SSU and
tef1 sequence data (Fig. 87) indicate that the three genera
cluster with high bootstrap support (100% ML, 1.0 PP), and
each genus is well-resolved. The divergence time for Valsariales is estimated as 224 MYA (stem age, Hongsanan
et al. 2020).
Accepted families: Valsariaceae.
Valsariaceae Jaklitsch, K.D. Hyde & Voglmayr, in Jaklitsch
et al., Fungal Diversity 73: 167 (2015).
211
Index Fungorum number: IF 811901; Facesoffungi number: FoF 06561, 41 species.
Saprobic and pathogens in bark of dicotyledons in terrestrial habitats or on culms of bamboo. Sexual morph:
Stromata eu- or pseudostromatic, immersed, erumpent
to superficial on bark, enclosed on top and/or at the sides
by a black pseudoparenchymatous. Ascomata perithecial,
immersed in the stromata, usually monostichous in valsoid or diatrypoid configuration, upright or oblique, with
several ostiolar necks fusing into one. Ostiole periphysate.
Peridium composed of several cell layers of brown to black
cells. Hamathecium comprising numerous paraphyses. Asci
4–8-spored, bitunicate, with or without obvious fissitunicate,
cylindrical, with short pedicel, an ocular chamber apex, and
a pulvinate ring staining in Congo Red. Ascospores (often
obliquely) uni-seriate, ellipsoid to subfusiform, dark brown,
1-septate, with a dark, usually not or slightly constricted at
the septum, budding in artificial culture, with surface ornamentation. Asexual morph: in nature coelomycetous, in culture hyphomycetous or coelomycetous. Conidia produced
on phialides, minute pegs or by budding of ascospores and
conidia, oblong or bullet-shaped, hyaline, 1-celled, smoothwalled, with inconspicuous guttules.
Type: Valsaria Ces. & De Not.
Notes: Based on bitunicate asci without obvious fissitunicate dehiscence, Ju et al. (1996) placed Valsaria in Dothideomycetes (as Loculoascomycetes). However, Kirk et al.
(2008) referred Valsaria to Diaporthales (Sordariomycetes),
based on the hamathecium consisting of true, apically free
paraphyses, a true ascomatal wall distinct from the surrounding pseudostroma and unitunicate asci (Barr 1978, 1990;
Glawe 1985; Huhndorf 1992). Recent phylogenetic analyses have shown Valsariaceae to be a distinct family within
Dothideomycetes (Jaklitsch et al. 2015; Pem et al. 2019b).
Valsariaceae was introduced by Jaklitsch et al. (2015) to
accommodate Bambusaria, Myrmaecium and Valsaria,
based on multi-gene phylogeny analyses of ITS, LSU, SSU,
rpb-2 and tef1 sequence data.
Valsaria Ces. & De Not., Comm. Soc. crittog. Ital. 1(fasc.
4): 205 (1863).
Index Fungorum number: IF 5704; Facesoffungi number:
33 morphological species (Species Fungorum 2020), 6 species with molecular data.
Type species: Valsaria insitiva (Tode) Ces. & De Not.
≡ Sphaeria insitiva Tode, Fung. mecklenb. sel.
(Lüneburg) 2: 36 (1791).
Notes: Valsaria contains plant pathogens and saprobes on
various hosts (Phillips 2000; Jaklitsch et al. 2015; Žežlina
et al. 2016). Morphological characteristic of Valsaria are
ovelapping and hence DNA sequence analyses are needed
to classify and delineate them. There are six epithets accommodated in Valsaria based on molecular data coupled with
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212
morphological characteristics (Jaklitsch et al. 2015; Wijayawardene et al. 2017a; Index Fungorum 2020).
Valsaria ostryae D. Pem, R. Jeewon, Camporesi & K.D.
Hyde, in Pem et al., PLoS ONE 14(6): e0217982, 5 (2019).
Index Fungorum number: IF 554758; Facesoffungi number: FoF 04614, Fig. 88
Description: see Pem et al. (2019b).
Material examined: Italy, Province of Forlì-Cesena [FC],
Santa Sofia, Camposonaldo, on dead branches of Ostrya
carpinifolia (Betulaceae), 18 March 2017, E. Camporesi,
IT 3290 (MFLU 17-0837, holotype).
Other genera included
Bambusaria Jaklitsch, D.Q. Dai, K.D. Hyde & Voglmayr, in
Jaklitsch et al., Fungal Diversity 73: 196 (2015).
Index Fungorum number: IF 811906; Facesoffungi number: FoF 06559; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Bambusaria bambusae (J.N. Kapoor &
H.S. Gill) Jaklitsch, D.Q. Dai, K.D. Hyde & Voglmayr, in
Jaklitsch et al., Fungal Diversity 73: 196 (2015).
≡ Valsaria bambusae J.N. Kapoor & H.S. Gill, Indian
Phytopath. 14(2): 152 (1962) [1961].
Notes: Based on a multi-locus phylogeny and morphological features, Jaklitsch et al. (2015) established a new
genus Bambusaria with a single species B. bambusae (synonym: Valsaria bambusae) as the type species, which is hostspecific for Bambusoideae. The genus is characterized by
eustromatic stromata, ascomata immersed in a stromata in
valsoid configuration, bitunicate asci without obvious fissitunicate dehiscence, ellipsoid to broadly fusiform or biconoid, brown, 1-septate ascospores with longitudinally ribbed
and produces asexual morph in culture.
Myrmaecium Nitschke ex Fuckel, Jb. nassau. Ver. Naturk.
23–24: 227 (1870) [1869–70].
Index Fungorum number: IF 3385; Facesoffungi number:
FoF 06560; – 7 morphological species (Species Fungorum
2020), 3 species with molecular data.
Type species: Myrmaecium rubricosum (Fr.) Fuckel, Jb.
nassau. Ver. Naturk. 23–24: 227 (1870) [1869–70].
≡ Sphaeria rubricosa Fr., Elench. fung. (Greifswald) 2:
63 (1828).
Notes: Previously Myrmaecium was treated as a synonym
of Valsaria (Ju et al. 1996, Jaklitsch et al. 2015). However,
a multi-gene molecular study established the monophyly of
Myrmaecium species, and their inclusion in Valsariaceae
(Jaklitsch et al. 2015). Myrmaecium can be distinguished
from other genera in Valsariaceae by the eu- or pseudostromata, showing ionomidotic reaction (KOH-extractable pigments). Myrmaecium has a worldwide distribution occurring
on sun-exposed, corticated logs and branches of coniferous
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Fungal Diversity (2020) 105:17–318
and broadleaf trees. In addition, Myrmaecium fulvopruinatum and M. rubrum are sources of secondary metabolites
(Aptroot 1995; Jaklitsch et al. 2015).
Economic and ecological significance
Species of Valsariaceae occur on a wide range of hosts as
saprobes, only Myrmaecium fulvopruinatum and Valsaria
insitiva were reported as pathogenic and play a negative role
in causing canker disease of Chinese chestnut (Castanea
mollissima) and stone fruit (Žežlina et al. 2016, Jiang et al.
2018a).
Venturiales Y. Zhang ter, C.L. Schoch & K.D. Hyde.
Index Fungorum number: IF 513386; Facesoffungi number: FoF 06345.
Venturiales was introduced by Zhang et al. (2011) using
morphological and ecological characteristics, as well as
strong support in multi-gene phylogenetic analyses. This
order presently contains Sympoventuriaceae and Venturiaceae (Zhang et al. 2011; Hyde et al. 2013). Some species of
this order are plant pathogens or saprobes, mostly on leaves
or stems of dicotyledons. Important characters of this order
are small to medium-sized ascomata, with or without setae,
a mostly evanescent hamathecium, 8-spored, broadly or
usually obclavate asci, usually lacking a pedicel, hyaline,
light greenish olivaceous to brown, 1-septate, symmetrical,
asymmetrical or apiosporous ascospores and hyphomycetous
asexual morphs (Zhang et al. 2011). The divergence time for
Venturiales is estimated as 184 MYA (stem age, Hongsanan
et al. 2020).
Accepted families: Sympoventuriaceae, Venturiaceae.
Sympoventuriaceae Y. Zhang ter, C.L. Schoch & K.D.
Hyde, Fungal Diversity 51: 255 (2011).
Index Fungorum number: IF 563117; Facesoffungi number: FoF 06346, 130 species.
Pathogenic or saprobic on plants. Sexual morph: Ascomata pseudothecia, solitary or aggregated, immersed in host
tissue, erumpent when mature, subglobose to globose, substomatal, subepidermal, inconspicuous; walls pigmented;
papillate, ostiolate. Peridium thick-walled, comprising of
2–3 layers of brown to dark brown cells of textura angularis.
Hamathecium comprising 2 μm wide, septate, hyaline, cellular pseudoparaphyses, anastomosing between and above the
asci, constricted at the septa. Asci 8-spored, bitunicate, fissitunicate, subcylindrical, pedicellate. Ascospores crowded
or partially overlapping in asci, symmetrical, fusoid-ellipsoidal, hyaline, medianly 1-septate, constricted at the septum,
widest in the middle of each cell, guttulate, smooth-walled
with slightly rough ends. Asexual morph: Hyphomycetous. “sympodiella”-like, “fusicladium”-like, Veronaeopsis.
Mycelium forming a superficial network with pigmented,
thin- to thick-walled, smooth, septate, anastomosing hyphae.
Fungal Diversity (2020) 105:17–318
213
Fig. 89 Sympoventuria capensis (redrawn from Crous et al.
2007b) a Pseudoparaphyses. b
Asci. c Ascospores. Scale bars:
a–c = 10 µm
Conidiogenous cells integrated, mono-blastic, holoblastic or
polyblastic, sympodial, with inconspicuous scars. Conidiophores septate, solitary, slightly thickened and darkened,
producing conidial chains with up to 20 conidia. Conidia
thin-walled, cylindrical, sometimes globose to broadly oval
at maturity, with truncate ends, hyaline to yellow brown, 1–5
septate to muriform, in unbranched chains on terminal portion of the conidiophores, smooth-walled, guttulate or not.
Type: Sympoventuria Crous & Seifert.
Notes: Species of Sympoventuriaceae have parasitic or
saprobic lifestyles and can be found on leaves or stems of
dicotyledons (Zhang et al. 2011). Species of Sympoventuriaceae are hyphomycetous and sexual morphs for most of
the species are not determined (Huanraluek et al. 2019b).
According to the outline of Ascomycota 2017, Acroconidiellina and Sympoventuria are included in this family (Wijayawardene et al. 2018).
Type species: Acroconidiellina loudetiae M.B. Ellis,
Mycol. Pap. 125: 23 (1971).
Notes: Species of Acroconidiellina can be distinguished
by having dark brown to blackish brown hairy colonies,
superficial hyphae arising setae, sclerotia, macronematous
conidiophores, polytretic conidiogenous cells, and broadly
ellipsoidal, brown, echinulate, 1- septate conidia (Ellis
1960).
Sympoventuria Crous & Seifert, Fungal Divers 25: 31
(2007).
Index Fungorum number: IF 501002; Facesoffungi number: FoF 06347; 3 morphological species (Species Fungorum 2020), 3 species with molecular data.
Type species: Sympoventuria capensis Crous & Seifert.
Notes: Sympoventuria is the type genus in the Sympoventuriaceae, distinguished from Venturiaceae by its saprobic habitat, pseudoparaphyses, and hyaline, symmetrical
ascospores (Fig. 89). The asexual morph of Sympoventuria
was described as “sympodiella”-like by Crous et al. (2007b).
Clavatispora Boonmee & K.D. Hyde, Phytotaxa 176(1): 95
(2014).
Index Fungorum number: IF 805923; Facesoffungi number: FoF 08829; –1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Clavatispora thailandica Boonmee & K.D.
Hyde, Phytotaxa 176(1): 96 (2014).
Notes: The genus is characterized by solitary, superficial
ascomata, covered by dark setae, with an ostiole, clavate,
multi-septate, yellowish to reddish brown ascospores (Boonmee et al. 2014a). It does not resemble other genera in Sympoventuriaceae. However, the LSU sequence data showed
that Clavatispora clusters with the hyphomycete genus
Scolecobasidiella in Sympoventuriaceae along with Ochroconis, Scolecobasidium, Fusicladium and Veronaeopsis
(Boonmee et al. 2014a). The asexual morph of Clavatispora
are unique in having 1–3-phragmoseptate, smooth-walled,
guttulate conidia, deeply constricted at septa, while other
genera have conidia and conidiogenous apparatus which are
mostly verrucose to denticulate (Boonmee et al. 2014a).
Other genera included
Acroconidiellina M.B. Ellis, Mycol. Pap. 125: 22 (1971).
Index Fungorum number: IF 7031; Facesoffungi number:
FoF 08828; – 4 morphological species (Species Fungorum
2020), 1 species with molecular data.
Echinocatena R. Campb. & B. Sutton, Trans. Br. mycol.
Soc. 69(1): 126 (1977).
Index Fungorum number: IF 8121; Facesoffungi number:
FoF 09212; –1 morphological species (Species Fungorum
2020), 1 species with molecular data.
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Fungal Diversity (2020) 105:17–318
Scolecobasidium podocarpi CPC 37078
86/1.00
73/0.94
Scolecobasidium terreum P043
Ochroconis mirabilis CBS 729.95
100/1.00
Acroconidiellina arecae NFCCI 3696
94/1.00
Ochroconis humicola CBS:116655
84/0.95
100/1.00 Verruconis gallopava CBS 437.64
Verruconis gallopava CBS:118.91
Scolecobasidiella avellanea CBS:772.73
92/0.91
Mycosisymbrium cirrhosum GUFCC 18012
Clavatispora thailandica MFLUCC 10-0107
100/1.00
Sympoventuriaceae
Echinocatena arthrinioides CPC 28754
72/1.00
Yunnanomyces panadanicola MFLUCC 17-2260
Yunnanomyces phoenicis MFLUCC 19-0253
Fusicladium cordae CBS:675.82
89/1.00
90/1.00
Fusicladium ramoconidii CBS :462.82
Matsushimaea fasciculata CBS:167.97
93/1.00 Matsushimaea monilioides FMR 16505
100/1.00
Fusicladium sicillianum CBS:105.85
77/Neocoleroa cameroonensis CBS:129041
Neocoleroa metrosideri PDD107531
88/0.97
Sympoventuria melaleucae CBS:144407
86/0.92
89/1.00
Sympoventuria capensis CBS:120136
100/1.00
Scolecobasidium excentricum CBS:469.95
Veronaeopsis simplex CBS:129155
100/1.00 Veronaeopsis simplex CBS:588.66
Sympoventuria regnans CBS:144506
Venturia inaequalis CBS:815.69
Venturia inaequalis CBS:594.70
Venturia inaequalis ATCC 60070
71/74/1.00
Venturia populina CBS:256.38
96/0.96
Coleroa circinans CBS:457.64
Coleroa robertiani CBS:458.64
Metacoleroa dickiei medipc
94/1.00
Metacoleroa dickiei Kruys 503 UPS
-/0.98
Venturiaceae
72/- Protoventuria alpina CBS:373.53
Tyrannosorus pinicola AFTOL ID-1235
73/- Tyrannosorus hystrioides CBS:117727
79/1.00 Caproventuria hanliniana ATCC960199
Magnohelicospora liberica FMR12414
92/1.00
Magnohelicospora fuscospora CBS:206.53
Dimerosporiopsis engleriana HPC 2591
-/0.98
99/1.00 Protoventuria barriae CBS:300.93
Apiosporina morbosa dimosp
Apiosporina collinsii CBS:118973
94/1.00 Pseudoanungitea vaccinii CBS:143164
Pseudoanungitea syzygii CBS:520.93
73/1.00
Tothia fuscella WU 31398
Cylindrosympodium variabile CBS 563.82
Sympodiella acicola CBS487.82
78/1.00
Sympodiella quercina CBS:145060
78/1.00
100/1.00
Repetophragma goidanichii HKUCC10828
100/1.00
Phaeotrichum benjaminii CBS 541.72
Trichodelitschia bisporula CBS 262.69
100/1.00
Phaeotrichales
Zeloasperisporium wrightiae MFLUCC 15-0225
Zeloasperisporium ficusicola MFLUCC 15-0221
100/1.00
Zeloasperisporiales
Natipusilla naponensis ILL AF217.1
Natipusilla decorospora ILL AF236 .1
100/1.00
Acrospermum longisporium MFLU 17-2849
0.06
13
Na pusillales
Acrospermum compressum M151
Out group
Fungal Diversity (2020) 105:17–318
◂Fig. 90 Phylogram generated from maximum likelihood analysis
(RAxML) of genera in Venturiales based on LSU, SSU and RPB2
sequence data. Maximum likelihood bootstrap values equal or above
70%, Bayesian posterior probabilities equal or above 0.90 (MLBS/
PP) are given at the nodes. An original isolate number is noted after
the species name. The tree is rooted to Acrospermum compressum
(M151) and Acrospermum longisporium (MFLU 17-2849). The extype strains are indicated in bold. Hyphen (-) represents support values below 70% MLBS and 0.90 PP
Type species: Echinocatena arthrinioides R. Campb. &
B. Sutton, Trans. Br. mycol. Soc. 69(1): 130 (1977).
Notes: The genus is characterized by micronematous and
mononematous conidiophores, consisting of branched, acropetal chains of polyblastic conidiogenous cells, with solitary,
sphaerical, brown, aseptate, echinulate conidia (Campbell
and Sutton 1977). Crous et al. (2018b) included this genus
in Sympoventuriaceae based on the sequence data of type
species (Echinocatena arthrinioides). Its polyblastic conidiogenous cells and straight and flexuous conidiophores are
reminiscent of the Venturiales (Shen et al. 2020). Therefore, we accept this genus in Sympoventuriaceae and note
that more sequence data is needed to resolve its taxonomic
placement.
Fusicladium Bonord., Handb. Allgem. mykol. (Stuttgart):
80 (1851).
Index Fungorum number: IF 8292; Facesoffungi number:
FoF 08830; – 81 morphological species (Species Fungorum
2020), 31 species with molecular data.
Type species: Fusicladium virescens Bonord., Handb.
Allgem. mykol. (Stuttgart): 80 (1851).
Note: Species of Fusicladium are characterised by having immersed, subcuticular to intra-epidermal mycelia, often
forming radiating strands or loose to dense stromatic aggregations, which give rise to conidiogenous cells or conidiophores penetrating the cuticle (Schubert et al. 2003).
Matsushimaea Subram., Kavaka 5: 96 (1978) [1977].
Index Fungorum number: IF 8865; Facesoffungi number:
FoF 08831; – 4 morphological species (Species Fungorum
2020), 3 species with molecular data.
Type species: Matsushimaea fasciculata Subram.,
Kavaka 5: 96 (1978) [1977].
Note: Species of Matsushimaea are characterised by having branched, septate, olive mycelium with hyphal coils, by
the production of sessile branched conidia arising directly
from vegetative hyphae and integrated, mono- or polyblastic, intercalary or terminal, elongated conidiogenous cells,
with irregular-shaped, septate, often branched filaments and
brown conidia. The production of sessile branched conidia
arising directly from vegetative hyphae is an important character of this genus (Crous et al. 2018b).
215
Mycosisymbrium Carris, Mycologia 86(1): 132 (1994).
Index Fungorum number: IF 27274; Facesoffungi number: FoF 06348; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Mycosisymbrium cirrhosum Carris, Mycologia 86(1): 132 (1994).
Note: According to Prathiba and Prabhugaonkar (2016),
Mycosisymbrium is well-supported as a genus in Sympoventuriaceae and is sister to Ochroconis and Verruconis
(Samerpitak et al. 2014; Machouart et al. 2014; Hyde et al.
2013; Zhang et al. 2011). This genus is similar to Ochroconis in having 2-celled conidia, slightly constricted at the
septa and polyblastic conidiogenous cells (Prathiba and
Prabhugaonkar 2016).
Ochroconis de Hoog & Arx, Kavaka 1: 57 (1974).
Index Fungorum number: IF 9136; Facesoffungi number:
FoF 06349; – 24 morphological species (Species Fungorum
2020), 14 species with molecular data.
Type species: Ochroconis constricta (E.V. Abbott) de
Hoog & Arx, Kavaka 1: 57 (1974) [1973].
≡ Scolecobasidium constrictum E.V. Abbott, Mycologia
19(1): 30 (1927).
Notes: Ochroconis species can be distinguished in their
slow to moderate growth, colonies of brown to olivaceous,
brownish, septate conidiophores and dark pigmented, and
rough-walled conidia produced by sympodial conidiogenesis
and liberated rhexolytically (Giraldo et al. 2014). Machouart
et al. (2014) included Ochroconis in Sympoventuriaceae
based on phylogeny. Pseudoparaphyses and hyaline, symmetrical ascospores and a saprobic nutritional mode are
considered to confirm its placement in Sympoventuriaceae.
Scolecobasidiella M.B. Ellis, Mycol. Pap. 125: 12 (1971).
Index Fungorum number: IF9839; Facesoffungi number:
FoF 09213; – 2 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Scolecobasidiella avellanea (Sappa &
Mosca) M.B. Ellis, Mycol. Pap. 125: 10 (1971).
Notes: Species of Scolecobasidiella are characterized by
superficial straw coloured to pale brown hyphae, septate,
pale brown, smooth conidiophores, 1–3 septate ramo-conidia
and broadly ellipsoidal or clavate, 1-septate, pale brown,
smooth conidia (Ellis 1971). This genus is closely related
to Clavatispora based on phylogenetic analyses (Boonmee
et al. 2014a). However, the type species of Scolecobasidiella
is related to Verruconis gallopava in our phylogenetic analyses (Fig. 90). The pale brown colonies with conidiophores
and conidia show a close affinity with Venturiales. Therefore, we accept this genus in Sympoventuriaceae.
Scolecobasidium E.V. Abbott, Mycologia 19(1): 30 (1927).
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Fungal Diversity (2020) 105:17–318
Fig. 91 Verruconis hevea (MFLU 17-1578, holotype). a Host specimen. b Colonies on the latex. c Arrangement of hyphal mass. d Hyphal
mass e Conidiogenesis cells. f Attachment of conidia to hypha. g–i Conidia. Scale bars: c = 100 μm, d–e = 50 μm, f = 20 μm, g–i = 5 μm
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Fungal Diversity (2020) 105:17–318
Index Fungorum number: IF9840; Facesoffungi number:
FoF 09214; – 34 morphological species (Species Fungorum
2020), 4 species with molecular data.
Type species: Scolecobasidium terreum E.V. Abbott,
Mycologia 19(1): 30 (1927).
Notes: Scolecobasidium are characterized by having T- or
Y-shaped or bilobed, 2- to many-celled conidia and ampulliform conidiogenous cells, possessing one to three conidiumbearing denticles at the apex of the conidiophores (Hao et al.
2013). Shen et al. (2020) included this genus in Sympoventuriaceae based on morphology of slow growing olivaceous
colonies. The type species of this genus also clusters inside
the family. Therefore, we accept this genus in this family.
Veronaeopsis Arzanlou & Crous, Stud. Mycol. 58: 91
(2007).
Index Fungorum number: IF 504571; Facesoffungi number: FoF 06350 – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Veronaeopsis simplex (Papendorf) Arzanlou & Crous, in Arzanlou et al., Stud. Mycol. 58: 91 (2007).
≡ Veronaea simplex Papendorf, Trans. Br. mycol. Soc.
52(3): 486 (1969).
Notes: Veronaeopsis is similar to Veronaea in its 1-septate conidia. Veronaeopsis differs from Veronaea based on
its much shorter, macronematous conidiophores and conidiogenous cells. Furthermore, Veronaeopsis has a geniculate rachis and Veronaea has a more or less straight rachis
(Arzanlou et al. 2007).
Verruconis Samerp., H.J. Choi, van den Ende, Horré & de
Hoog, Fungal Diversity 65: 117 (2014).
Index Fungorum number: IF 519169; Facesoffungi number: FoF 06351; – 9 morphological species (Species Fungorum 2020), 9 species with molecular data.
Type species: Verruconis gallopava (W.B. Cooke)
Samerp. & de Hoog, in Samerpitak, Van der Linde, Choi,
Gerrits van den Ende, Machouart, Gueidan & de Hoog, Fungal Diversity 65: 117 (2013) [2014].
≡ Diplorhinotrichum gallopavum W.B. Cooke, in Georg,
Bierer & Cooke, Sabouraudia 3: 242 (1964).
Notes: Verruconis are dematiaceous hyphomycetes
characterized by clavate to cylindrical, two-celled conidia
(Samerpitak et al. 2014). Verruconis hevea is illustrated
below.
Verruconis heveae Huanraluek., Senwanna., Jayaward.
& K.D. Hyde, in Huanraluek et al., Phytotaxa 403(1): 50
(2019).
Index Fungorum number: IF 555289; Facesoffungi number: FoF 04838, Fig. 91
Description: see Huanraluek et al. (2019b).
217
Material examined: Thailand, Phayao Province, on
dried latex of rubber, Hevea brasiliensis (Euphorbiaceae),
1 November 2016, C. Senwanna M2 (MFLU 17-1578,
holotype).
Yunnanomyces Tibpromma & K.D. Hyde, Fungal Diversity:
92: 75 (2018).
Index Fungorum number: IF 555334; Facesoffungi number: FoF 04518; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Yunnanomyces pandanicola Tibpromma
& K.D. Hyde, in Tibpromma et al., Fungal Diversity 93:75
(2018).
Notes: Yunnanomyces is characterized by globose to
broadly oval and yellow–brown, muriform conidia (Zang
et al. 2019). It is similar to Pseudocoleodictyospora reported
from teak (Doilom et al. 2017), but phylogenetic analysis of
Yunnanomyces does not support its placement in Pseudocoleodictyosporaceae. Yunnanomyces differs from Fusicladium and Verruconis based on characters of conidiophores
and conidia (Tibpromma et al. 2018b).
Economic and ecological significance
Ochroconis species are opportunistic pathogens which cause
diseases of vertebrate animals and humans. Ochroconis
humicola is the causal agent of muscular black spot disease
of Atlantic salmon (Schaumann and Priebe 1994). Verruconis gallopava was considered to be the source of several
zoonoses in broiler house chickens and trumpeters. Horré
et al. (1999) have listed 13 brain infections in young birds
and one in a cat brain (Samerpitak et. al 2014).
Venturiaceae E. Müll. & Arx ex M.E. Barr, Mycologia
71(5):947 (1979).
Index Fungorum number: IF 81516; Facesoffungi numbers: FoF 06352, 217 species.
Saprobic or parasitic on leaves or stems usually on dicotyledons, sometimes on monocotyledons. Sexual morph:
Ascomata superficial or immersed or slightly erumpent at
maturity, scattered or gregarious, single locules in a stroma
or below a blackened clypeus, globose to subglobose, membraneous or carbonaceous, usually with seta or without seta,
ostiolate, sometimes occurring on a well-developed subiculum with ascomata. Peridium comprising a pigmented single
cell layer of textura angularis. Hamathecium comprising
branched, cellular pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, evanescent, broadly or usually obclavate
or cylindrical, with or without short pedicel, ocular chamber distinct. Ascospores uni-seriate or overlapping 2-seriate,
cylindrical to ellipsoidal, with broadly rounded apex, hyaline
or light greenish, or pale yellow to brown, 1-septate upper
cell shorter and broader than lower one. Asexual morph:
Hyphomycetous. Conidiophores pigmented, singly or in
13
218
clusters, simple or branched. Conidiogenous cells integrated,
terminal or sometimes intercalary, proliferating sympodially
or percurrently, with conspicuous annellations Conidia solitary or arranged as chains, pigmented, aseptate or septate.
Type: Venturia Sacc.
Notes: Venturiaceae comprises 12 genera (Wijayawardene
et al. 2018). The members of this family are characterized
by having sphaerical superficial or erumpent ascomata
comprising elongated or oblong or nearly cylindrical, bitunicate, 8-spored asci and 2-celled, symmetric or asymmetric
ascospores, which are hyaline or pale green when immature and olive-brown or greyish green, or rarely dark brown
at maturity (Li et al. 2014). Crous et al. (2018b) included
Cylindrosympodium, Pseudoanungitea, Sympodiella, Tothia,
and Repetophragma in Venturiaceae based on their phylogenetic analyses without including all genera of the family in
their analyses. In our analyses (Fig. 90), we included all genera that have sequence data and found that the main clade of
Venturiaceae comprises species of Apiosporina, Caproventuria, Coleroa, Dimerosporiopsis, Metacoleroa, Protoventuria, Tyrannosorus, and Venturia with high support (99%
ML, 1.0 BYPP). Cylindrosympodium, Pseudoanungitea,
Sympodiella, Tothia, and Repetophragma cluster together
as a sister clade with the main clade of Venturiaceae with
low support, but sometimes in other analyses, these genera
did not cluster with members of the family (data not shown).
We presume that this separate clade can be a new family
in Venturiales, and tentatively accept Cylindrosympodium,
Pseudoanungitea, and Sympodiella in Venturiaceae as their
type species cluster within this clade. More sequence data
are needed to confirm this clade as a new family. The placement of Repetophragma is not clear as the sequence data of
type species is unavailable, thus we follow Wijayawardene
et al. (2020) and place it in Pleosporales genera incertae
sedis. We exclude Tothia from Venturiaceae base on its morphology (thyriothecial ascomata), and place the genus in
Microthyriales genera incertae sedis.
Venturia Sacc., Syll. fung. (Abellini) 1: 586 (1882).
Index Fungorum number: IF 5717; Facesoffungi number:
FoF 06353; 123 morphological species (Species Fungorum
2020), 58 species with molecular data.
Type species: Venturia inaequalis (Cooke) G. Winter.
≡ Sphaerella inaequalis Cooke, J. Bot., Lond. 4: 248
(1866).
Notes: Venturia species can be distinguished by separate
or aggregated ascomata with or without stroma, or forming
below a blackened clypeus, symmetrical or asymmetrical,
olive green, pale brown or dark brown ascospores and being
parasitic on leaves or stems of dicotyledons (Zhang et al.
2011).
Other genera included
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Apiosporina Höhn., Sber. Akad. Wiss. Wien, Math.- naturw.
Kl., Abt. 1 119: 439 (1910).
Index Fungorum number: IF 266; Facesoffungi number:
FoF 06354; – 3 morphological species (Species Fungorum
2020), 2 species with molecular data.
Type species: Apiosporina collinsii (Schwein.) Höhn.
Notes: Apiosporina species are characterized by
erumpent, subglobose, black ascomata, 8-spored bitunicate,
subcylindrical or broadly clavate asci and hyaline to pale
brown, clavate apiospores (Zhang et al. 2011). The asexual
morph of A. collinsii was reported as Cladosporium sp.
(Sivanesan 1984) and it was assigned as Fusicladium sensu
lato by Braun et al. (2003). Dibotryon had been treated as a
synonym of Apiosporina (von Arx and Müller 1975; Crous
et al. 2007b). Although the black knots on branches of Prunis cultis formed by Dibotryon morbosum can readily be
distinguished from the black spots on leaves formed by Apiosporina, their congeneric relationship was confirmed by
molecular data (Winton et al. 2007). Sequence data are available for Apiosporina collinsii and Apiosporina morbosa.
Apiosporina collinsii (Schwein.) Höhn., Sber. Akad. Wiss.
Wien, Math.-naturw. Kl., Abt. 1 119: 439 (1910).
≡ Sphaeria collinsii Schwein., Trans. Am. phil. Soc.,
New Series 4(2): 211 (1832) [1834].
Index Fungorum number: IF 122234; Facesoffungi number: FoF 06355; Fig. 92
Parasitic on leaves, stems petioles, buds and flowers
appearing as black dots like areas. Sexual morph: Ascomata
150–200 high× 150–200 diam. μm ( x̄ = 194 × 195 μm, n
= 10) small, gregarious, superficial, globose to subglobose,
developing on a dark hyphal mass, easily removed from the
substrate, small papillate and ostiolate. Hypha 4 μm wide,
pale brown to dark brown, branched, septate. Peridium thinwalled, composed of three layers, dark brown to hyaline cells
of textura angularis. Hamathecium comprising 4 μm wide,
rare, septate, cellular pseudoparaphyses. Asci 7–9 ×55–65
μm ( x̄ = 7.8 × 62.2 μm, n = 10), 8-spored, bitunicate to
narrowly obclavate, cylindrical, with a short, thick, furcate
or knob-like pedicel or pedicel lacking. Ascospores 4–6 ×
12–15 μm ( x̄ = 5.1 ×13.4 μm, n = 10), uni-seriate and partially overlapping, ovoid to narrowly ovoid, hyaline to pale
brown, apiosporous, 1-septate near the lower end, barely
constricted at the septum, verrucose. Asexual morph: Cladosporium sp. (Sivanesan 1984).
Material examined: Canada, Saskatchwan, on leaves of
Cotoneaster sp. (K(M):158702, ex herb. Broome).
Atopospora Petr., Annls mycol. 23(1/2): 100 (1925).
Index Fungorum number: IF 450; Facesoffungi number:
FoF 06356; – 4 morphological species (Species Fungorum
2020), molecular data unavailable.
Fungal Diversity (2020) 105:17–318
219
Fig. 92 Apiosporina collinsii (K(M):158702). a, b Herbarium specimens. c Ascomata on host substrate. d Cross section of ascoma. e
Peridium. f Wall of ascoma composed darkly pigmented cells. g
Hyphal mass. h Arrangement of hamathecium. i Pseudoparaphyses.
j–l Asci. m, n Ascospores. Scale bars: d = 100 μm, e, g, h = 20 μm,
f, i–l = 10 μm, m, n = 5 μm
Type species: Atopospora betulina (Fr.) Petr., Annls
mycol. 23(1/2): 101 (1925).
≡ Xyloma betulinum Fr., Observ. mycol. (Havniae) 1: 198
(1815).
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220
Notes: The multiloculate ascostromata, fusoid asci and
lightly pigmented, bi-seriate, apiospores are the main characteristics that allows its placement in the family Venturiaceae (Müller and von Arx 1962).
Caproventuria U. Braun, Monogr. Cercosporella, Ramularia Allied Genera (Phytopath. Hyphom.) 2: 396 (1998).
Index Fungorum number: IF 27882; Facesoffungi number: FoF 06357; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Caproventuria hanliniana (U. Braun &
Feiler) U. Braun, Annls mycol. 23(1/2): 101 (1925).
≡ Capronia hanliniana U. Braun & Feiler, Microbiol.
Res. 150(1): 90 (1995).
Notes: Species of Caproventuria are characterized by
astromatic, papillate, ostiolate ascomata with aseptate dark
brown setae and 8-spored bitunicate asci with yellowish
olivaceous monostichous or distichous and ellipsoidovoid
ascospores (Zhang et al. 2011). Crous et al. (2007b) and
Seifert et al. (2011) suggested Pseudocladosporium as a
synonym of Fusicladium. However, Zhang et al. (2011)
concluded Pseudocladosporium to be asexual morph of
Caproventuria. We accept Pseudocladosporium as asexual morph of Caproventuria, but this needs confirmation
by phylogenetic analyses. In our phylogenetic analysis,
Caproventuria hanliniana (CBS 588.93, type material) was
observed to be phylogenetically distant from Venturiaceae
(data not shown), while the strain ATCC 96019 is related
to Tyrannosorus in Venturiaceae. We tentatively place this
genus in Venturiaceae until more collections and sequence
data are available to resolve its taxonomic placement.
Coleroa Rabenh., Klotzschii Herb. Viv. Mycol., Edn 1: no.
1456 (in sched.) (1850).
Index Fungorum number: IF 1174; Facesoffungi number:
FoF 06358; – 30 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Coleroa chaetomium (Kunze) Rabenh.,
Rabenh. Krypt.-Fl., Edn 2 (Leipzig) 1: 198 (1850).
≡ Dothidea chaetomium Kunze, Syst. mycol. (Lundae)
2(2): 563 (1823).
Notes: The scattered, setose ascomata, deliquescing pseudoparaphyses, fusoid to obclavate asci and 1-septate, constricted ascospores are similar characters to other genera of
Venturiaceae. However, the 1-septate ascospores with a pale
brown to greenish tint, separates this genus (Zhang et al.
2011).
Cylindrosympodium W.B. Kendr. & R.F. Castañeda, in
Castañeda Ruiz & Kendrick, Univ. Waterloo Biol. Ser. 32:
9 (1990).
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Index Fungorum number: IF11257; Facesoffungi number:
FoF 09215; – 12 morphological species (Species Fungorum
2020), 2 species with molecular data.
Type species: Cylindrosympodium variabile (de Hoog)
W.B. Kendr. & R.F. Castañeda, in Castañeda Ruiz & Kendrick, Univ. Waterloo Biol. Ser. 32: 10 (1990).
≡ Subulispora variabilis de Hoog, Stud. Mycol. 26: 56
(1985).
Notes: Species of Cylindrosympodium are characterized
by cylindrical, hyaline or sub hyaline, erect conidiophores
with cylindrical, septate hyaline conidia. Conidiogenous
cells are conspicuous, flat-topped denticles and extended
sympodially reduced from conidiophores (Paulus et al.
2003). In our analyses (Fig. 90), we tentatively accept this
genus in Venturiaceae based on the sequence data of type
species. However, the clade contains Cylindrosympodium
is unstable within Venturiales, thus more sequence data are
needed to clarify its placement within Venturiales.
Dimeriella Speg., Revta Mus. La Plata 15(2): 12 (1908).
Index Fungorum number: IF 1575; Facesoffungi number:
FoF 06359; – 25 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Dimeriella hirtula Speg., Revta Mus. La
Plata 15(2): 12 (1908).
Notes: Dimeriella is a plant pathogenic genus characterized by brown ascospores and brown, superficial mycelium.
This mycelium forms pale-coloured hyphal clumps and penetrate via hyaline hyphae through sub-stomatal chamber to
the mesophyll (Farr 1965).
Dimerosporiopsis Henn., Hedwigia 40(Beibl.): (173)
(1901).
Index Fungorum number: IF 1588; Facesoffungi number:
FoF 08832; – 2 morphological species (Species Fungorum
2020; Crous et al. 2019b), 1 species with molecular data.
Type species: Dimerosporiopsis engleriana (Henn.)
Henn., Hedwigia 40(Beibl.): (173) (1901).
≡ Dimerosporium englerianum Henn., Bot. Jb. 17: 31
(1893).
Notes: Dimerosporiopsis is similar to other genera such
as Proventuria in having superficial pseudothecia with pseudoparaphyses, cylindrical asci and pale olivaceous 1-septate
ascospores. However, this genus differs from other genera
in having pluri-guttulate ascospores (Crous et al. 2019b).
Magnohelicospora R.F. Castañeda, Hern.-Restr., Gené &
Guarro, Mycotaxon 121: 172 (2013).
Index Fungorum number: IF 563847; Facesoffungi number: FoF 06360; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.
Fungal Diversity (2020) 105:17–318
Type species: Magnohelicospora iberica R.F. Castañeda,
Hern.-Restr., Gené & Guarro, in Castañeda-Ruiz et al.,
Mycotaxon 121: 172 (2013) [2012].
Notes: Members of Magnohelicospora can be distinguished by polyblastic, integrated, sympodial conidiogenous
cells and solitary, doliiform or conical, multi-euseptate,
brown conidia, tightly coiled in three planes (CastañedaRuíz et al. 2013).
Metacoleroa Petr., Annls mycol. 25(3/4): 332 (1927).
Index Fungorum number: IF 3139; Facesoffungi number:
FoF 06361; – 1 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Metacoleroa dickiei (Berk. & Broome)
Petr. [as ‘dieckiei’], Annls mycol. 25(3/4): 332 (1927) [as
‘dieckiei’], Annls mycol. 25(3/4): 332 (1927).
≡ Sphaeria dickiei Berk. & Broome, Ann. Mag. nat. Hist.,
Ser. 2 9: 317 (1852).
Notes: Metacoleroa is a monotypic genus characterized
by ascomata produced on well-defined subiculum, not cupulate when dry and olivaceous brown ascospores at maturity
(Zhang et al. 2011). No mature ascomata were found on the
specimen examined by Crous et al. (2007b).
Neocoleroa Petr., Hedwigia 74: 38 (1934).
Index Fungorum number: IF 3446; Facesoffungi number:
FoF 06362; – 3 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Neocoleroa sibirica Petr., Hedwigia 74:
38 (1934).
Notes: Neocoleroa is characterized by lobed to dichotomously branched, blunt-tipped setae and persistent pseudoparaphyses. Recent authors have used morphology to
accomodate Neocoleroa in Pseudoperisporiaceae (e.g. Barr
1997, Kirk et al. 2008), although it was noted that these taxa
are similar to Venturiaceae (Johnston and Park 2016).
Protoventuria Berl. & Sacc., Atti Soc. Veneto-Trent. Sci.
Nat. 10(1): 174 (1887).
Index Fungorum number: IF 4399; Facesoffungi number:
FoF 06363; – 19 morphological species (Species Fungorum
2020), 3 species with molecular data.
Type species: Proventuria rosae De Not., G. bot. ital.
1(1): 332 (1844).
Notes: Proventuria is characterized by superficial ascomata developing on dead leaf tissues, sparse aerial mycelium, a subcuticular hypostroma and penetrating, intramaterical hyphae (Carris and Poole 1993).
Pseudoanungitea Crous, in Crous et al., Fungal Systematics
and Evolution 1: 199 (2018).
221
Index Fungorum number: IF 824789; Facesoffungi number: FoF 08833; – 2 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Pseudoanungitea syzygii (Crous, W.B.
Kendr. & M.J. Wingf.) Crous, in Crous, et al., Fungal Systematics and Evolution 1: 199 (2018).
≡ Anungitea syzygii Crous, W.B. Kendr. & M.J. Wingf.,
Can. J. Bot. 73(2): 225 (1995).
Notes: Species of Pseudoanungitea are similar in characters to Anungitea in having dark, solitary conidiophores with
flattened denticle heads and 1-septate, subhyaline conidia
forming as a cylindrical chain with apical and basal scars
(Crous et al. 2018b).
Pseudoparodiella F. Stevens, Illinois Biol. Monogr.
(Urbana) 11(2): 166 (1927).
Index Fungorum number: IF 4467; Facesoffungi number:
FoF 06364; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Pseudoparodiella vernoniae F. Stevens,
Illinois Biol. Monogr. (Urbana) 11(no. 2): 166 (1927).
Notes: Pseudoparodiella is similar with other genera in
having smaller ascomata with or without pseudoparaphyses,
obclavate asci and 1-septate, olivaceous brown ascospores
(Zhang et al. 2011). However, this genus differs from the
similar genus Acantharia in having mature pale brown
ascospores (Zhang et al. 2011). Ascomata consisting of rare
pseudoparaphyses and obclavate asci 1-septate, olivaceous
brown ascospores showing a close affinity with Venturiaceae
(Zhang et al. 2011).
Sympodiella W.B. Kendr., Trans. Br. mycol. Soc. 41(4): 519
(1958).
Index Fungorum number: IF10151; Facesoffungi number:
FoF 09216; – 7 morphological species (Species Fungorum
2020), 4 species with molecular data.
Type species: Sympodiella acicola W.B. Kendr., Trans.
Br. mycol. Soc. 41(4): 519 (1958).
Notes: Sympodiella is characterized by superficial networks of darkly pigmented, septate, anastomosing hyphae.
Conidiophores are simple, septate, and darkly pigmented
except at the extreme apex and conidia are cylindrical,
smooth-walled and hyaline (Kendrick 1958).
Tyrannosorus Unter. & Malloch, Mycol. Res. 99(8): 910
(1995).
Index Fungorum number: IF 11631; Facesoffungi number: FoF 06365; – 1 morphological species (Species Fungorum 2020), 1 species with molecular data.
Type species: Tyrannosorus pinicola (Petrini & P.J.
Fisher) Unter. & Malloch, Mycol. Res. 99(8): 910 (1995).
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Fungal Diversity (2020) 105:17–318
Zeloasperisporium wrightiae MFLUCC 15-0214
100/1.0
100/1.0
Zeloasperisporium wrightiae MFLUCC 15-0225
Zeloasperisporium eucalyptorum CBS 124809
99/1.0
Zeloasperisporium hyphopodioides CBS 218.95
100/1.0
100/1.0
100/1.0
Zeloasperisporium cliviae CPC 25145
Zeloasperisporium ficusicola MFLUCC 15-0221
Zeloasperisporiales
Fig. 93 Phylogram generated
from maximum likelihood
analysis (RAxML) of genera in
Zeloasperisporiales based on
LSU and SSU sequence data.
Maximum likelihood bootstrap
values equal or above 70%,
Bayesian posterior probabilities
equal or above 0.90 (MLBS/
PP) are given at the nodes. An
original isolate number is noted
after the species name. The tree
is rooted to Venturia inaequalis
(CBS 594.70, CBS 815.69). The
ex-type strains are indicated in
bold. Hyphen (-) represents support values below 70% MLBS
and 0.90 PP
Zeloasperisporium ficusicola MFLUCC 15-0222
100/1.0
Zeloasperisporium siamense IFRDCC 2194
100/1.0
100/1.0
Natipusilla limonensis L AF286 1A
Natipusilla decorospora L A236 1A
Natipusilla naponensis L AF217 1A
Natipisillales
100/1.0
Trichodelitschia munkii Kruys 201 UPS
Trichodelitschia bisporula CBS 262.69
100/1.0
Phaeotrichales
100/1.0
Phaeotrichum benjaminii CBS 541.72
100/1.0
0.02
Venturia inaequalis CBS 594.70
Venturia inaequalis CBS 815.69
≡ Capronia pinicola Petrini & P.J. Fisher, in Müller et al.,
Trans. Br. mycol. Soc. 88(1): 68 (1987).
Notes: Tyrannosorus was introduced as a monotypic
genus. Helicodendron is considered as the asexual morph
(Untereiner et al. 1995). This genus differs from other genera in having a helicosporous asexual morph and longitudinally ridged ascospores with multiple germ slits (Zhang
et al. 2011).
Economic and ecological significance
Most species in Venturiaceae are saprobic or parasitic. They
are capable of growing in living or dead leaves, stalks, or
twigs of various higher plants, rarely on Sphagnum or dung.
Some are important plant pathogens which have a cosmopolitan distribution causing diseases of perennial plants, such
as apple scab (Venturia inaequalis), pear scab (V. pyrina),
poplar shoot blight (V. populina), soybean black leaf blight
(Arkoola nigra), and the widespread black knot disease of
cherry (Dibotryon morbosum) (von Arx and Müller 1975;
Talbot 1971). Apiospora mabosa is another important pathogenic species causing black knot disease of Prunus, mainly
affecting cultivated plum, prune and cherry and to a lesser
extent wild plum and cherry (Snover and Arneson 2002).
Zeloasperisporiales Hongsanan & K.D. Hyde.
Index Fungorum number: IF 551336; Facesoffungi number: FoF 551336.
13
Venturiales (Outgroup)
The order was established by Hongsanan et al. (2015b) to
accommodate Zeloasperisporiaceae. Previous phylogenetic
analyses using LSU and SSU sequence data indicated that
this order is a distinct lineage from Microthyriales within
Dothideomycetes, and formed as a sister clade to freshwater order Natipusillales. Morphologically, this order is very
different from Natipusillales. Therefore, the order status of
Zeloasperisporiales is strongly supported by both phylogeny
and morphology (Figs. 93 and 94). The divergence time for
Zeloasperisporiales is estimated as 180 MYA (stem age,
Hongsanan et al. 2020).
Accepted families: Zeloasperisporiaceae.
Zeloasperisporiaceae Crous, Persoonia 34: 215 (2015).
Index Fungorum number: IF 812487; Facesoffungi numbers: FoF 08834, 8 species.
Epiphytic on the surface of living and dead fallen leaves,
appearing as small black dots, or in air. Superficial hyphae
absent. Sexual morph: Thyriothecia superficial, solitary,
circular, flattened, brown to dark brown, poorly developed at the base, ostiole lacking. Upper wall comprises
ellipsoid angular cells, radiating from the center to the
outer rim. Hamathecium pseudoparaphyses not observed.
Asci 8-spored, bitunicate, fissitunicate, globose to ovoid
or clavate, apedicellate, with an apical ocular chamber.
Ascospores 2–3-seriate, obovoid to clavate, hyaline, 1-septate, slightly constricted at the septum, wider in upper cell,
Fungal Diversity (2020) 105:17–318
223
Fig. 94 Zeloasperisporium wrightiae (MFLU 15-1308, holotype).
a Thyriothecia on host surface. b Section through thyriothecium
and close-up of margin. c Thyriothecium in squash mount. d Upper
wall of thyriothecium in squash mount. e Ascus. f Ascus in Melzer’s
reagent. g Ascospores. h Crowded conidia on conidiogenous cells.
i Conidia on conidiogenous cell with sympodial proliferation. j
Conidia. k Micronematous conidiogenous cells. Scale bars: b, c = 50
μm, d–f, h, i, k = 10 μm, g, j =5 μm
smooth-walled or verrucose, with or without thin mucilaginous sheath (Wu et al. 2011b; Crous et al. 2015b). Asexual
morph: Hyphomycetous. Hyphae branched, septate, and
swollen by constrictions at the septa, pale brown to brown,
occasionally hyaline, smooth-walled, micronematous conidiogenous present or absent. Conidiophores reduced to conidiogenous cells, arising as lateral hyphal branches, cylindrical
to subcylindrical, straight or slightly curved, unbranched,
slightly tapering towards the apex, brown, slightly thickwalled. Conidial proliferation sympodial, with one to several
conidiogenous loci, mostly crowded at the apex, protuberant of thickened, refractive, conidial scars. Conidia fusiform
to obclavate or cylindrical, straight to curved, pale brown
to brown, 1–3-septate, distinctly or slightly constricted at
the septum, tapered towards the apex, towards a protruding
scar, somewhat thickened and darkened, refractive, smoothwalled or verrucose (Hongsanan et al. 2015b).
Type: Zeloasperisporium R.F. Castañeda.
Notes: Zeloasperisporiaceae was established by Crous
et al. (2015b) to accommodate Neomicrothyrium (sexual
13
224
genus Microthyrium-like) and Zeloasperisporium (asexual genus). This family was poorly resolved in molecular
analysis as it is related to freshwater taxa in Natipusillales,
and morphology of Zeloasperisporiaceae is very different
from Natipusillales (Ferrer et al. 2011; Wu et al. 2011b).
Hongsanan et al. (2015b) linked morphs and synonymized
Neomicrothyrium under Zeloasperisporium. Moreover, they
introduced the new order, Zeloasperisporiales to accommodate Zeloasperisporiaceae based on its distinct clade within
Dothideomycetes.
Zeloasperisporium R.F. Castañeda, Mycotaxon 60: 284
(1996).
Index Fungorum number: IF 27808; Facesoffungi number: FoF 08835; 8 mophological species (Species Fungorum
2020), 8 species with molecular data
Type species: Zeloasperisporium hyphopodioides R.F.
Castañeda.
Notes: Zeloasperisporium was introduced as genus incertae sedis by Castañeda-Ruíz et al. (1996), with the type
species Z. hyphopodioides isolated from air in Cuba. Crous
et al. (2007b) placed this genus in Venturiaceae based on
phylogenetic placement basal to Venturiaceae. Two species
from leaves of Eucalyptus and Clivia were introduced based
on asexual characters and molecular data (Cheewangkoon
et al. 2009; Crous et al. 2015b). Hongsanan et al. (2015b)
linked the sexual morph genus (Neomicrothyrium) with
Zeloasperisporium and added two new species to this genus.
Phylogenetic analyses strongly support species of Zeloasperisporium as a distinct family and order (Hongsanan et al.
2015b).
Zeloasperisporium wrightiae Hongsanan & K.D. Hyde, in
Hongsanan et al., Cryptog. Mycol. 36(3): 310 (2015).
Index Fungorum number: IF 551313; Facesoffungi number: FoF 551313, Fig. 94
Description: see Hongsanan et al. (2015b).
Material examined: Thailand, Chiang Rai, Tasud, Mae
Fah Luang University, on leaves of Wrightiar eligiosa Benth.
(Apocynaceae), 15 January 2015, S. Hongsanan MOK01
(MFLU 15-1308, holotype).
Economic and ecological significance
Species of Zeloasperisporiaceae are usually found on living and dead fallen leaves, and rarely sampled in air. Species found on leaves sometimes produce organs to attach the
surface of host plants at the ascomatal margin. However, it
does not reveal any penetration of the host (Hongsanan et al.
2015b). The appearance of Zeloasperisporiaceae on leaves
as a few small black dots is not a serious problem.
Dothideomycetes, families incertae sedis
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Fungal Diversity (2020) 105:17–318
In this section we include families that cannot be accommodated in orders of Dothideomycetes, mainly because they
lack molecular data. However, since they have a suite of
characters that exclude them from other families, they are
accepted as distinct families.
Alinaceae Boonmee & K.D. Hyde, in Boonmee et al.,
Mycosphere 8(10): 1699 (2017).
Index Fungorum number: IF 553826; Facesoffungi numbers: FoF 03661, 1 species.
Parasitic or epiphytic on living leaves. Sexual morph:
Ascomata superficial, developed on hyphae, globose to
subglobose, brown, covered by brown hyphae. Peridium
comprises brown cells of textura angularis, lacking pseudoparaphyses. Asci 8-spored, bitunicate, subglobose to clavate,
sessile, thick-walled. Ascospores overlapping, ovoid to obovoid, apex wider, narrower towards the lower and rounded
end, brown, 1-septate. Asexual morph: Hyphomycetous,
Septoidium sp., Shivomyces sp. (adapted from Boonmee
et al. 2017).
Type: Alina Racib.
Notes: Boonmee et al. (2017) re-examined the ex-isotype
of Alina jasmini, and concluded that the morphology of
Alina has a set of characters that can differentiate it from
other families in Dothideomycetes. Therefore, they introduced Alinaceae to accommodate the monotypic genus,
Alina (Boonmee et al. 2017). Molecular data is unavailable
to confirm its taxonomic placement.
Alina Racib., Bull. int. Acad. Sci. Lett. Cracovie, Cl. sci.
math. nat. Sér. B, sci. nat. 3: 374 (1909).
Index Fungorum number: IF 125; Facesoffungi number:
FoF 07866; 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Alina jasmini Racib.
Notes: Alina was introduced from living leaves of Jasminum sp. Alina is similar to Tretospora (the asexual morph
of Balladynopsis) based on some hyphomycetous conidial
structures found on the ex-isotype of Alina jasmini (Sivanesan 1981). By re-examining the same specimen, Boonmee
et al. (2017) reported that they were unable to find some
structures e.g. setae, hyphopodia-like structures or even
conidia as previously shown in Raciborski (1909) and
Sivanesan (1981). They concluded that Alina has unique
characters and could not be placed in any families in Dothideomycetes (Boonmee et al. 2017). Thus, they established
Alinaceae to accommodate this genus.
Alina jasmini Racib., Bull. int. Acad. Sci. Lett. Cracovie,
Cl. sci. math. nat. Sér. B, sci. nat. 3: 375 (1909).
Index Fungorum number: IF 205013; Facesoffungi number: FoF 03662; Fig. 95
Description: see Boonmee et al. (2017).
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225
Fig. 95 Alina jasmini (IMI 214481, ex-isotype). a Squash mount of ascomata. b Mycelia. c–e Asci. e–i Ascospores. Scale bars: a = 20 µm, b =
50 µm, c–e = 10 µm
Material examined: Indonesia, Java, Djasinga, on living
leaves of Jasminum sp. (Oleaceae), 1900, Raciborski M.
(K(M) 177969 = IMI 214481, ex-isotype).
Economic and ecological significance
Species in Alinaceae are parasitic or epiphytic on living
leaves, however they do not cause serious problem on plants.
Argynnaceae Shearer & J.L. Crane, Trans. Br. mycol. Soc.
75(2): 193 (1980).
Index Fungorum: IF 80470; Facesoffungi numbers: FoF
06386, 3 species.
Saprobic on plant stems or wood and in freshwater and
terrestrial habitats. Sexual morph: Ascomata apothecial or
perithecial, solitary, superficial to slightly immersed, discoid, dark brown to black, coriaceous, lacking ostioles, lacking a stipe. Excipulum thick, ectal excipulum composed of
brown to dark brown tissues mixed with host cells, medullary excipulum composed of hyaline cells of textura porrecta. Hymenium composed of wide, septate, branching,
hyaline, cellular pseudoparaphyses. Epithecium lacking. Asci
8-spored, bitunicate, fissitunicate, clavate, globose to subglobose, pedicellate, thick-walled, lacking an ocular chamber, exposed or not. Ascospores 2–3-seriate, or multi-seriate,
papilionaceous to broadly reniform, light brown, orange to
light orange-brown, 1-septate, smooth-walled, 2- to multiguttulate, with a dark band at the septum. Asexual morph:
Undetermined.
Type: Argynna Morgan.
Notes: Argynnaceae was introduced to accommodate two
genera of Argynna and Lepidopterella (Hyde et al. 2013;
Kirk et al. 2013; Wijayawardene et al. 2018). The species
in Argynnaceae are saprobes and occurring on stems, twigs
and woody debris from dicotyledonous and monocotyledonous plants. Additional collections and molecular study are
required to clarify the phylogeny of this family.
Argynna Morgan, J. Cincinnati Soc. Nat. Hist. 18: 41
(1895).
Index Fungorum number: IF 306; Facesoffungi numbers:
FoF 06387; 1 morphological species (Species Fungorum
2020), molecular data unavailable.
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Fig. 96 Argynna polyhedron (BPI 618534). a Material label. b
Ascoma on the host substrate. c Ascoma. d Section of ascoma and
peridium. e Hymenium. f Pseudoparaphyses. g–j Immature and
mature asci. k–p Papilionaceous ascospores. Scale bars: c = 1000
µm, d = 200 µm, e = 100 µm, f–j = 20 µm, k–p = 5 µm
Type species: Argynna polyhedron (Schwein.) Morgan.
Notes: Argynna comprises a single species, Argynna
polyhedron (Morgan 1895). The genus is characterized by
solitary, superficial apothecia and 1-septate papilionaceous,
light brown, smooth-walled ascospores with a dark band at
the septum. Argynna polyhedron is a rare species and lacks
molecular data to allow its taxonomic placement.
Material examined: USA, Missouri, Big Spring, on
stalks of Zea mays, 15 April 1939, G.W. Martin (3922),
BPI618534.
Argynna polyhedron (Schwein.) Morgan [as ‘polyedron’],
J. Cincinnati Soc. Nat. Hist. 18: 41 (1895).
≡ Physarum polyhedron Schwein. [as ‘polyaedron’],
Trans. Am. phil. Soc., New Series 4(2): 257 (1832) [1834].
Index Fungorum: IF 140004; Facesoffungi number: FoF
06388; Fig. 96
Description: see Hyde et al. (2013).
13
Other genera included
Lepidopterella Shearer & J.L. Crane, Trans. Br. mycol. Soc.
75(2): 194 (1980).
Index Fungorum number: IF 2746; Facesoffungi numbers:
FoF 06389; – 2 morphological species (Species Fungorum
2020), 2 species with molecular data.
Type species: Lepidopterella palustris Shearer & J.L.
Crane, Trans. Br. mycol. Soc. 75(2): 194 (1980).
Notes: Lepidopterella was introduced by Shearer and
Crane (1980) and placed in Argynnaceae. The asexual
morph has not been reported. Species are saprobic fungi in
aquatic habitats (Raja and Shearer 2008; Shearer et al. 2009;
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227
Fig. 97 Ascoporia lateritia (BPI 1109903). a Material, habit on
decorticated wood and appearance of ascostroma on wood substrate.
b Section of ascostroma showing arrangement of locules. c Hamathe-
cium pseudoparaphyses. d, e Asci changed to pinkish in 3% KOH.
f–h Ascospores. Scale bars: a = 5 mm, b = 200 μm, c = 5 μm, d, e =
50 μm, f–h = 20 μm
Hyde et al. 2013). Sequence data are available for the type
species, Lepidopterella palustris, and L. tangerina. Phylogenetic analysis indicated that the type species represents a
basal taxon to Mytilinidiales in Dothideomycetes with moderate support (Shearer et al. 2009). Lepidopterella shares
butterfly-shaped ascospores with Argynna, but differs in having globose to subglobose ascomata, and papilionaceous to
broadly reniform, brown to orange-brown ascospores (Hyde
et al. 2013).
Saprobic on decorticated wood in terrestrial habitats. Sexual morph: Ascostromata relatively large, superficial, circular or cupulate, slightly raised, pedicellate, solitary, exterior
surrounded by brown mycelium, orange to red-brown at the
margin or edge, dark brown to black in the centre, cells of
ascostromata comprising a maze-like arrangement of mixed,
brown-walled cells of textura angularis to subglobosa or
epidermoidea, multi-loculate. Locules in a single layer at the
periphery of the ascostromata, subcylindric-elongate, obpyriform to ovoid, crowded, opening by small, narrow, vertical ostiole. Hamathecium comprising numerous filiform,
branched, septate, cellular pseudoparaphyses, embedded in
a gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate,
narrowly cylindric-clavate, pedicellate, apically rounded
with a small ocular chamber. Ascospores 2-seriate, fusiform,
hyaline to yellowish when immature, olive-brown to dark
brown when mature, 1-septate, constricted at the septum.
Asexual morph: Coelomycetous.
Type: Ascoporia Samuels & A.I. Romero.
Economic and ecological significance
Members of Argynnaceae are saprobic on plant stems or
wood and in freshwater and terrestrial habitats and play a
role in recycling organic matter.
Ascoporiaceae Kutorga & D. Hawksw., Syst. Ascom. 15(1–
2): 25 (1997).
Index Fungorum number: IF 81923, Facesoffungi number: FoF 07867, 1 species.
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228
Notes: Ascoporiaceae is presently treated in Dothideomycetes as family incertae sedis (Wijayawardene et al. 2018).
The family is monotypic with a single species Ascoporia
lateritia (Hyde et al. 2013). Ascoporiaceae lacks molecular
data to establish phylogenetic placement.
Ascoporia Samuels & A.I. Romero, Bolm Mus. paraense
‘Emílio Goeldi’, sér. bot. 7(2): 264 (1993) [1991].
Index Fungorum number: IF 26458, Facesoffungi number: FoF 07945; − 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Ascoporia lateritia Samuels & A.I.
Romero.
Notes: Ascoporia was introduced to accommodate a single species A. lateritia that occurs on terrestrial decorticated
wood (Samuels and Romero 1991).
Ascoporia lateritia Samuels & A.I. Romero, Bolm Mus.
paraense ‘Emílio Goeldi’, sér. bot. 7(2): 264 (1993) [1991].
Index Fungorum number: IF 360813, Facesoffungi number: FoF 07946; Fig. 97
Description: see Hyde et al. (2013).
Material examined: Brazil, Belem, Iha do Combu, Estacao Experimental Combu, Para. 01d30’s, 48d27’w, on
decorticated wood, January 1989, G.J. Samuels; K.F. Rodrigues, 6207 (BPI1109903, isotype of Ascoporia lateritia).
Economic and ecological significance
Ascoporiaceae is a small family only known from Brazil.
Species in this family are saprobes on decorticated wood
(Samuels and Romero 1991).
Aulographaceae Luttrell ex P.M. Kirk et al., in Kirk et al.,
Ainsworth & Bisby’s Dictionary of the Fungi, Edn 9 (Wallingford): ix (2001).
Index Fungorum number: IF 80503; Facesoffungi numbers: FoF 06201, 50 species.
Saprobic on fallen leaves. Mycelium sparse, not radially
arranged. Sexual morph: Ascomata forming as small black
dots on host surfaces, solitary or clustered, nearly superficial, easily removed or breaking from the substrate, globose
when immature, becoming sub-linear or v-shaped at maturity, rounded, carbonaceous, dark brown to black, absence of
ostiole at the maturity, with slit-like opening. Hamathecium
comprising septate, anastomosing and branched, trabeculate
pseudoparaphyses, embedded in a gelatinous matrix with
asci. Asci 8-spored, bitunicate, fissitunicate, oval to ellipsoid, apex thickened, short pedicellate or lacking, and with
a wide indistinct ocular chamber. Ascospores overlapping
2–3-seriate, oblong to ovate, narrow at both ends, hyaline,
1-septate at the center, the upper cell often broader than
lower cell. Asexual morph: Undertermined.
Type: Aulographum Lib.
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Notes: Aulographaceae was established by Libert (1834)
with the type genus Aulographum. They are characterized
by superficial elongate thyriothecia, opening by longitudinal or slit-like, X- or Y-shaped dehiscence, with walls composed of dark cells that are pale at the margin of the scutellum, and lack of appressoria (Hyde et al. 2013; Hongsanan
et al. 2014b). Hongsanan et al. (2014b) provided notes on
this family and excluded the family from Asterinales based
on morphology and phylogeny. Only a single species of
Aulographum has sequence data and its placement is uncertain. Thus, more collections and sequence data are needed
to clarify the taxanomic placement of this family within
Dothideomycetes. Crous et al. (2019b) provided sequence
data of Lembosina species and excluded this genus from
Aulographaceae. Aulographaceae comprises three genera Aulographum, Echidnodes, and Lembosiella (Wijayawardana et al. 2018; Crous et al. 2019b).
Aulographum Lib., Pl. crypt. Arduenna, fasc. (Liège) 3 (nos
201–300): no. 272 (1834)
Index Fungorum number: IF 461; Facesoffungi number:
FoF 06202; 18 morphological species (Species Fungorum
2020), 1 species with molecular data
Type species: Aulographum hederae Lib.
Notes: Aulographum is similar to Aulographina (incertae
sedis genus in Capnodiales) and Lembosia (Asterinaceae)
and these genera may be synonyms. However, only a single
sequence is available for this genus, and its phylogenetic
placement is uncertain. Molecular data of fresh specimens
are needed to resolve its placement (Hyde et al. 2013). Several of the species however, were transferred to other families, such as Asterinaceae, Elsinoaceae, Leptopeltidaceae,
or Lophiostomataceae (Hongsanan et al. 2014b).
Aulographum hederae Lib., Pl. crypt. Arduenna, fasc.
(Liège) 3: no. 272 (1834).
Index Fungorum number: IF 161393; Facesoffungi number: FoF 07947; Fig. 98
Saprobic on fallen leaves of Hedera helix. Mycelium
sparse, not radially arranged. Sexual morph: Ascomata
52-76 μm diam, 35–42 μm high ( x̄ = 72 × 40 μm, n = 5),
forming as small black dots on host surfaces, solitary or
clustered, nearly superficial, easily removed or breaking
from the substrate, globose when immature, becoming sublinear or v-shaped at maturity, rounded, carbonaceous, dark
brown to black, with slit-like opening, ostiolar canal filled
with hyaline cells, with cells comprising parallel, radiating lines from center of the ostiole to outer cells, becoming pale brown at the margin. Hamathecium comprising
1 µm wide, septate, anastomosing, branched, trabeculate
pseudoparaphyses, embedded in a gelatinous matrix with
asci. Asci 24–30 × 10–15 μm ( x̄ = 28–12 μm, n = 10)
8-spored, bitunicate, fissitunicate, oval to ellipsoid, apex
Fungal Diversity (2020) 105:17–318
Fig. 98 Aulographum hederae (MFLUCC 12-0397). a Habit, ascomata on host substrate. b Ascomata with slit-like opening. c Section through ascoma. d Walled cells of ascoma. e, f Hamathecium
in Melzer’s reagent and Cotton blue reagent respectively. g–i Asci. j
229
Asci in Melzer’s reagent. k Asci in Cotton blue reagent. l Ascospore
germinated. m–o Ascospore in water, Melzer’s reagent and Cotton
blue reagent respectively. Scale Bars: b = 50 µm, c, g–k = 20 µm, d–f
= 10 µm, l–o = 5 µm
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230
thickened, short pedicellate or lacking, and with a wide,
indistinct ocular chamber. Ascospores 11–14 × 3–5 μm
( x̄ = 12–3 μm, n = 10) overlapping 2–3-seriate, oblong
to ovate, narrow at both ends, hyaline, 1-septate at the
center, constricted at the septum, the upper cell often
broader than lower cell, smooth-walled. Asexual morph:
Undertermined.
Material examined: Germany, Frankfurt, on fallen leaves
of Hedera helix, in 2012, Meike Piepenbring MFLU, culture
MFLUCC 12-0397 (MFU) = CPC21373 (CBS).
Notes: Aulographum hederae is the type species of
Aulographaceae. It is characterized by longitudinally splitting ascomata and a perithecial wall appressed by mycelium with bright coloured margin (Hongsanan et al. 2014b).
Sequence data of this specimen was provided by Hongsanan
et al. (2014b), but without a plate and description. Therefore, we provide a photoplate of the same specimen with
descriptions.
Other Genera included
Echidnodes Theiss. & Syd., Annls mycol. 15(6): 422 (1918).
Index Fungorum number: 1732; Facesoffungi number:
FoF 07949; – 29 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Echidnodes liturae (Cooke) Theiss. & Syd.
[as ‘litsurae’], Annls mycol. 15(6): 422 (1918).
≡ Aulographum liturae Cooke [as ‘Ailographium’], Grevillea 12(no. 61): 38 (1883).
Notes: Hongsanan et al. (2014b) placed Echidnodes in
Aulographaceae based on its superficial hyphae, lack of
appressoria and similar morphological characters of its asci
and ascospores. However, molecular data is needed to confirm the placement of Echidnodes in Asterinales.
Lembosiella Sacc., Syll. fung. (Abellini) 9: 1101 (1891).
Index Fungorum number: IF 2726; Facesoffungi number:
FoF 07948; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Lembosiella polyspora (Pat.) Sacc., Syll.
fung. (Abellini) 9: 1101 (1891)
≡ Lembosia polyspora Pat., Bull. Soc. mycol. Fr. 3(2):
125 (1887).
Notes: Wu et al. (2011b) transferred Lembosiella to
Aulographaceae based on the elongate thyriothecia with a
slit-like opening and lack of appressoria on the superficial
hyphae of Lembosiella polyspora (Hongsanan et al. 2014b).
Thyriopsis Theiss. & Syd., Annls mycol. 13(3/4): 369
(1915).
Index Fungorum number: IF 5468; Facesoffungi number:
FoF 07950; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
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Type species: Thyriopsis halepensis (Cooke) Theiss. &
Syd., Annls mycol. 13(3/4): 369 (1915).
≡ Dothidea halepensis Cooke, Grevillea 7(no. 41): 35
(1878).
Notes: Lumbsch and Huhndorf (2010) placed Thyriopsis
in Asterinaceae. However, the thyriothecia of Thyriopsis
differs from Asterinaceae in having elongate thyriothecia,
globose asci, hyaline ascospores without surface mycelia bearing appressoria. While Asterinaceae has circular
thyriothecia, surface mycelia with appressoria and brown
ascospores at maturity (Hongsanan et al. 2014b). Therefore,
Thyriopsis is replaced in Aulugraphaceae (Hongsanan et al.
2014b). However, Thyriopsis is somewhat different from
other genera in this family as its thyrothecia lack a fibrous
dark brown margin and has dark brown elongate peridium
cells. Molecular data of fresh collection is needed to confirm
its placement in this family.
Economic and ecological significance
The members of this family differ from other species in
Asterinales as these species can grow in artificial media
indicating that the growth of the species is not reliant on the
host (Hongsanan et.al 2014b). They are saprobic and play a
role in recycling organic matter.
Balladynaceae Boonmee & K.D. Hyde, in Boonmee et al.,
Mycosphere 8(10): 1701 (2017).
Index Fungorum number: IF 553827; Facesoffungi numbers: FoF 03663, 47 species.
Parasitic on living leaves. Sexual morph: Mycelium network-like, branched, septate, with hyphopodia, with or without setae. Hyphopodia numerous, knob-like, dark brown.
Ascomata superficial, formed on hyphae, with a subiculum,
solitary, scattered, globose to subglobose, with apical pore,
minutely pedicellate at the base, olivaceous brown to dark
brown, with transparent wall, surrounded by mycelium network. Peridium comprises 1–2 layers of olivaceous-brown
cells of textura angularis, lacking pseudoparaphyses.
Asci 8-spored, bitunicate, globose to subglobose, sessile.
Ascospores multi-seriate, ellipsoid, upper cell wider, hyaline
to light brown when immature, fawn to olivaceous-brown or
brown at maturity, 1-septate. Asexual morph: Hyphomycetous. Clasterosporium sp., Tretospora sp. (adapted from
Boonmee et al. 2017).
Type: Balladyna Racib.
Notes: Balladynaceae was established by Boonmee et al.
(2017) to accommodate Balladyna (type), Balladynocallia
and Balladynopsis. Balladynaceae can be distinguished from
other families of Dothideomycetes based on its parasitic lifestyle, hyphae with hyphopodia, ascomata formed on hyphae,
with or without setae, subglobose asci and 1-septate, hyaline
to brown ascospores. No molecular data is available to confirm its phylogenetic placement within Dothideomycetes.
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231
Fig. 99 Balladyna gardenia (BPI 691469, isotype). a, b Herbarium specimen. c Colonies on leaf surface. d Hyphae with hyphopodia. e Setae.
f–h Asci when immature and mature. i–l Ascospores. Scale bars: c = 200 µm, d = 30 µm, e = 100 µm, f–l = 20 µm
Balladyna Racib., Parasit. Alg. Pilze Java’s (Jakarta) 2: 6
(1900).
Index Fungorum number: IF 502; Facesoffungi number:
FoF 07868; 35 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Balladyna gardeniae Racib.
Notes: Seifert et al. (2011) reported the asexual morphs
as Clasterosporium sp. found in Balladyna vanderystii var.
ferulae-foetidae and Tretospora sp. found in Balladyna
negrii. Boonmee et al. (2017) re-examined the isotype
specimen (BPI 691469), but were unable to find asexual
characters. They also established Balladynaceae to accommodate Balladyna, Balladynocallia and Balladynopsis based
on morphology.
Balladyna gardeniae Racib., Parasit. Alg. Pilze Java’s
(Jakarta) 2: 6 (1900).
Index Fungorum number: IF 218036; Facesoffungi number: FoF 03664; Fig. 99
Description: see Boonmee et al. (2017).
Material examined: Indonesia, Java, Bogor, on living
leaves of Gardenia lucida Roxb. (Rubiaceae), M. Raciborski
Nr. 88 (BPI 691469, isotype).
Other genera included
Balladynocallia Bat., in Batista, Silva & Bezerra, Atas Inst.
Micol. Univ. Pernambuco 2: 216 (1965).
Index Fungorum number: IF 505; Facesoffungi number:
FoF 07951; – 3 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Balladynocallia glabra (Hansf.) Bat., in
Batista, Silva & Bezerra, Atas Inst. Micol. Univ. Recife 2:
216 (1965).
≡ Balladynastrum glabrum Hansf., Proc. Linn. Soc. London 157(3): 157 (1946) [1944–45].
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Fig. 100 Cleistosphaera macrostegia (S-F10853, holotype). a, b Herbarium specimen. c Habit on leaf surface. d Sections of ascomata. e Peridium. f Hyphae with hyphopodia. g–i Asci. j–m Ascospores. Scale bars: c = 1000 µm, d, g–i = 100 µm, e = 30 µm, f = 50 µm, j–m = 20 µm
Notes: The genus was introduced to accommodate B.
amazonensis and the type species B. glabra. Boonmee et al.
(2017) re-examined the type specimen of B. glabra, but it
was not in good condition. However, they used descriptions
and illustrations from Sivanesan (1981) to place this Balladynocallia in Balladynaceae.
Type species: Balladynopsis philippinensis Syd. & P.
Syd., Annls mycol. 15(6): 476 (1918) [1917].
≡ Henningsomyces philippinensis Syd. & P. Syd., Philipp.
J. Sci., C, Bot. 9(2): 161 (1914)
Notes: Descriptions and illustrations can be seen in Boonmee et al. (2017).
Balladynopsis Theiss. & Syd., Annls mycol. 15(6): 475
(1918) [1917].
Index Fungorum number: IF 506; Facesoffungi number:
FoF 07952; – 9 morphological species (Species Fungorum
2020), molecular data unavailable.
Economic and ecological significance
Species in Balladynaceae are parasitic on living leaves. Currently there is no report concerning species economic significance. By considering their appearance on leaves which
is somewhat similar to Asterinaceae, they probably reduce
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respiration and photosynthesis of plants and probably cause
leaf stunt disease or host tissues to become pale.
Cleistosphaeraceae Boonmee & K.D. Hyde, in Boonmee
et al., Mycosphere 8(10): 1708 (2017).
Index Fungorum number: IF 553828; Facesoffungi numbers: FoF 03669, 1 species.
Parasitic or saprobic on leaves. Mycelium superficial,
branched, septate, with hyphopodia. Hyphopodia formed on
mycelium, subglobose, dark brown Sexual morph: Ascomata superficial, grouped to solitary, globose to subglobose,
dark brown, covered with mycelium. Hamathecium comprising sparse, septate, cellular pseudoparaphyses. Peridium comprises dark brown cells of textura angularis. Asci
8-spored, bitunicate, ovoid to broadly clavate, sessile or with
knob-like pedicel. Ascospores 2–3-seriate, ellipsoid-oblong,
hyaline to pale grey, aseptate, smooth-walled, with granular
contents, gelatinous sheath. Asexual morph: Coelomycetous (adapted from Boonmee et al. 2017).
Type: Cleistosphaera Syd. & P. Syd.
Notes: Cleistosphaeraceae was established to accommodate the monotypic genus Cleistosphaera. The morphology
of this family is quite similar to Botryosphaeriaceae, however, the generic type of Cleistosphaeraceae, Cleistosphaera
differs from Botryosphaeriaceae by its ascomata features,
mycelium with hyphopodia and sparse pseudoparaphyses.
Cleistosphaera Syd. & P. Syd., Annls mycol. 14(1/2): 74
(1916).
Index Fungorum number: IF 1100; Facesoffungi number:
FoF 07869; 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Cleistosphaera macrostegia Syd. & P. Syd.
Notes: Cleistosphaera is similar to taxa in Botryosphaeriaceae based on its asci and ascospore features, however,
it differs from Botryosphaeriaceae by mycelium with hyphopodia. The asexual morph of C. macrostegia are phoma-like
(coelomycetous) which differentiates it from all genera in
Perisporiopsidaceae (hyphomycetous).
Cleistosphaera macrostegia Syd. & P. Syd. [as ‘macrostegiae’], Annls mycol. 14(1/2): 75 (1916).
Index Fungorum number: IF 161307; Facesoffungi number: FoF 03670; Fig. 100
Description: see Boonmee et al. (2017).
Material examined: Peru, Amazonas, Rio Acre, Seringal Auristela, on leaves of Piptadenia sp. (Leguminosae),
August 1911, E.H.G Ule No. 3502 (S-F10853, holotype).
Economic and ecological significance
Species of Cleistosphaeraceae can be plant parasitic or play
a role in recycling organic matter.
233
Coccoideaceae P. Henn. ex Sacc. & D. Sacc., Henn. ex Sacc.
& D. Sacc., Syll. fung. (Abellini) 17: 860 (1905).
Index Fungorum number: IF 80616, Facesoffungi number: FoF 07953, 14 species.
Parasitic on living leaves. Sexual morph: Ascostromata
large, up to 2 mm diam., single to solitary, or scattered,
mostly superficial, slightly immersed in host tissue, circular
to subcircular, discoid, cushion-like, semicircular in section,
edge entire, slightly convex at the top, raised from the base,
black, thickened at the base, soft, multi-loculate, in a layer
in the upper part the ascostromata. Cells of ascostromata
comprising heavily, pigmented cells of textura angularis.
Locules completely immersed in ascostromata, globose,
subglobose or obpyriform, with apical ostiole. Hamathecium comprising filiform, anastomosing, branched, septate,
hyaline, relatively wide, cellular pseudoparaphyses. Asci
8-spored, bitunicate, fissitunicate, cylindrical to subclavate,
pedicellate, apically rounded, with a distinct, wide, ocular
chamber. Ascospores 2-seriate, apiosporous, ellipsoidal-obovoid, slightly oval, subglobose, tapering towards the narrow,
lower end, yellowish to light brown, 1-septate at the lower
end, not constricted at the septum, darkened at the septa,
smooth-walled. Asexual morph: Undetermined.
Type: Coccoidea Henn.
Notes: Coccoideaceae includes Coccoidea, Coccoidella
and Englerodothis, and is treated in Dothideomycetes as
family incertae sedis (Hyde et al. 2013; Boonmee et al.
2017; Wijayawardene et al. 2018; Pem et al. 2019c). The
characteristic feature of the family is ascostromata which
are formed as circular to discoid or cushion-like with dark
pigment, which is rather unique of its members. Boonmee
et al. (2017) revisited Coccoidella and provided a description and illustration of type species C. scutula and accepted
this genus in Coccoideaceae. Pem et al. (2019c) included
Englerodothis in Coccoideaceae. The family lacks molecular
data to establish phylogenetic placement, especially as the
type species is yet to be sequenced and no additional new
taxa in this family.
Coccoidea Henn., Bot. Jb. 28(3): 275 (1900).
Index Fungorum number: IF 1144, Facesoffungi number:
FoF 07954; 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Coccoidea quercicola Henn. & Shirai.
Notes: Coccoidea was introduced by Hennings (1900)
and typified by C. quercicola, which is parasitic on living
leaves. The genus lacks molecular data to indicate its generic
placement.
Coccoidea quercicola Henn. & Shirai, in Hennings, Bot. Jb.
28(3): 275 (1900).
Index Fungorum number: IF 167300, Facesoffungi number: FoF 07955; Fig. 101
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Fig. 101 Coccoidea quercicola (BPI 643971). a Material, habit on
leaf and appearance of ascostromata on lower leaf surface. b Section
of ascostroma showing arrangement of multilocule ascomata. c Close
up of peridial wall. d Hamathecium of trabeculate pseudoparaphyses.
e, f Asci. g–i Ascospores. Scale bars: a = 1 mm, b = 100 μm, c = 40
μm, d = 5 μm, e, f = 20 μm, g–i = 10 μm
Description: see Hyde et al. (2013).
Material examined: India, Sukliapokhari, Alt. 7,000 ft.,
on living leaves of Quercus lamellosa Sm., 11 May 1967,
M.K. Maity PCC 1252, (BPI 643971).
Notes: Coccoidella was revisited by Boonmee et al.
(2017) with description and illustration under a specimen
named Coccoidella scutula (BPI642219) to represent the
genus. This species was reported as parasitic on living leaves
of Persea palustris (Lauraceae) from the USA. Coccoidella
is placed in Coccoideaceae based on ascomata features
(Boonmee et al. 2017). However, collections and sequence
data are required to confirm the placement of this genus.
Other genera included
Coccoidella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
Kl., Abt. 1 118: 847 (1909).
Index Fungorum number: IF 1145; Facesoffungi number:
FoF 07956; – 9 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Coccoidella scutula (Berk. & M.A. Curtis)
Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1
118: 847 (1909).
≡ Dothidea scutula Berk. & M.A. Curtis, Grevillea 4(no.
31): 105 (1876).
13
Englerodothis Theiss. & Syd., Annls mycol. 13(3/4): 285
(1915).
Index Fungorum number: IF1818; Facesoffungi number:
FoF 06240; – 3 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Englerodothis kilimandscharica (Henn.)
Theiss. & Syd., Annls mycol. 13(3/4): 285 (1915).
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235
Fig. 102 Cookella microscopica (PAD: no. 2509) a Material
and habit on leaves. b Ascostroma with multilocule asci.
c, d Young and mature asci. e
Ascospores
≡ Cocconia kilimandscharica Henn., in Engler,
Pflanzenw. Ost-Afrikas Nachbarg., Teil C: 31 (1895).
Notes: Pem et al. (2019c) re-examined the type species
Englerodothis kilimandscharica (S-F203689). They included
Englerodothis in Coccideaceae based on its superficial, circular or discoid, multi-loculate ascomata, saccate-oblong
to cylindric-cavate asci, and ellipsoidal-oblong to clavulate,
1-septate, light pigmented ascospores. Species of Englerodothis are reported as parasitic or saprobic on living and dead
leaves. Fresh collections and sequence data are required to
indicate phylogenetic placement.
Economic and ecological significance
Coccoideaceae includes three genera with 14 species and is
distributed in temperate and tropical regions. Its members
are parasitic or saprobic on living leaves of dicotyledons and
angiosperms (Hyde et al. 2013; Boonmee et al. 2017; Pem
et al. 2019c).
Cookellaceae Höhn.ex Saccardo & Trotter, in Saccardo,
Syll. Fung. (Abellini) 22: 585 (1913).
Index Fungorum number: IF 80637; Facesoffungi numbers: FoF 07870, 6 species.
Parasitic (or epiphytic) on leaves in terrestrial habitats,
some possibly fungicolous. Sexual morph: Ascostromata
superficial, subglobose or irregular, pulvinate to effuse,
cushion-shaped, surface rough, flat or applanate at the base,
scattered, pigmented, containing locules with individual
asci, apapillate, opening by breaks in the upper surface; soft
textured, cells of ascostromata comprising textura angularis to subglobulosa, with a basal hypostroma, developing
in the host tissue, appearing as stromal blocks and swollen cells, of compressed packages of textura angularis to
prismatica. Locules scattered throughout ascostromata,
small, globose to subglobose, containing single asci, wall
not obvious. Hamathecium lacking pseudoparaphyses. Asci
8-spored, bitunicate, fissitunicate, globose to subglobose or
saccate, apedicellate, lacking an ocular chamber. Ascospores
scattered in 3–5-seriate overlapping rows, muriform, ellipsoid to oblong, hyaline when immature, becoming brown
at maturity, straight to slightly curved, ends rounded, with
3-transverse septa and 1-longitudinal septum in each cell,
constricted at the septa, smooth-walled. Asexual morph:
Coelomycetous, in genus Stigmella.
Type: Cookella Sacc.
Notes: The family includes only two genera Cookella and
Pycnoderma (Wijayawardene et al. 2018), while Uleomyces
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236
was transferred to Myriangiaceae by Boonmee et al. (2017).
Cookellaceae is treated in Dothideomycetes as family incertae sedis (Wijayawardene et al. 2018). The family is parasitic
or epiphytic on leaves of dicotyledons and monocotyledons,
with cushion-shaped to irregular thyriothecial single to multiloculate, ascostromata or ascomata, absent of pseudoparaphyses, 8-spored, bitunicate asci and muriform, light to dark
pigmented ascospores (Hyde et al. 2013). The family lacks
molecular data to establish its phylogenetic placement, especially as the type species is yet to be sequenced and there are
no additional new taxa in this family.
Cookella Sacc., Michelia 1(no. 4): 407 (1878).
Index Fungorum number: IF 1225, Facesoffungi number:
FoF 07957; 4 morphological species (Species Fungorum
2020), molecular data unavailable.
Type: Cookella microscopica Sacc.
Notes: Cookella was introduced to accommodate C.
microscopica found on lower surface of leaves of Quercus
pedunculatum (oak) in Italy (Saccardo 1878). The genus
needs sequence data to resolve its generic placement.
Cookella microscopica Sacc., Michelia 1(no. 4): 407 (1878).
Index Fungorum number: IF 218808, Facesoffungi number: FoF 07958; Fig. 102
Description and Material examined: see Hyde et al.
(2013).
Other genera included
Pycnoderma Syd. & P. Syd., Annls mycol. 12(6): 563
(1914).
Index Fungorum number: IF 4565, Facesoffungi number:
FoF 07959; – 2 morphological species, (Species Fungorum
2020), molecular data unavailable.
Type species: Pycnoderma bambusinum Syd. & P. Syd.,
Annls mycol. 12(6): 563 (1914).
Notes: Pycnoderma was introduced to accommodate P.
bambusinum found on bamboo leaves (Sydow and Sydow
1914a). Only two species are accepted in the genus (Hongsanan et al. 2014a; Wijayawardene et al. 2017a). Fresh collections of its members are needed to clarify the generic
placement.
Economic and ecological significance
Cookellaceae includes two genera with six species. Its genera are parasitic or epiphytic on leaves of dicotyledons and
monocotyledons (Hyde et al. 2013; Hongsanan et al. 2014a).
Dimeriaceae E. Müll. & Arx ex Arx & E. Müll., Stud.
Mycol. 9: 104 (1975).
Index Fungorum: IF 80704; Facesoffungi number: FoF
06390, 73 species.
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Parasitic on living leaves, colonies dark brown, mycelium superficial, dark brown, amphigenous. Sexual morph:
Ascomata superficial, subglobose, with hyphal appendages,
flexuous, cylindrical, septate, dark brown, roughened. Asci
8-spored, bitunicate, fissitunicate, saccate-oblong to ellipsoidal, apically thickened. Ascospores conglobate, ellipsoidfusiform to subclavate, ends rounded, olivaceous-brown to
brown, 1-septate, darkly pigmented at the septum, upper
cell wider than lower cell, septum supramedian. Asexual
morph: Hyphomycetous.
Type: Dimerium (Sacc. & P. Syd.) McAlpine.
Notes: Dimeriaceae accommodates taxa occurring on the
surface of living leaves (Hawksworth and Eriksson 1986;
von Arx and Müller 1975; Barr 1987b, 1997). Its members
are characterized by solitary, superficial ascomata with
hyphal appendages and 1-septate ellipsoidal, dark brown
ascospores darkly pigmented at the septum (Boonmee et al.
2017). Dimerium olivaceum was assigned as the type species
(Boonmee et al. 2017). However, the family lacks molecular
data to confirm the taxonomic placement.
Dimerium (Sacc. & P. Syd.) McAlpine, Proc. Linn. Soc.
N.S.W. 28: 98 (1903).
≡ Dimerosporium subgen. Dimerium Sacc. & P. Syd.,
Syll. fung. (Abellini) 16: 410 (1902).
Index Fungorum number: IF 25887; Facesoffungi number: FoF 06391; 73 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Dimerium olivaceum Syd. & P. Syd.
Notes: Dimerium is a single genus in Dimeriaceae
(Wijayawardene et al. 2018). Many species were transferred to other genera based on shared characters (Boonmee
et al. 2017). More collections of the members in this genus
and sequence data are needed to clarify their taxonomic
placement.
Dimerium olivaceum Syd. & P. Syd., Annls mycol. 2(2):
169 (1904).
Index Fungorum number: IF 234107; Facesoffungi number: FoF 06392, Fig. 103
Description: see Boonmee et al. (2017).
Material examined: Chile, Anden bei Villarica, on living leaves of Cynoctonum nummulariaefolium (Asclepiadaceae), February 1897, F.W. Neger (SF11956, holotype).
Economic and ecological significance
The members in Dimeriaceae are reported as parasitic on
living leaves of various hosts.
Dubujianaceae D. Pem, Doilom & K.D. Hyde, Mycosphere
10(1): 1115–1246 (2019).
Index Fungorum number: IF 557065; Facesoffungi number: FoF 06679, 1 species.
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237
Fig. 103 Dimerium olivaceum (SF11956, holotype). a Material
label. b Herbarium specimen and habit on leaves. c Appearance of
colonies on the leaves surface. d Ascomata, marked by arrows e Section of ascoma. f Peridium. g Hamathecium comprising pseudopara-
physes. h–m Asci. n–v Ascospores. w Mycelium with septate and
hyphopodia-like structures (red arrows). x–z Conidiophores in different stages of developing conidia. Scale bars: c, d = 500 µm, e = 50
µm, f–m = 20 µm, n–z = 10 µm
Endophytic, saprobic or pathogenic on leaves. Sexual
morph: Undetermined. Asexual morph: Mycelium at
first subcuticular, developing onto leaf surface from leaf
glands forming scattered to sporadically confluent superficial thalli, circular in outline, composed of a basal layer
of flattened, parallel, branching, brown hyphae with setae.
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Fig. 104 Dubujiana glandulifera (UC AUS399, holotype). a Details of herbarium material. b Habit and appearance of conidiomata on host surface. c Section of conidioma. d Conidiogenesis. e–g Conidia. Scale bars: b = 2 mm, c = 40 µm, d = 100 µm, e, f = 5 µm, g = 10 µm
Lacking hyphopodia, Hyphal strands radiate from base of
central, raised pycnidium over the basal layer and onto the
substrate. Conidiomata wall composed of thick dark-brown
walled cells of textura globulosa. Conidiophores densely
aggregated, slender, subulate, simple, frequently branched
above, reduced to conidiogenous cells, or with 1–2 supporting cells. Conidiogenous cells annellidic, simple, tapering, hyaline, smooth, rarely with percurrent proliferations.
Conidia ellipsoidal, fusiform to lunate, hyaline becoming
pale brown at maturity, 1-septate, punctate, smooth-walled.
Type: Dubujiana D.R. Reynolds & G.S. Gilbert.
Notes: Pem et al. (2019c) introduced Dubujianaceae with
Dubujiana as the type genus. Species of Dubujianaceae have
an epifoliar lifestyle with pycnidia arising from leaf glands
in the outer portions of the hyphal system with individual
hyphal strands extending from the large central pycnidium.
Dubujiana D.R. Reynolds & G.S. Gilbert, Aust. Syst. Bot.
18(3): 282 (2005).
Index Fungorum number: IF 28993; Facesoffungi number: FoF 06238; 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Dubujiana glandulifera D.R. Reynolds &
G.S. Gilbert.
Notes: The monotypic genus Dubujiana is characterized by raised pycnidia over the basal layer of the substrate,
densely aggregated, slender conidiophores, and tapering,
hyaline conidiogenous cells giving rise to pale brown,
1-septate, punctate conidia. No cultures or sequence data
are available for this genus.
13
Dubujiana glandulifera D.R. Reynolds & G.S. Gilbert,
Aust. Syst. Bot. 18(3): 282 (2005).
Index Fungorum number: IF 344434; Facesoffungi number: FoF 08038; Fig. 104
Description: see Pem et al. (2019c).
Material examined: Australia, Queensland, Cape Tribulation Coastal Boardwalk Dubuji, 145o28’ E, 16 o4’S., on
leaves, 2 February 2002, D.R. Reynolds (UC AUS399,
holotype).
Economic and ecological significance
Species of Dubujiana are epifoliar and maintain a commensal relationship with the host (Reynolds and Gilbert 2005).
Dysrhynchaceae Boonmee & K.D. Hyde [as ‘Dysrhynchisceae’], in Boonmee et al., Mycosphere 8(10): 1714 (2017).
Index Fungorum: IF 628246; Facesoffungi number: FoF
06393, 4 species.
Saprobic on dead leaves. Sexual morph: Ascomata
superficial, globose to subglobose, dark brown, solitary,
aggregated, with central pore, with a protruding synnematous neck, comprised of tightly compacted hyphae, sparse
superficial mycelium with hyphopodia, brown to dark
brown, with partially immersed mycelia, lacking setae, lacking pseudoparaphyses. Peridium membranous, comprising
brown cells of textura angularis. Asci 8-spored, bitunicate,
broadly obclavate to ovoid, sessile, apically thickened.
Ascospores multi-seriate, conglobate, ellipsoidal, clavate
to obpyriform, hyaline to light brown. Asexual morph:
Hyphomycetous.
Type: Dysrhynchis Clem.
Fungal Diversity (2020) 105:17–318
Notes: Boonmee et al. (2017) introduced Dysrhynchisceae as a new family to accommodate Dysrhynchis. The
morphological features of sexual and asexual morphs were
described in Boonmee et al. (2017). Additional collections
and molecular study are required to further clarify the phylogeny of this family.
Dysrhynchis Clem., Gen. fung. (Minneapolis): 32 (1909).
Index Fungorum number: IF 1728; Facesoffungi number:
FoF 06394; 4 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Dysrhynchis pulchella (Sacc.) Clem.
Notes: Dysrhynchis is the type genus of Dysrhynchaceae
and both morphs of the life cycle have been reported. The
genus lacks molecular data to establish phylogenetic placement and no additional new taxa have been introduced.
Dysrhynchis pulchella (Sacc.) Clem., in Clements & Shear,
Gen. fung., Edn 2 (Minneapolis): 253 (1931).
≡ Henningsomyces pulchellus Sacc. & D. Sacc., Syll.
fung. (Abellini) 17: 689 (1905).
Index Fungorum: IF 254191; Facesoffungi number: FoF
06395; Fig. 105
Description: see Boonmee et al. (2017).
Material examined: Brazil, Rio de Janeiro, Mauá, on dead
leaves of Byrsonima sericae A. Juss. (Malpighiaceae), 21
July 1899, E.H.G. Ule, Mycotheca brasiliensis no. 71 (PAD
No. 71, holotype).
Economic and ecological significance
Dysrhynchisceae are saprobes, occurring on plants from terrestrial habitats (Wijayawardene et al. 2017a). This family
has no report as causing plant diseases.
Endosporiaceae D. Pem, Doilom & K.D. Hyde, Mycosphere 10(1): 1115–1246 (2019).
Index Fungorum number: IF 557066; Facesoffungi number: FoF 06681, 2 species.
Pathogenic on bud of Populus tremuloides. Sexual
morph: Undetermined. Asexual morph: Colonies (conidiomata) on PDA, OA and CA black, raised, cerebriform, comprising numerous cellular clumps in a thin covering, subcircular to irregular in outline, reverse black. Margin irregular
with scarce, occasionally aggregated hyphae on PDA, even
with adpressed or submerged hyphae on MEA. On PDA,
cells more broad, swollen, subhyaline and dumbbell-shaped
when two-celled, becoming multicellular by splitting in all
directions, becoming darkly pigmented, irregular in shape,
frequently separating into smaller clumps. Hyphae cylindrical or toruloid, light to dark brown, branched or unbranched,
aerially determinate in growth, mostly elongated, hyaline
at apex, forming muriform and darkly pigmented bodies.
239
Endoconidia developing from the separation of adjacent
daughter cells in cellular clumps through septum schizolysis, ellipsoidal to subglobose becoming broadly ellipsoidal
to globose, hyaline, aseptate. Blastic conidia abundant in
mature colonies, arising from cells of cellular clumps or
seldom from sides of hyphae, cylindrical to ellipsoidal,
regularly truncate at the base, sometimes globose, obovoid,
fusiform, hyaline or light brown, aseptate.
Type: Endosporium Tsuneda.
Notes: Pem et al. (2019c) introduced Endosporiaceae to
accommodate Endosporium. The family is characterized by
cylindric hyphae, ellipsoidal, subglobose to globose endoconida and cellular clumps, globose, obovoid, fusiform blastic
conidia. Endosporiaceae is unique in producing endoconidia
from cellular clumps.
Endosporium Tsuneda, in Tsuneda et al., Botany 86(9):
1022 (2008).
Index Fungorum number: IF 536893; Facesoffungi number: FoF 06239; 2 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Endosporium populi-tremuloides Tsuneda.
Notes: Endosporium is characterized by cylindrical
hyphae, ellipsoidal, subglobose to globose endoconida and
cellular clumps, globose, obovoid, fusiform blastic conidia.
The sexual morph is unknown. Two species are known, E.
aviarium and E. populi-tremuloides.
Endosporium populi-tremuloides Tsuneda, in Tsuneda
et al., Botany 86(9): 1023 (2008).
Index Fungorum number: IF 536901; Facesoffungi number: FoF 08836; Fig. 106
Description: see Pem et al. (2019c).
Material examined: Canada, Whitemud Creek, Edmonton, Lansdowne, ca. 1 km west of the Northern Forestry
Centre, Alberta, on bud of Populus tremuloides (Salicaceae),
3 November 2002, A. Tsuneda (UAMH 10529, holotype).
Economic and ecological significance
Two species in Endosporiaceae were found as pathogens
which can uptake nutrients from plants.
Englerulaceae Henn., Hedwigia 43: 353 (1904).
= Schiffnerulaceae Hosag., Pl. Pathol. Quarant. 1(2): 132
(2011).
Index Fungorum number: IF 80736; Facesoffungi number: FoF 07960, 125 species.
Parasitic or epiphytic on living leaves, primarily tropical. Colonies superficial, thin to dense, brown to dark
brown, confluent, velvety, with unicellular appressoria.
Hyphae straight to flexuous, septate, irregularly to reticulate
branched, brown. Hyphopodia globose, brown, thick-walled,
irregular or absent. Sexual morph: Ascomata relatively
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240
small, superficial on the hyphae or on a short stalk-cell, scattered, ellipsoid to globose, cupulate, brown to black, membranous, containing few asci, lacking ostioles, dissolving
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at center to release asci. Hamathecium comprising brown,
septate, cellular pseudoparaphyses or pseudoparaphyses
lacking. Asci 2 to 8 spored, bitunicate, thick-walled, ovate,
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241
◂Fig. 105 Dysrhynchis pulchella (PAD No. 71, holotype). a, b Mate-
Index Fungorum number: IF 1820; Facesoffungi number:
FoF 07961; 10 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Englerula macarangae Henn.
Notes: This genus is characterized by brown to black colonies with angled hyphae, globose to subglobose ascomata
with a thin peridium and obovoid to globose asci (von Arx
and Müller 1975; Eriksson 1981; Hosagoudar et al. 2011;
Hyde et al. 2013; Dai et al. 2014b). The species of this genus
needs to be recollected.
globose to subglobose, pedicellate, with an ocular chamber.
Ascospores uni- to multi-seriate, oblong to ellipsoid, ellipsoid to ovate, or fusiform, hyaline to dark brown, 1-septate,
smooth-walled, constricted at the septum. Asexual morph:
Conidiomata pycnidial, similar to ascomata, superficial,
globose, thin-walled. Conidiogenous cells holoblastic, phialidic, discrete, hyaline, smooth, 1-celled, nearly ampulliform, alternate or absent. Conidia oblong to ovate, globose
or nearly globose, hyaline to brown, aseptate.
Type: Englerula Henn.
Notes: Englerulaceae was introduced by Hennings
(1904a, b) for taxa with brown to dark brown colonies on
host leaves, with superficial hyphae, scattered, ascomata
lacking ostioles. Lumbsch and Huhndorf (2007, 2010)
included seven genera (Englerula, Goosia, Parenglerula,
Rhizotexis, Rhytidenglerula, Schiffnerulaand and Thrauste)
in the family, while Hyde et al. (2011) considered Schiffnerulaceae (Hosagoudar 2011) to be a synonym. Schiffnerula
was introduced by von Höhnel (1909b) and placed in Englerulaceae based on its globose ascomata and globose to ovate
asci. Hosagoudar (2011) included four asexual genera in the
family with more than 100 species. Schiffnerula has dark
colonies on the leaf surface, with brown, superficial, septate
mycelium, 8-spored, bitunicate, cylindrical to globose asci,
and brown, 1-septate, ascospores which are constricted at the
septa. With such characters Schiffnerula can be accommodated in Englerulaceae (Hyde et al. 2013). Dai et al. (2014b)
redescribed and illustrated type species of all genera in this
family. They suggested that Rhizotexis should be excluded
from this family and placed into Dothideomycetes genera
incertae sedis based on having stromatic ascomata and lack
of hyphae.
There is no molecular data for any species of Englerulaceae in GenBank. Fresh collections and molecular data
are required to clarify the relationships in Englerulaceae.
Englerulaceae are closely related to the Asterinaceae, but
differ in having sphaerical ascomata without radiate wall
and growing on brown mycelium (von Arx and Müller 1975;
Eriksson 1981; Hyde et al. 2013).
Englerula macarangae Henn., Bot. Jb. 34: 49 (1904).
Index Fungorum number: IF 140464; Facesoffungi number: FoF 07962; Fig. 107
Description: see Dai et al. (2014b).
rial label and herbarium specimen. b Appearance of ascomata on
the upper leaf surface. c Close up the appearance of ascomata on the
upper leaf surface. d Appearance of ascomata on the lower leaf surface. e Close up the appearance of ascomata on the lower leaf surface. f Squash mount of ascoma with protruding synnematous necks.
g Appearance of ascoma at from the side. h Appearance of ascomamfrom at below. i Peridium. j, k Asci. l, m Ascospores. n Synnema
with apical conidia. o Hyphopodia and appressoria (red arrows). p–s
Conidia. Scale bars: f, g = 50 µm, h–k, n, o = 20 µm, l, m, p–s = 10
µm
Englerula Henn., Bot. Jb. 34: 49 (1904).
= Anatexis Syd., Annls mycol. 26(1/2): 90 (1928).
Other genera included
Allosoma Syd., Annls mycol. 24(5/6): 353 (1926).
Index Fungorum number: IF 143; Facesoffungi number:
FoF 08927; – 4 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Allosoma cestri Syd., Annls mycol.
24(5/6): 353 (1926).
Notes: The genus is characterized by superficial, globose to subglobose ascomata, subglobose asci, and oblongellipsoid to ellipsoid, 1-septate ascospores. This genus was
placed this genus in Englerulaceae by Thambugala et al.
(2014b) and Wijayawardene et al. (2017a, 2020). Sequence
data is needed to confirm its placement.
Digitosarcinella S. Hughes, Can. J. Bot. 62(11): 2208
(1984).
Index Fungorum number: IF 11070; Facesoffungi number: FoF 08928; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Digitosarcinella caseariae S. Hughes, Can.
J. Bot. 62(11): 2208 (1984).
Notes: The genus is characterized by cheiroid conidia,
hyphae with sessile hyphopodia and as asexual state of
Schiffnerula (Hughes 1984). Wijayawardene et al. (2017a,
2020) placed this genus in Englerulaceae.
Goosia B. Song, Mycotaxon 87: 413 (2003).
Index Fungorum number: IF 28768; Facesoffungi number: FoF 07963; – 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Goosia melastomatis B. Song, Mycotaxon
87: 413 (2003).
Notes: The monotypic genus is characterized by dark
brown phialides developing directly on the hyphae, and obovoid asci containing two ascospores (Bin 2003). This genus
is similar to Thrauste in its superficial, ellipsoid ascomata
and globose hyphopodia formed on dark brown hyphae. Bin
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Fig. 106 Endosporium populi-tremuloides (UAMH 10529, holotype). a–c Details of herbarium material. d Squash mount of mycelium. e–g
Hyphae giving rise to cellular clumps. h–k Blastic conidia. Scale bars: d, e = 20 µm, f, g = 30 µm, h–k = 5 µm
(2003) did not observe mature ascomata and considered that
Goosia differs from Thrauste with the former having phialides and smaller ascospores. Fresh collections are needed
to confirm if they are different and also show its natural
taxonomic relationships. Dai et al (2014b) redrew the main
characters from Bin (2003).
having superficial, globose to subglobose, dark ascomata
containing obovoid asci and broadly ellipsoid to subobovoid
ascospores with a single septum, but differs by darker ascomata and ascospores (von Höhnel 1910; Dai et al. 2014b).
The genus has not been studied recently and thus needs fresh
collections.
Parenglerula Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
Kl., Abt. 1 119: 465 [73 repr.] (1910).
Index Fungorum number: IF 3727; Facesoffungi number:
FoF 07964; – 6 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Parenglerula macowaniana (Thüm.)
Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1
119: 465 [73 repr.] (1910).
≡ Meliola macowaniana Thüm., Mycoth. Univ., cent. 6:
no. 568 (1876).
Notes: Parenglerula was erected by von Höhnel (1910)
with seven species. Parenglerula is similar to Englerula in
Rhytidenglerula Höhn., Sber. Akad. Wiss. Wien, Math.naturw. Kl., Abt. 1 127: 386 [58 repr.] (1918).
Index Fungorum number: IF 4738; Facesoffungi number:
FoF 07965; – 9 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Rhytidenglerula carnea (Ellis & G. Martin) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt.
1 127(4): 386 [58 repr.] (1918).
≡ Asterina carnea Ellis & G. Martin, Am. Nat. 17(2):
1285 (1883).
Notes: Rhytidenglerula has small ascomata forming on
dark hyphae, a character that distinguishes it from other
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243
Fig. 107 Englerula macarangae (redrawn from Dai et al. 2014b). a Squash mount of ascoma. b–d Asci with ascospores. e–f Ascospores with
one septum. Scale bars: a–d = 50 µm, e, f = 20 µm
genera with large and dark ascomata in the family. Rhytidenglerula is similar to Schiffnerula in having hyphopodia
and globose ascomata, obovoid asci and hyaline ascospores
with a single septum. Rhytidenglerula is linked to a coelomycetous asexual morph, Capnodiastrum (Kirk et al. 2008;
Wijayawardene et al. 2012) which is characterized by holoblastic, doliiform, conidiogenous cells and ellipsoid, pale
brown, 1-celled conidia with a conic apex (Sutton 1980).
Schiffnerula has been linked to four different hyphomycetous
asexual morphs, Digitosarcinella, Mitteriella, Questieriella
and Sarcinella (Hosagoudar 2011; Hosagoudar et al. 2011;
Seifert et al. 2011). These four genera are characterized by
monoblastic to polyblastic conidiogenous cells and sigmoid to ellipsoid, dark brown to reddish brown, or black,
0–4-celled conidia (Hosagoudar 2011).
Schiffnerula Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
Kl., Abt. 1 118: 867 [55 repr.] (1909).
Index Fungorum number: IF 4886; Facesoffungi number:
FoF 07966; – 96 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Schiffnerula mirabilis Höhn., Sber. Akad.
Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 868 [56 repr.]
(1909).
Notes: Hosagoudar (2011) considered Schiffnerula as a
new family, because Schiffnerula has young ascomata which
look like a shield-plate on hyphae. According to von Höhnel
(1909b, 1910), both Rhytidenglerula and Schiffnerula have
thin, shield-like immature ascomata and should belong to
Englerulaceae. In addition, Schiffnerula has globose, ascomata, developing on brown hyphae and contain a few globose to ovate asci with multi-seriate, ellipsoid, 1-septate
ascospores, with Digitosarcinella, Mitteriella, Sarcinella,
Questieriella as its asexual morph. Therefore, we retain
Schiffnerula as a genus in Englerulaceae.
Thrauste Theiss., Verh. zool.-bot. Ges. Wien 66: 337 (1916).
Index Fungorum number: IF 5455; Facesoffungi number:
FoF 07967; – 3 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Thrauste medinillae (Racib.) Theiss., Verh.
zool.-bot. Ges. Wien 66: 338 (1916).
≡ Balladyna medinillae Racib., Bull. int. Acad. Sci. Lett.
Cracovie, Cl. sci. math. nat. Sér. B, sci. nat. 3: 373 (1909).
Notes: This genus is similar to Goosia in its dark hyphopodia and ellipsoid immature ascomata growing directly on
hyphae (Theissen 1916; Bin 2003). However, Thrauste differs in its 8-spored, pyriform to clavate asci with an ocular
chamber, while Goosia has 2-spored, obovoid asci and the
ocular chamber is not well-developed. Dai et al. (2014b)
examined a collection (F: 111745), labelled T. medinillae,
obtained from S. There are marked differences in the shape
and size of asci and ascospores between the original description of the species (Saccardo 1913) under the genus. Based
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Fig. 108 Homortomyces tamaricis (a–g = MFLU 14-0595; h–l =
MFLU 14-0167). a Appearance of ascomata on host surface b Vertical section through ascoma c Peridium d Pseudoparaphyses and
mature ascus e–g Ascospores h Vertical sections through conidioma.
i, j Developing conidia and paraphyses k–l Conidia. Scale bars: b =
100 μm, c, h = 50 μm, d, i = 20 μm, e–g, j =10 μm
on these differences, they proposed a new species for this
specimen (Thrauste parvii).
layers, of lightly pigmented to hyaline cells of textura angularis. Hamathecium comprising cylindrical, smooth, flexuous, apex obtuse, hyaline, sparingly septate, paraphyses,
intermingled among conidiogenous cells. Conidiophores
reduced to conidiogenous cells or one supporting cell. Conidiogenous cell with supporting cell, hyaline, percurrently
proliferating at the tip of the supporting cells. Conidia ellipsoid to subcylindrical, straight to slightly curved, golden
brown to dark brown, 3(–4)-euseptate, apex obtuse, base
truncate with a visible scar.
Type: Homortomyces Crous & M.J. Wingf.
Notes: Thambugala et al. (2017) introduced the monotypic family Homortomycetaceae to accommodate its type
Homortomyces, as its phylogenetic placement is uncertain.
Thambugala et al. (2017) placed Homortomycetaceae in
Dothideomycetes, families incertae sedis.
Economic and ecological significance
Englerulaceae is a poorly known family, most species are
parasitic on leaves. There is no sequence data for any members of this family in GenBank and more fresh collections
are required to clarify the relationships in Englerulaceae.
Homortomycetaceae Thambug., A.J.L. Phillips & K.D.
Hyde, in Thambugala et al., Fungal Diversity 82: 45 (2016).
Index Fungorum number: IF 552086; Facesoffungi number: FoF 02150, 2 species.
Foliicolous, associated with leaf spots or saprobic on
dead twigs or branches. Sexual morph: Ascomata scattered,
immersed to erumpent, black, globose to subglobose, ostiolate. Peridium composed of a few layers of brown to dark
brown, thick-walled cells of textura angularis. Hamathecium comprising septate, cellular pseudoparaphyses. Asci
2–6-spored, bitunicate, fissitunicate, cylindrical, apically
rounded. Ascospores 1–2-seriate, fusiform with broadly to
narrowly rounded ends, yellowish brown to brown, 3-septate, smooth-walled. Asexual morph: Coelomycetous.
Conidiomata pycnidial, solitary or gregarious, immersed or
slightly erumpent, uniloculate or multi-loculate, globose to
subglobose, ostiolate. Conidiomatal wall comprising a few
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Homortomyces Crous & M.J. Wingf., in Crous et al., IMA
Fungus 3(2): 110 (2012).
Index Fungorum number: IF 801349; Facesoffungi number: FoF 07968; 2 morphological species (Species Fungorum 2020), 2 species with molecular data.
Type species: Homortomyces combreti Crous & M.J.
Wingf.
Notes: Crous et al. (2012a) established Homortomyces
in Dothideomycetes genera incertae sedis to accommodate
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H. combreti, which was associated with leaf spots on Combretum erythrophyllum. The second species, H. tamaricis
was introduced by Wijayawardene et al. (2014b) from dead
branches of Tamarix gallica. Thambugala et al. (2017)
described the sexual morph of H. tamaricis associated with
the same host. Homortomyces is similar to Stilbospora,
which is classified in Stilbosporaceae, Diaporthales, Sordariomycetes (Wijayawardene et al. 2014b; Senanayake et al.
2017), while the genera Endocoryneum, Hendersoniopsis,
Angiopomopsis and Ceratopycnis show similar conidial
characteristics of Homortomyces. However, all these genera
have been classified in incertae sedis due to lack sequence
data (Crous et al. 2012a; Thambugala et al. 2017).
Homortomyces tamaricis Wijayaw., Camporesi & K.D.
Hyde, in Wijayawardene et al., Phytotaxa 176(1): 160
(2014).
Index Fungorum: IF 550192; Facesoffungi number: FoF
02151; Fig. 108
Saprobic on Tamarix gallica L. Sexual morph: Ascomata
190–280 μm high × 280–350 μm diam. ( x̄ = 234 × 310
μm, n = 6), scattered, sometimes aggregated, immersed to
partially erumpent through the host tissues, black, globose
to subglobose, unilocular, ostiolate. Peridium 20–40 μm
wide, comprising several layers of brown to dark brown,
thick-walled cells of textura angularis. Hamathecium comprising 2–4 μm wide, septate, cellular pseudoparaphyses.
Asci 80–100 × 18–23 μm ( x̄ = 87 × 21 μm, n = 6), mostly
2–6-spored, bitunicate, fissitunicate, cylindrical, short pedicellate, apically rounded, with a distinct ocular chamber.
Ascospores 23–30 × 8–12 μm ( x̄ = 25 × 10 μm, n = 25),
1–2-seriate, fusiform with broadly to narrowly rounded ends,
yellowish brown to brown, 3-septate, smooth-walled, without a mucilaginous sheath. Asexual morph: Coelomycetous.
Conidiomata 150–250 μm high × 300–350 μm diam. ( x̄ =
200 × 330 μm, n = 5), solitary or gregarious, immersed
to slightly erumpent, pycnidial to irregular, uniloculate to
multi-loculate, subglobose, ostiolate. Conidiomatal wall
10–22 μm), comprising a few layers of lightly pigmented to
hyaline cells of textura angularis. Hamathecium comprising
numerous, aseptate, cylindrical, guttulate, paraphyses. Conidiophores reduced to conidiogenous cells. Conidiogenous
cell 6–25 × 2.5–5.5 μm ( x̄ = 15 × 3.6 μm, n = 20), with
supporting cell, hyaline, percurrently proliferating at the tip
of the supporting cells. Conidia ellipsoid to subcylindrical,
straight to slightly curved, 20–30 × 9–12.5 μm ( x̄ = 25 ×
11 μm, n = 30), golden brown to dark brown, becoming
coloured before release from conidiogenous cells, smoothwalled, initially asptate to 1-septate, becoming at maturity
3-septate, apex obtuse, base truncate, smooth-walled.
Material examined: Italy, Province of Forlì-Cesena [FC]),
Ravaldino in Monte Forlì, on dead branches of Tamarix
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gallica L. (Tamaricaceae), 22 November 2012, Erio Camporesi IT 922 (MFLU 14-0595 & MFLU 14-0167).
Notes: Wijayawardene et al. (2014b) introduced Homortomyces tamaricis from Tamarix gallica based on its asexual
morph, while Thambugala et al. (2017) described the sexual
morph of this species from the same host based both morphological traits and molecular sequence data.
Economic and ecological significance
As the species of Homortomyces are associated with ornamental plants such as Combretum erythrophyllum and Tamarix gallica, they reduce the value of those plants.
Hyalomeliolinaceae Boonmee & K.D. Hyde, in Boonmee
et al., Mycosphere 8(10): 1718 (2017).
Index Fungorum: IF 553830; Facesoffungi number: FoF
06396, 2 species.
Parasitic on living leaves. Sexual morph: Colonies
form on a lower surface of leaves, black, subcircular, with
outwardly radiating mycelium, with hyphae that are long
hairy, flexible, dark brown to black, densely, fluffy, partially erect, unbranched, and septate. Ascomata superficial,
globose to subglobose, black, seated on dark hyphae, covered by dense hyphae, superficial mycelium, unbranched,
septate, dark to black. Peridium comprising dark brown to
black cells of textura angularis. Hamathecium comprising
elongate-filiform, septate, unbranched, filamentous, cellular
pseudoparaphyses, extending over asci, constricted at the
septa. Asci 8-spored, bitunicate, fissitunicate, saccate-oblong
to ellipsoidal, sessile, lower median widest, with the small
ocular chamber. Ascospores 2–3-seriate, ellipsoid-fusiform,
end narrow, pigmented, multi-septate. Asexual morph:
Undetermined.
Type: Hyalomeliolina F. Stevens.
Notes: Boonmee et al. (2017) re-examined the type species Hyalomeliolina guianensis and concluded the unique
suite of characters and could not place it in any families
in Dothideomycetes. Therefore, Hyalomeliolinaceae was
established to accommodate Hyalomeliolina (Boonmee
et al. 2017). No molecular data is available to confirm its
phylogenetic placement within Dothideomycetes.
Hyalomeliolina F. Stevens, Illinois Biol. Monogr. (Urbana)
8(no. 3): 27 (1924).
Index Fungorum: IF 553830; Facesoffungi number: FoF
06397; 2 morphological species (Species Fungorum 2020),
molecular data unavailable.
Type species: Hyalomeliolina guianensis F. Stevens.
Notes: Hyalomeliolina is a single genus in Hyalomeliolinaceae, and it was established by Stevens (1923)
based on morphology. The genus is characterized by a life
mode as parasites on living leaves, forming black colonies
on leaf surfaces, ascomata covered by dense hyphae and
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Fig. 109 Hyalomeliolina guianensis (BPI688907, holotype). a
Material label. b Herbarium specimen. c Ascomata covered by
hyphae on the leaf surface. d Section of ascoma. e Peridium. f, g
Hairy hyphae. h Pseudoparaphyses. i–k Asci. l–o Ascospores. Scale
bars: c = 5 mm, d, f = 500 µm, e, g, i–k = 50 µm, h, l–o = 20 µm
ellipsoid-fusiform, 1–3-septate, light to dark pigmented
ascospores. The genus comprises Hyalomeliolina guianensis
and H. linderi, while H. costaricensis were transferred to
Nematostoma by Hansford (1946).
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Hyalomeliolina guianensis F. Stevens, Illinois Biol. Monogr. (Urbana) 8(no. 3): 28 (1924) [1923].
Index Fungorum number: IF 266643; Facesoffungi number: FoF 06398, Fig. 109
Description: see Boonmee et al. (2017).
Material examined: Guyana (= British Guiana), Rockstone, on living leaves of Licania Aubl. (Chrysobalanaceae),
17 July 1922, F.L. Stevens No. 454 (BPI688907, holotype).
Economic and ecological significance
Members in Hyalomeliolinaceae play roles in host plant
penetration by being parasitic.
Leptopeltidaceae Höhn. ex Trotter [as ‘Leptopeltineae’],
Syll. fung. (Abellini) 24(2): 1255 (1928).
Index Fungorum: IF 81594; Facesoffungi number: FoF
07969, 15 species.
Epiphytic on ferns (Aspidium), Aruncus, Potentilla and
other flowering herbaceous plants as well as Connarus
suberosus Planch. Sexual morph: Ascomata thyriothecial,
superficial, visible as black dots or irregular on host surface,
producing vegetative mycelium penetrating the host, solitary
to gregarious, flattened, circular, round, elongate or irregular, or Y-shaped, easily removed from the host, brown to
dark brown, waxy, in section lenticular, scutate, conical, or
quadrilateral trapezoid, dehiscence by slit-like opening, or
splitting of the upper wall. Peridium thin-walled, composed
of reddish-brown to dark brown, isodiametric or polygonal
upper-walled cells with a parallel or irregular arrangement,
radiating from the centre, basal layer poorly developed,
occasionally forming basal cell layers, arranged in a textura
angularis in verical section. Hamathecium comprising
broadly filamentous, septate, unbranch, tapering towards
the apex, paraphyses, embedded in gelatinous matrix. Asci
8-spored, bitunicate, broadly cylindrical to cylindric-clavate,
or subglobose to ampulliform, with obtuse to truncate or
acutate apex, sesille to subsessile, with truncate base, ocular
chamber indistinct, clearly visible when young. Ascospores
overlapping 1–3-seriate, twisted or fasciculate, varied in
shape, ellipsoidal, broadly fusiform, cylindrical, lunate,
or muriform, with rounded to acute ends, hyaline, aseptate
or septate, smooth-walled, with or without small guttules.
Asexual morph: Hyphomycetous idriella-like (asexual
morph of Dothiopeltis) and coelomycetous leptothyriumlike (asexual morph of Leptopeltis) (Wijayawardene et al.
2017b).
Type: Leptopeltis Höhn.
Notes: von Höhnel (1917) invalidly introduced Leptopeltidaceae as ‘Leptopeltineen’ and treated the family in
Phacidiales. However, the family was validly introduced
by Saccardo (1928). The familial concept contained highly
heterogenous taxa which were described as epiphytic fungi
having excipular or subcuticular, uni-loculate ascomata with
247
longitudinal irregular scratch-like openings and hyaline,
fusoid, two-celled ascospores with paraphyses (von Höhnel
1917; Holm and Holm 1977; Hyde et al. 2013). von Höhnel
(1917) included 12 genera in this family, Bifusella, Coccomyces, Duplicaria, Entopeltis, Haplophyse, Leptopeltella,
Leptopeltis, Lophodermina, Phacidina, Schizothyrioma,
Thyriopsis and Vizella. The family has a long historical discussion by various authors and many genera were included
and excluded from this family (Petrak 1947a, b; von Arx
and Müller 1975; Holm and Holm 1977; Eriksson 1981;
Lumbsch and Huhndorf 2010; Hyde et al. 2013). Holm and
Holm (1977) mentioned that taxa in Leptopeltidaceae have
unitunicate asci as they could not find the “Jack in the box”
dehiscence exhibited by Leptopeltis asci based on their
observation under the transmission electron microscope
(TEM). Eriksson (1981) disagreed with Holm and Holm
(1977) and demonstrated bitunicate asci with “Jack in the
box” dehiscence in Leptopeltidaceae as he had found the
endotunica and ectotunica layers of the asci. However, there
was no sharp delimitation between endotunica and ectotunica in ascomycetes. Hence, it is difficult to name the bitunicate or unitunicate asci in many cases of which Leptopeltis is
one of these cases (Eriksson 1981; Hyde et al. 2013).
Hyde et al. (2013) re-circumscribed genera in Leptopeltidaceae following the lists of Lumbsch and Huhndorf
(2010). Based on the generic type studies, Hyde et al. (2013)
accepted five genera in Leptopeltidaceae viz. Leptopeltis,
Dothiopeltis, Nannfeldtia Ronnigeria and Staibia and classified the family in Dothideomycetes, family incertae sedis.
Hyde et al. (2013) excluded Phacidina from Leptopeltidaceae as the type genus, P. gracilis has unitunicate asci,
with J+, subapical ring. However, Wijayawardene et al.
(2018) listed the genus in Leptopeltidaceae. We re-examine
the type of genera in Leptopeltidaceae and exclude Nannfeldtia from Leptopeltidaceae. The generic type of Nannfeldtia, N. atra has morphological similarity with taxa in Leotiomycetes in having apothecial ascomata, apically swollen
paraphyses and unitunicate asci, with J+ apex (Phookamsak,
pers. comm.). Therefore, we accept four genera: Leptopeltis,
Dothiopeltis, Ronnigeria and Staibia in Leptopeltidaceae.
However, these genera lack molecular data to clarify their
phylogenetic affinities, and their morphological characteristics are heterogeneous. A taxonomic revision of genera in
this family based on molecular data awaits study and will
resolve whether these taxa are Dothideomycetes or belong
in another class.
Leptopeltis Höhn., Ber. dt. bot. Ges. 35: 418 (1917).
Index Fungorum number: IF 2786; Facesoffungi number:
FoF 07970; 9 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Leptopeltis filicina (Lib.) Höhn., Ber. dt.
bot. Ges. 35: 422 (1917).
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Fig. 110 Morphological characteristics of genera in Leptopeltidaceae
based on the type studies (a, g, k, n, r–u = Leptopeltis filicina (≡
Aulographum filicinum); b, e, h, l, o, v, w = Dothiopeltis arunci; c, j,
p, x, y = Ronnigeria arctica (≡ Microthyrium arcticum Oudem.); d,
f, i, m, q, z, aa = Staibia connari). a–d Appearance of ascomata on
host surface. e, f Upper view of ascomata. g–i Upper view of perid-
ium showing cell arrangement. j–m Section through ascomata. n–q
Asci (n, o = stained by cotton blue). r Paraphyses stained by cotton
blue. s–aa Ascospores (s, t, x, y = stained by Melzer’s reagent, aa =
stained by cotton blue). Scale bars: e, l, m = 50 µm, f, h, i–k = 20
µm, g, n–r = 10 µm, s–aa = 5 µm
≡ Aulographum filicinum Lib., Pl. crypt. Arduenna, fasc.
(Liège) 3(nos 201–300): no. 275 (1834).
Notes Leptopeltis was introduced by von Höhnel (1917)
and is typified by L. filicina (≡ Aulographum filicinum 1834).
The genus is characterized by subcuticular or superficial,
elongate or Y-shaped thyriothecia, opening with slit-like
ostiole. The peridium is composed of a single layer of dark
brown to black, isodiametric cells, which are paler at the
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base. Asci are 8-spored, bitunicate, cylindrical to cylindricclavate, ellipsoidal to oblong, sessile, apically rounded, with
an indistinct ocular chamber, raising in between broad, aseptate paraphyses and ascospores are fusiform, oblong to cylindrical, or sometimes falcate to reniform, hyaline, 0–3-septate
(Hyde et al. 2013). The genus lacks a modern taxonomic
treatment and molecular data to investigate its phylogenetic
placement. The type species, L. filicina was collected from
Asplenium felix-mas in Belgium and is currently treated as
a synonym of Fouragea filicina (≡ Opegrapha filicina 1845)
in Opegraphaceae (Index Fungorum 2020, accessed 8 June
2020). However, the basionym of Leptopeltis filicina was
introduced earlier than the basionym of Fouragea filicina.
We therefore, reinstate Leptopeltis based on its generic type
being introduced earlier. The asexual morph of Leptopeltis
was reported as coelomycetous, leptothyrium-like, forming
in dimidiate or scutate pycnidial cavities, with small, onecelled conidia (von Arx and Müller 1975; Hyde et al. 2013;
Wijayawardene et al. 2017b). However, the connection of
the sexual and asexual morhs has not been proven (Fig. 110).
Other genera included
Dothiopeltis E. Müll., Sydowia 10(1–6): 197 (1957) [1956].
Index Fungorum number: IF 1700; Facesoffungi number:
FoF 07971; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Dothiopeltis arunci E. Müll., Sydowia
10(1–6): 198 (1957) [1956].
Notes: Dothiopeltis arunci was collected from Aruncus
silvester in Switzerland. The genus is epiphytic on dry stems.
Ascomata are superficial, rounded, lenticular, scattered to
clustered, shiny, lacking ostioles, forming brown, septate,
vegetative hyphal webs at the margin, immersed in host tissue and lack paraphyses. The upper wall of the peridium is
composed of thickened, dark, radially globose or angular
cells, with well-developed basal walls. Asci are 8-spored,
bitunicate, ellipsoidal to clavate, sessile to subsessile and
ascospores are hyaline, muriform, ellipsoidal to cylindrical
and occasionally clavate to obclavate (Müller 1956). Holm
and Holm (1977) found that D. arunci has longitudinal slitlike openings and they did not see the basal layers of the
peridium. The second species, D. cicerbitae Granmo &
Math. was found on stalks of Cicerbita alpina in Norway
and is distinct from D. arunci in several aspects (Granmo
and Mathiassen 2013). The asexual morph of Dothiopeltis
was reported as hyphomycetous, idriella-like which is characterized by hyaline or brown hyphae, brown, simple, aseptate geniculate conidiophores, tapering towards the tip and
lunate or falcate conidia with acuminate tips, produced in
dry heads (Nelson and Wilhelm 1956; Hyde et al. 2013;
Wijayawardene et al. 2017b).
Ronnigeria Petr., Sydowia 1(4–6): 310 (1947).
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Index Fungorum number: IF 4782; Facesoffungi number:
FoF 07972; – 1 morphological species (Species Fungorum
2020), molecular data unavailable
Type species: Ronnigeria arctica (Oudem.) Petr.,
Sydowia 1(4–6): 310 (1947).
≡ Microthyrium arcticum Oudem., Ned. kruidk. Archf,
2 sér. 3: 160 (1886).
Notes: Petrak (1947b) introduced a monotypic genus
Ronnigeria to accommodate Ronnigeria arctica (≡ Microthyrium arcticum), epiphytic on Potentilla fragiformis in
Novaya Zemlya, Russia (Holm and Holm 1977). The genus
is characterized by perithecial, subcuticular, gregarious,
amphigenous, orbicular to elliptical ascomata, lacking ostioles, opening by peridial cracks, with a thin-walled peridium, composed of a single layer of dark brown, membranous, pseudoparenchymatous cells and with paraphyses.
Asci are 8-spored, clavate, with rounded to truncate bases
and ascospores are hyaline, oblong to clavate, or subfusoid
(Petrak 1947b; Holm and Holm 1977). The asexual morph
of this genus is undetermined. Petrak (1947b) accommodated the genus in Leptopeltidaceae and this was followed
by various subsequent authors (von Arx and Müller 1975;
Holm and Holm 1977; Lumbsch and Huhndorf 2010; Hyde
et al. 2013; Wijayawardene et al. 2014a, 2018). The genus
has an arctic-alpine distribution and is commonly found on
Potentilla spp. in northern Scandinavia and central Europe
(Holm and Holm 1977; Farr and Rossman 2020). However,
the genus lacks modern taxonomic treatment and molecular
data to confirm its phylogenetic affinity. Recollection based
on type material is required for a better understanding of
this genus.
Staibia Bat. & Peres, in Batista et al., Atas Inst. Micol. Univ.
Pernambuco 3: 142 (1966).
Index Fungorum number: IF 5181; Facesoffungi number:
FoF 07973; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Staibia connari Bat. & Peres, in Batista
et al., Atas Inst. Micol. Univ. Recife 3: 142 (1966).
Notes: Staibia was introduced as a monotypic genus to
accommodate an epiphytic fungus occurring on leaves of
Connarus suberosus in Brazil and is characterized by subcuticular, dark brown, orbicular, dimidiate, membranous
ascomata, lacking ostioles and paraphyses. The upper wall
of the peridium is radiate, with longitudinally isodiametric
cells, endowed with multiple hymenium, black at the margin and with a poorly-developed base. Asci are 8-spored,
bitunicate and globose and ascospores are hyaline, oblong
to subovoid and 1-septate (Batista et al. 1966). The asexual
morph of this genus is undetermined. Batista et al. (1966)
treated the genus in Leptopeltidaceae and this was followed
by subsequent authors (von Arx and Müller 1975; Lumbsch
and Huhndorf 2010; Hyde et al. 2013; Wijayawardene et al.
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Fig. 111 Macrovalsaria megalospora (= M. leonensis PDD 14987,
co-extype). a Details of herbarium material. b Habit and appearance of ascomata on host surface. c Section of ascoma. d Peridium. e
Hamathecium. f Asci. g–i Ascospores. Scale bars: b = 1 mm, c, d =
200 µm, e = 10 µm, g–i = 20 µm
2014a, 2018). The genus is poorly known and lacks molecular data to reveal its taxonomic status, hence, it was tentative
placed in this family pending further studies.
apically rounded with a small ocular chamber. Hamathecium
comprising unbranched, tapering upwards, apically free,
paraphyses. Ascospores uni-seriate to irregularly uni-seriate, elliptical-fusoid, brown, 1-septate, slightly constricted
at septum, with skull cap-like germ apparatus at the lower
end, transverse striation near the center of lower cell with
longitudinal striations from transverse striation to the end
cell surface smooth, granular to verrucose. Asexual morph:
Undetermined.
Type: Macrovalsaria leonensis (Deighton) Petr.
Notes: Pem et al. (2019c) introduced Macrovalsariaceae
to accommodate Macrovalsaria. The family is characterized
by dark brown to black ascostromata, cylindro-clavate asci,
with a short fine pedicel and elliptical to fusoid ascospores
with skull cap-like germ apparatus.
Economic and ecological significance
Taxa in Leptopeltidaceae are commonly as epiphytic occurring on stems or leaves of ferns and some dicotyledonous
plants. Species in this family are mainly distributed in
Europe, except Staibia connari which was found on leaves
of Connarus suberosus in Brazil (Batista et al. 1966; von
Arx and Müller 1975; Holm and Holm 1977; Farr and Rossman 2020).
Macrovalsariaceae D. Pem, Doilom & K.D Hyde, in Pem
et al., Mycosphere 10(1): 1149 (2019).
Index Fungorum number: IF 557067; Facesoffungi number: FoF 06682, 2 species.
Saprobic on dead twigs, wood, bamboo and culms of
a wide range of hosts. Sexual morph: Ascostromata dark
brown to black, immersed to erumpent, solitary to a few
in a group, carbonaceous, oblate, sphaeroid to subsphaerical, with a central ostiole. Peridium comprising brown and
small-celled textura angularis. Asci 8-spored, bitunicate,
fissitunicate, cylindro-clavate, with a short fine pedicel,
13
Macrovalsaria Petr., Sydowia 15(1–6): 298 (1962) [1961].
Index Fungorum number: IF 2971; Facesoffungi number:
FoF 06251; 2 morphological species (Species Fungorum
2020), 2 species with molecular data.
Type species: Macrovalsaria megalospora (Mont.) Sivan.
Notes: Macrovalsaria is unique in producing brown, uniseptate ascospores that are constricted at the septum and the
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251
Fig. 112 Meliolina cladotricha (K 176503). a Herbarium packet. b The host leaf. c Ascomata on the leaf surface. d Squash mount of ascoma. e
Hyphae with dichotomously branched phialophores. f Ascus. g–i Ascospores. Scale bars: c = 400 μm, d = 100 μm, f = 20 μm, e, g–i = 10 μm
skull cap-like germ apparatus at the base (Sivanesan 1975).
The asexual morph is unknown.
This family is saprobic and thus involved in nutrient
recycling.
Macrovalsaria megalospora (Mont.) Sivan. Trans. Br.
Mycol. Soc. 65: 400 (1975).
≡ Sphaeria megalospora Mont., Annls Sci. Nat., Bot.,
sér. 2 14: 324 (1840).
Index Fungorum number: IF 317110; Facesoffungi number: FoF 01868; Fig. 111
Description: see Pem et al. (2019c).
Material examined: Africa, Sierra Leone, Njala (Kori), on
dead branches of Caesalpinia sappan (Fabaceae), 12 July
1949, F.C Deighton (PDD 14987, co-extype).
Meliolinaceae S. Hughes, Mycol. Pap. 166: 176 (1993).
Index Fungorum number: IF 81959; Facesoffungi numbers: FoF 06889; 50 species.
Parasitic on living leaves. Colonies superficial, dense
to subdense, black. Hyphae superficial, brown, straight to
substraight, branched, septate, darker at septa, reticulate,
with setiform phialides. Sexual morph: Ascomata globose
to subglobose, dense, gregarious, with a central ostiole, verrucose. Hamathecium lacking paraphyses. Asci 4–8-spored,
fissitunicate, ellipsoid to broadly clavate, evanescent.
Ascospores overlapping 2–4-seriate, cylindrical, hyaline
when young, becoming brown at maturity, 3-septate, end
Economic and ecological significance
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252
cells shorter and smaller, with hyaline subterminal bands.
Asexual morph: Undetermined.
Type: Meliolina Syd. & P. Syd.
Notes: Meliolinaceae comprises a single genus Meliolina
and 38 species, all but one of which are confined to members
of Myrtaceae. Reynolds (1989) described Briania fruticetum
as the mitosporic state of Meliolina, and this was accepted
by Kirk et al. (2008). The only phylogenetic study of Meliolinaceae was carried out by Saenz and Taylor (1999), illustrating that it is a separate family from Meliolaceae. Lumbsch and Huhndorf (2010) and Wijayawardene et al. (2018)
placed the family as Dothideomycetes incertae sedis.
Meliolina Syd. & P. Syd., Annls mycol. 12(6): 553 (1914).
Index Fungorum number: IF 3105; Facesoffungi number
FoF06890; 49 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Meliolina cladotricha (Lév.) Syd. & P. Syd.
Notes: Meliolina species produce thick, spongy, hypophyllous colonies that predominantly infect Myrtaceae
plants. Stevens (1927, 1928) included the genus in Meliolaceae. Eriksson (1981) postulated that Meliolaceae and
Meliolina may share a common ancestry, but the result from
Saenz and Taylor (1999) demonstrated that Meliolina is phylogenetically distant from Meliolaceae. However, Hyde et al.
(2013) treated this genus as a member of Meliolales based
on morphology. There are several morphological differences
between Meliola and Meliolina. Meliola species produce
haustoria from capitate hyphopodia, while Meliolina species penetrate the host by stomatopodia (Hansford 1946;
Hughes 1993) and develop internal hyphae. The superficial
mycelium of Meliola develops directly from an ascospore,
while the superficial mycelium of Meliolina is the result of
egress cells which grow out of leaf stomata and produce
mini-colonies of superficial hyphae (Hughes 1993). Meliola produce phialospores by possessing sessile phialides
(mucronate hyphopodia) which are produced directly from
the superficial hyphae, while Meliolina produces setiform
phialides which are produced from arborescent branches of
the superficial hyphae (Hughes 1981; Reynolds 1989; Mueller et al. 1991). Meliolina usually forms eight ascospores per
ascus (Hughes 1993), whereas Meliola never produces eight
mature ascospores within an ascus (Stevens 1925; Hansford
1961; Hongsanan et al. 2015d). The ascospores of Meliolina
usually display hyaline subterminal bands, which are never
found in Meliola.
Meliolina cladotricha (Lév.) Syd. & P. Syd., Annls mycol.
12(6): 553 (1914).
≡ Meliola cladotricha Lév., Annls Sci. Nat., Bot., sér. 3
5: 266 (1846).
Index Fungorum number: IF 120177; Facesoffungi numbers: FoF 06891; Fig. 112
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Parasitic on living leaves. Colonies hypophyllous, superficial, dense to subdense, black. Hyphae superficial, brown,
straight to substraight, branched, septate, darker at septa,
closely reticulate. Sexual morph: Ascomata globose to
subglobose, dense, gregarious, with a central ostiole, verrucose. Hamathecium lacking paraphyses. Asci 6–8-spored,
fissitunicate, ellipsoid to broadly clavate, with short pedicel,
evanescent. Ascospores overlapping 2–4-seriate, cylindrical
with somewhat flattened ends, hyaline when young, becoming brown at maturity, 3-septate, constricted and darken at
septa, end cells shorter and smaller than central cells, with
hyaline bands near the septum, with roughened walls. Asexual morph: Undetermined.
Material examined: Australia, Queensland, Cape York
Peninsula, on leaves of Agonis longifolia, L.J. Brass, 25
August 1948 (K 176503).
Other genus included
Briania D.R. Reynolds, Pacific Sci. 43(2): 161 (1989).
Index Fungorum number: IF 11017; Facesoffungi numbers: FoF 06892; 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Briania fruticetum D.R. Reynolds, Pacific
Sci. 43(2): 161 (1989).
Notes: Briania is a monotypic genus introduced as the
asexual morph of Meliolina sydowiana found on leaves of
Metrosideros polymorpha. The link between the sexual and
asexual morph are questionable.
Economic and ecological significance
Meliolinaceae is a poorly understood family with limited
taxa. Similar to black mildews and sooty moulds, their colonies cover the leaf surface and may reduce photosynthesis
and increase the temperature and respiration in those areas
(Hongsanan et al. 2015c).
Mesnieraceae Arx & Müller, Stud. Mycol. 9: 94 (1975).
Index Fungorum number: IF 80998, Facesoffungi numbers: FoF 07871, 6 species.
Synonym: Stegasphaeriaceae Syd. & P. Syd., Annls
mycol. 14(5): 364 (1916).
Parasitic on leaves, causing necrotic symptoms on leaves,
or saprobic. Sexual morph: Mycelium hyaline, developing
within host tissue. Ascomata densely gregarious, immersed,
sphaerical, flattened or globose to subglobose, yellowishgreen, fleshy, light with peridium composed of many cells,
uni-loculate, with numerous asci and pseudoparaphyses.
Ostiole present or lacking, usually opening apically with
a pore, or with wide and large hole in mature ascomata,
erumpent through the upper epidermis, comprising elongated columnar cells. Peridium thin-walled, hyaline, composed of many layers of flattened cells of textura angularis,
or comprising a few layers of elongate cells. Hamathecium
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253
comprising numerous, hypha-like, filamentous, septate
pseudoparaphyses, embedded in a gelatinous matrix. Asci
4–24-spored, bitunicate, cylindrical to clavate, or elongateellipsoidal, short-pedicellate, inner membrane thickened
in the upper part, apex rounded, with or without a welldeveloped ocular chamber. Ascospores 2- to multi-seriate,
ellipsoidal, dark brown to brown, or dark reddish-brown,
0–1-septate, thick-walled, constricted at the septum,
with or without a mucilaginous sheath. Asexual morph:
Undetermined.
Type: Mesniera Sacc. & P. Syd.
Notes: Mesnieraceae was introduced by von Arx and
Müller (1975) and type species is Mesniera rottlerae.
Mesnieraceae includes the genera Bondiella, Mesniera and
Stegasphaeria (Kirk et al. 2008). Hyde (1996) considered
Bondiella as a good member in Mesnieraceae (Pirozynski
1972; Eriksson 1981) and observed, redescribed and illustrated Bondiella palmicola from leaf blades of dried fallen
palm fronds, while Lumbsch and Huhndorf (2010) included
four genera with the addition of Helochora. Helochora was
isolated from Puya sp. in Chile, and described as a new
genus by Sherwood (1979) and was accommodated in Polystigmataceae. It has ornamented spores which are uni-seriate
in the asci. Hyde et al (2013) observed the type specimen
of Helochora hypertropha and considered that Helochora
does not belong to Mesnieraceae therefore Helochola was
transferred to Sordariomycetes genera incertae sedis.
Notes: Barr (1987b) placed Bondiella in Pleosporales,
but did not mention about Bondiella or Mesnieraceae. Hyde
(1996) observed type species Bondiella palmicola and considered Bondiella is a member of Mesnieraceae which supports the findings of Pirozynski (1972) and Eriksson (1981).
Mesniera Sacc. & Syd., Syll. Fung. 16: 440 (1902).
Index Fungorum number: IF 3134; Facesoffungi number:
FoF 07872; 3 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Mesniera rottlerae (Racib.) Sacc. & P. Syd.
[as ‘rotlerae’], Syll. fung. (Abellini) 16: 441 (1902).
≡ Anthostomella rottlerae Racib., Parasit. Alg. Pilze
Java’s (Jakarta) 2: 11 (1900).
Notes: Eriksson (1981) examined several collections
of Mesniera and all had concentric rings of ascomata in
necrotic patches, wide pores usually containing mature asci
with ascospores, and mature ascomata present on both of the
upper and lower side of the leaves. Unfortunately, the type
species of Mesniera rottlerae (≡ Anthostomella rottlerae)
may be lost (Hyde et al. 2013) and need recollecting and
neotypifying.
Stegasphaeria pavonina Syd. & P. Syd., Annls mycol.
14(5): 362 (1916).
= Mesniera pavonina (Syd. & P. Syd.) Petr., Annls mycol.
39(4/6): 345 (1941).
Index Fungorum number: IF 139864; Facesoffungi number: FoF 07876; Fig. 114
Description: see Hyde et al (2013).
Material examined: Philippines, Laguna, Mount Maquiling, near Los Baños, on living leaf of Macaranga sp., 7
March 1914 (C.F. Baker no.4032, holotype)
Other genera included
Bondiella Piroz., Mycol. Pap. 129: 6 (1972).
Index Fungorum number: IF 625; Facesoffungi number:
FoF 07873; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Bondiella palmicola Piroz., Mycol. Pap.
129: 6 (1972).
Bondiella palmicola Piroz., Mycol. Pap. 129: 6 (1972).
Index Fungorum number: IF 309834; Facesoffungi number: FoF 07874; Fig. 113
Description: see Hyde et al (2013).
Material examined: Tanzania, Kigoma, Kakombe, on
fallen fronds of Elaeis guineensis, 19 December 1963, K.A.
Pirozynski M26c, IMI 105789c, holotype).
Stegasphaeria Syd. & P. Syd., Annls mycol. 14(5): 362
(1916).
Index Fungorum number: IF 5190; Facesoffungi number:
FoF 07875; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Stegasphaeria pavonina Syd. & P. Syd.,
Annls mycol. 14(5): 362 (1916).
Notes: Stegasphaeria is pathogenic forming very rough
round spots on leaves, consisting of many individual ascomata arranged in narrow rings. The mature ascospore are
released through breakdown of the upper part of ascomata
as in Mesniera; this is the main character that places them
in Mesnieraceae.
Economic and ecological significance
Parasitic on leaves causing necrotic patches, this is important
as it can cause serious problems to economic plants. Fungi
can be saprobic over the winter on dead wood or leaves,
forming the sexual morph and waiting to infect the plant in
the next crop.
Naetrocymbaceae Höhn. [as ‘Naetrocymbeen’], Sber. Akad.
Wiss. Wien, Math.-naturw. Kl., Abt. 1 118: 1200 (1909).
Index Fungorum number: IF 81056; Facesoffungi number: FoF 07877, 41 species.
Epiphytic or Lichenicolous on twigs, wood, or on upper
and lower surface of living leaves, rarely on stone. Mycelium
on host surface, branched, septate, brown to grayish. Sexual
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254
Fig. 113 Bondiella palmicola (IMI 105789c, holotype). a Herbarium
specimens. b Ascomata semi-immerged on host. c Peridium. d Vertical section of ascoma. e Ostiolar canal with hyaline periphyses. f,
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h Asci. g Immature bitunicate asci. i, j Ascospores with 2 cells. k, l
Ascospores strained in India ink. Scale bars: d = 100 μm, e = 50 μm,
c, f–h = 20 μm, i, j = 10 μm, k, l = 5 μm
Fungal Diversity (2020) 105:17–318
255
Fig. 114 Stegasphaeria pavonina (C.F. Baker no.4032, holotype). a, b Herbarium specimen labels. c Ostiolate ascomata on leaf of Macaranga
sp. d Vertical section through ascoma. e, f Peridium. g, h Ascospores. Scale bars: d = 100 μm, e, f = 20 μm, g, h = 10 μm
morph: Ascomata perithecial or thyriothecial, superficial
to immersed, globose to subglobose, comprises a single to
multiple chambers, dark brown to black, ostiolate. Peridium
thin, comprises hyaline to dark brown cells of textura angularis to epidermoidea. Hamathecium comprising coarse,
branching pseudoparaphyses, hyaline, often with re- fractive or differentially-staining, cellular pseudoparaphyses
tips. Asci 6–8-spored, bitunicate, fissitunicate, obpyriform,
globose-subglobose or oblong, thick-walled near apex, with
a short pedicellate, apically rounded and endotunica, lacking
an ocular chamber. Ascospores 2–3 to multi- seriate, oblong
to broad ellipsoid, elongate to ellipsoid, or fusiform, hyaline,
olivaceous-brown when mature, 1- to multi-septate, slightly
constricted at septa, wall slightly ornamented. Asexual
morph: Microconidia short and rod-shaped, macroconidia
known from some Leptorhaphis species (Hyde et al. 2013).
Type: Naetrocymbe Körb.
Notes: The family (as Naetrocymbeen) was originally
described by von Höhnel (1909b), but without Latin or in
accordance with ICBN rules. Harris (1995) validated the
family name and included four genera in this family, Jarxia,
Leptorhaphis, Naetrocymbe, and Tomasellia. This was followed by Lumbsch and Huhndorf (2010), Hyde et al. (2013),
Doilom et al. (2018) and Wijayawardene et al. (2018). Pem
et al. (2019c) included Bonaria in this family based on morphological characters. Sequence data is not available for this
family.
Naetrocymbe Körb., Parerga lichenol. (Breslau) 5: 441
(1865).
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Fig. 115 Tomasellia arthonioides (F217044). a Herbarium materials. b Black stromata on the host. c, d Section of astroma. e Psuedoparaphyses. f, g Asci with ascospores. Scale bars: b = 500 µm, c, d = 50 µm, e–g = 10 µm
Index Fungorum number: IF 3410; Facesoffungi number:
FoF 07878; 25 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Naetrocymbe fuliginea Körb.
Notes: Most species in Naetrocymbe were transferred
to Limacinula (Reynolds 1971), however, Luttrell (1973)
noted that Naetrocymbe has questionable status. Naetrocymbe species are characterized by immersed, globose, black
ascomata, surrounded by dark brown fungal tissue, thickwalled, clavate to obovoid asci, and 1-septate, oblong to long
ellipsoid, hyaline ascospores. Naetrocymbe was accepted as
a distinct genus by Barr (1979a) and Eriksson (1981). Species in this genus were found as non-lichen-forming fungi
(Harris 1995) and found associated with Trentepohlia algae
(Roux 2009; Coppins 2002). Hyde et al. (2013) noted that
the concept of this genus is debatable.
Other genera included
Bonaria Bat., Publicações Inst. Micol. Recife 56: 438
(1959).
13
Index Fungorum number: IF 624; Facesoffungi number:
FoF 05164; – 3 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Bonaria lithocarpi (V.A.M. Mill. & Bonar)
Bat., Publicações Inst. Micol. Recife 56: 439 (1959).
≡ Protopeltis lithocarpi V.A.M. Mill. & Bonar, University of Calif. Publ. Bot. 19: 412 (1941).
Notes: Bonaria was treated as a genus incertae sedis in
Dothideomycetes by (Wijayawardene et al. 2018). Pem et al.
(2019c) re-examined the isotype specimen of B. lithocarpi
(S-F 3573) anf concluded that Bonaria should be placed
in Naetrocymbaceae based on its immersed, subglobose,
black ascomata, 8-spored, thick-walled, obpyriform asci,
and multi-seriate, oblong to long ellipsoid, hyaline, 1-septate
ascospores (Pem et al. 2019c).
Jarxia D. Hawksw., Stud. Mycol. 31: 93 (1989).
Index Fungorum number: IF 25310; Facesoffungi number: FoF 07879; – 2 morphological species (Species Fungorum 2020), molecular data unavailable.
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257
Type species: Jarxia thelenula (Müll. Arg.) D. Hawksw.,
Stud. Mycol. 31: 95 (1989).
≡ Microthelia thelenula Müll. Arg., Bot. Jb. 6: 416
(1885).
Notes: Jarxia contains two non lichen-forming species
(Doilom et al. 2018; Index Fungorum 2020). Doilom et al.
(2018) re-examined the syntype of J. thelenula which has the
same morphology as the holotype (Hawksworth 1989), thus
they agree to place this genus in Naetrocymbaceae.
Tomasellia arthonioides (A. Massal.) A. Massal., Flora,
Regensburg 39: 284 (1856).
≡ Arthopyrenia arthonioides A. Massal., Ric. auton. lich.
crost. (Verona): 169 (1852).
Index Fungorum number: IF 124738; Facesoffungi number: FoF 04652; Fig. 115
Description: see Doilom et al. (2018).
Material examined: Italy, Trentino-Alto Adige Province,
on the bark of Fraxinus ornus L., J. Milde. (S, F217044).
Leptorhaphis Körb., Syst. lich. germ. (Breslau): 371 (1855).
Index Fungorum number: IF 2795; Facesoffungi number:
FoF 07880; – 8 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Leptorhaphis oxyspora (Nyl.) Körb., Syst.
lich. germ. (Breslau): 371 (1855).
≡ Verrucaria oxyspora Nyl., Bot. Notiser: 179 (1852).
Notes: Leptorhaphis occurs on bark, with a high level
of host-specificity (Aguirre-Hudson 1991, 2009; AguirreHudson et al. 2002). This genus has been considered as saprotrophic and non lichen-forming (Vainio 1921a; Swinscow
1965a; Harris 1973, 1995; Aguirre-Hudson and Hawksworth
1987). One species is lichenicolous (Kalb et al. 1995), while
some species are associated with algae (Swinscow 1965a;
Aguirre-Hudson 1991, 2009; Aguirre-Hudson and Fiol
1993; Aguirre-Hudson et al. 2002). This genus has been
placed in Naetrocymbaceae by Harris (1995), AguirreHudson et al. (2002), and Doilom et al. (2018). However,
Harris (1995) mentioned that Leptorhaphis differs from
other genera in Naetrocymbaceae based on the characters
of hamathecium, asci, and macroconidia.
Economic and ecological significance
Species in Naetrocymbaceae are not harmful to plants or
animals. However, they have lichenicolous lifestyle which
is a parasitic fungus living on lichens as the host.
Tomasellia A. Massal., Flora, Regensburg 39: 283 (1856).
Index Fungorum number: IF 5489; Facesoffungi number:
FoF 07881; – 6 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Tomasellia arthonioides (A. Massal.) A.
Massal., Flora, Regensburg 39: 284 (1856).
Notes: The morphology of Tomasellia has often been discussed with reference to Mycoporum based on compound
ascomata containing several locules, each producing their
own ostiole (Harris 1995). These two genera can be distinguished based on ascus, ascospore, conidial and hamathecial characters (Harris 1995). Harris (1995) transferred most
species of this genus to Mycoporum (Mycoporaceae), and
accepted only five non lichen-forming species in Tomasellia. The generic delimitations in Harris (1995) have been
accepted by Sanderson and Coppins (2009a, b), while others accept a different circumscription (Aptroot 2002b, c).
Although, sequence data is not available for Tomasellia, it
is accepted as a genus in Naetrocymbaceae in many studies (Hyde et al. 2013; Wijayawardene et al. 2017a; Doilom
et al. 2018).
Nematotheciaceae Boonmee & K.D. Hyde, in Boonmee
et al., Mycosphere 8(10): 1728 (2017).
Index Fungorum number: IF 553836; Facesoffungi numbers: FoF 03700, 12 species.
Parasitic on lower surface of living leaves. Mycelium covered ascomata, with or without hyphopodia-like structures.
Sexual morph: Ascomata superficial, with a subiculum,
gregarious or solitary, globose to subglobose, dark brown,
with apical ostiole, covered by dark brown mycelium. Peridium comprises multilayers of dark brown cells of textura
angularis. Hamathecium comprising numerous, cylindrical, filiform, branched, septate, cellular pseudoparaphyses.
Asci 8-spored, bitunicate, cylindrical-subclavate, sessile.
Ascospores fasciculate, multi-seriate, elongate fusiform,
curved or flexuous, tapering towards the acute ends, brown,
multi-septate. Asexual morph: Undetermined (adapted from
Boonmee et al. 2017).
Type: Nematothecium Syd. & P. Syd.
Notes: Nematotheciaceae was established based on morphology by Boonmee et al. (2017) to accommodate three
parasitic genera Nematothecium (type genus), Nematostigma
and Ophioparodia. Boonmee et al. (2017) mentioned that
Nematotheciaceae is similar to Tubeufiaceae in the features
of ascomata, asci and ascospores, however, it differs from
Tubeufiaceae by being parasitic and occurring on living
leaves. Sequence data is not available for this family.
Nematothecium Syd. & P. Syd., Leafl. of Philipp. Bot. 5(no.
76): 1534 (1912).
Index Fungorum number: IF 3444; Facesoffungi number:
FoF 07882; 5 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Nematothecium vinosum Syd. & P. Syd.
Notes: The genus was introduced from the Island of Palawan in the Philippines. Nematothecium is characterized by
large mycelium colonies, irregular, superficial ascomata
with dark red ascomata, bitunicate asci and elongate fusiform, curved or flexuous, multi-septate, hyaline to red brown
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258
Fig. 116 Nematothecium vinosum (S-F10269, holotype). a Herbarium specimen and habit on leaves. b Appearance of ascomata on
leaf surface. c Section of ascoma. d Peridium. e Pseudoparaphyses. f
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Arrangement of asci. g–j Asci. k, l Ascospores. Scale bars: b = 500
μm, c = 200 μm, f–l = 50 μm, d = 40 μm, e = 5 μm
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ascospores. Nematothecium was placed in Pseudoperisporiaceae by Kirk et al. (2008), Lumbsch and Huhndorf (2010),
and Hyde et al. (2013). Boonmee et al. (2017) re-examined
the holotype and lectotype of N. vinosum and concluded
that the shape and septation of ascospores are typical of
Tubeufiaceae, however, its habitat can differentiate them
from Tubeufiaceae. Therefore, Nematothecium is placed in
the new family Nematotheciaceae (Boonmee et al. 207).
Nematothecium vinosum Syd. & P. Syd., Leafl. of Philipp.
Bot. 5(no. 76): 1534 (1912).
Index Fungorum number: IF 249022; Facesoffungi number: FoF 03243, Fig. 116
Description: see Boonmee et al. (2017).
Material examined: Philippines, Palawan, Island of Palawan: Puerto Princesa, Mt Pulgar, on living leaves of Eugenia
incarnata Elmer. (Myrtaceae), May 1911, A.D.E. Elmer (no.
13232), (S-F10269, holotype, BPI 699632, lectotype).
Other genera included
Nematostigma Syd. & P. Syd., Annls mycol. 11(3): 262
(1913).
Index Fungorum number: IF 3442; Facesoffungi number:
FoF 07883; – 6 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Nematostigma obducens Syd. & P. Syd.,
Annls mycol. 11(3): 262 (1913).
Notes: The genus is characterized by dark brown, sparse
setae, sphaerical ascomata, and hyaline, multi-septate
ascospores. Nematostigma was transferred to Dimeriaceae,
Parodiopsidaceae (= Parodiellinaceae), and Pseudoperisporiaceae (Hansford 1946; von Arx and Müller 1975; Eriksson
and Hawksworth 1993; Hawksworth et al. 1995; Barr 1997;
Hyde et al. 2013). However, Boonmee et al. (2017) included
this genus in Nematotheciaceae based on morphology.
Ophioparodia Petr. & Cif., Annls mycol. 30(3/4): 223
(1932).
Index Fungorum number: IF 3610; Facesoffungi number:
FoF 07884; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Ophioparodia pulchra Petr. & Cif., Annls
mycol. 30(3/4): 223 (1932).
Notes: The genus was introduced to accommodate a single species Ophioparodia pulchra, with septoidium as its
asexual morph. Ophioparodia was placed in Parodiopsidaceae by Sivanesan (1984) based on its septoidium-like
conidia. However, Boonmee et al. (2017) re-examinded
the type specimens and could not find pseudoparaphyses
or the asexual morph. They included Ophioparodia in a
new family Nematotheciaceae as it has superficial, globose
to subglobose, uniloculate ascomata, cylindrical asci and
259
long, multi-septate ascospores and a hyphomycetous asexual
morph (Boonmee et al. 2017).
Economic and ecological significance
Species in Nematotheciaceae are found as parasitic on leaves
which can uptake nutrients from plants.
Neoparodiaceae Boonmee & K.D. Hyde, in Boonmee et al.,
Mycosphere 8(10): 1734 (2017).
Index Fungorum number: IF 553832; Facesoffungi numbers: FoF 03683, 1 species.
Epiphytic or parasitic on living leaves. Sexual morph
Colonies up to 2 mm diam. appear as black spots, superficial, subcircular, outwardly radiating, dense, branched, septate, with dark brown mycelia. Ascostromata epiphyllous or
hypophyllous, superficial, crustose, on a subiculum, solitary,
scattered, dark brown, multi-loculate, comprises subglobose
locules, thickened at the base. Peridium comprises hyaline
to brown cells of textura angularis. Hamathecium lacking pseudoparaphyses. Asci 8-spored, bitunicate, broadly
clavate, oblong to obovoid, sessile or with short pedicel,
apically thickened, with an acute to subacute ocular chamber. Ascospores overlapping 2-seriate, conglobate, broadly
ellipsoid, ends rounded, each cell subglobose, reddish-brown
to dark brown, separated by light brown regions, 1-septate.
Asexual morph: Undetermined (adapted from Boonmee
et al. 2017).
Type: Neoparodia Petr. & Cif.
Notes: Neoparodiaceae was established to accommodate
a single genus Neoparodia based on morphology by Boonmee et al. (2017). This family shares some characters with
Cookellaceae and Myriangiaceae (Hyde et al. 2013; Dissanayake et al. 2014; Boonmee et al. 2017). However, Neoparodiaceae differs from Cookellaceae and Myriangiaceae
in having ascostromata, broadly clavate asci and broadly
ellipsoid, 1-septate, dark brown ascospores (Boonmee et al.
2017).
Neoparodia Petr. & Cif., Annls mycol. 30(3/4): 219 (1932).
Index Fungorum number: IF 3471; Facesoffungi number:
FoF 07885; 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Neoparodia ekmanii Petr. & Cif.
Notes: Neoparodia was placed in Perisporiopsidaceae by
Müller and von Arx (1962), but later it was transferred to
Parodiopsidaceae (Kirk et al. 2008; Lumbsch and Huhndorf 2010; Hyde et al. 2013; Wijayawardene et al. 2014a).
Boonmee et al. (2017) examined the holotype of N. ekmanii
and its asexual morph, and introduced a new family Neoparodiaceae to accommodate this genus based on its distinct
morphological characters.
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260
Fig. 117 Neoparodia ekmanii (W Krypto 1977–0019515, holotype).
a Herbarium specimen and habit on leaves. b Appearance of colony
and ascostromata seated on mycelium. c Close up of ascostroma. d,
e Squash mount of ascostroma. f Section of ascomal locules. g, h
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Close up of ascomal locules. i Anastomosing and branched mycelium
and apical conidium-like spores. j–l Immature and mature asci. m–o
Immature and mature ascospores. Scale bars: b = 1 mm, c–h = 100
μm, i–l = 50 μm, m–o = 10 μm
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Neoparodia ekmanii Petr. & Cif. Annls mycol. 30(3/4): 219
(1932).
Index Fungorum number: IF 263814; Facesoffungi number: FoF 03684, Fig. 117
Description: see Boonmee et al. (2017).
Material examined: Dominican Republic, Santiago, El
Cerrazo, on living leaves of Trema micrantha (Ulmaceae),
19.53°N, 70.62°W elev. 800 msl, 20 February 1930, P.L.
Clemen No. 3518 (W Krypto 1977–0019515, holotype).
Economic and ecological significance
The family is poorly studied which comprises a single species and it is epiphytic or parasitic on living leaves. Its habitats and distributions have not been reported.
Palawaniaceae Mapook & K.D. Hyde, in Mapook et al.,
Mycosphere 7(11): 1737 (2016).
Index Fungorum: IF 552528; Facesoffungi number: FoF
02653, 3 species.
Saprobic on surface of dead rachides and spines of Arecaceae. Sexual morph: Ascomata superficial, solitary or
scattered, coriaceous, appearing as circular, scattered, flattened, dark brown to black spots, covering the host, without
a subiculum, with a poorly developed basal layer and an
irregular margin. Ostiole central. Peridium comprising of
dark brown or black to light brown cell of textura epidermoidea at top with light brown cells of textura angularis at
side. Hamathecium comprising cylindrical to filiform, septate, cellular pseudoparaphyses. Asci 8-spored, bitunicate,
inequilateral to ovoid, pedicellate, straight or slightly curved,
without small ocular chamber. Ascospores overlapping,
irregularly arranged, broadly fusiform, hyaline, 1-septate,
constricted at the septum, guttulate, surrounded by hyaline
gelatinous sheath, observed clearly when mounted in Indian
ink. Asexual morph: Undetermined.
Type: Palawania Syd. & P. Syd.
Notes: The family was placed in Muyocopronales, based
on phylogeny and morphology, together with the use of
divergence times as additional evidence for the introduction of this family (Mapook et al. 2016c). Palawania species share some morphological characters with Muyocopron species, but differ in its peridium, shape of asci and
ascospores which are surrounded by a hyaline gelatinous
sheath, while the character was not found in Muyocopron.
In the analyses of Hongsanan et al. (2020, Fig. 1), Palawaniaceae has uncertain phylogenetic placement. Thus, we place
this family in Dothideomycetes families incertae sedis, until
sequence of type species is available.
Palawania (Niessl) Syd. & P. Syd., Philipp. J. Sci., C, Bot.
9(2): 171 (1914).
261
Index Fungorum number: IF 3682; Facesoffungi number:
FoF 06426; 3 morphological species (Species Fungorum
2020), 1 species with molecular data.
Type species: Palawania grandis (Niessl) Syd. & P. Syd.,
Philipp. J. Sci., C, Bot. 9(2): 171 (1914).
≡ Microthyrium grande Niessl, in Rabenhorst, Fungi
europ. exsicc.: no. 2467 (1876).
Notes: Species in this genus are known from the genera
Calamus, Cocos and Dypsis (Arecaceae). There are eight
species epithets listed in Index Fungorum (2020), while five
species were transferred to Parmulariaceae. The type material was re-examined by Wu et al. (2011b). Recently, a fresh
collection was described and illustrated by Mapook et al.
(2016c) from Thailand.
Palawania thailandensis Mapook & K.D. Hyde, in Mapook
et al., Mycosphere 7(11): 1740 (2016).
Index Fungorum number: IF 552527; Facesoffungi number: FoF 02654; Fig. 118
Description: see Mapook (2016c).
Material examined: Thailand, Chiang Rai, Chiang Rai
Horticultural Research Center, on dried petiole and rachis
of Phoenix roebelenii (Arecaceae), 09 September 2014, A.
Mapook, (MFLU 16-1873, paratype).
Notes: Palawania thailandensis was introduced as a new
species in Mapook et al. (2016c) with molecular data to
confirm the phylogenetic placement of the genus. The species clusters with the Muyocopronaceae clade and diverged
at 172 MYA in the Jurassic period (Mapook et al. 2016c).
Economic and ecological significance
Members in this family are usually common on dried petioles and rachiches of palms (Arecaceae), and are mostly saprobes (Batista and Vital 1960; Mapook et al. 2016c; Sydow
and Sydow 1914b; Wu et al. 2011b).
Paranectriellaceae Boonmee & K.D. Hyde, in Hyde et al.,
Fungal Diversity 63: 186 (2013).
Index Fungorum number: IF 804521, Facesoffungi number: FoF 07886, 24 species.
Parasitic (or biotrophic, hyperparasitic) on leaves causing slight discoloration around fruiting bodies. Sexual
morph: Ascostromata superficial, crustose, solitary, scattered, white to light orange, fleshy, soft in texture, surrounded by colourless, sparse hairs, multi-loculate, with
ascomata arranged in a peripheral outer layer. Locules
globose to subglobose, with individual ostioles. Peridium
relatively thin, composed of light yellow, thick-walled cells
of textura angularis. Hamathecium comprising filiform,
septate, branched, pseudoparaphyses, extending over asci
and embedded in a gelatinous layer. Asci 8-spored, bitunicate, fissitunicate, cylindrical, oblong-clavate, apedicellate,
rounded at the apex. Ascospores 2-seriate, broadly fusiform,
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262
Fig. 118 Palawania thailandensis (MFLU 16-1873, paratype). a–c
Superficial ascomata on substrate. d, e Squash mounts showing ascomata walls. f Section of ascoma. g Ostiole h Peridium. i Pseudoparaphyses. j–m Asci. n–r Ascospores. s Ascospores surrounded by hya-
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line gelatinous sheath in Indian ink. Scale bars: b = 500 µm, c = 200
µm, d = 100 µm, f, j–m = 50 µm, g = 20 µm, e, h, n–s = 10 µm, i =
5 µm
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Fig. 119 Paranectriella juruana (BPI 632134). a Material and habit
on leaf. b Appearance of ascostromata on lower leaf surface. c Section of ascostroma. d Close up of peridium. e Pseudoparaphyses. f
263
Asci. g, h Ascospores. Scale bars: a, e = 5 mm, b = 500 μm, c = 200
μm, d, f = 20 μm, g, i = 10 μm
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264
hyaline, narrowed at the ends, multi-septate, slightly constricted at the septa, with apical spine-like appendages.
Asexual morph: Hyphomycetous, staurosporous.
Type: Paranectriella (Henn. ex Sacc. & D. Sacc.)
Magnus.
Notes: Paranectriellaceae was introduced to accommodate two hyperparasitic genera Paranectriella and Puttemansia (Hyde et al. 2013; Boonmee et al. 2014b). Rossman et al.
(2015) proposed to protect Paranectriella over Araneomyces
because Paranectriella was more widely reported and most
species were not linked with Araneomyces. Currently, Paranectriellaceae is placed in Dothideomycetes families incertae sedis (Wijayawardene et al. 2017a, 2020). Sequence data
are needed to confirm the placement of taxa in this family.
Paranectriella (Henn. ex Sacc. & D. Sacc.) Magnus., Botan.
Zbl. 98: 587 (1905).
= Araneomyces Höhn., Sber. Akad. Wiss. Wien, Math.naturw. Kl., Abt. 1 118: 894 (1909).
= Paranectria subgen. Paranectriella Henn. ex Sacc. &
D. Sacc., Syll. fung. (Abellini) 17: 812 (1905).
Index Fungorum number: IF 3708, Facesoffungi number:
FoF 07887, 8 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Paranectriella juruana (Henn.) Höhn.,
Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 119:
899 (1910).
Notes: Paranectriella was introduced to accommodate
Paranectriella juruana a tropical hyperparasitic. Araneomyces was accepted as the asexual state of Paranectriella
(Sutton 1984; Wu et al. 1997). Wijayawardene et al. (2014a)
synonymized Araneomyces under Paranectriella.
Paranectriella juruana (Henn.) Höhn., Sber. Akad. Wiss.
Wien, Math.-naturw. Kl., Abt. 1 119: 899 (1910).
≡ Paranectria juruana Henn., Hedwigia 43(4): 245
(1904).
Index Fungorum number: IF 319198, Facesoffungi number: FoF 07888, Fig. 119
Description: see Hyde et al. (2013).
Material examined: Haiti, Dept. de la Grand’ anse, Massif de la Hotte, “Geffrard” 44 km south of Roseaux Road
to Camp Perrin, alt. 780m, 18°25′N73°53′W, on leaves of
Miconia sp., William R. Buck (9169), 14 November 1982,
determined by A.Y. Rossman, (BPI 632134).
Other genera included
Puttemansia Henn., Hedwigia 41: 112 (1902).
Index Fungorum number: IF 4562, Facesoffungi number:
FoF 00217; – 16 morphological species, (Species Fungorum
2020), molecular data unavailable.
Type species: Puttemansia lanosa Henn., Hedwigia 41:
112 (1902).
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Notes: Puttemansia was introduced to accommodate parasitic or hyperparasitic taxa occurring on leaves (Hennings
1902). Its asexual morphs are hyphomycetous in the genera
Guelichia and Tetranacrium and may need synonymizing
(Hyde et al. 2011, 2013). Puttemansia is presently placed in
Paranectriellaceae (see morphological description in Boonmee et al. 2014b). The genus needs sequence data to indicate
the generic placement.
Economic and ecological significance
Paranectriellaceae comprises two genera and their type species occurring on living leaves of dicotyledonous and monocotyledonous plants and are distributed widely in tropical
regions, mostly in Africa and South America (Hyde et al.
2013; Boonmee et al. 2014b).
Parodiellaceae Theiss. & Syd. ex M.E. Barr, Mycotaxon
29: 503 (1987).
Index Fungorum number: IF 81844; Facesoffungi numbers: FoF 07889, 23 species.
Biotrophic on upperside of leaves. Sexual morph:
Ascomata superficial, scattered, gregarious between veins,
uniloculate, subglobose to globose, with short stalk when
mature, smooth or verrucose, black, with blackened fungal
tissue in the leaf below stalk. Central ostiole with periphyses. Peridium thick-walled, comprising brown to dark brown
cells of textura angularis at outer layer, and pale brown,
flattened cells towards the inner layer. Hamathecium comprising irregular pseudoparaphyses. Asci 8-spored, bitunicate, cylindrical to clavate, with broad, with lobed pedicel,
apically rounded with ocular chamber. Ascospores overlapping 1–2-seriate, fusoid to ellipsoidal, brown to dark
brown, 1-septate, constricted at septum, inequilateral, upper
cell broader and longer than lower cell, slightly narrowed
at the ends, smooth-walled or striate, lacking a mucilaginous sheath. Asexual morph: Coelomycetous, assumed as
Ascochytopsis. Conidiomata thick-walled, black. Conidiogenous cell cylindrical, proliferating percurrently. Conidia
straight to falcate, hyaline, aseptate (Sivanesan 1984; Cannon and Kirk 2007; Hyde et al. 2013).
Type: Parodiella Speg.
Notes: Parodiellaceae was invalidly introduced by Theissen and Sydow (1918) and was validated by Barr (1987b).
Hyde et al. (2013) provided an updated description based
on the syntype of Parodiella hedysari (K(M) 176017) from
Kew collection to represent Parodiellaceae. Parodiellaceae
comprises a single genus, Parodiella and is treated as Dothideomycetes families incertae sedis (Wijayawardene et al.
2018). Sequence data is not available for this family.
Parodiella Speg., Anal. Soc. cient. argent. 9(4): 178 (1880).
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265
Fig. 120 Parodiella hedysari (K(M) 176017, syntype). a Herbarium
specimens. b, c Ascomata on host. d–f Vertical section through the
ascoma and peridium. g, h Bitunicate asci. i Pedicel of ascus. j–l
Ascospores. Scale bars: d =200 μm, c = 100 μm, e, f =50 μm, g–i =
20 μm, j–l =10 μm
Index Fungorum number: IF 3749 Facesoffungi numbers:
FoF 07890; 23 morphological species (Species Fungorum
2020), molecular data unavailable.
Type speices: Parodiella perisporioides (Berk. & M.A.
Curtis) Speg.
Notes: Parodiella is monotypic genus in Parodiellaceae (Lumbsch and Huhndorf 2010), typified by P. perisporioides. Barr (1987b) and Kirk et al. (2008) included
Pododimeria and Neopeckia in Parodiellaceae, respectively.
However, Wijayawardene et al. (2018) accepted only a single
genus in this family. The type specimen was not located,
therefore, Hyde et al. (2013) illustrated Parodiella hedysari.
Parodiella hedysari (Schwein.) S. Hughes, Can. J. Bot.: 75
(1958).
≡ Coryneum hedysari Schwein., Trans. Am. phil. Soc.,
New Series 4(2): 306 (1832) [1834].
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Fig. 121 Perisporiopsis struthanthi (S-F9814, holotype). a Material
and habit on leaves. b Appearance of ascomata. c Superficial mycelium. d Sections of ascoma. e Peridium. f Hamathecium pseudopara-
physes. g–i Asci. j, k Ascospores. Scale bars: b = 200 μm, c, j, k =
20 μm, d = 100 μm, e = 40 μm, f = 10 μm, g–i = 50 μm
Index Fungorum number: IF 302296; Facesoffungi numbers: FoF 07891; Fig. 120
Description: see Hyde et al. (2013).
Material examined: USA, South Carolina, on leaves
of Rhynchosia sp., 1879, H.W Ravenel s.n. [ex herb M.J.
Berkeley] (K(M) 176017, syntype).
colonize leaf surfaces and this may produce some compound
against other fungus/microbe.
Economic and ecological significance
Parodiella is a biotrophic fungus found in several hosts especially Leguminosae, and in different areas such as Australia,
Brazil, Columbia, India, Japan, Kenya and Philippines.
Based on the type specimen of Parodiella hedysari, it can
13
Perisporiopsidaceae E. Müll. & Arx ex R. Kirschner & T.A.
Hofm., in Kirschner et al., Sydowia 62(2): 238 (2012).
= Perisporiopsidaceae E. Müll. & Arx, Beitr. Kryptfl.
Schweiz 11(no. 2): 167 (1962).
Index Fungorum number: IF 567164, Facesoffungi humber: FoF 07892, 23 species.
Parasitic, hyperparasitic or pathogenic on living leaves
of various plants. Sexual morph: Ascomata or ascostromata, superficial, gregarious, solitary, uni to multi-locules,
Fungal Diversity (2020) 105:17–318
seated on a subiculum, sometimes surrounded by aerial
mycelium, globose to subglobose, obovoid, bright to dark
pigmented, with or without apical ostiole. Peridium relatively thick-walled, comprising hyaline and bright to dark
brown cells of textura angularis. Hamathecium comprising
branched, septate, anastomosed, hyaline, cellular pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, broadly
ellipsoid, apically thickened, with or without ocular chamber, short pedicellate. Ascospores 2–3-seriate, ellipsoidaloblong or fusiform widest in the middle to upper part of the
apical cell, with broadly rounded apex and tapering towards
the ends, slightly curved, 0–4-septate, hyaline to light pigmented, sometimes pale-yellow brown, smooth- to roughwalled. Asexual morph: Hyphomycetous.
Type: Perisporiopsis Henn.
Notes: Perisporiopsidaceae includes only five genera
Asteronia, Byssocallis, Chevalieropsis, Parodiellina and
Perisporiopsis and is treated in Dothideomycetes as family
incertae sedis (Boonmee et al. 2017; Wijayawardene et al.
2018; Index Fungorum 2020). Taxa in this family are parasitic on the surface of living leaves. Phylogenetic analyses
are known from putative Perisporiopsis strains (endophytic
taxa) and place the strains close to some genera in Pleosporales, but with no detail of morphology (Chaverri and Gazis
2010; Hyde et al. 2013). Furthermore, other sequence strains
in GenBank are unpublished. Therefore, Perisporiopsidaceae is of uncertain taxonomic placement.
Perisporiopsis Henn., Hedwigia 43(2): 83 (1904).
Index Fungorum number: IF 3827, Facesoffungi number:
FoF 07893; 17 morphological species, (Species Fungorum
2020), molecular data unavailable.
Type species: Perisporiopsis struthanthi Henn.
Notes: Perisporiopsis was revisited by Boonmee et al.
(2017) and provided with a description and illustration of
the type species P. struthanthi from the holotype (S-F9814).
267
Type species: Asteronia sweetiae Henn., Hedwigia 34:
104 (1895).
= Parodiopsis sweetiae (Henn.) G. Arnaud, Annls Épiphyt. 7: 53 (1921).
Notes: Wu et al. (2010) revisited Asteronia and provided
a description and illustration of the holotype of A. sweetiae
(K (M) 159800) and suggested to place this genus in families Asterinaceae or Meliolaceae more suitable than place
in Microthyriaceae. Later, Wu et al. (2011b) transferred
Asteronia to Venturiales incertae cedis. Pem et al. (2019c)
provided description and illustration of this genus based on
the isotype of A. sweetiae (S-F46114). Its morphology and
substrate are consistent with genera in Perisporiopsidaceae.
Therefore, Asteronia was transferred to Perisporiopsidaceae
by Pem et al. (2019c). Furthermore, species in Asteronia
need sequence data to indicate the generic placement.
Byssocallis Syd., Annls mycol. 25(1/2): 14 (1927).
Index Fungorum number: IF 700, Facesoffungi number:
FoF 06224; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Byssocallis phoebes Syd., Annls mycol.
25(1/2): 14 (1927).
Notes: Boonmee et al. (2011) excluded this genus from
Perisporiopsidaceae based on morphology of isolectotype specimen (F10891). Pem et al. (2019c) re-examined
the syntype specimen of Byssocallis phoebes (E00455471)
and found that it shares common characters with genera in
Perisporiopsidaceae such as occur on leaf surface, superficial ascomata with surrounding mycelium and ellipsoidal
oblong, multi-septate, hyaline ascospores. Thus, they transferred Byssocallis to Perisporiopsidaceae.
Perisporiopsis struthanthi Henn., Hedwigia 43(2): 83
(1904).
Index Fungorum number: IF 216818, Facesoffungi number: FoF 03687; Fig. 121
Description: see Boonmee et al. (2017).
Material examined: Brazil, Rio de Janeiro, Estado de
Rio de Janeiro, Serra dos Orgaos, on leaves of Struthanthus sp. (Apocynaceae), August 1899, E.H.G. Ule No. 2631
(S-F9814, holotype).
Chevalieropsis G. Arnaud, Annls Épiphyt. 9: 2 (1923).
Index Fungorum number: IF 992, Facesoffungi number:
FoF 07894; – 1 morphological species (Speices Fungorum
2020), molecular data unavailable.
Type species: Chevalieropsis ctenotricha (Pat. & Har.)
G. Arnaud, Annls Épiphyt. 9: 2 (1923).
≡ Dimerosporium ctenotrichum Pat. & Har., J. Bot., Paris
14: 242 (1900).
Notes: Chevalieropsis is placed in Perisporiopsidaceae
based on its parasitic habit on leaves and shares some characteristics that unite the genus into this family, such as ascomata (or ascostromata) features, thick-walled bitunicate asci
and lightly pigmented ascospores.
Other genera included
Asteronia (Sacc.) Henn., Hedwigia 34: 104 (1895).
Index Fungorum number: IF425, Facesoffungi number:
FoF 06215; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Parodiellina Henn. ex G. Arnaud, Annals d’École National
d’Agric. de Montpellier, Série 2 16(1–4): 21 (1918) [1917].
Index Fungorum number: IF 3750, Facesoffungi number:
FoF 07895; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
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268
Type species: Parodiellina manaosensis (Henn.) G.
Arnaud, Annals d’École National d’Agric. de Montpellier,
Série 2 16(1–4): 21 (1918) [1917].
≡ Parodiella manaosensis Henn., Hedwigia 43(6): 358
(1904).
Notes: Boonmee et al. (2017) revisited Parodiellina and
provided a description and illustration of the holotype of
P. manaosensis (PC0084492). Parodiellina is placed in
Perisporiopsidaceae based on its parasitic habit on leaves
and shares some characteristics that unite the genus into
Perisporiopsidaceae.
Economic and ecological significance
Taxa in this family are parasitic or hyperparasitic on living
leaves and occur on upper and lower leaf surfaces. They are
widespread in temperate to subtropical and tropical regions
such as in Central America and South America (Boonmee
et al. 2011, 2017; Hyde et al. 2013; Pem et al. 2019c).
Phaeodimeriellaceae Boonmee, Mapook & K.D. Hyde, in
Boonmee et al., Mycosphere 8(10): 1748 (2017).
Index Fungorum number: IF 553833; Facesoffungi numbers: FoF 03689, 27 species.
Epiphytic on living leaves, or associated with other
fungi on leaves. Sexual morph: Ascomata superficial, solitary or scattered, coriaceous, globose to subglobose, dark
brown, surrounded by brown to dark brown mycelium at the
base, apical ostiole, with hook-like and dark brown setae.
Peridium brown to dark brown cells of textura angularis.
Hamathecium comprising cylindrical to filiform, branched,
septate, cellular pseudoparaphyses. Asci 8-spored, bitunicate, oval to ellipsoidal, apex rounded, short pedicellate,
with a small ocular chamber. Ascospores 2–3-seriate, oblong
to broadly fusiform, hyaline when immature and pale brown
to brown at maturity, 1-septate, with a mucilaginous sheath.
Asexual morph: Coelomycetous. Conidiomata pycnidial,
superficial, globose to subglobose. Conidiomatal setae long,
hook-like, aseptate, dark brown, with blunt apex. Peridium
comprises brown cells of textura angularis. Conidiophores
reduced to conidiogenous cells. Conidiogenous cells enteroblastic, monophialidic. Conidia fusiform, hyaline, aseptate,
guttulate (adapted from Boonmee et al. 2017).
Type: Phaeodimeriella Speg.
Notes: Mapook et al. (2016a) provided sequence data
for Phaeodimeriella cissampeli and P. dilleniae, and indicated that both species clustered in a distinct clade within
Dothideomycetes. Although, Mapook et al. (2016a) treated
Phaeodimeriella as a genus in Pseudoperisporiaceae,
Boonmee et al. (2017) excluded this genus and placed it in
Phaeodimeriellaceae.
Phaeodimeriella Speg., Revta Mus. La Plata 15(2): 13
(1908).
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Index Fungorum number: IF 3908; Facesoffungi number:
FoF 01927; 27 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Phaeodimeriella parvula (Cooke) Hansf.
Notes: Phaeodimeriella was lectotypified by P. occulta
(Theissen 1912). However, P. occulta was synonymized
under P. parvula by Hansford (1946), therefore the current
name of the type species is P. parvula. Mapook et al. (2016a)
provided molecular data of P. cissampeli and P. dilleniae.
In their analyses, the clade of Phaeodimeriella is related to
the clade of Lentitheciaceae with moderate support. However, Phaeodimeriella differs from genera in Lentitheciaceae
(see Mapook et al. 2016a). Boonmee et al. (2017) included
this genus in the new family Phaeodimeriellaceae based on
its morphology and phylogeny. Phaeodimeriella comprises
three species, P. parvula, P. dilleniae and P. cissampeli.
Phaeodimeriella parvula (Cooke) Hansf., Mycol. Pap. 15:
64 (1946).
≡ Dimerosporium parvulum Cooke, Grevillea 20(no. 93):
5 (1891).
Index Fungorum number: IF 289280; Facesoffungi number: FoF 07896; Fig. 122
Desctiption: see Boonmee et al. (2017).
Material examined: Australia, Queensland, on leaves of
Trema tomentosa var. viridis (Cannabaceae). Coll. Bailey
902, Det. A. Sivanesan, (K(M) 181461, holotype).
Economic and ecological significance
Species in Phaeodimeriellaceae are epiphytic on living
leaves, however, they do not cause serious problem on
plants. Their life style is unclear.
Pododimeriaceae Boonmee & K.D. Hyde, in Boonmee
et al., Mycosphere 8(10): 1750 (2017).
Index Fungorum number: IF 553834; Facesoffungi numbers: FoF 03692, 5 species.
Parasitic on living coniferous leaves. Sexual morph:
Ascomata superficial, with or without hypostroma, solitary, scattered, easily removed, black, with apical ostiole.
Peridium comprises multi-layers of brown to dark brown
cells of textura angularis. Hamathecium comprising dense,
branched, septate, cellular pseudoparaphyses. Asci 8-spored,
bitunicate, cylindrical to broadly clavate or ellipsoid, sessile
or with knob-like pedicel, rounded at apex, with an ocular chamber. Ascospores 2–3-seriate, irregularly arranged,
ellipsoid-fusiform to subclavate, rounded at both ends, olivaceous-brown, 1-supramedian-septate, constricted at the
septum, upper cell wider than lower cell. Asexual morph:
Undetermined.
Type: Pododimeria E. Müll.
Notes: Pododimeriaceae was introduced by Boonmee
et al. (2017) based on morphology to accommodate two
Fungal Diversity (2020) 105:17–318
Fig. 122 Phaeodimeriella parvula (K(M) 181461, holotype). a Herbarium specimen and habit on leaves. b Appearance of ascoma on
leaf surface. c Setae. d Section of ascoma. e Peridium. f Hamathe-
269
cium of pseudoparaphyses. g, h Asci. i–k Ascospores. l Section of
conidioma. m Setae. n Conidia. Scale bars: b, d = 40 μm, c, e, j–l =
10 μm, f, n = 5 μm, g, i = 20 μm, m = 50 μm
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Fig. 123 Pododimeria gallica (Z 33992, holotype). a Herbarium
specimen and habit on coniferous leaves. b Appearance of ascomata
on leaf surface. c Section of ascoma. d Close up of apical ostiole. e
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Pseudoparaphyses. f–i immature to mature asci. j–n Immature and
mature ascospores. Scale bars: b = 500 μm, e = 5 μm, c, d, f–i = 40
μm, k–n = 10 μm
Fungal Diversity (2020) 105:17–318
genera Chaetoscutula and Pododimeria. No sequence data
is available for this family.
Pododimeria E. Müll., Sydowia 12(1–6): 193 (1959) [1958].
Index Fungorum number: IF 4279; Facesoffungi number:
FoF 07897; 4 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Pododimeria gallica E. Müll.
Notes: Pododimeria was placed in different families,
Capnodiaceae, Dimeriaceae, Parodiellaceae, and Pseudoperisporiaceae based on its ascomata and ascospores
features and its epiphytic habitat (Gäumann 1964; von
Arx and Müller 1975; Barr 1987b; Jaklitsch et al. 2002;
Kirk et al. 2008; Lumbsch and Huhndorf 2010; Hyde et al
2013; Wijayawardene et al. 2014a). However, Boonmee
et al. (2017) studied the holotype specimen of P. gallica
and concluded that Pododimeria differs from the species
in these families mentioned above. Thus, they introduced a
new family Pododimeriaceae to accommodate two genera,
Chaetoscutula and Pododimeria.
Pododimeria gallica E. Müll., Sydowia 12(1–6): 195 (1959)
[1958].
Index Fungorum number: IF 304077; Facesoffungi number: FoF 03693; Fig. 123
Description: see Boonmee et al. (2017).
Material examined: France, Province Gorges du Verdon,
above the bridge over the Artuby River, on living leaves of
Juniperus phoenicea (Cupressaceae), 24 June 1956, E. Müller 22813/11 (Z 33992, holotype).
Other genera included
Chaetoscutula E. Müll., Sydowia 12(1–6): 190 (1959)
[1958].
Index Fungorum number: IF 967; Facesoffungi number:
FoF 07898; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Chaetoscutula juniperi E. Müll., Sydowia
12(1–6): 191 (1959) [1958].
Notes: Chaetoscutula was placed in Pseudoperisporiaceae by Tian et al. (2014) based on morphology of the
herbarium specimen, C. juniper (F225899). Boonmee et al.
(2017) included this genus with Pododimeria in the new
family Pododimeriaceae, and suggested that molecular
data can clarify the relationship between Chaetoscutula and
Pododimeria.
Economic and ecological significance
Pododimeriaceae are parasitic on living leaves of coniferous plants (Butin 1973; Luttrell and Barr 1978; Boonmee
et al. 2017).
271
Polyclypeolinaceae Boonmee & K.D. Hyde, in Boonmee
et al., Mycosphere 8(10): 1754 (2017).
Index Fungorum number: IF 553835; Facesoffungi numbers: FoF 03695, 1 species.
Saprobic, epiphyllous on dried leaves. Sexual morph:
Thyriothecium superficial, irregular, scattered to loosely
clustered, dark brown to black, slightly shield-like or shallow-convex, epithecium membranaceous cells, edge entire,
with apical ostiole. Peridium comprises multi-layers of
dark brown cells of textura angularis. Hamathecium comprising branched, septate, cellular pseudoparaphyses. Asci
8-spored, bitunicate, fissitunicate, saccate, cylindric-clavate,
with pedicellate. Ascospores irregularly arranged, ellipsoid
to fusiform, narrowly obovoid, hyaline, 1-septate, upper cell
wider, tapering towards narrow end (adpated from Boonmee
et al. 2017). Asexual morph: Undetermined.
Type: Polyclypeolina Bat. & I.H. Lima.
Notes: Boonmee et al. (2017) established Polyclypeolinaceae to accommodate a single genus Polyclypeolina based
on its distinct morphology from other families in Dothideomycetes. No sequence data is available for this family.
Polyclypeolina Bat. & I.H. Lima, in Batista, Publicações
Inst. Micol. Recife 56: 457 (1959).
Index Fungorum number: IF 4307; Facesoffungi number:
FoF 07899; 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Polyclypeolina brideliae (Hansf.) Bat.
Notes: The genus was placed in Aulographaceae based
on its branched colonies, superficial, irregular, multi-loculate ascomata, with slit-like opening, and 1-septate, hyaline
ascospores (Luttrell 1973). However, Boonmee et al. (2017)
introduced Polyclypeolinaceae to accommodate this genus
based on its thyriothecium and ascospore features.
Polyclypeolina brideliae (Hansf.) Bat., Publicações Inst.
Micol. Recife 56: 457 (1959).
≡ Polyclypeolum brideliae Hansf., Proc. Linn. Soc. London 157: 37 (1945) [1944–45].
Index Fungorum number: IF 304116; Facesoffungi number: FoF 03696; Fig. 124
Description: see Boonmee et al. (2017).
Materials examined: Uganda, Entebbe, on dried leaves of
Brideliae micranthae (Euphorbiaceae), 1945, G.C. Hansford
3309 (K (M) 177972, holotype).
Economic and ecological significance
Polyclypeolinaceae comprises a single species found on
dried leaves. There is no report concerning its economic
and ecological significance. More collections and sequence
data will help to understand its roles in the environment.
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272
Fig. 124 Polyclypeolina brideliae (K (M) 177972, holotype). a,
b Herbarium specimen and habit on leaf. c Section of ascoma. d
Peridium e Pseudoparaphyses. f–i Immature and mature asci. j–n
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Ascospores. Scale bars: b = 2 mm, c = 100 μm, d = 50 μm, e = 10
μm, f–i = 20 μm, j–n = 5 μm
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273
Fig. 125 Dothidella australis (LPS 318, holotype). a Type material. b Appearance of black ascomata on the host. c Section of ascomata. d–f
Asci g–l Ascospores. Scale bars: a = 1 cm, b = 1 mm, c = 50 µm, d, e = 15 µm, g–l = 10 µm
Polystomellaceae Theiss. & P. Syd., Annls mycol. 13(3/4):
158 (1915).
Index Fungorum: IF 81205; Facesoffungi number: FoF
06489, 43 species.
= Munkiellaceae (Theiss. & Syd.) Luttr., in Ainsworth
et al., The fungi (London) 4a: 154 (1973).
Biotrophic or parasitic on leaves, typical tropical and subtropical distribution. Sexual morph: Stromata superficial or
subcuticular, black, erumpent, pulvinate, and circular to suborbicular, elliptical or elongate, rugulose, containing mostly
fungal tissue, globose to subglobose, uniloculate or multiloculate (with 5 to numerous locules), hyphae flattened and
ribbon-like with discrete central ostiole. Cells of ascostromata murky brown-walled textura angularis. Peridium of
locules soft, often lightly pigmented thick-walled cells of
textura angularis and in side view pseudoparenchymatous,
darken above locules. Hamathecium comprising numerous, hyaline, septate or aseptate cellular pseudoparaphyses without branching, often deliquescent at maturity. Asci
8-spored, bitunicate, fissitunicate, relatively short, oblong to
cylindro-clavate or occasionally obclavate, often ventricose,
with a short pedicel, apically rounded, with a small ocular
chamber. Ascospores 1–2-seriate, fusiform to ellipsoidal or
obpyriform, hyaline, yellowish to olive brown, 0–1-septate,
upper cell wider and shorter than the lower cell, euseptate,
septate near the lower end or middle, with or without constrictions at the septum, asymmetric, lacking a sheath, wall
smooth or verruculose with guttulate contents, sometimes
crowded in ascus. Asexual morph: Coelomycetous, acervular, linked to Lasmenia and Stictochorella (Wakefield
1940; Swart 1987; Kirk et al. 2008); conidiogenesis often
holoblastic.
Type: Dothidella Speg.
Notes: Polystomellaceae was introduced by Theissen and
Sydow (1915) for Munkiella, Parmularia and Polystomella
(the type genus) based upon a foot or hypostroma which
anchors the thyrothecium within the host (von Arx and
Müller 1975). Other names used for this family are Stigmateaceae (Theissen 1916) and Munkiellaceae (Luttrell
1973). Polystomellaceae consists of species with superficial ascomata, while in Munkiellaceae ascomata are subcuticular, yet the two families were combined by Eriksson
(1981). Barr (1987a) included Polystomellaceae as a family
in Pleosporales with the genera Atopospora, Hormotheca
(currently referred to Venturiaceae) and Ellisiodothis (currently referred to Microthyriaceae). Lumbsch and Huhndorf
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274
(2010) included three genera Dothidella (= Polystomella)
with ascospores having a median septum, Munkiella with
apiospores and Parastigmatea in Polystomellaceae under
Dothideomycetes, family incertae sedis, and this was followed by Index Fungorum (2020). All share similar characteristics such as a stromatic ascomata resulting from ascolocular ontogeny, with uni- to multi-locular ascostromata, and
hyaline ascospores, with one or two cells. Several other
genera with amerospores and two with phragmospores have
been included in Munkiellaceae (Luttrell 1973). von Arx
and Müller (1954) placed Parastigmatea under Botryosphaeriaceae, while in a recent review of the family, Liu et al.
(2012) did not include the genus in this family. The three
genera placed in Polystomellaceae were described before
1965 (Index Fungorum 2020) and no molecular sequence
data is available. Thus, fresh collections are needed to establish the phylogenetic relationships of the genera and Polystomellaceae. Munkiella and Parastigmatea are presently listed
in Polystomellaceae, but their inclusion is uncertain for the
purpose of keying out the genera.
Dothidella Speg., Anal. Soc. Cient. Argent. 9: 9 (1880).
Index Fungorum number: IF 4327; Facesoffungi number:
FoF 06490, 34 morphological species (Species Fungorum
2020), molecular data unavailable.
= Polystomella Speg., Anal. Soc. cient. argent. 26(1): 53
(1888).
Type species: Dothidella australis Speg.
Notes: Dothidella is the type genus of Polystomellaceae
and was introduced by Spegazzini (1880). Many species of
Dothidella were transferred to Endodothella, Microcyclus,
Phyllachora, Rehmiodothis and Stigmochora which belong
to Mycosphaerellaceae and Phyllachoraceae (Index Fungorum 2017). They share similar characters such as being
biotrophic or parasitic on leaves and have 1–2-celled, hyaline
ascospores. Dothidella differs from other genera by its multiloculate ascostromata and ellipsoidal hyaline two celled
ascospores, while Phyllachora (Phyllachoraceae) has unilocular ascostromata, unitunicate asci and one celled, hyaline ascospores, and Microcyclus (Mycosphaerellaceae) has
multi-loculate ascostromata with two celled hyaline obovoid
ascospores (Monkai et al. 2013).
Dothidella australis Speg., Anal. Soc. Cient. Argent. 10:
21 (1880).
≡ Polystomella pulcherrima Speg., Anal. Soc. cient.
argent. 26(1): 53 (1888).
Index Fungorum number: IF 233978; Facesoffungi number: FoF 06491; Fig. 125
Biotrophic or parasitic on leaves. Sexual morph: Ascostromata black, superficial to erumpent, pulvinate, and circular to suborbicular, elliptical or elongate, globose to
subglobose, with five to numerous locules, with individual
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central ostioles, cells of ascostromata compsed of dark
brown-walled cells of textura angularis. Peridium of locules composed of small heavily pigmented thick-walled
cells of textura angularis. Hamathecium comprising numerous, hyaline, septate or aseptate pseudoparaphyses. Asci
8-spored, bitunicate, fissitunicate, oblong, cylindro-clavate
or obclavate with a short pedicel, apically rounded, with a
small ocular chamber. Ascospores 2-seriate, ellipsoidal or
obpyriform, hyaline, septate, 1-septate, upper cell wider
and shorter than the lower cell, euseptate, constricted at the
septum, lacking a sheath. Asexual morph: linked to Stictochorella (Wakefield 1940; Swart 1987).
Material examined: Argentina, Buenos Aires, Tigre; on
the leaves of Solanum boerhaviaefolium (Solanaceae), O.
Schnyder, April1880 (LPS 318, holotype).
Other genera included
Dermatodothella Viégas, Bragantia 4(1–6): 150 (1944).
Index Fungorum number: IF 1472; Facesoffungi number:
FoF 06492; – 1 species (Species Fungorum 2020), molecular
data unavailable.
Type species: Dermatodothella multiseptata Viegas, Bragantia 4(1–6): 150 (1944).
Notes: Dermatodothella shares similar characteristics to
Dothidella in having multi-loculate, globose to subglobose
ascostromata, with widely porate ostioles. Dermatodothella
differs from Dothidella in having filiform pseudoparaphyses
and multi-septate ascospores, while Dothidella has bi-celled
ascospores and lacks pseudoparaphyses (Hyde et al. 2013).
Dermatodothella was placed in Polystomellaceae by Ariyawansa et al. (2013b) because of its similarities to other
genera in the family.
Munkiella Speg., Anal. Soc. Cient. Argent. 19(6): 248
(1885).
Index Fungorum number: IF 3288; Facesoffungi number:
FoF 06493; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Munkiella caa-guazu Speg., Anal. Soc.
cient. argent. 19: 248 (1885).
Notes: The asexual morph of this genus is undetermined.
The type species of the genera Munkiella, Coscinopeltis and
Apiotrabutia were found to be the same fungus, which was
named Munkiella caa-guazu. Munkiella was accepted in
Polystomellaceae, Dothideomycetes families incertae sedis
(Lumbsch and Huhndorf 2010; Hyde et al. 2013; Wijayawardene et al. 2014a).
Parastigmatea Doidge, Bothalia 1(1): 22 (1921).
Index Fungorum number: IF 3723; Facesoffungi number:
FoF 06494; – 7 morphological species (Species Fungorum
2020), molecular data unavailable.
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275
Fig. 126 Protoscypha pulla (BPI663043, type). a Material and habit on leaves. b Appearance of ascostromata. c Sections of ascostroma. d
Hamathecium pseudoparaphyses. e, f Asci. g, h Ascospores. Scale bars: b = 2 mm, c = 200 μm, d = 5 μm, e–h = 20 μm
Type species: Parastigmatea nervisita Doidge, Bothalia
1(1): 22 (1921).
Notes: Parastigmatea nervisita was isolated from leaves
of Stephania hernandifolia in South Africa. Parastigmatea
is characterised by epiphyllous ascomata, ovate or broadly
ellipsoid, 8-spored asci and conglobate, hyaline ascospores.
Parastigmatea was accepted in Polystomellaceae, Dothideomycetes families incertae sedis (Lumbsch and Huhndorf
2010; Hyde et al. 2013; Wijayawardene et al. 2018). The
sexual morph is undetermined.
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276
Economic and ecological significance
Dothidella include many species which are of economic
importance. They play a negative role by causing crop diseases. The Amazon’s loss of the rubber market was caused
by species from Dothidella making the more efficient plantation cropping impossible in the Amazon (Bunker 1985)
Protoscyphaceae Kutorga & D. Hawksw., Syst. Ascom.
15(1–2): 70 (1997).
Index Fungorum number: IF 81924, Facesoffungi number: FoF 07900, 2 species.
Parasitic on living leaves of Miconia thomasiana. Sexual
morph: Ascostromata superficial, solitary, scattered, occurring
on the lower leaf surface, discoid or cup-shaped, slightly convex or raised pulvinate, semi-immersed, flattened and thickened
at the base, occurring in lesions of light brown regions and
surrounded by external black mycelial, multi-locular, opening
by rupturing or cracking of the apical ascostromata. Peridium
thick, composed of dark cells arranged in a textura angularis.
Hamathecium comprising hyaline, relatively wide, anastomosing, septate, cellular pseudoparaphyses, often with swollen
cells, intermixed with reddish brown colouration, and surrounded by a gelatinous matrix. Asci 8-spored, bitunicate, fissitunicate, subglobose oblong, broadly cylindrical to subclavate,
somewhat thickened at the apex, short pedicellate, with a wide,
but short ocular chamber. Ascospores 2–3-seriate, muriform,
oblong to oval-sub ellipsoid, hyaline, pale brown or light brown,
multi-septate, constricted at the septum, surrounded by a thin
gelatinous sheath. Asexual morph: Undetermined.
Type: Protoscypha Syd.
Notes: The family currently comprises a monotypic genus
Protoscypha with two species and is treated in Dothideomycetes as family incertae sedis (Hyde et al. 2013; Wijayawardene
et al. 2018). The family lacks molecular data to establish phylogenetic placement, especially as the type species is yet to be
sequenced and no additional new taxa in this family.
Protoscypha Syd., Annls mycol. 23(3/6): 402 (1925).
Index Fungorum number: IF 4394; Facesoffungi number:
FoF 07901; – 2 species (Species Fungorum 2020), molecular
data unavailable.
Type species: Protoscypha pulla Syd.
Notes: Protoscypha is a tropical genus and comprises
only two parasitic species on living leaves (Index Fungorum 2020). The genus is characterized by discoid to cup-like
ascostromata, asci in a single locule and muriform, hyaline
to lightly pigmented ascospores (Hyde et al. 2013).
Protoscypha pulla Syd., Annls mycol. 23(3/6): 403 (1925).
Index Fungorum number: IF 275651; Facesoffungi number: FoF 07902; Fig. 126
Description: see Hyde et al. (2013).
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Material examined: Costa Rica, Los Angeles de San
Ramon, on leaves of Miconia thomasiana, 30 January 1925,
H. Sydow, BPI663043 (type).
Economic significance
Protoscyphaceae is a parasitic family occurring on living
leaves and distributed in tropical countries such as Costa
Rica and Brazil.
Pseudoperisporiaceae Toro, in Seaver & Palacios Chardon,
Scient. Surv. P. Rico 8(1): 40 (1926).
Index Fungorum number: IF 81227, Facesoffungi number: FoF 00122, 48 species.
Parasitic, biotrophic or saprobic on leaves, or other
fungi, primarily in tropical regions. Sexual morph: Ascomata superficial, solitary to gregarious, scattered, globose
to subglobose, collapsed when dry, surrounded by brown
mycelium at the base, ostiole central and surrounded by
brown, septate setae, tapering to subacute apex. Peridium
composed of cells of textura angularis, brown to red brown,
thin-walled, sometimes collapsed when dry. Hamathecium
comprising cylindrical, filiform, hyaline, branched, septate,
cellular pseudoparaphyses, in a gelatinous matrix. Asci
8-spored, bitunicate, fissitunicate, oblong-clavate, slightly
curved, thickened at the apex, sessile, or with knob-like
pedicel, with small ocular chamber. Ascospores 2-seriate, fusoid-ellipsoid, rounded and subacute ends, hyaline,
becoming brownish at maturity, 1-septate, slightly constricted at the septum, wall minutely verrucose. Asexual
morph: Coelomycetous.
Type: Lasiostemma Theiss., Syd. & P. Syd.
Notes: Pseudoperisporiaceae comprises only four genera
Bryomyces, Eudimeriolum, Lasiostemma and Nematostoma
and is treated in Dothideomycetes as family incertae sedis
(Boonmee et al. 2017; Wijayawardene et al. 2018). Taxa in
this family are reported as parasitic or saprobic on leaves
(Hyde et al. 2013, 2017; Boonmee et al. 2017). The family
lacks molecular data to establish phylogenetic placement,
especially as the type species is yet to be sequenced.
Lasiostemma Theiss., Syd. & P. Syd., in Sydow & Sydow,
Annls mycol. 15(3/4): 218 (1917).
Index Fungorum number: IF 2660, Facesoffungi number:
FoF 07903, 6 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Lasiostemma melioloides (Berk. & Ravenel) Theiss., Syd. & P. Syd.
Notes: The genus is characterized by superficial, redbrown to dark brown ascomata, with long hyphae, oblongclavate asci, fusoid-ellipsoid, 1-septate, light brown
ascospores. Lasiostemma comprises eight species on living leaves and is distributed in tropical regions e.g., Brazil,
Cuba, Malaysia, Uganda (Hyde et al. 2013; Boonmee et al.
Fungal Diversity (2020) 105:17–318
277
Fig. 127 Lasiostemma melioloides (BPI699548). a Material and habit on leaves. b Appearance of ascomata. c Squash mount of ascoma showing ostiole. d Hamathecium pseudoparaphyses. e, f Asci. g–j Ascospores. Scale bars: b, c = 100 μm, d = 5 μm, e, f = 20 μm, g–j = 10 μm
2017). Hyde et al. (2013) examined a specimen of Lasiostemma melioloides (BPI699548). The type species Lasiostemma melioloides and its members need sequence data
to establish taxonomic placement of this genus.
Lasiostemma melioloides (Berk. & Ravenel) Theiss., Syd.
& P. Syd., in Sydow & Sydow, Annls mycol. 15(3/4): 218
(1917).
≡ Sphaeria melioloides Berk. & M.A. Curtis, in Berkeley,
J. Linn. Soc., Bot. 10(no. 46): 387 (1868) [1869].
Index Fungorum number: IF 102249, Facesoffungi number: FoF 03698; Fig. 127
Description: see Hyde et al. (2013) and Boonmee et al.
(2017).
Material examined: Venezuela, Federal district, Puerto La
Cruz Valley, El Limón, on leaves of Erigeron bonariensis,
16 January 1928, H. Sydow, Fungi venezuelani No. 249
(BPI699548).
Other genera included
Bryomyces Döbbeler, Mitt. bot. StSamml., Münch. 14: 233
(1978).
Index Fungorum number: IF 664, Facesoffungi number:
FoF 07904; – 9 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Bryomyces scapaniae Döbbeler, Mitt. bot.
StSamml., Münch. 14: 254 (1978).
Notes: Hyde et al. (2017) revisited Bryomyces and
accepted this genus in Pseudoperisporiaceae based on its
parasitic or biotrophic habit on living hairy leaves of mosslike plants (liverworts or hornworts). Taxa in this genus are
widely distributed and have been found in Austria, Bolivia,
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278
Japan and Sweden. Hyde et al. (2017) provided description
and illustration of the holotype specimen Bryomyces scapaniae, habit on living hairy leaves of Scapania undulata from
Austria.
Eudimeriolum Speg., Anal. Mus. nac. Hist. nat. B. Aires
23: 36 (1912).
Index Fungorum number: IF 1922, Facesoffungi number:
FoF 07905; – 16 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Eudimeriolum elegans Speg., Anal. Mus.
nac. Hist. nat. B. Aires 23: 36 (1912).
Notes: Boonmee et al. (2017) revisited Eudimeriolum and
accepted this genus in Pseudoperisporiaceae based on its
parasitic and saprobic habit on leaves. Taxa in this genus
are widely distributed such as in Brazil, Germany, Philippines, Puerto Rico, Tanzania and other tropical countries.
Boonmee et al. (2017) provided a description and illustration of the type species Eudimeriolum elegans based on the
protologue of Spegazzini (1912).
Nematostoma Syd. & P. Syd., Annls mycol. 12(2): 161
(1914).
Index Fungorum number: IF 3443, Facesoffungi number:
FoF 07906; – 17 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Nematostoma artemisiae Syd. & P. Syd.,
Annls mycol. 12(2): 161 (1914).
Notes: Boonmee et al. (2017) revisited Nematostoma and
accepted this genus in Pseudoperisporiaceae based on parasitic and saprobic habit on leaves. Taxa in this genus are
widely distributed such as in Argentina, Japan, Mexico, the
USA and other tropical countries. Boonmee et al. (2017)
provided a description and illustration of the holotype specimen Nematostoma artemisiae, habit on living leaves of Artemisia vulgaris var. indica from Japan.
Economic and ecological significance
Pseudoperisporiaceae is rather a large family and comprises
48 taxa which are parasitic, biotrophic or saprobic habit on
leaves. All taxa in this family are widespread in distribution
and usually associated on living leaves, but some taxa are
found on dead leaves. Bryomyces is commonly associated
with moss-like plants such as liverwort and hornworts (Döbbeler 1978; Hyde et al. 2017).
Pseudorobillardaceae Crous, Fungal Systematics and Evolution 3: 108 (2019).
Index Fungorum number: IF 829342; Facesoffungi numbers: FoF 07907, 18 species.
Saprobic, pathogenic or endophytic on soil or decaying
living and dead leaves, stems. Sexual morph: Undetermined. Asexual morph: Conidiomata immersed, globose,
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unilocular, with central ostiole. Conidiogenous cells thinwalled with of 3–6 layers, composed of flattened cells of
textura angularis. Hamathecium numerous, hyphae-like,
aseptate, flexuous, paraphyses, intermingled among conidiophores. Conidiogenous cells sub-cylindrical to ampulliform,
hyaline, smooth. Conidia contain micro- or macroconidia,
subcylindrical, hyaline, aseptate, smooth-walled, guttulate,
apex obtuse, base truncate; apical appendages hair-like,
flexuous, unbranched, fragile, flexuous, unbranched.
Type: Pseudorobillarda M. Morelet.
Notes: Pseudorobillardaceae (Dothideomycetes) was
informally introduced by Crous et al. (2018a) based on morphology and phylogeny. Later Pseudorobillardaceae was
formally introduced by Crous et al. (2019a) to accommodate
Pseudorobillarda. In previous studies this genus was placed
in Dothideomycetes genera incertae sedis (Tangthirasunun
et al. 2014; Wijayawardene et al. 2018).
Pseudorobillarda M. Morelet, Bull. Soc. Sci. nat. Arch.
Toulon et du Var 175: 5 (1968).
Index Fungorum number: IF 9607; Facesoffungi number:
FoF 07559; 18 morphological species (Species Fungorum
2020), 7 species with molecular data.
Type species: Pseudorobillarda phragmitis (Cunnell) M.
Morelet.
Notes: The members of this genus can be saprobic, pathogenic and endophytic and are known from temperate and
tropical countries such as Argentina, Canada, Cuba, Germany, India, Nigeria, South Africa, Thailand, UK, Ukraine,
and the USA (Petrini 1986; Nag-Raj 1993; Bianchinotti
1997; Vujanovic and St-Arnaud 2003; Plaingam et al. 2005;
Tangthirasunun et al. 2014; Crous et al. 2018a). This genus
occurs on both living and dead leaves, stems or bark and is
also commonly found in soil (Petrini 1986; Vujanovic and
St-Arnaud 2003; Kadowaki et al. 2014). Keys to species in
Pseudorobillarda were provided in Sutton (1980), Nag-Raj
(1993) and Vujanovic and St-Arnaud (2003). The revised
descriptions and illustrations of Pseudorobillarda, based
on morphological characteristics are available in Plaingam
et al. (2005). No sexual morph is linked to Pseudorobillarda (Hyde et al. 2011; Wijayawardene et al. 2012, 2017a;
Tangthirasunun et al. 2014).
Pseudorobillarda phragmitis (Cunnell) M. Morelet, Bull.
Soc. Sci. nat. Arch. Toulon et du Var 175: 6 (1968).
≡ Robillarda phragmitis Cunnell, Trans. Br. mycol. Soc.
41(4): 405 (1958).
Index Fungorum number: IF 321865; Facesoffungi number: FoF 07565, Fig. 128
Description: see Li et al. (2020).
Materials examined: UK, England, Middlesex, near
Staines, on dead stems of Phragmites communis (Poaceae),
July 1955, G. J. Cunnell (IMI 70768, holotype).
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279
Fig. 128 Pseudorobillarda phragmitis (IMI 70768, holotype). a Herbarium packet. b Dried culture. c Colony characteristics in culture. d
Section of conidioma. e Conidiogenous cells with developing conidia.
f–h Conidiogenous cells. i, j Conidia. Scale bars: c = 200 μm, d =
100 μm, e = 50 μm, f–j = 5 μm,
Economic and ecological significance
Pseudorobillardaceae is a small family containing only a
single genus and they are important for nutrient cycling as
they are saprobic on a wide variety of plant substrates. On
the other hand, some species of Pseudorobillarda are known
as one of the top ten most abundant soil fungi (Kadowaki
et al. 2014).
Lichenicolous on various substrates such as bark, rock,
soil, bryophytes or living leaves. Sexual morph: Ascomata
perithecioid, densely grouped or dispersed over large areas
of the thallus, sessile or immersed in the host thallus, black,
in longitudinal section, subglobose to broadly ovoid or
pyriform, often with light greenish-blue tint in the ostiolar
canal. Peridium with isodiametric to elongated cells, brown
to dark brown. Hymenium hyaline, I-, KI-. Hamathecium of
numerous, persistent, branched, anastomosing pseudoparaphyses; apical cells not enlarged; non-branching periphyses.
Asci 4–8-spored, bitunicate, short pedicellate, I-, KI-, ocular chamber not distinct. Ascospores 1–2-seriate, oblong to
ellipsoidal, or fusiform, dark brown, often paler brown at
Pyrenidiaceae Zahlbr. in Engler, syllabus der Pflanzenfamilien, Edn 2 (Berlin): 46 (1898).
Index Fungorum number: IF 81790; Facesoffungi numbers: FoF 07908, 13 species.
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Fig. 129 Pyrenidium actinellum (Ertz. 16557 (BR)). a
Appearance of ascomata on
host surface. b Vertical section of host with perithecioid
ascoma visible. c, d Asci. e–g
Ascospores. Scale bars: b = 50
µm, c, d = 10 µm, e–g = 5 µm
the spore tips, 3-septate, smooth-walled. Asexual morph:
Undetermined (adapted from Huanraluek et al. 2019a).
Type: Pyrenidium Nyl.
Notes: Huanraluek et al. (2019a) provided sequence
data for Pyrenidium species and indicated that Pyrenidium
formed a distinct clade outside Dacampiaceae. However,
Huanraluek et al. (2019a) did not assign Pyrenidiaceae to
any order. Pyrenidiaceae formed a distinct clade outside Pleosporales in personal analyses of Hongsanan et al. (2020,
data not showed), and formed within Pleosporales as a distinct family (data not showed) in the dataset which includes
all representative families in Dothideomycetes (data not
13
showed). Therefore, we agree with Huanraluek et al. (2019a)
to treat this family as incertae sedis in Dothideomycetes
until more collections with multi-loci are available to clarify
its phylogenetic placement.
Pyrenidium Nyl., Flora, Regensburg 48: 210 (1865).
Index Fungorum number: IF 4578; Facesoffungi numbers:
FoF 07909; 13 morphological species (Species Fungorum
2020), 3 species with molecular data.
Type: Pyrenidium actinellum Nyl.
Notes: Pyrenidium was placed in Dacampiaceae based
on morphology and the environmental habitats (Hyde et al.
Fungal Diversity (2020) 105:17–318
2013; Wijayawardene et al. 2014a, 2018; Jaklitsch et al.
2016). Crivelli (1983) suggested that Pyrenidium has close
relationship with Dacampia in Dacampiaceae in its similar
ascomatal structures and wall tissues (Crivelli 1983). However, sequence data of three species, Pyrenidium actinellum,
P. aff. aggregatum and P. borbonicum, were provided by
Huanraluek et al. (2019a). These sequence data formed a
distint clade far from Dacampiacceae. Thus, Pyrenidiaceae
was resurrected to accommodate species of Pyrenidium.
Pyrenidium actinellum Nyl., Flora, Regensburg 48: 210
(1865).
Index Fungorum number: IF 403001; Facesoffungi numbers: FoF 04634; Fig. 129
Description: see Huanraluek et al. (2019a).
Materail examined: Belgium, Aywaille, versant droit de
l’Amblève, Heid des Gattes, 200 m elev. Paroi de grès à
Anema, on the lichen Scytinium plicatile, 2011, (Ertz. 16557
(BR)).
Economic and ecological significance
Species of Pyrenidium are lichenicolous developing on
lichen thalli, with parasymbionts living in symbiosis (Hawksworth 1982; Diederich et al. 2018). The ascomata of Pyrenidium are immersed to erumpent in host thalli with the
upper part exposed and sometimes causing gall-like malformations of the host (Eriksson and Hawksworth 1993;
Matzer 1996; Pirogov 2014; Lendemer 2017; Muscavitch
et al. 2017).
Seynesiopeltidaceae K.D. Hyde, in Hyde et al., Fungal
Diversity 63: 232 (2013).
Index Fungorum number: IF 804042; Facesoffungi number: FoF 06522, 1 species.
Foliar epiphytes on leaves, causing black spots, usually
confluent in groups, forming black colonies. Sexual morph:
Mycelium superficial, sparse, light, septate, olive-green.
Thyriothecia solitary or gregarious, superficial, circular,
shield-like, shallow, conical, dome-like, flattened, upper
region, carbonaceous, unilocular, brown to black, basal layer
poorly developed, lacking a distinct central ostiole. Upper
wall comprising radiating arranged parallel cells, cells at
margin branching. Hamathecium comprising 1–3 µm wide,
comprising numerous, cellular, branched pseudoparaphyses.
Asci 8-spored, bitunicate, fissitunicate, saccate, with a knoblike pedicel, and conspicuous ocular chamber (in Congo
red). Ascospores irregularly 2–3-seriate, dark brown when
mature, 1-septate, strongly constricted at septum, upper
cells longer and wider, basal cell shorter and narrower with
an acute base, with a thick, structured, hyaline, cell wall.
Asexual morph: Undetermined.
Type: Seynesiopeltis F. Stevens & R.W. Ryan.
281
Notes: Seynesiopeltidaceae was introduced by Hyde
et al. (2013) based on the unique brown ascospores with
thick structured walls and thick ascus wall when stained in
Congo red. The cells of the peridium consist of two strata of
textura angularis cells. Ascomata are described as thyriothecia-like and they resemble some genera in Parmulariaceae
(e.g. Polycyclus andinus, Inácio and Cannon 2008). Genera
such as Dothidasteroma and Cocconia (also Parmulariaceae) may also be related as they have brown ascospores,
but the asci are significantly different (Inácio and Cannon
2008). Notably, Seynesiopeltis differs from Parmularia, the
type genus of Parmulariaceae. In Parmularia styracis, the
thyriothecia have a similar appearance to shells on the host
surface, the asci are obclavate and ascospores are hyaline.
Seynesiopeltis F. Stevens & R.W. Ryan, in Stevens, Bulletin
of the Bernice P. Bishop Museum, Honolulu, Hawaii 19: 69
(1925).
Index Fungorum number: IF 5020; Facesoffungi number:
FoF 07910, 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Seynesiopeltis tetraplasandrae F. Stevens
& R.W. Ryan.
Notes: Seynesiopeltis is the type genus of Seynesiopeltidaceae and it was established by Stevens (1925) as
a monotypic genus with S. tetraplasandrae as the type and
has remained monotypic. This genus was earlier assigned
to Microthyriaceae by Stevens (1925) which was followed
by Muller and von Arx (1962) and Lumbsch and Huhndorf
(2010). This genus was moved to Dothideomycetes genera
incertae sedis by Wu et al. (2011b) since it was unlike any
other member of the class Dothideomycetes. It is characterised by unfree mycelium, round and radiate ascomata with
thallus being setose. Stevens (1925) mentioned that species
in this genus resembles Seynesia but are different in that they
possess one to several ascomata in one fungus body and that
this bears setae which intensify the uniqueness of the genus
and family. However, no setae in the type specimen was
observed by Hyde et al. (2013). A plausible reason might
be loss of setae during storage as the latter is described as
being fragile. The genus is monotypic and warrants further
investigation.
Seynesiopeltis tetraplasandrae F. Stevens & R.W. Ryan,
Bulletin of the Bernice P. Bishop Museum, Honolulu,
Hawaii 19: 70 (1925).
Index Fungorum number: IF 158371; Facesoffungi number: FoF 06523; Fig. 130
Foliar epiphyte on leaves, causing black spots, typically
confluent, forming black colonies. Sexual morph: Mycelium superficial, sparse, septate, olive-green. Thyriothecia
347–483 µm diam. 438–565 µm high, solitary or gregarious, superficial, with a circular, shield-like, shallow, conical,
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Fig. 130 Seynesiopeltis tetraplasandrae (BISH 1089, holotype).
a Details of herbarium material. b, c Habit and appearance of thyriothecia on the host surface. d Section of ascoma. e Peridium. f
Hamathecium. g, h Asci. i–l Ascospores. Scale bars: b = 2000 µm,
c = 500 µm, d = 25 µm, e = 35 µm, f = 10 µm, g, h = 50 µm, i–l =
20 µm
dome-like, flattened, upper region, carbonaceous, unilocular, brown to black, basal layer poorly developed, lacking
a distinct central ostiole. Upper wall comprising radiating
parallel cells, cells at margin branching. Hamathecium
comprising 1–3 µm wide, numerous, cellular, branched
pseudoparaphyses. Asci 117–142 × 59–68 µm ( x̄ = 127 ×
65 µm, n = 20), 8-spored, bitunicate, fissitunicate, saccate,
with a knob-like pedicel and conspicuous ocular chamber.
Ascospores 36–41 × 13–15 µm ( x̄ = 39 × 14 µm, n = 20)
irregularly 2–3-seriate, dark brown when mature, 1-septate,
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283
Fig. 131 Stomatogene agaves (NY 02977047). a Herbarium specimen and habit on leaves. b Appearance of ascomata on leaf surface. c Section
of ascoma. d Peridium. e Hamathecium. f, g Asci. h–k Ascospores. Scale bars: b = 200 μm, c = 50 μm, f, g = 20 μm, h–k = 10 μm
strongly constricted at septum, upper cells longer and
wider, basal cell shorter and narrower with an acute base,
with a thick, structured, hyaline, cell wall. Asexual morph:
Undetermined.
Material examined: USA, Hawaii, Oahu, Hamakua, upper
ditch trail, on leaves of Tetraplasandra (Araliaceae), 31 May
1921, Stevens F.L. (BISH 1089, holotype).
Notes: Seynesiopeltis tetraplasandrae was introduced by
Stevens and Ryan (1925). It was found on leaves of Tetraplasandra in Hawaii and was described as having unfree mycelium, round ascomata consisting several locules and lacking
paraphyses. Further collections are required to resolve this
genus.
Economic and ecological significance
Seynesiopeltis is a monotypic genus and foliar epiphytes on
leaves. Its life style is unclear as to whether it is saprobic or
parasitic.
Stomatogeneceae Boonmee & K.D. Hyde, in Boonmee
et al., Mycosphere 8(10): 1762 (2017).
Index Fungorum number: IF 553837; Facesoffungi numbers: FoF 03703, 4 species.
Parasitic on living leaves of Agave sp. (Asparagaceae)
and mosses. Colonies large, superficial, subcircular, irregular, with numerous, dark brown mycelium, radiating outwards, flexuous, septate. Sexual morph: Ascomata superficial, semi-immersed at the base, with a basal hypostroma
developing in the host tissue, gregarious, or solitary, black,
with apical pore, surrounded by dark brown mycelium.
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Fig. 132 Morphology of Thyrinula spp. (a–d = Thyrinula
eucalypti redrawn from Crous
et al. 2019c; e–g = Thyrinula
parasitica redrawn from Summerell et al. 2006). a Ascomata
on lesion surface. b Vertical
section through ascomata. c
Asci. d Ascospores. e–g Conidiophores and conidia. Scale bars
= 10 μm
Peridium 3–4 layers, dark pigmented cells of textura angularis, lacking pseudoparaphyses. Asci 8-spored, bitunicate,
fissitunicate, subglobose, broadly ovoid to subclavate, apically thickened, with an ocular chamber at apex, knob-like
pedicellate. Ascospores multi-seriate, ellipsoidal oblong,
hyaline when immature, brown at maturity, 1-septate, upper
cell slightly broader and shorter than lower cell, verrucose
or smooth-walled. Asexual morph: Undetermined (adapted
from Boonmee et al. 2017).
Type: Stomatogene Theiss.
Notes: This family was introduced by Boonmee et al.
(2017) to accommodate the single genus, Stomatogene.
Stomatogeneceae can be differentiated from other families
in Dothideomycetes by its colonies, ascomata, asci and
ascospore features. However, no sequence data is available
in GenBank.
Stomatogene Theiss., Annls mycol. 14(6): 406 (1917)
[1916].
Index Fungorum number: IF 5270; Facesoffungi number:
FoF 03704; 4 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Stomatogene agaves (Ellis & Everh.)
Theiss.
Notes: The genus was placed in Parodiopsidaceae by
Eriksson et al. (2001), Kirk et al. (2008), Lumbsch and
Huhndorf (2010) and Hyde et al. (2013). Boonmee et al.
13
(2017) concluded that Stomatogene should be placed in its
own family based on its unique morphological characters.
Stomatogene agaves (Ellis & Everh.) Theiss., Annls mycol.
14(6): 406 (1918) [1917].
≡ Asterina agaves Ellis & Everh., Bull. Torrey bot. Club
27: 571 (1900).
Index Fungorum number: IF 121867; Facesoffungi number: FoF 03705; Fig. 131
Description: see Boonmee et al. (2017).
Material examined: Mexico, Sonora, in the vicinity of
Álamos, on living leaves of Agave sp. L (Asparagaceae),
March 1910, J.N. Rose No.13063 (NY 02977047).
Economic and ecological significance
Species in Stomatogeneceae are parasitic on living leaves of
Agave sp. (Asparagaceae) and mosses.
Thyrinulaceae X.Y. Zeng, S. Hongsanan & K.D. Hyde,
fam. nov.
Index Fungorum number: IF 557945; Facesoffungi number: FoF 06898, 10 species.
Pathogens on surface of leaves or petioles. Leaf spots
epiphyllous or hypophyllous, circular to irregular, single to
confluent, brown. Sexual morph: Ascomata superficial, dark
brown, hysterothecial, opening by elongated, longitudinal
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285
Fig. 133 Toroa dimerosporioides (BPI 692120). a Herbarium specimen. b Ascomata on leaf surface. c Section of ascoma. d Peridium. e
Hamathecium. f, g Asci. h–j Ascospores. Scale bars: b = 100 μm, c, f, g = 50 μm, d, h–j = 20 μm, e = 10 μm
or Y-shaped slits, margin without restricted brown, superficial hyphae. Asci 8-spored, bitunicate, broadly ellipsoid
to somewhat obovoid, intermixed among hyphae-like,
septate, branched pseudoparaphyses, with ocular chamber. Ascospores cylindrical to ellipsoid, hyaline, 1-septate, constricted at septum, ends obtusely rounded, guttulate, smooth-walled. Asexual morph: Coelomycetous or
hyphomycetous. Coelomycetous: Mycelium brown, septate, branched, verrucous. Conidiomata brown, acervular,
solitary to gregarious or confluent, immersed to erumpent,
depressed globose, unilocular. Ostiole absent, dehiscence
by irregular splits of apical wall in the middle part. Conidiomatal wall composed of a single layer of brown cells in
the basal part. Conidiophores arising from inner wall layer
of basal stroma, brown, branched at the base, septate. Conidiogenous cells brown, holothallic, cylindrical, integrated
or discrete, determinate, verrucous. Conidia brown, arthric,
formed by disarticulation of the conidial chain, produced in
simple unbranched chains with the youngest conidium at the
base, surrounding by mucoid sheath, ellipsoidal to globose
or subcylindrical, truncate at the base, aseptate, verrucose,
guttulate (adapted from Giraldo et al. 2017). Hyphomycetous asexual morph: Conidiophores single or sporodochial,
if conidiophores single, micronematous, mononematous,
straight to slightly flexuous, brown, septate, smooth; if conidiophores sporodochial, brown, verrucose, thick-walled,
13
286
Fig. 134 Asterina labecula (PC 0084645, syntype). a Specimens.
b Ascomata on host substrate. c Ascomata with thin and superficial
thallus in 70% lactic acid. d Walled cells of ascoma and ostiole. e
Walled cells of thallus composed darkly pigmented cells. f Hamathecium in Melzer’s reagent. g Asci when immature in Melzer’s reagent.
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h Asci in Melzer’s reagent. i Asci in cotton blue reagent. j Ascospore
2-septate in 70% lactic acid. k Ascospore uniseptate in Melzer’s reagent. l Ascospore 2-septate in cotton blue reagent. Scale bars: c = 100
μm, e–i = 20 μm. j–l = 10 μm
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Fig. 135 Trichothyrium sarciniferum (LPS1441, holotype). a The
leaf substrate. b Colonies on the leaf surface. c Squash mount of the
ascoma. d The upper wall of ascomata. e Superficial mycelium forming thalli on hyphae of Meliolaceae species. f, g Cross section of
287
the ascoma. h–j Asci mounted in Melzer’s reagent. k, l Ascospores
mounted in Melzer’s reagent. Scale bars: b = 200 µm, c = 50 µm, e–f
= 20 µm, d, g–j = 10 µm, k–l = 5 µm
13
288
Fungal Diversity (2020) 105:17–318
Fig. 136 Vizella oleariae (Redrawn from Swart 1971). a Section through ascoma. b Surface of ascoma. c Proliferating crosiers. d Asci. e
Ascospores. f Surface view of hypha. g–l Conidia and spermatia. Scale bars: a–g, j = 10 µm, h, i, k, l = 5 µm
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branched, septate. Appressoria when present, numerous,
entire, globose to cylindrical, alternate to unilateral, aseptate, brown, penetration peg central on the appressorial cell.
Conidiogenous cells holoblastic, monoblastic, integrated,
intercalary, cylindrical to elliptical, light brown to brown,
smooth, sometimes with indistinct scars. Conidial secession schizolytic. Conidia solitary or in branched or simple
acropetal chains, ellipsoidal to cylindrical, aseptate to multiseptate, medium brown to brown, smooth or verruculose,
sometimes with rounded ends.
Type: Thyrinula Petr. & Syd.
Notes: This family is introduced to accommodate species
that are phylogenetically related to Asterinales sensu lato, a
clade that is typified by the type sequence data of Asterina
and Lembosia obtained by Guatimosim et al. (2015). However, the genetic distance between the two genera is less
than the two Asterina species in their phylogeny, indicating
that these two sequences are questionable. There are three
genera included in this family based on phylogeny, and they
are sister to Parmulariaceae (Parmulariales).
Thyrinula Petr. & Syd., Annls mycol. 22(3/6): 373 (1924).
= Alysidiella Crous, in Summerell et al., Fungal Diversity
23: 325 (2006).
Index Fungorum number: IF 10218; Facesoffungi number: FoF 06728; 5 morphological species (Crous et al.
2019c), 5 species with molecular data.
Type species: Thyrinula eucalyptina Petr. & Syd., Annls
mycol. 22(3/6): 373 (1924)
Notes: Thyrinula is characterized by hysterothecia opening by elongated, longitudinal or Y-shaped slits, and 1-septate ascospores. Its asexual morph Alysidiella was introduced to accommodate a single species with both aseptate
and multi-septate conidia. Crous et al. (2019c) provided
sequence data of all members of the genus from fresh collections, and synonymized Alysidiella under Thyrinula based
on morphology (Fig. 132).
Other genera included
Blastacervulus H.J. Swart, Trans. Br. mycol. Soc. 90(2):
289 (1988).
Index Fungorum number: IF 11029; Facesoffungi number: FoF 06733; – 4 morphological species (Species Fungorum 2020), 3 species with molecular data.
Type species: Blastacervulus eucalypti H.J. Swart, Trans.
Br. mycol. Soc. 90(2): 289 (1988).
Notes: Blastacervulus is a coelomycetous genus characterised by acervular conidiomata and thallic, 1-septate
conidia with truncate, flat hila. Wijayawardene et al. (2016)
showed that B. eucalypti clustered in Planistromellaceae,
however, it forms within the Thyrinulaceae in our analyses
(Fig. 5).
289
Paraopeba V.P. Abreu, A.A.M. Gomes, Firmino & O.L.
Pereira, in Crous et al., Persoonia 39: 349 (2017).
Index Fungorum number: IF 821209; Facesoffungi number: FoF 06740; – 1 morphological species (Species Fungorum 2020), with molecular data.
Type species: Paraopeba schefflerae V.P. Abreu, A.A.M.
Gomes, Firmino & O.L. Pereira, in Crous et al., Persoonia
39: 349 (2017).
Notes: Paraopeba is hyphomycetous characterized by
schizolytic conidiogenous cells, 2–13-septate conidia and
the present of appressoria. Crous et al. (2017) introduced it
as a monotypic genus in Asterinales based on the phylogenetic analysis.
Economic and ecological significance
Members of this family are important plant pathogens that
can cause leaf and petiole lesions.
Toroaceae Boonmee & K.D. Hyde, in Boonmee et al.,
Mycosphere 8(10): 1769 (2017).
Index Fungorum number: IF 553838; Facesoffungi numbers: FoF 03708, 1 species.
Saprobic on dried leaves. Sexual morph: Ascomata
superficial, globose to subglobose, solitary to gregarious,
scattered, black, with a subiculum, collapsing when dry,
with apical ostiole. Peridium comprising red brown to dark
brown cells of textura angularis. Asci 4-spored, bitunicate, oblong-ellipsoid to ovoid, sessile, apically thickened.
Ascospores 2-seriate, ellipsoid-fusiform, narrow towards
the ends, pale brown to orange brown, light at both ends,
smooth-walled. Asexual morph: Undetermined (adapted
from Boonmee et al. 2017).
Type: Toroa Syd.
Notes: Boonmee et al. (2017) established this family
to accommodate the genus Toroa. Toroaceae differs from
other families in the Dothideomycetes in having only four
ascospores in the ascus and a combination of other characters (Boonmee et al. 2017). No sequence data is available
for this family.
Toroa Syd., in Toro, J. Dept. Agric. Porto Rico 10(2): 19
(1926).
Index Fungorum number: IF 5500; Facesoffungi number:
FoF 07911; 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Toroa dimerosporioides (Speg.) Syd.
Notes: Toroa was established to accommodate the type
species T. dimerosporioides which is the current name of
Asteridium dimerosporioides, Zukalia dimerosporioides,
Gibberella dimerosporoides, Perisporium bromeliae and
Chaetosphaeria bromeliae (Spegazzini 1888; Saccardo
1891; von Höhnel 1907; Stevens 1917; González-Fragoso
and Ciferri 1925; Toro 1926; Boonmee et al. 2017). Barr
13
290
(1997) transferred Toroa to Pseudoperisporiaceae, however,
Toroa can be distinguished from Pseudoperisporiaceae and
other families in Dothideomycetes in having 4-spored asci,
and 2–3-septate, pale brown to orange brown ascospores
which are light at both ends (Boonmee et al. 2017).
Toroa dimerosporioides (Speg.) Syd. [as ‘dimerosporoides’], J. Dept. Agric. Porto Rico 2: 20 (1926).
≡ Asteridium dimerosporioides Speg., Anal. Soc. cient.
argent. 26(1): 19 [no. 50] (1888).
Index Fungorum number: IF 263219; Facesoffungi number: FoF 03245; Fig. 133
Description: see Boonmee et al. (2017).
Material examined: Puerto Rico, La Vega, on dried leaves
Bromelia pinguin L. (Bromeliaceae), 19 March 1919, Kern
and R.A. Toro (No. 207 / No. I.T.A 5413, BPI 692120).
Economic and ecological significance
Toroaceae comprises a single genus which was found as
saprobic on dead leaves (Boonmee et al. 2017). It plays a
role in recycling organic matter.
Trichopeltinaceae Bat., C.A.A. Costa & Cif., Atti
dell’Istituto Botanico della Università e Laboratorio Crittogamico di Pavia 15: 37 (1958).
Index Fungorum number: IF 81489; Facesoffungi numbers: FoF 06203, 29 species.
Epiphytic on living leaves, appearing as rounded or linear, or “root-like” darkened areas. Thallus very thin on the
surface of the host plant, superficial, comprising neatly or
irregularly arranged, angular or cylindrical to cuboid cells,
with brown to dark brown walls. Sexual morph: Thyriothecia developing under thallus tissue, globose to subglobose, flattened, with a porate, central ostiole. Hamathecium
usually lacking pseudoparaphyses, sometimes comprising
hyaline pseudoparaphyses. Asci 6–8-spored, bitunicate, fissitunicate, subglobose to clavate, short pedicellate or apedicellate, long pedicellate when immature in some species,
apically rounded, sometimes with small ocular chamber or
apical ring. Ascospores 2–4-seriate, oblong to ellipsoidal,
or clavate, hyaline to pale brown, 1–3-septate or muriform,
slightly constricted at the septa in some species, smoothwalled. Asexual morph: Undetermined.
Type: Trichopeltina Theiss.
Notes: Trichopeltinaceae are epiphytes on leaves of
various plants with a worldwide distribution. The name
Trichopeltinaceae was predated by Trichopeltaceae (Theissen 1914b). Batista et al. (1958) renamed the family to
Trichopeltinaceae based on Trichopeltina, and accommodated genera that have flattened thyriothecia developing
under a thallus viz. Brefeldiella, Petrakiopeltis, Phragmoscutella, Trichopeltella, Trichopeltheca, Trichopeltina, Trichopeltospora and Trichopeltum. Genera of
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Fungal Diversity (2020) 105:17–318
Trichopeltinaceae with thyriothecia were transferred to
Stephanothecaceae (Petrak 1947c; Luttrell 1951). However, Stephanothecaceae (type genus Stephanotheca)
has a hypostroma, and was considered to be distinct from
Trichopeltinaceae (Bessey 1952; Batista et al. 1958). This
was supported by Jayawardena et al. (2014) who illustrated
the thyriothecia of Stephanotheca as individual and not
covered by a thallus. Hyde et al. (2013) referred the name
Brefeldiellaceae to this group, and divided Brefeldiellaceae
into Brefeldiineae and Trichopeltineae. However, the older
name Trichopeltinaceae should be used to accommodate this
group (Hongsanan et al. 2014a). In the following account,
Acrogenotheca, Brefeldiella, Saccardinula, Trichopeltella, Trichopeltheca and Trichopeltina (Trichopeltum) are
included in Trichopeltinaceae based on their thyriothecia
being covered by a “skin-like” thallus.
Trichopeltina Theiss., Beih. bot. Zbl., Abt. 2 32: 3 (1914).
Index Fungorum number: IF 5566; Facesoffungi number:
FoF 06204; 5 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Trichopeltina labecula (Mont.) Theiss.
Notes: Trichopeltina was established by Theiss (1914a)
based on Trichopeltina labecula (≡ Asteroma labecula). Clements and Shear (1931) placed fossil material with a thin
thallus and 1-septate ascospores in Trichopeltina. Species
of Trichopeltina are characterized by a superficial, irregularly arranged thin thallus, globose to subglobose flattened
ostioles, thyriothecia, 8-spored, clavate, fissitunicate asci
with hyaline, long clavate, 1–3-septate ascospores (Hongsanan et al. 2014a). However, Dilcher (1965) disagreed that
the fossil material cited by Clements and Shear (1931) is
Trichopeltina.
Trichopeltina labecula (Mont.) Theiss., Zentbl. Bakt. ParasitKde, Abt. II 39: 630 (1914).
≡ Asteroma labecula Mont., Annls Sci. Nat., Bot., sér. 2
14: 328 (1840).
Index Fungorum number: IF 220570; Facesoffungi numbers: FoF 07912, Fig. 134
Epiphytic on leaves, appearing as “root-like” darkened
areas. Thallus very thin, superficial, comprising neatly or
irregularly arranged, angular or cylindrical to cuboid cells,
with brown to dark brown walls. Sexual morph: Thyriothecia 65–78 μm diam. × 33–48 μm high ( x̄ = 74 × 39 μm, n =
10), developing under thallus tissue, globose to subglobose,
flattened, with a porate, central ostiole. Hamathecium comprising branched, aseptate pseudoparaphyses. Asci 20–29 ×
14–17 μm ( x̄ = 26 × 15 μm, n = 10), 8-spored, bitunicate,
fissitunicate, clavate, short pedicellate or apedicellate, long
pedicellate when immature, apically rounded, usually with
a small ocular chamber. Ascospores 15–17 × 3–4 μm ( x̄
= 16 × 3 μm, n = 10), 2–3-seriate, long clavate, hyaline,
Fungal Diversity (2020) 105:17–318
sometimes pale brown in the lower cells, 1–2-septate,
slightly constricted at each septum, upper cells wider and
shorter than lower cells, smooth-walled. Asexual morph:
Undetermied.
Material examined: France, Guiana, Gabaret River, on
coriaceous leaf of Desfontainia, September 1835, Collec no.
664 (PC 0084645, syntype of Asterina labecula).
Other genera included
Acrogenotheca Cif. & Bat., in Batista & Ciferri, Saccardoa
2: 51 (1963).
Index Fungorum number: IF 49; Facesoffungi number:
FoF 07913; – 3 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Acrogenotheca pulcherrima Bat. & Cif,
Saccardoa 2: 52 (1963).
Notes: Acrogenotheca comprises three species, A. elegans, A. ornata and A. pulcherrima. Hongsanan et al.
(2014a) examined the type material of A. pulcherrima and
found that it has clavate asci, with clavate, 3-celled, greyish
brown ascospores which differ from those of Trichopeltina.
Brefeldiella Speg., Boln Acad. nac. Cienc. Córdoba 11(4):
558 (1889).
Index Fungorum number: IF 648; Facesoffungi number:
FoF 07914; – 4 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Brefeldiella brasiliensis Speg., Boln Acad.
nac. Cienc. Córdoba 11(4): 558 (1889).
Notes: The type collection of this genus was examined by
Eriksson (1981), but specimens were considered immature
(Hyde et al. 2013). Species of Brefeldiella are characterized
by a superficial, irregularly arranged thin thallus, globose to
subglobose flattened ostioles, thyriothecia, 8-spored, cylindric or clavate, fissitunicate asci with hyaline, oblong to
ellipsoidal, 1-septate ascospores (Hongsanan et al. 2014a).
Brefeldiella and Trichopeltella need molecular studies to
investigate their separation (Hongsanan et al. 2014a).
Saccardinula Speg., Anal. Soc. cient. argent. 19(6): 257
(1885).
Index Fungorum number: IF 4809; Facesoffungi number:
FoF 07915; – 12 morphological species (Species Fungorum
2020), molecular data unavailable.
Type Species: Saccardinula guaranitica Sacc., Anal.
Soc. Cient. Argent. 19:257 (1885).
Notes: Saccardinula was introduced by Spegazzini (1885)
and the type species (Saccardinula guaranitica) was illustrated in Li et al. (2011). Saccardinula guaranitica has a
less well-developed thallus than Elsinoaceae species which
covers the thyriothecia. Asci are saccate and typical of
Trichopeltinaceae, while ascospores differ from Elsinoceae
as they are muriform (Hyde et al. 2013; Li et al. 2011). Li
291
et al. (2011) and Jayawardena et al. (2014) placed Saccardinula in Brefeldiellaceae and thus the genus should belong
in Trichopeltinaceae (Hongsanan et al. 2014a).
Trichopeltella Höhn., Sber. Akad. Wiss. Wien, Math.naturw. Kl., Abt. 1 119: 458 (1910).
Index Fungorum number: IF 5564; Facesoffungi number:
FoF 07916; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Trichopeltella montana (Racib.) Höhn.,
Sber. Akad. Wiss. Wien, Math.-naturw. Kl., Abt. 1 119:
458 (1910).
≡ Trichopeltis montana Racib., Bull. int. Acad. Sci. Lett.
Cracovie, Cl. sci. math. nat. Sér. B, sci. nat. 3: 378 (1909).
Notes: Wu et al. (2014) examined the holotype of T. montana, but the specimen only comprised the asexual morph.
Thus, the comparison with Brefeldiella and Trichopeltella
is unclear. Hyde et al. (2013) mentioned that Trichopeltella
may be a synonym of Brefeldiella. Based on its distinct asexual morph, with a “root-like” thallus and aseptate hyaline
conidia it is accommodated as a separate genus in Trichopeltinaceae (Hongsanan et al. 2014a).
Trichopeltheca Bat., C.A.A. Costa & Cif., Atti Ist. bot.
Univ. Lab. crittog. Pavia, Ser. 5 15: 44 (1958).
Index Fungorum number: IF 5565; Facesoffungi number:
FoF 07917; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Trichopeltheca asiatica Bat., C.A.A. Costa
& Cif., Publicações Inst. Micol. Recife 90: 13 (1958).
Notes: Trichopeltheca is similar to Saccardinula based
on its muriform ascospores, but differs in ascus shape and
thallus form. Saccardinula has globose to ovoid asci and the
thallus is rounded (Spegazzini 1885), while Trichopeltheca
has clavate to sub-cylindrical asci and the thallus appears
“root-like” (Batista et al. 1958).
Trichothyrinula Petr. Sydowia 4(1–6): 171 (1950).
Index Fungorum number: IF 5595; Facesoffungi number:
FoF 07918; – 2 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Trichothyrinula sydowii Petr., Sydowia
4(1–6): 171 (1950).
Notes: Müller and von Arx (1962) introduced the second
species, Trichothyrinula petrakii and assigned it to Trichothyriaceae. Lumbsch and Huhndorf (2010) placed Trichothyrinula in Microthyriaceae. However, Trichothyrinula is
similar to Brefeldiella, but differs by its smaller globose to
subglobose, apedicellate asci (Brefeldiella has cylindricclavate, short-pedicellate asci), and rough ornamented spore
wall, with spores becoming brownish at maturity (Wu et al.
2014).
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Fungal Diversity (2020) 105:17–318
Economic and ecological significance
Members of this family are epiphytic on living leaves. However, there is no record on penetration organs which can
uptake nutrients from plants. The thallus covers host surface
and can reduce respiration and photosynthesis of its plant
host.
upper covering cells. Hawksworth et al. (1983) included Trichothyrium under Microthyriaceae, Wu et al. (2011b) transferred Trichothyrium back to Trichothyriaceae based on its
fungicolous life mode and a well-developed basal peridium.
This was followed in subsequent studies (Hyde et al. 2013;
Wijayawardene et al. 2018).
Trichothyriaceae Theiss., Beih. bot. Zbl., Abt. 2 32: 3
(1914).
Index Fungorum number: IF 81497; Facesoffungi numbers: FoF 06893; 75 species.
Parasitic on Diatrypaceae, Meliolaceae and lichens, or
on their conidial morphs. Mycelium superficial, forming
thalli on hyphae of parasitized fungus, membranous. Sexual
morph: Thyriothecia superficial, solitary flattened, circular,
brown, with a central ostiole. Upper wall comprising radial
arranged cells in parallel rows of textura epidermoidea,
anastomosing at the ostiole. Basal wall well-developed,
with a single-cell thick layer. Asci 8-spored, bitunicate, fissitunicate, obclavate, arranged at the periphery and converge
towards the ostiole. Ascospores overlapping, hyaline or light
brown, 1-septate, not constricted at the septum, sometimes
with ciliate appendages. Asexual morph: Hyphomycetous.
reported as Hansfordiella and Isthmospora (Hyde et al.
2011).
Type: Trichothyrium Speg.
Notes: Trichothyriaceae was introduced by Theissen
(1914a) accommodating Loranthomyces, Trichothyriella,
Trichothyriopsis and the type genus Trichothyrium, Hawksworth et al. (1983) included genera of Trichothyriaceae
under Microthyriaceae, and this was followed by Lumbsch
and Huhndorf (2010). Wu et al. (2011b) transferred them
back to Trichothyriaceae based on its fungicolous life mode
and a well-developed basal peridium, and included another
parasitic genus Lichenopeltella in the family. This was
followed by Hyde et al. (2013) and Wijayawardene et al.
(2018). Trichothyriaceae differs from other thryriothecial
families by a well-developed peridium, but there are too few
morphological characters to delimit those similar genera.
Sequence data is not available for this family.
Trichothyrium sarciniferum Speg., Boln Acad. nac. Cienc.
Córdoba 11(4): 556 (1889).
Index Fungorum number: IF 161963; Facesoffungi number: FoF 07919; Fig. 135
Description: see Hyde et al. (2013).
Material examined: Brazil, Apizhy, on Meliolaceae
on unidentified leaves, 1881, C. Spegazzini (LPS1441,
holotype).
Trichothyrium Speg., Boln Acad. nac. Cienc. Córdoba
11(4): 555 (1889).
Index Fungorum number: IF 5598; Facesoffungi number:
FoF 06894; 25 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Trichothyrium sarciniferum Speg.
Notes: Trichothyrium is characterized by flattened ascomata with an upper and lower, single-cell thick layer occurring on the surface of parallel hyphal bands. Another genus
forming a similar cellular hypostroma with lobed thalli is
Brefeldiella, which is characterized by ascomata dispersed
beneath the band-like thallus and opening by a fissure of the
13
Other genera included:
Lichenopeltella Höhn., Sber. Akad. Wiss. Wien, Math.naturw. Kl., Abt. 1 128: 553 (1919).
Index Fungorum number: IF 2850; Facesoffungi number:
FoF 06895; 48 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Lichenopeltella maculans (Zopf)
Höhn. Sber. Akad Wiss. Wien, Math.-naturw.Kl.,
Abt.1128(7–8):553 (1919).
≡ Microthyrium maculans Zopf, Nova Acta Acad. Caes.
Leop.-Carol. German. Nat. Cur. 70(4): 258 (1898).
Notes: Müller and von Arx (1962) and von Arx and Müller (1975) placed Lichenopeltella in Leptopeltidaceae, and
this was followed in Kirk et al. (2008) who accept Leptopeltidaceae as one of three families in Microthyriales.
However, this is not followed in the more recent classifications in Lumbsch and Huhndorf (2010), and Wu et al.
(2011b) placed it in Trichothyriaceae based on its fungicolous life mode.
Macrographa Etayo, in Etayo & Sancho, Biblthca Lichenol.
98: 116 (2008).
Index Fungorum number: IF 532867; Facesoffungi number: FoF 06896; 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Macrographa antarctica Etayo, in Etayo &
Sancho, Biblthca Lichenol. 98: 116 (2008).
Notes: Macrographa was introduced as a monotypic
genus occurring on lichens. This genus is characterised by
multiloculate ascostromata, ellipsoidal-fusiform asci and
1-septate ascospores (Wu et al. 2011b). Lumbsch and Huhndorf (2010) accepted it as a member of Microthyriaceae.
Wu et al. (2011b) placed it in Trichothyriaceae, along with
Lichenopeltella, due to their fungicolous life mode.
Pachythyrium G. Arnaud ex Spooner &P.M. Kirk, Mycol.
Res. 94(2): 228 (1990).
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Index Fungorum number: IF 25378; Facesoffungi number: FoF 06897; 1 morphological species (Species Fungorum 2020), molecular data unavailable.
Type species: Pachythyrium parasiticum (Fabre) G.
Arnaud ex Spooner & P.M. Kirk, Mycol. Res. 94(2): 228
(1990).
≡ Bertia parasitica Fabre, Annls Sci. Nat., Bot., sér. 6 9:
95 (1879) [1878].
Notes: Pachythyrium was introduced as a monotypic
genus, which is synonymised from Bertia parasitica occurring on Eutypa lata. Hawksworth et al. (1995) and Lumbsch
and Huhndorf (2010) accepted it as a member of Microthyriaceae. Wu et al. (2011b) placed it in Trichothyriaceae, and
noted that the placement of this genus in Trichothyriaceae
may be warranted.
Economic and ecological significance
Trichothyriaceae is a poorly studied family parasitic on
fungi.
Vizellaceae H.J. Swart, Trans. Br. mycol. Soc. 57: 456
(1971).
Index Fungorum number: IF 81519; Facesoffungi number: FoF 07920, 31 species.
Parasitic or saprophytic on leaves, tropical in distribution, developing in or beneath the cuticle, often forming an
intracuticular mycelium, without superficial hyphae. Sexual
morph: Ascostromata dimidiate, scutate or crustose, roundish or elongated in outline, with a thin wall composed of
small, dark cells, opening by a fissure or dehiscence, central
part of loculus may be sterile. Hamathecium comprising filamentous, pseudoparaphyses, often filling the central part of
the loculus or may be sparse or absent. Asci 6–8-spored,
bitunicate, probably evanescent, broadly clavate or nearly
cylindrical. Ascospores 2–3-seriate, overlapping, hyaline
to brown, 1–2-celled or with small appendage cell at the
base, with a light transverse band. Asexual morph: linked
to Chrysogloeum and Manginula. Pycnidia disc-shaped with
upper wall of irregularly arranged cells, subglobose, dark
brown to black, scutate or crustose. Conidiogenous cells
hyaline, cylindrical, holoblastic. Conidia ovoid to ellipsoidal or rarely subglobose, brown to dark brown, with transverse hyaline band, formed on phialidic, hyaline cells lining
the inner cavity of the upper wall (Hughes 1953a, b; Petrak
1953).
Type: Vizella Sacc.
Notes: Vizellaceae was introduced to include Blasdalea
and Vizella (Swart 1971). Entopeltis was synonymized under
Vizella (Swart 1971). Hughes (1953a) described the mycelium of the type species and confirmed mycelium composed
of hyaline to light brown hyphae with thick, dark brown
cross walls. Swart (1971) suggested that the mycelium might
well be of diagnostic value, but several authors could not
293
observe this because mycelium is entirely hyaline and thus
difficult to see. In most species of Vizellaceae, the mycelium
branches frequently at fairly wide angles, and so forms a network without radial orientation (Hyde et al. 2013). von Arx
and Müller (1975) noted that Vizella and Entopeltis can be
distinguished by the arrangement of the asci and ascospores
characters. van Wyk et al. (1976) studied the type species
and other Entopeltis and Vizella species and suggested that
they could not be distinguished. Therefore, Entopeltis should
be synonymised under Vizella.
The asexual morph of most species in this family are
undetermined, except for three Vizella species and Blasdalea disciformis. Hughes (1953a) described a conidial form
associated with Vizella hendrickxii. Swart (1971) introduced
two new species to this genus from Australia with conidial
morphs. The asexual morph of Blasdalea is Chrysogloeum
(Petrak 1953). Wijayawardene et al. (2012) listed the asexual
morphs as Chrysogloeum and Manginula. Chrysogloeum
was established to accommodate the coelomycetous asexual morphs of Blasdalea (as Singeriella). Sivanesan (1984)
considered that Vizella has Manginula or “Manginula”-like
asexual morphs.
Vizella Sacc., Syll. Fung. (Abellini) 2: 662 (1883).
Index Fungorum number: IF 5747; Facesoffungi number:
FoF 07921; 30 morphologocal species (Species Fungorum
2020), molecular data unavailable.
Type species: Vizella conferta (Cooke) Sacc.
≡ Micropeltis conferta Cooke, Grevillea 6(no. 39): 118
(1878)
Notes: The confusion between Entopeltis and Vizella
is discussed in the family notes and in Hyde et al. (2013).
The ascomata of Vizella have thin walls as in Entopeltis
(Fig. 136). This intermediate form between Vizella and
Entopeltis further indicates they all two genera may be
congeneric. Mycerema was introduced by Batista et al.
(1963). It was considered to resemble Vizella in ascospores
and pseudoparaphyses characters (Hyde et al. 2013). However, Phookamsak et al. (2016) indicated that the characters
described in Hyde et al. (2013) is Vizella bingervilliana
(see more detail in the note of Mycerema, Hongsanan et al.
2020).
Other genera included
Acarella Syd., Annls mycol. 25(1/2): 123 (1927).
Index Fungorum number: IF 7005; Facesoffungi number:
FoF 06207; – 3 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Acarella costaricensis Syd., Annls mycol.
25(1/2): 123 (1927).
Notes: Acarella is characterized by scattered, small, completely superficial pycnidia, with a thin peridium, and ellipsoid to cylindrical conidia. The sexual morph of Acarella is
13
294
undetermined. Perizomella was synonymised under Acarella
based on the host relationship and morphological characters
(Pem et al. 2019c). Acarella is placed in Vizellaceae due to
the asexual species have ovoid to ellipsoidal or rarely subglobose, brown to dark brown, with transverse hyaline band,
formed on phialidic, hyaline cells lining the inner cavity of
the upper wall (Hyde et al. 2013; Pem et al. 2019c), pending
fresh collections and molecular data.
Blasdalea Sacc. & P. Syd., Syll. Fung. (Abelini) 16: 634
(1902).
Index Fungorum number: IF 592; Facesoffungi number:
FoF 07922; – 1 morphological species (Species Fungorum
2020), molecular data unavailable.
Type species: Blasdalea disciformis (Rehm) Sacc. & P.
Syd., Syll. Fung. (Abellini) 16: 634 (1902).
≡ Vizella disciformis Rehm, Hedwigia 39(4): 227 (1900).
Notes: von Arx and Müller (1954) illustrated the type
species which has cylindrical asci and abundant pseudoparaphyses and ascomata opening by a large ostiole. Hyde
et al. (2013) were unable to find the sexual morph in the
holotype which was described in Rehm (1900). They found
one-celled, brown conidia with a hyaline transverse band,
some two-celled, hyaline to brown, with a medium lighter
septum. Blasdalea may be related with Fasciatispora which
has unitunicate asci, but the nature of the asci in the illustration provided in von Arx and Müller (1954) is not clear.
In this study, Blasdalea is maintained as a distinct genus in
Vizellaceae pending fresh collections and molecular studies.
Economic and ecological significance
Most species in Vizellaceae are parasitic on leaves which can
uptake nutrients from plant issue.
Acknowledgements Sinang Honsanan would like to thank National
Natural Science Foundation of China for supporting the project Biodiversity, Taxonomy, Phylogeny, Evolution and Phytogeography of
phytopathogens in Dothideomycetes from Southern China (Grant No.
31950410548) for funding this research. Ning Xie would like to thank
Project of DEGP (2019KTSCX150). Kevin D Hyde thanks the Thailand Research Fund for the grant RDG6130001 entitled “Impact of
climate change on fungal diversity and biogeography in the Greater
Mekong Subregion”. Rungtiwa Phookamsak thanks CAS President’s
International Fellowship Initiative (PIFI) for young staff (Grant
No. Y9215811Q1), the Yunnan Provincial Department of Human
Resources and Social Security (Grant No. Y836181261), and National
Science Foundation of China (NSFC) project code 31850410489
(Grant No. Y81I982211) for financial support. Dhanushka Wanasinghe
would like to thank CAS President’s International Fellowship Initiative
(PIFI) for funding his postdoctoral research (number 2019PC0008) and
the 64th batch of China Postdoctoral Science Foundation (Grant No.
Y913083271). Vemuri V. Sarma would like to thank SERB, Department of Science and Technology, Government of India, for funding
a project (SERB/SB/SO/PS/18/2014 dt.19.5.2015) and Ministry of
Earth Sciences (MOES), Govt. of India for funding a project (Sanction order: MOES/36/OO1S/Extra/40/ 2014/PC-IV dt. 14.01.2015); the
Department of Biotechnology, Pondicherry University for facilities;
13
Fungal Diversity (2020) 105:17–318
forest departments of Andaman and Nicobar Islands and Tamil Nadu,
India are thanked for providing permission to collect samples. Saranyaphat Boonmee would like to thank the National Research Council
of Thailand (projects no. 61215320013 and No. 61215320023), the
Thailand Research Fund (project no. TRG6180001) and Plant Genetic
Conservation Project under the Royal Initiation of Her Royal Highness Princess Maha Chakri Sirindhorn - Mae Fah Luang University.
Alan JL Phillips acknowledges the support from UIDB/04046/2020
and UIDP/04046/2020 Centre grants from FCT, Portugal (to BioISI). Saowaluck Tibpromma would like to thank the International
Postdoctoral Exchange Fellowship Program (number Y9180822S1),
CAS President’s International Fellowship Initiative (PIFI) (number
2020PC0009), China Postdoctoral Science Foundation, and the Yunnan Human Resources and Social Security Department Foundation
for funding her postdoctoral research. Shu-Hua Jiang would like to
thank the National Natural Science Foundation of China (Project
Nos. 31800010 and 31750001) for financial support. Nalin N. Wijayawardene thanks the National Natural Science Foundation of China (No.
NSFC 31950410558) and Guizhou Medical University (grant number
FAMP201906K). Dong-Qin Dai would like to thank tthe National Natural Science Foundation of China (No. NSFC 31760013), the Scientific
Research Foundation of Yunnan Provincial Department of Education
(2017ZZX186) and the Thousand Talents Plan, Youth Project of Yunnan Provinces for finance support. R Jeewon would like to thank the
University of Mauritius and Mae Fah Luang University for support.
Mingkwan Doilom thanks the 5th batch of Postdoctoral Orientation
Training Personnel in Yunnan Province (Grant No. Y934283261) and
the 64th batch of China Postdoctoral Science Foundation (Grant No.
Y913082271). M Niranjan thanks SERB, Govt. of India for a fellowship. Huang Zhang would like to thank Natural Science Foundation
of China (NSF 31500017). Jadson DP Bezerra thanks the Conselho
Nacional de Desenvolvimento Científico e Tecnológico (CNPq),
the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior
(CAPES, Finance Code 001), and the Fundação de Amparo à Ciência
e Tecnologia de Pernambuco (FACEPE) for fellowship. B. Devadatha
thanks MOES, Govt. of India for a fellowship. Hai-Xia Wu would
like to the Fundamental Research Funds for the Central Non-profit
Research Institution of CAF (Grant No. CAFYBB2019QB005), and the
Ten Thousand Talents Plan, Youth Top Project of Yunnan Provinces
for finance support. Ausana Mapook thanks to Research and Researchers for Industries (RRI) under Thailand Research Fund for a personal
grant (PHD57I0012). Putarak Chomnunti would like to thank Mae
Fah Luang University (Grant No. DR256201012003) and DiversityBased Economy Development Office and National Research Council
of Thailand Research (Grant No. T2561022) for the financial support.
Satinee Suetrong thanks the collaborative project between BIOTEC and
Department of Marine and Coastal Resources (DMCR), Ministry of
Natural Resources and Environmentunder a project: Marine Microbes
for National Reserves: Alternative Ways of State Property. N. Chaiwan would like to thank the Thailand Research Fund (PHD60K0147).
Jianchu Xu thanks the Strategic Priority Research Program of Chinese Academy of Sciences. We would like to thank Huzefa Raja who
provide the plate of Minutisphaera parafimbriatispora, Naruemon
Huanraluek to provide plate of Pyrenidium actinellum, LiYan Mei to
provide plate of Meliolina cladotricha, Wen-Jing Li to provide pictures
of Pseudorobillarda phragmitis, Felix Schumm to provide ascospore
images of Bogoriella isthmospora and Schummia angulate, Nick Turland for assistance in resolving the nomenclatural status of the name
in Strigulaceae. We thank Supalak Yacharoen for providing part of the
image plate of Trypetheliopsis boninensis.
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317
Affiliations
Sinang Hongsanan1,2,6 · Kevin D. Hyde2,3,7 · Rungtiwa Phookamsak2,3,4,5,8,35 · Dhanushka N. Wanasinghe3,4,5,35 ·
Eric H. C. McKenzie9 · V. Venkateswara Sarma10 · Robert Lücking12 · Saranyaphat Boonmee2,11 ·
Jayarama D. Bhat13 · Ning‑Guo Liu2,14,15 · Danushka S. Tennakoon2,11 · Dhandevi Pem2,11 ·
Anuruddha Karunarathna2,3,4,6,50 · Shu‑Hua Jiang17 · Gareth E. B. Jones18,19 · Alan J. L. Phillips20 ·
Ishara S. Manawasinghe2,7,21 · Saowaluck Tibpromma3,4,5,35 · Subashini C. Jayasiri2 · Diana Sandamali2,11,50 ·
Ruvishika S. Jayawardena2,11 · Nalin N. Wijayawardene22 · Anusha H. Ekanayaka2 · Rajesh Jeewon23 ·
Yong‑Zhong Lu24 · Chayanard Phukhamsakda2 · Asha J. Dissanayake16 · Xiang‑Yu Zeng2,25,26 · Zong‑Long Luo27 ·
Qing Tian2 · Kasun M. Thambugala28 · Dongqin Dai22 · Milan C. Samarakoon2,48 · K. W. Thilini Chethana2,11 ·
Damien Ertz29,30 · Mingkwan Doilom3,4,5,35 · Jian‑Kui (Jack) Liu16 · Sergio Pérez‑Ortega31 · Ave Suija32 ·
Chanokned Senwanna2,6 · Subodini N. Wijesinghe2,11,33 · Mekala Niranjan10,36 · Sheng‑Nan Zhang2,6 ·
Hiran A. Ariyawansa34 · Hong‑Bo Jiang2,3,4,11 · Jin‑Feng Zhang2,37 · Chada Norphanphoun2 · Nimali I. de Silva3,48 ·
Vinodhini Thiyagaraja2,3,4,6 · Huang Zhang38 · Jadson D. P. Bezerra39 · Ricardo Miranda‑González40 ·
André Aptroot41 · Hiroyuki Kashiwadani42 · Dulanjalee Harishchandra2,11,21 · Emmanuël Sérusiaux43 ·
Pranami D. Abeywickrama2,11,21 · Dan‑Feng Bao2,6,27 · Bandarupalli Devadatha2,10 · Hai‑Xia Wu44 ·
Kwang Hee Moon45 · Cecile Gueidan46 · Felix Schumm47 · Digvijayini Bundhun2,6 · Ausana Mapook2 ·
Jutamart Monkai2 · Chitrabhanu S. Bhunjun2,11 · Putarak Chomnunti11 · Satinee Suetrong49 · Napalai Chaiwan2,3 ·
Monika C. Dayarathne33 · Jing Yang2,15 · Achala R. Rathnayaka2,11,50 · Jian‑Chu Xu3,4,5,35 · Jiesheng Zheng1 ·
Gang Liu1 · Yao Feng51 · Ning Xie1
* Ning Xie
ning.xie@szu.edu.cn
1
2
3
Guangdong Provincial Key Laboratory for Plant Epigenetics,
Shenzhen Key Laboratory of Microbial Genetic Engineering,
College of Life Science and Oceanography, Shenzhen
University, Shenzhen 518060, People’s Republic of China
Center of Excellence in Fungal Research, Mae Fah Luang
University, Chiang Rai 57100, Thailand
CAS Key Laboratory for Plant Diversity and Biogeography
of East Asia (KLPB), Kunming Institute of Botany,
Chinese Academy of Science, Kunming 650201, Yunnan,
People’s Republic of China
11
School of Science, Mae Fah Luang University,
Chiang Rai 57100, Thailand
12
Botanischer Garten und Botanisches Museum, Freie
Universität Berlin, Königin-Luise-Straße 6–8, 14195 Berlin,
Germany
13
128/1-J, Azad Co-Op Housing Society, Curca,
P.O. Goa Velha 403108, India
14
Faculty of Agriculture, Natural Resources and Environment,
Naresuan University, Phitsanulok 65000, Thailand
15
Guizhou Key Laboratory of Agricultural Biotechnology,
Guizhou Academy of Agricultural Sciences,
Guiyang 550006, People’s Republic of China
4
East and Central Asia Regional Office, World
Agroforestry Centre (ICRAF), Kunming 650201, Yunnan,
People’s Republic of China
16
School of Life Science and Technology, University
of Electronic Science and Technology of China,
Chengdu 611731, People’s Republic of China
5
Centre for Mountain Futures (CMF), Kunming
Institute of Botany, Kunming 650201, Yunnan,
People’s Republic of China
17
State Key Laboratory of Mycology, Institute
of Microbiology, Chinese Academy of Sciences,
Beijing 100101, People’s Republic of China
6
Department of Entomology and Plant Pathology, Faculty
of Agriculture, Chiang Mai University, Chiang Mai 50002,
Thailand
18
Department of Botany and Microbiology, College of Science,
King Saud University, P.O Box 2455, Riyadh 11451,
Kingdom of Saudi Arabia
7
Institute of Plant Health, Zhongkai University of Agriculture
and Engineering, Haizhu District, Guangzhou 510225,
People’s Republic of China
19
Nantgaredig, 33B St Edwards Road, Southsea,
Hants PO5 3DH, UK
20
8
World Agroforestry Centre, East and Central Asia,
132 Lanhei Road, Kunming 650201, Yunnan,
People’s Republic of China
Faculdade de Ciências, Biosystems and Integrative Sciences
Institute (BioISI), Universidade de Lisboa, Lisbon, Portugal
21
Beijing Key Laboratory of Environment Friendly
Management on Fruit Diseases and Pests in North China,
Institute of Plant and Environment Protection, Beijing
Academy of Agriculture and Forestry Sciences, Beijing,
China
22
Center for Yunnan Plateau Biological Resources Protection
and Utilization, College of Biological Resource and Food
Engineering, Qujing Normal University, Qujing 655011,
Yunnan, People’s Republic of China
9
Manaaki Whenua-Landcare Research, Private Mail Bag
92170, Auckland, New Zealand
10
Fungal Biotechnology Lab, Department of Biotechnology,
School of Life Sciences, Pondicherry University, Kalapet,
Pondicherry 605014, India
13
318
Fungal Diversity (2020) 105:17–318
23
Dept of Health Sciences, Faculty of Medicine and Health
Sciences, University of Mauritius, Reduit, Mauritius
24
25
38
School of Food and Pharmaceutical Engineering,
Guizhou Institute of Technology, Guiyang 550003,
People’s Republic of China
Faculty of Agriculture and Food, Kunming University
of Science & Technology, Kunming 650500,
People’s Republic of China
39
State Key Laboratory Breeding Base of Green Pesticide
and Agricultural Bioengineering, Key Laboratory of Green
Pesticide and Agricultural Bioengineering, Ministry
of Education, Guizhou University, Guiyang 550025,
People’s Republic of China
Setor de Micologia, Departamento de Biociências e
Tecnologia, Instituto de Patologia Tropical e Saúde Pública,
Universidade Federal de Goiás, Rua 235 – s/n – Setor
Universitário, CEP: 74605-050 Goiânia, GO, Brazil
40
Department of Botany and Plant Pathology, Oregon State
University, 2082 Cordley Hall, Corvallis, OR 97331-2902,
USA
26
The Engineering Research Center of Southwest
Bio-Pharmaceutical Resource, Ministry
of Education, Guizhou University, Guiyang 550025,
People’s Republic of China
41
Instituto de Biociências, Universidade Federal de
Mato Grosso do Sul, Avenida Costa e Silva, s/n
Bairro Universitário, CEP 79070-900 Campo Grande,
Mato Grosso do Sul, Brazil
27
College of Agriculture and Biological Sciences, Dali
University, Dali 671003, Yunnan, People’s Republic of China
42
28
Genetics and Molecular Biology Unit, Faculty
of Applied Sciences, University of Sri Jayewardenepura,
Gangodawila, Nugegoda, Sri Lanka
Department of Botany, National Museum of Nature
and Science, 4-1-1 Amakubo, Tsukuba 305-0005, Ibaraki,
Japan
43
Department Research, Meise Botanic Garden, Nieuwelaan
38, 1860 Meise, Belgium
Evolution and Conservation Biology, InBios Research
Center, University of Liège, Sart Tilman B22, Quartier
Vallée 1, Chemin de la Vallée 4, 4000 Liège, Belgium
44
International Fungal Research and Development Centre, The
Research Institute of Resource Insects, Chinese Academy
of Forestry, Kunming 650224, People’s Republic of China
45
Biological Resources Research Department, National
Institute of Biological Resources, Gyoungser-dong, Seo-gu,
Incheon 404-170, Korea
46
Australian National Herbarium, National Research
Collections Australia, CSIRO-NCMI, Canberra, ACT 2601,
Australia
47
Mozartstraße 9, 73117 Wangen, Germany
48
Department of Biology, Faculty of Science, Chiang Mai
University, Chiang Mai 50200, Thailand
49
National Biobank of Thailand (NBT), National Science
and Technology Development Agency (NSTDA), Thailand
Science Park, Thanon Phahonyothin, Tambon Khlong Nueng,
Amphoe Khlong Luang, Pathum Thani 12120, Thailand
50
Department of Plant Medicine, National Chiayi University,
300 Syuefu Road, Chiayi City 60004, Taiwan
51
College of Agriculture, Guizhou University,
Guiyang 550025, Guizhou, People’s Republic of China
29
30
Fédération Wallonie-Bruxelles, Service Général de
l’Enseignement Supérieur et de la Recherche Scientifique,
Rue A. Lavallée 1, 1080 Bruxelles, Belgium
31
Dept. Micología, Real Jardín Botánico (CSIC), Plaza Murillo
2, 28014 Madrid, Spain
32
Institute of Ecology and Earth Sciences, University of Tartu,
Lai 40, Tartu 51005, Estonia
33
Department of Plant Pathology, Agriculture College,
Guizhou University, Guiyang 550025, Guizhou Province,
People’s Republic of China
34
35
36
37
Department of Plant Pathology and Microbiology, College
of Bio-Resources and Agriculture, National Taiwan
University, No.1, Sec. 4, Roosevelt Road, Taipei 106, Taiwan
Honghe Center for Mountain Futures, Kunming
Institute of Botany, Chinese Academy of Sciences,
Honghe County 654400, Yunnan, People’s Republic of China
Department of Botany, Rajiv Gandhi University, Rono Hills,
Doimukh, Itanagar, Arunachal Pradesh 791112, India
Guizhou Tea Research Institute, Guizhou Academy
of Agricultural Sciences, Guiyang 550006,
People’s Republic of China
13