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ARTICLE
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© 2013 International Mycological Association
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volume 4 · no. 2
INTRODUCTION
Cercospora-like fungi are one of the largest groups of
mostly plant pathogenic, leaf-spotting hyphomycetes,
comprising more than 2000 names (Crous & Braun
2003). Numerous economically relevant plant diseases
are involved, as for instance angular leaf spot of bean
(Pseudocercospora griseola), black leaf streak of banana
(Pseudocercospora jiensis), fruit and leaf spot disease of
citrus (Pseudocercospora angolensis), leaf spot disease of
celery (Cercospora apii), sugarbeet (Cercospora beticola),
etc. Cercosporoid fungi are known from all parts of the world,
but they are especially abundant and diverse in tropical
and subtropical areas of Africa, Asia, Australia and Central
to South America (e.g. Beilharz et al. 2002, Braun & Freire
2004, Hernández-Gutiérrez & Dianese 2008, 2009). They
are dematiaceous hyphomycetes characterised by having
macronematous conidiophores, formed singly, in fascicles,
sporodochia or synnemata, with integrated, terminal to
intercalary conidiogenous cells, holoblastic (mono- to
polyblastic) conidiogenesis, percurrent to sympodial
proliferation, forming amero- to scolecosporous conidia,
solitary or in acropetal chains. Cercosporoid hyphomycetes
are asexual fungi with relation to mycosphaerella-like sexual
morphs as far as meiotic states are formed at all. Many or
probably most species are asexual holomorphs, i.e. they
have lost the ability to form sexual morphs. Mycosphaerella
sexual morphs within this complex are morphologically
rather uniform and provide few characters to support a
further splitting into smaller generic units, whereas the
asexual morphs are morphologically strongly differentiated,
which is reected in numerous asexual genera introduced
for particular morphological groups in this complex.
Furthermore, the genus Mycosphaerella proved to be
polyphyletic (Crous et al. 2007). Based on comprehensive
phylogenetic examinations using analyses of DNA sequence
data, the Mycosphaerella complex has been separated
into several families and monophyletic genera (Crous et
al. 2007, 2009b, c, 2013, Groenewald et al. 2013). Due to
the changes of the International Code of Nomenclature for
doi:10.5598/imafungus.2013.04.02.12 IMA FUNGUS · VOLUME 4 · NO 2: 265–345
Cercosporoid fungi (Mycosphaerellaceae) 1. Species on other
fungi, Pteridophyta and Gymnospermae*
Uwe Braun1, Chiharu Nakashima2, and Pedro W. Crous3
1Martin-Luther-Universität, Institut für Biologie, Bereich Geobotanik und Botanischer Garten, Herbarium, Neuwerk 21, 06099 Halle (Saale),
Germany; corresponding author e-mail: uwe.braun@botanik.uni-halle.de
2Graduate School of Bioresources, Mie University, 1577 Kurima-machiya, Tsu, Mie 514-8507, Japan
3CBS-KNAW, Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands
Abstract: Cercosporoid fungi (former Cercospora s. lat.) represent one of the largest groups of hyphomycetes
belonging to the Mycosphaerellaceae (Ascomycota). They include asexual morphs, asexual holomorphs or species
with mycosphaerella-like sexual morphs. Most of them are leaf-spotting plant pathogens with special phytopathological
relevance. The only monograph of Cercospora s. lat., published by Chupp (1954), is badly in need of revision.
However, the treatment of this huge group of fungi can only be accomplished stepwise on the basis of treatments of
cercosporoid fungi on particular host plant families. The present rst part of this series comprises an introduction, a
survey on currently recognised cercosporoid genera, a key to the genera concerned, a discussion of taxonomically
relevant characters, and descriptions and illustrations of cercosporoid species on other fungi (mycophylic taxa),
Pteridophyta and Gymnospermae, arranged in alphabetical order under the particular cercosporoid genera, which
are supplemented by keys to the species concerned. The following taxonomic novelties are introduced: Passalora
austroplenckiae comb. nov., P. backmanii comb. nov., P. condensata comb. nov., P. gymnocladi comb. nov., P.
thalictri comb. nov., Pseudocercospora davalliicola sp. nov., P. chamaecyparidis comb. nov., P. cratevicola nom.
nov., P. gleicheniae comb. nov., P. lygodiicola sp. nov., P. lygodiigena nom. nov., P. nephrolepidigena sp. nov., P.
paraexosporioides sp. nov., P. pini-densiorae var. montantiana comb. et stat. nov., P. pteridigena sp. nov., P. ptisanae
sp. nov., P. sciadopityos sp. nov., P. subramanianii nom. nov., P. thujina comb. nov., and Zasmidium australiense
comb. nov.
Article info: Submitted: 22 August 2013; Accepted: 21 November 2013; Published: 3 December 2013.
Key words:
Ascomycota
Cercospora s. lat.
conifers
ferns
fungicolous
hyphomycetes
*In memoriam Charles D. Chupp (1886–1967), the author of the rst
monograph of Cercospora.
Braun et al.
ARTICLE
266 ima fUNGUS
Algae, Fungi and Plants (ICN) accepted during the Botanical
Congress in Sydney 2011, above all the abolishment of the
previous Article 59 dealing with pleomorphic fungi, asexual
morph-typied genera are now nomenclaturally equivalent
to sexual morph-typied ones (Hawksworth 2011, Norvell
2011, Braun 2012). Hence, several asexual morph genera
within the Mycosphaerella complex can be used as generic
holomorph names for particular groups segregated from this
complex based on morphology and molecular sequence
analyses. Mycosphaerella punctiformis, the type species
of Mycosphaerella, has Ramularia endophylla as asexual
morph (Verkley et al. 2004). The genus Mycosphaerella is
however a facultative synonym of Ramularia, since the latter
genus is older and has priority.
Fries (1849) introduced Passalora, the rst generic name
within the complex of cercosporoid hyphomycetes, followed
by Cercospora introduced by Fresenius (in Fuckel 1863).
Authors of the rst half of the 19th century usually applied broad
generic concepts of Cercospora (e.g. Viégas 1945). Chupp
(1954) published the rst and only monograph of cercosporoid
hyphomycetes in which he followed a very broad generic
concept. He referred most cercosporoid fungi to Cercospora,
reduced several genera to synonymy with the latter genus, e.g.
Ragnhildiana and Cercoseptoria, but maintained Passalora as
separate genus (with Cercosporidium as synonym) for species
with (0–)1(–3)-septate conidia. His general understanding
of “genera confused with Cercospora”, reected in his key
(Chupp 1954: 17–20) in which he recognised genera like
Cercodeuterosporium, Vellosiella and Pseudocercospora, is,
however, vague and confusing. Soon after Chupp’s (1954)
treatment of this fungal group, the erosion of his broad generic
concept set in. Above all Deighton’s numerous treatments
of cercosporoid fungi (Deighton 1967, 1971, 1973, 1974,
1976, 1979, 1983, 1987, 1990) strongly inuenced the further
taxonomy of this fungal group at generic and species rank.
The strong proliferation and splitting of cercosporoid genera
was later questioned due to numerous morphologically
intermediate taxa and a continuum in several previously
applied generic characters as well as the rst phylogenetic
results based on DNA sequence data (Crous et al. 2000,
Crous & Braun 2003), which again lead to a reduction of
recognised cercosporoid genera. Most of these changes in
generic circumscriptions have recently been conrmed during
the course of various molecular DNA studies, although a few
newly segregated cercosporoid genera have been introduced
to reect monophyletic, morphologically separated clades, e.g.
Phaeocercospora (Crous et al. 2012), Pallidocercospora and
Phaeomycocentrospora (Crous et al. 2013). Other genera,
previously sometimes considered to be synonyms, have been
conrmed as separate genera, e.g. Pantospora (Minnis et
al. 2011), Paracercospora s. str. and Scolecostigmina s. str.
(Crous et al. 2013).
The history and the evolution of concepts of cercosporoid
genera was comprehensively discussed in Deighton (1976),
Braun (1995a) and Crous & Braun (2003) as well as recently in
Crous et al. (2013) and Groenewald et al. (2013) with special
emphasis on Pseudocercospora and Cercospora, respectively.
These treatments may be consulted for further details.
Beside numerous, partly controversial treatments of
cercosporoid hyphomycetes at generic level, a very large
number of additional species have been described since
Chupp’s (1954) monograph, and numerous comprehensive
taxonomic treatments and works dealing with particular areas,
including large countries, have been published, e.g. Viégas
(1945, Brazil), Muntañola (1960), Deighton (1967, 1971, 1974,
1976, 1979, 1983, 1987, 1990), Katsuki (1965, Japan), Yen &
Lim (1980, Singapore and the Malay Peninsula), Hsieh & Goh
(1990, Taiwan), Guo & Hsieh (1995, China, Pseudocercospora),
Crous & Braun (1996, South Africa), Braun & Mel’nik (1997,
Russia and adjacent countries), Shin & Kim (2001, Korea),
Guo et al. (2005, China, Cercospora), Guo et al. (2003, China,
Passalora s. lat.), Kamal (2010, India) and Phengsintham et al.
(2013a, b, Laos, Thailand). Hence, Chupp’s (1954) monograph
is badly out of date. There is no modern comprehensive
treatment of cercosporoid species, and taxonomic work on this
fungal group is complicated and time-consuming due to the
large number of species involved, and descriptions scattered
thoughout the mycological and phytopathological literature.
Therefore, a new “Chupp” summarising all taxa of and data
on cercosporoid hyphomycetes is urgently required. To
accomplish a comprehensive treatment of all taxa concerned
in a monograph as in Chupp (1954), we have decided to treat
the cercosporoids occurring on particular host families, i.e., in
principle following Chupp’s (1954) concept, but supplemented
by keys to the species and illustrations for most taxa. In
the present rst contribution, hyperparasitic cercosporoid
species and taxa on hosts belonging to Pteridophyta and
Gymnospermae are treated. After comprehensive treatments
of cercosporoids of all host families, it will be possible
to supplement the individual publications and compile a
monograph.
MATERIAL AND METHODS
The present work is a compilation based on papers and
unpublished data of the authors as well as global literature.
Details on methods are to be found in the papers cited. As
far as new examinations are concerned, fungal structures
have been examined on the base of standard methods of light
microscopy, using an Olympus BX50 microscope, with distilled
water and lactic acid as media, but without any staining. If
possible, measurements of 30 conidia and other structures
have been made at a magnication of ×1000. The following
abbreviations are used: author names follow Brummit & Powell
(1992), journals Bridson (2004a, b), and exsiccatae http://
www.botanischestaatssammlung.de/DatabaseClient/IndExs/
index.jsp (IndExs – Index of Exsiccatae). Taxonomy and
nomenclature of plant families, genera and species are based
on the “Angiosperm Phylogeny Website” (http://www.mobot.
org/mobot/research/apweb/), Tropicos database (http://www.
Tropicos.org/), and The Plant List (http://www.theplantlist.org).
Taxonomically relevant characters
Basic characters at generic rank (for the discrimination of
genera) are mainly conned to colour of conidiophores (hyaline
or pigmented) and the structure of the conidiogenous loci and
conidial hila (conspicuous by being thickened and darkened
versus inconspicuous, neither thickened nor darkened, or at
least unthickend). Other features previously used to distinguish
Cercosporoid fungi 1
ARTICLE
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volume 4 · no. 2
smaller generic units (segregated genera) proved to be of little
relevance at generic level, e.g. absence or formation of external
mycelium in vivo, arrangement and formation of conidiophores
in vivo (formed singly on supercial hyphae, fasciculate, in
sporodochia or synnemata). However, these characters are
useful at species rank, i.e. for the differentiation of allied,
morphologically similar species. The following additional
characters are useful and applicable for the characterisation
of species and their discrimination from allied taxa: shape
and size, septation, pigmentation (pale to dark brown), wall
thickness and surface structure (smooth to verrucose) of
conidiophores; shape, size, integration and proliferation mode
of conidiogenous cells; location, size (diameter) and shape of
conidiogenous loci (scars) (e.g. inconspicuous, on shoulders
caused by sympodial proliferation, denticle-like, etc.);
formation of conidia (solitary or in simple or branched chains),
shape, size, septation (eu- or distoseptate, transversely or
longitudinally to obliquely septate, amero- to scolecosporous),
colourless or pigmented, wall smooth or sculptured; shape and
size, number and location of hila.
SEXUAL MORPHS AND CURRENTLY
RECOGNISED CERCOSPOROID GENERA
Mycosphaerella and mycosphaerella-like
sexual morphs
Sexual morphs of various cercosporoid hyphomycetes were
traditionally referred to as Mycosphaerella. Ascomata of the
latter genus are morphologically rather uniform and provide few
characters to support further splitting. The associated asexual
morphs are, in contrast, morphologically strongly differentiated
and better reect genotypic variation. The type species of
Mycosphaerella, M. punctiformis, has an asexual morph
described as Ramularia endophylla and clusters together
with other Ramularia species in a monophyletic Ramularia
clade within the Mycosphaerellaceae (Verkley et al. 2004,
Crous et al. 2007), so that Mycosphaerella s. str. becomes a
facultative synonym of Ramularia according to the new rules
of the ICN (Art. 59). The older name Ramularia has priority and
should be applied as holomorph name for this monophyletic
unit composed of asexual or sexual morphs, or Ramularia
species with Mycosphaerella sexual morphs in their life cycles.
The further splitting of ascomycetes with mycosphaerella-
like sexual morphs is closely connected with and based
on assemblages of certain species with particular asexual
morphs for which asexual morph-typied generic names are
available. Ramularia (Mycosphaerella) as mucedinaceous
genus, characterised by having colourless conidiophores
and conidia, does not belong to the complex of cercosporoid
genera, and is hence not treated in this series. Ramularia and
allied mucedinaceous genera have been monographed by
Braun (1995a, 1998), and a detailed molecular phylogeny will
be published elsewhere (Videira et al., in prep.).
CERCOSPORA S. STR.
The genus Cercospora was introduced by Fresenius (in Fuckel
1863) for passalora-like species with pluriseptate conidia.
The typication of a genus is essential for further taxonomic
treatments, i.e. for changed generic circumscriptions s. str. or
s. lat. However, the typication of Cercospora was previously
controversially handled and discussed. Cercospora apii was
usually considered as type species (or lectotype species) of
Cercospora (Solheim 1930, Clements & Shear 1931, Ellis
1971, etc.), based on the assumption that the rst description
of this genus dates back to Fresenius (1863). This typication
was accepted by most subsequent authors, including Sutton &
Pons (1988), but proved to be wrong, which was pointed out
by Braun (1995a) who showed that the rst valid description of
Cercospora was published somewhat earlier in Fuckel (1863)
and simultaneously in Fuckel, Fungi Rhen. Exs., Fasc. II, No.
117, 1863 with Cercospora penicillata (C. depazeoides) as
only and hence type species. Fuckel (1863) only added a new
forma (f. apii), which is, however, irrelevant for the typication.
Early circumscriptions of Cercospora comprised Cercospora
ferruginea, a species with thickened, darkened conidiogenous
loci and pigmented conidia, which was later reallocated to
Mycovellosiella and belongs now to Passalora emend. During
the course of the following 100 years, the application of
Cercospora had been continuously widened (Saccardo 1880,
Solheim 1930, etc.). All kinds of supercially similar species,
with or without conspicuous conidiogenous loci, with hyaline or
pigmented conidia, formed singly or in chains, were assigned
to this genus. This broad generic concept was used and
applied by Chupp (1954) in his monograph of Cercospora s.
lat., in which he treated 1 419 species. The number of species
assigned to Cercospora increased rapidly in the following
decades. Pollack (1987) listed more than 3 000 names. Several
early attempts to split Cercospora s. lat. into smaller generic
units had been made, e.g. by Earle (1901) (Cercosporidium),
Ferraris (1909) (Phaeoisariopsis), Spegazzini (1910)
(Cercosporina, Pseudocercospora), Maublanc (1913a, b)
(Asperisporium), Rangel (1917) (Mycovellosiella), Miura (1928)
(Cercosporiopsis), Sydow (1930) (Stenella), Petrak (1951)
(emendation of Chaetotrichum to include some cercosporoid
taxa), Muntañola (1960) (Phaeoramularia), etc. However, the
genera concerned did not gain wide acceptance and application
until Deighton’s (1967, 1971, 1974, 1976, 1979, 1983, 1987,
1990) comprehensive studies on cercosporoid hyphomycetes,
and his new generic concepts and splitting of Cercospora s. lat.
Ellis (1971, 1976) followed Deighton’s concepts and accepted
Cercosporidium, Mycovellosiella, Phaeoramularia and other
cercosporoid genera segregated from Cercospora. Deighton
(1976) re-introduced Pseudocercospora and widened its
circumscription to all kinds of cercosporoid hyphomycetes with
inconspicuous (unthickened, not darkened) conidiogenous
loci, i.e. he nished the gradual reduction of the application
of the genus Cercospora and conned it (s. str.) to species
resembling C. apii, i.e. to taxa with pigmented conidiophores,
thickened and darkened conidiogenous loci (scars) and
colourless, pluriseptate, scolecosporous conidia formed
singly, a concept accepted up to now and conrmed by
molecular sequence analyses in which Cercospora s. str.
proved to represent a monophyletic clade (Crous et al. 2000,
Groenewald et al. 2013). Numerous molecular examinations
of Cercospora species have been carried out, based on
rDNA ITS data as well as multilocal approaches (Stewart et
al. 1999, Crous et al. 2000, 2004a, 2009a, b, Goodwin et al.
Braun et al.
ARTICLE
268 ima fUNGUS
2001, Tessmann et al. 2001, Pretorius et al. 2003, Groenewald
et al. 2005, 2006a, b, 2010, 2013, Montenegro-Calderón
et al. 2011). Crous & Braun (2003) published an annotated
list of Cercospora and Passalora emend. names, including
659 recognised species of Cercospora s. str. and further
281 species names reduced to synonymy with C. apii s. lat.
since they were morphologically not or barely distinguishable
from C. apii s. str. on celery. This concept was inuenced by
assumptions as well as proven results suggesting that C.
apii s. lat. comprises specialised as well as plurivorous races
(Ellis 1971). Crous & Braun (2003) provided a morphological
circumscription of Cercospora apii s. lat. and recommended to
assign new collections on hosts of new families or genera to C.
apii s. lat. if morphologically indistinguishable and if not proven
by means of molecular sequence analyses and/or inoculation
experiments that distinct species are involved. However, the
speciation within Cercospora s. str., above all with regard to C.
apii s. lat., is more complicated than previously assumed and
far from being comprehensively perceived. Detailed molecular
examination of Cercospora s. str. based on a multilocus DNA
approach (ITS, actin, calmodulin, histone H3, and translation
elongation factor 1-alpha genes) has recently been carried out
by Groenewald et al. (2013). The monophyly of Cercospora s.
str. was veried, but the speciation within this genus proved
to be very complicated. Cercospora apii s. lat. as plurivorous
monophyletic species could not be conrmed. Cercospora apii
s. lat. is rather a complicated complex composed of specialised
as well as numerous species with wide host range, often with
overlapping host ranges between particular taxa. The problem
is that most of the taxa involved, including various plurivorous
ones, are morphologically barely distinguishable. Certain hosts
or host ranges are also not reliable for taxonomic purposes
within C. apii s. lat. Cercospora collections on certain hosts
with agreeing morphology found in different geographical
regions do often not belong to a single species, i.e. it is usually
not possible to simply apply old names of Cercospora species
described from Europe or North America to taxa in Africa,
Asia, Australia or South America and vice versa. Therefore,
descriptions of new species on new hosts morphologically
indistinguishable from C. apii s. lat. should be avoided without
support by molecular sequence analyses. The collections
concerned should be simply referred to as C. apii s. lat. (C. apii
complex) as recommended by Crous & Braun (2003).
Attempts to separate Cercospora into several
subgeneric units have been made by Penzes (1927)
(three sections: Brachycercosporae, Macrocercosporae
and Mediocercosporae) and Solheim (1930) (21 sections
based on mycelium internal/external, conidiophores simple/
branched, stroma and conidium shape), which are, however,
barely practicable and not useful since these classications
were derived from a wide range of species of Cercospora s.
lat. that now belong to different genera. Therefore, it is not
surprising that these subgeneric concepts have never been
applied by other authors.
A special problem concerns Cercospora species with
colourless but passalora-like conidia, i.e. broadly ellipsoid-
ovoid to obclavate-cylindrical, only with few septa, e.g.
Cercospora chenopodii (Passalora dubia) and C. sojina (P.
sojina). In this case, the generic relevance of the conidial
shape versus conidial colour was unclear and differently
evaluated. However, this problem has recently been clearly
solved on the basis of molecular sequence analyses
(Groenewald et al. 2012), which showed that species
with hyaline conidia, independent of the conidial shape,
cluster within the Cercospora clade. Other Cercospora
species are characterised by having colourless or very pale
conidiophores. They are supercially similar to Cercosporella
species, but species of the latter genus have quite distinct
conidiogenous loci (Braun 1995a). Braun (1993) introduced
Cercospora subgen. Hyalocercospora for such species.
Molecular data for Cercospora cercosporelloides, the type
species of this subgenus, or any other species assigned to it
in Braun (1995a) are not yet available, i.e. it is still unproven
if they cluster in the monophyletic Cercospora clade.
The following description circumscribes Cercospora
as currently perceived and phylogenetically conrmed as
monophyletic genus:
Cercospora Fresen., in Fuckel, Hedwigia 2: 133
(1863) [and in Fuckel, Fungi Rhen. Exs., Fasc. II, No.
117, 1863].
Type species: Cercospora penicillata (Ces.) Fresen. 1863 (C.
depazeoides (Desm.) Sacc. 1876).
Synonyms: Virgasporium Cooke, Grevillea 3: 182 (1875)
[type species: V. maculatum Cooke 1875].
Cercosporina Speg., Anales Mus. Nac. Buenos Aires 20: 424
(1910) [type species: C. asparagicola Speg. 1910].
Literature: Chupp (1954), Vasudeva (1963), Katsuki (1965),
Ellis (1976: 244), Yen & Lim (1980: 152–166), Hsieh & Goh
(1990), Braun (1995a: 40), Shin & Kim (2001: 20), Guo et al.
(2005), Kamal (2010: 11), Seifert et al. (2011: 128–130).
Description [emend. Crous & Braun (2003: 16)]: Hyphomycetes
(asexual morphs, asexual holomorphs) or Cercospora with
mycosphaerella-like sexual morphs; Mycosphaerellaceae.
Saprobic, secondary invaders or mostly plant pathogenic,
symptomless or usually causing distinct lesions (leaf-spotting
fungi). Mycelium internal, rarely external; hyphae branched,
septate, colourless or almost so to usually pigmented, thin-
walled, smooth, rarely faintly rough-walled. Stromata lacking
to well-developed, substomatal, intaepidermal to deeply
immersed, mostly pigmented, composed of textura angulata
or globosa. Conidiophores mononematous, macronematous,
solitary or fasciculate, in small to large fascicles, rarely in
sporodochial conidiomata, emerging through stomata or
erumpent, very rarely arising from supercial hyphae, erect,
continuous to pluriseptate, hyaline or almost so (subgen.
Hyalocercospora) to mostly pigmented, pale olivaceous to
dark brown (subgen. Cercospora), wall smooth to somewhat
rough, thin to moderately thick; conidiogenous cells
integrated, terminal or intercalary, sometimes conidiophores
aseptate, i.e. reduced to conidiogenous cells, monoblastic,
determinate to usually polyblastic, proliferation sympodial,
rarely percurrent, conidiogenous loci (scars) conspicuous,
thickened and darkened-refractive, planate with minute
central pore. Conidia solitary, rarely in short chains (mainly
under high humidity), mostly scolecosporous, obclavate-
cylindrical, acicular, liform and plurieuseptate, rarely amero-
to phragmosporous, broadly ellipsoid-ovoid to broadly
Cercosporoid fungi 1
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volume 4 · no. 2
obclavate-cylindrical, but always hyaline or subhyaline (with
a pale greenish tinge), thin-walled, smooth or almost so, hila
thickened and darkened, conidial secession schizolytic.
DISTOCERCOSPORA
The genus Distocercospora is passalora-like (thickened and
darkened loci and hila and pigmented conidia), but differs in
having distoseptate conidia. The meaning of distoseptation
as character on generic level within the complex of
cercosporoid genera is, however, unclear. Distosepta may
occasionally occur intermixed between genuine septa in
some Passalora species (Crous & Braun 2003) as well as
in Pseudocercospora cryptomeriicola (Nakashima et al.
2007). It is possible that distosepta mixed with eusepta are
more common in cercosporoid fungi than hitherto known as
it is often rather difcult to discern such septa in taxa with
thin walls. However, cultures and molecular data based on
Japanese material of the type species of Distocercospora, D.
pachyderma, and D. livistonae have recently been examined
by C. Nakashima (unpubl. data), which showed that these
species cluster within the Mycosphaerellaceae clade in a
more basal and separate position. These results indicate
that Distocercospora has to be maintained as separate
genus, distinguished from Passalora by having consistently
distoseptate conidia.
Distocercospora Pons & B. Sutton, Mycol. Pap. 160:
60 (1988).
Type species: Distocercospora pachyderma (Syd. & P. Syd.)
Pons & B. Sutton 1988.
Literature: Braun (1995a: 40), Braun & Mel’nik (1997: 15),
Crous & Braun (2003: 26), Seifert et al. (2011: 187).
Description: Foliicolous, plant pathogenic, leaf spotting
hyphomycetes (asexual morphs), sexual morphs unknown.
Mycelium in vivo internal; hyphae branched, septate,
subhyaline to pigmented, thin-walled, smooth. Stromata
lacking to well-developed, pigmented, textura angulata to
globosa. Conidiophores macronematous, mononematous,
simple to branched, often strongly branched, septate,
pigmented, thin-walled, smooth to rough-walled;
conidiogenous cells integrated, terminal, occasionally
intercalary, proliferation sympodial, conidiogenous loci
conspicuous, somewhat thickened and darkened. Conidia
formed singly, rarely in short chains, scolecosporous, mostly
obclavate to cylindrical, with a single to several transverse
distosepta, subhyaline to pigmented, wall smooth to rough,
hila somewhat thickened and darkened, conidial secession
schizolytic.
PASSALORA EMEND. CROUS & BRAUN (2003)
Passalora, described by Fries (1849), was the rst genus
introduced for cercosporoid hyphomycetes. Fresenius
(in Fuckel 1863) introduced Cercospora for passalora-
like species with pluriseptate conidia. Later the concept of
Cercospora was considerably widened, and culminated in
Chupp’s (1954) monograph of Cercospora s. lat. with very
broad generic concept. Several attempts to segregate smaller
generic units from Cercospora s. lat. for cercosporoids with
thickened and darkened conidiogenous loci were made in the
20th century, but attracted little attention and were barely used,
e.g. Cercosporidium (Earle 1901), Mycovellosiella (Rangel
1917), Ragnhildiana (Solheim 1930), Cercodeuterospora
(Curzi 1932), Berteromyces (Ciferri 1954), Fulvia and
Oreophylla (Ciferri 1954), Phaeoramularia (Muntañola 1960),
Tandonella (Prasad & Verma 1970).
Deighton (1967) reintroduced Cercosporidium for
cercospora-like hyphomycetes with internal mycelium,
usually well-developed stromata, fasciculate conidiophores
and solitary, non-scolecosporous, mostly pale conidia and
maintained Passalora, although very similar, as separate
genus based on “lacking” stromata and pale brown conidia.
For similar species characterised by conidia formed in chains,
Ellis (1971, 1976) and Deighton (1979) accepted and used
Phaeoramularia, and for those with solitary conidiophores
arising from supercial mycelium the old name Mycovellosiella
was reintroduced (Deighton 1974, 1979, 1983; Ellis 1971,
1976). This splitting of the Passalora complex had been the
generally recognised genus concept in the following about
25 years, recognised by most subsequent authors (e.g., von
Arx 1983, Pons & Sutton 1988, Hsieh & Goh 1990, Braun
1995a, Braun & Mel’nik 1997, Shin & Kim 2001, Guo et al.
2003, etc.).
First doubts with regard to the acceptance and rationality
of the differentiation between Passalora and Cercosporidium,
as made in Deighton (1967), were expressed in Deighton’s
(1990) reassessment of the synnematous genus
Phaeoisariopsis in which he conned the latter genus to a few
species around P. griseola, the type species, characterised
by having subconspicuous, non-protuberant conidiogenous
loci on non-geniculate conidiogenous cells. Other former
Phaeoisariopsis species with inconspicuous (unthickened,
not darkened) loci were assigned to Pseudocercospora,
and the remaining species with thickened, darkened loci on
distinctly geniculate conidiogenous cells were proposed to
be candidates for Passalora. Braun (1995b) reassessed the
Passalora complex and proposed to reduce Cercosporidium,
Mycovellosiella and Phaeoramularia to synonymy with
Passalora. Crous & Braun (2003) discussed this generic
complex in detail and came to the conclusion that these
genera and various additional ones should better be merged
under the oldest name Passalora. There is a continuum of
all relevant morphological characters between species of the
genera involved. Numerous examples of intermediate species
were cited and discussed in Crous & Braun (2003). Neither
the conidial formation, solitary or in chains, nor the presence
or absence of supercial mycelium represent reliable features
on generic rank. With regard to morphological features reliably
applicable for the discrimination on generic level, Passalora is
comparable with other cercosporoid and ramularioid genera
like Pseudocercospora, Pseudocercosporella, Ramularia
and Zasmidium. Thus, the new emended circumscription of
Passalora (according to Crous & Braun 2003) encompassed
cercosporoid species with internal and external or only internal
mycelium, solitary, fasciculate to synnematous conidiophores
Braun et al.
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270 ima fUNGUS
and conidia formed singly or in chains, but in all cases with
conspicuous (thickened and darkened) conidiogenous loci
(scars) and mostly non-scolecosporous, pigmented conidia.
This new concept was also supported by rst molecular
sequence analyses (Crous et al. 2000, 2001). Hernández-
Gutiérrez & Dianese (2009) divided Passalora in sections,
viz. sect. Passalora, Mycovellosiella, Phaeoramularia, and
Pseudophaeoisariopsis, which can be seen as morphological,
non-monophyletic groups, which are, however, connected by
numerous morphologically intermediate species. Sequences
of Passalora fulva (Cladosporium fulvum, Fulvia fulva),
type species of the genus Fulvia, with a special habit of
conidiophores, are close to those of some other Passalora
species in the Mycosphaerellaceae clade (Thomma et al.
2005), which supported the reduction of Fulvia to synonym
with Passalora as proposed in Crous & Braun (2003). In
this wide concept of Passalora, some species with hyaline
passalora-like conidia (non-scolecosporous, broad, only
with few septa) were included, as for instance Passalora
dubia and P. janseana. However, in molecular sequence
analyses, such species clustered within the Cercospora
clade (Groenewald et al. 2012), i.e. the colour of the conidia
(absence or presence of pigmentation) is more important
than the conidial shape, although most Passalora species
are characterised by having non-scolecosporous conidia and
most Cercospora species by scolecosporous ones.
The currently applied wide concept of Passalora introduced
by Braun (1995b) and Crous & Braun (2003) is not generally
accepted and was questioned by some authors. For instance,
Guo et al. (2003) treated Chinese species of the Passalora
s. lat. complex and recognised three genera, namely
Mycovellosiella, Passalora (including Cercosporidium) and
Phaeoramularia. Baker et al. (2000) discussed the intricated,
controversial history of Passalora and Cercosporium in detail,
admitted a morphological continuum between the two genera,
but considered the merging of these genera as undesirable.
They discussed morphological differences between these
genera, and considered them to be sufciently distinct to be
maintained until more profound examinations will be available.
With the increasing application of molecular methods in the
Cercospora complex, more profound approaches are now
given, but corresponding analyses of sequence data do not
support a clear separation of Cercosporidium, Passalora and
other genera involved.
The phylogenetic structure of Passalora is, however,
complicated and causes severe problems. Numerous available
phylogenetic examinations based on rDNA ITS data and other
markers indicate that Passalora s. lat. is not monophyletic
(Crous et al. 2000, 2001, 2009b, c, 2013; Thomma et al.
2005). Taxa within the Mycosphaerellaceae are at least
paraphyletic, if not polyphyletic. Comprehensive phylogenetic
analyses based on a much broader sampling, including the
type species of Passalora and its synonymised genera, are
necessary for a better understanding of the generic structure
of Passalora s. lat. However, a severe problem is that clades
or subclades spread within the Mycosphaerellaceae are not
clearly connected with morphological groups within Passalora,
i.e. mycovellosiella-like, phaeoramularia-like groups of species
and other morphological types are not reected in phylogenetic
assemblages. Hence, a comprehensive phylogenetic
reassessment and further splitting of Passalora is currently not
possible based on available data, and it has to be accepted as
a para- or polyphyletic genus, at least for the present.
Passalora Fr., Summa Veg. Scand. 2: 500 (1849).
Type species: Passalora bacilligera (Mont. & Fr.) Mont. & Fr.
1849.
Synonyms: Cercosporidium E. Earle, Muhlenbergia 1: 16
(1901) [type species: C. helleri E. Earle 1901].
Vellosiella Rangel, Bol. Agric. (São Paulo), Ser. 16 A, 2: 151
(1915), nom. illeg. (Art. 53.1).
Mycovellosiella Rangel, Arch. Jard. Bot. Rio de Janeiro 2: 71
(1917) [type species: M. cajani (Henn.) Rangel ex Trotter
1931].
Passalora sect. Mycovellosiella (Rangel) A. Hern.-Gut. &
Dianese, Mycotaxon 108: 3 (2009).
Ormathodium Syd., Ann. Mycol. 26: 138 (1928) [type species:
O. styracis Syd. 1928], de Muntañola (1960).
Ragnhildiana Solheim, Mycologia 23: 365 (1931) [type
species: R. agerati (F. Stevens) F. Stevens & Solheim
1931].
Cercodeuterospora Curzi, Boll. Staz. Patol. Veg. Roma, Ser.
2, 12: 149 (1932) [type species: C. trichophila Curzi 1932].
Fulvia Cif., Atti Ist. Bot. Univ. Lab. Critt. Pavia, Ser. 5, 10: 246
(1954) [type species: F. fulva (Cooke) Cif.1954].
Mycovellosiella subgen. Fulvia (Cif.) U. Braun, Monogr.
Cercosporella, Ramularia 1: 39 (1995).
Berteromyces Cif., Sydowia 8: 267 (1954) [type species: B.
aeneus Cif. 1954].
Oreophyllum Cif., Sydowia 8: 253 (1954) [type species: O.
angelaemariae Cif. 1954].
Phaeoramularia Munt.-Cvetk., Lilloa 30: 182 (1960) [type
species: P. gomphrenicola (Speg.) Munt.-Cvetk. 1960].
Passalora sect. Phaeoramularia (Munt.-Cvetk.) A. Hern.-Gut.
& Dianese, Mycotaxon 108: 3 (2009).
Tandonella S.S. Prasad & R.A.B. Verma, Indian Phytopathol.
23: 111 (1970) [type species: T. ziziphi S.S. Prasad &
R.A.B. Verma 1970].
Walkeromyces Thaung, Trans. Brit. Mycol. Soc. 66: 213
(1976) [type species: W. grewiae Thaung 1976].
Passalora sect. Pseudophaeoisariopsis U. Braun, Dianese &
A. Hern.-Gut., Mycotaxon 108: 3 (2009).
Literature: Deighton (1967), Ellis (1971, 1976, as
Cercosporidium), Hsieh & Goh (1990), Braun (1995a: 41),
Braun & Mel’nik (1997: 16–17), Shin & Kim (2001: 135),
Crous & Braun (2003: 21), Guo et al. (2003: 65), Kamal
(2010: 101), Seifert et al. (2011: 331–332).
Description [emend. Crous & Braun (2003: 21)]:
Hyphomycetes (asexual morphs or asexual holomorphs)
or Passalora species with mycosphaerella-like sexual
morphs; Mycosphaerellaceae. Foliicolous, occasionally also
caulicolous or on fruits, usually phytopathogenic, causing
leaf-spots or other lesions, occasionally symptomless, rarely
hyperparasitic or saprobic. Mycelium in vivo internal and
external, supercial; hyphae branched, septate, colourless
or almost so to pigmented, thin-walled, smooth or almost
so. Stromata absent to well-developed, substomatal,
intraepidermal to deeply immersed, applanate to subglobose,
Cercosporoid fungi 1
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subhyaline to usually pigmented. Conidiophores in vivo
solitary, arising from supercial hyphae, lateral, occasionally
terminal, or loosely to densely fasciculate, arising from
internal hyphae or stromata, sometimes in sporodochia or
synnemata, macronematous, cylindrical, liform to strongly
geniculate-sinuous, unbranched or sometimes branched, pale
to distinctly pigmented, olivaceous to medium dark brown,
continuous to pluriseptate, wall thin to somewhat thickened,
smooth, occasionally somewhat rough; conidiogenous
cells integrated, terminal, occasionally intercalary or
pleurogenous, or conidiophores aseptate, i.e. conidiophores
reduced to conidiogenous cells, mono- to mostly polyblastic,
proliferation sympodial, rarely percurrent, with a single to
numerous conspicuous conidiogenous loci, non-protuberant
to distinctly so, somewhat thickened and darkened-refractive,
more or less cercosporoid, i.e. planate. Conidia solitary or
catenate, in simple or branched acropetal chains, amero-
to scolecosporous, aseptate to plurieuseptate, rarely with
additional distosepta, pale olivaceous to distinctly pigmented,
wall thin to slightly thickened, smooth to nely rough, hila
conspicuous, somewhat thickened and darkened-refractive,
conidial secession schizolytic.
PSEUDOCERCOSPORA EMEND. CROUS
ET AL. (2013), PALLIDOCERCOSPORA AND
PHAEOCERCOSPORA
The genus Pseudocercospora was introduced by Spegazzini
(1910) in order to accommodate Septonema vitis (Cercospora
vitis). This genus name was little used until Deighton (1976)
reintroduced it in the course of his splitting of Cercospora s.
lat. into smaller generic units. He used Pseudocercospora for
cercosporoid hyphomycetes with unthickened “conidial scars”
(i.e. conidiogenous loci), widened its circumscription and added
a large number of species. Unthickened conidiogenous loci are
characterised by having locus walls not thicker than the walls
of the surrounding conidiogenous cells. Such loci are often
inconspicuous, but may be more rigid and conspicuous in other
species by being more or less denticle-like. The unthickened
nature of the conidiogenous loci is, indeed, the fundamental
character of Pseudocercospora, although in almost all species
of this genus the loci are in addition not darkened-refractive,
i.e. not darker than the surrounding wall of the conidiogenous
cells (the darker pigmentation of conidiogenous loci, which
is diagnostic for species of other cercosporoid genera like
Cercospora, Passalora and Zasmidium, is generally caused
by a combination of darker pigmentation and sometimes
structural differences that may cause altered refraction of the
light). The basic importance of unthickened conidiogenous loci
as generic character for Pseudocercospora is supported by
several “abnormal” species with more conspicuous loci which
are unthickened but darker by being somewhat refractive or
even slightly darkened (Crous & Braun 2003). Nevertheless
such species belong in Pseudocercospora, which has been
conrmed in several studies based on molecular sequence
analyses (e.g., Crous et al. 2001, 2012).
Another problem concerns conidial formation. In almost
all Pseudocercospora species, the conidia are consistently
formed singly, but conidial catenation may occasionally
occur, e.g. in Pseucocercospora nyctanthis, P. millettiae and
P. noveboracensis (Hsieh & Goh 1990, Braun 1995a, Braun
& Mel’nik 1997, Crous & Braun 2003, Crous et al. 2013).
Some other pseudocercospora-like species, characterised
by subconspicuous conidiogenous loci (unthickened or
almost so, but somewhat refractive or darkened-refractive)
giving rise to short conidial chains or disarticulating conidia,
were considered to be intermediate between Passalora
and Pseudocercospora and assigned to the new genus
Pseudophaeoramularia (Braun & Mel’nik 1997). However, the
phylogenetic position of species with Pseudophaeoramularia
morphology within the Pseudocercospora (s. str.) clade
revealed that catenate conidia and somewhat darkened-
refractive loci and hila are acceptable variation within
Pseudocercospora.
Prathigada is another example for a misinterpreted
and confused genus. Due to at least partly conspicuous
conidiogenous loci, it has previously been interpreted as close
to Passalora, but distinct by having obclavate, relatively thick-
walled, often apically rostrate conidia. Beside type material,
a collection on Crateva formosensis, collected in Japan,
has been examined and compared. The Japanese material
agrees perfectly with the type material. The conidia do not
exceed 50 µm in length and three septa in both specimens.
Results of molecular sequence analyses based on the
Japanese material (C. Nakashima, unpubl. data) showed
that Prathigada crataevae, the type species of Prathigada,
clusters within the Pseudocercospora s. str. clade near to P.
jiensis, i.e. Prathigada has to be reduced to synonym with
Pseudocercospora. The molecular data are supported by
morphological characters of P. crataevae. The conidiogenous
loci and conidial hila are somewhat intermediate between
the Passalora and Pseudocercospora type by ranging from
inconspicuous to conspicuous by being denticle- or peg-like or
by being somewhat refractive to slightly darkened-refractive,
but on the other hand they are always unthickened and thus
not in conict with the current concept of Pseudocercospora
s. str. (Crous et al. 2013). Unthickened conidiogenous loci
which are more conspicuous by being somewhat darkened-
refractive are known in other proven Pseudocercospora
species, e.g. in Pseudophaeoramularia, which has been
reduced to synonymy with the latter genus (Crous et al.
2013). Due to the discussed results, Prathigada crataevae
has to be reallocated to Pseudocercospora. The following
redescription is based on the re-examination of type material
and an additional Japanese sample:
Pseudocercospora cratevicola C. Nakash. & U.
Braun, nom. nov.
MycoBank MB805517
(Fig. 1)
Basionym: Napicladium cratevae Syd. & P. Syd. (“crataevae”),
Ann. Mycol. 11: 329 (1913), non Pseudocercospora
cratevae Phengsintham et al. (2013).
Synonyms: Macraea cratevae (Syd. & P. Syd.) Subram.
(“crataevae”), Proc. Indian Acad. Sci, B, Biol. Sci., 36(4):
164 “1952” (1953).
Prathigada cratevae (Syd. & P. Syd.) Subram. (“crataevae”),
in Subramanian & Ramakrishnan, J. Madras Univ., B, 26:
367 (1956).
Braun et al.
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272 ima fUNGUS
Description: Leaf spots almost lacking to well-developed,
amphigenous, subcircular to angular-irregular, 1–10 mm
diam or conuent and larger, yellowish, ochraceous, pale to
medium brown, later dark brown by abundant fructication,
lesions uniform or with a paler centre, nally sometimes
greyish white, surrounded by a darker border, often with a
diffuse yellowish to ochraceous halo. Caespituli amphigenous,
scattered, punctiform, sometimes pustulate, later conuent
and dense, forming larger layers, dark brown. Mycelium
internal. Stromata lacking or almost so to well-developed,
substomatal to intraepidermal or deeply immersed,
subcircular in outline, 10–80 µm diam, or oblong, up to 150 ×
50 µm, olivaceous-brown, yellowish brown to medium brown,
composed of swollen hyphal cells, subcircular to angular-
irregular in outline, 3–8 µm diam, wall somewhat thickened.
Arrangement of conidiophores variable, ranging from small
fascicles arising from internal hyphae or small stromata,
rarely even solitary, to well-developed, large, dense fascicles,
forming sporodochial conidiomata, emerging through
stomata or erumpent, erect, straight to somewhat curved,
subcylindrical to mostly attenuated towards the tip, conical
or somewhat irregularly shaped, unbranched, 10–35 × 5–10
µm, 0–1-septate, pale olivaceous to olivaceous-brown or light
brown, wall thin, about 0.5–0.75 µm, smooth or almost so to
verruculose; conidiophores mostly reduced to conidiogenous
cells, occasionally integrated, terminal, 10–30 µm long,
proliferation sympodial, rarely percurrent, with a single or few
conidiogenous loci, inconspicuous to conspicuous by being
denticle-like (peg-like) or by being somewhat refractive or
slightly darkened-refractive, but always unthickened, 1.5–3
µm diam. Conidia solitary, 15–50 × 5–9 µm, 0–3-septate,
young small conidia ellipsoid-ovoid, obovoid or subcylindrical,
0–1-septate, thin-walled, subhyaline or pale, uniformly
pigmented, mostly verruculose, fully developed older conidia
obclavate, straight to often curved, above all at the apex,
2–3-septate, pale olivaceous to medium dark brown, above
all in the lower part, uniformly pigmented or often distinctly
paler towards the tip, wall thickened, up to 1.5 µm, smooth
or almost so to verruculose, apex obtuse to somewhat
pointed, base short obconically truncate, sometimes abruptly
attenuated, peg-like, 1.5–3 µm diam, hila neither thickened
nor darkened, at most somewhat refractive.
Material examined: India: Madras: Coimbatore, Government Farm,
on Crateva religiosa (Capparaceae), 5 Feb. 1912, W. McRae,
No. 9 (S, F42112, holotype). – Japan: Shizuoka, Ito, on Crateva
formosensis, 29 Sep. 1999, T. Kobayashi & C. Nakashima (CNS 797
and HAL 2597 F).
Notes: Subramanian (1953) introduced Micaraea for
Napicladium cratevae as type species and added the new
species M. punjabensis distinguished from the latter species
by having longer conidia, about 30–70 µm, with mostly 3–6
septa. Later he assigned the two species to Prathigada
Fig. 1. Pseudocercospora cratevicola (S, F42112).
A. Conidiophore fascicles. B. Conidiophores.
C. Conidia. Bar = 10 µm.
Cercosporoid fungi 1
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introduced as replacement name since Micaraea proved to
be a younger homonym. The source for the description and
illustration of Prathigada cratevae in Ellis (1971) is unclear,
but undoubtedly covers P. cratevae as well as P. punjabensis,
although the latter name is not cited as synonym. The re-
examination of type material of Napicladium cratevae (S,
F42112) showed that this collection perfectly coincides with
Subramanian’s (1953) description and illustration. Mature,
fully developed conidia do not exceed a length of 50 µm
and are consistently 0–3-septate, and the examined and
sequenced Japanese collection on Crateva formosensis
agrees completely. Therefore, we prefer to maintain Macraea
punjabensis (Prathigada punjabensis) as separate, easily
distinguishable species and transfer it to Pseudocercospora
as well:
Pseudocercospora subramanianii U. Braun & C.
Nakash., nom. nov.
MycoBank MB805518
Basionym: Macraea punjabensis Subram., Proc. Indian
Acad. Sci., B, Biol. Sci., 36: 166 “1952” (1953), non
Pseudocercospora punjabensis (Syd.) U. Braun &
Bagyan., 1999.
Synonym: Prathigada punjabensis (Subram.) Subram., in
Subramanian & Ramakrishnan, J. Madras Univ., B, 26:
367 (1956).
Notes: It is not surprising that the structure of the conidiogenous
loci of Prathigada was previously misinterpreted and
confused with scars of genera with thickened and darkened
loci. Due to at least partly conspicuous conidiogenous loci,
Prathigada has previously been interpreted as a genus close
to Passalora, but distinct by having obclavate, relatively thick-
walled, often apically rostrate conidia (Braun 1995a, Crous &
Braun 2003). As discussed above, the loci may be visible by
being more rigid and somewhat darkened-refractive, but they
are always unthickened, i.e. they fall into the morphological
range of Pseudocercospora. Due to the discussed confusion,
additional prathigada-like cercosporoid hyphomycetes have
been assigned to Prathigada, including several species with
true passalora-like conidiogenous loci, i.e. with conspicuously
thickened and darkened scars, which cannot be reallocated
to Pseudocercospora. They belong in Passalora in its current
circumscription. The latter genus is, as far as known, not
monophyletic, but a comprehensive phylogenetic revision is
not yet available. Hence, the species concerned can currently
only be assigned to Passalora s. lat.:
Passalora austroplenckiae (A. Hernández-Gutierrez
& Dianese) U. Braun, comb. nov.
MycoBank MB805519
Basionym: Prathigada austroplenckiae A. Hernández-
Gutierrez & Dianese, Mycotaxon 106: 57 (2009).
Passalora backmanii (Furlan. & Dianese) U. Braun,
comb. nov.
MycoBank MB805520
Basionym: Prathigada backmanii Furlan. & Dianese, Mycol.
Res. 103: 1203 (1999).
Passalora condensata (Ellis & Kellerm.) U. Braun,
comb. nov.
MycoBank MB805521
Basionym: Cercospora condensata Ellis & Kellerm., J. Mycol.
1(1): 2 (1885).
Synonym: Prathigada condensata (Ellis & Kellerm.) U. Braun,
Cryptog. Mycol. 20: 166 (1999).
Passalora gymnocladi (Ellis & Kellerm.) U. Braun,
comb. nov.
MycoBank MB805522
Basionym: Cercospora gymnocladi Ellis & Kellerm., Bull.
Torrey Bot. Club 11: 121 (1884).
Synonym: Prathigada gymnocladi (Ellis & Kellerm.) U. Braun,
Sydowia 48: 209 (1996).
Passalora thalictri (Thüm.) U. Braun, comb. nov.
MycoBank, MB805523
Basionym: Cercospora thalictri Thüm., Contr. Fl. Mycol.
Lusat. 2: 5 (1879).
Synonym: Cercospora thalictri var. thalictri-avi Thüm.,
Mycoth. Univ.: 1470 (1886).
Prathigada thalictri (Thüm.) U. Braun, in Braun & Mel’nik,
Trudy Bot. Inst. Im. V.L. Komarova 20: 97 (1997).
Notes: Other species assigned to Prathigada do not agree
with Passalora, belong elsewhere or they are doubtful and
unclear. The status of Prathigada bauhiniae (Rao et al.
1975) is unclear. Stenellopsis shoreae was reallocated to
Prathigada by Kamal (2010), but due to unthickened hila it
is possible that this species belongs to Pseudocercospora.
Prathigada tamarindi (Muthappa 1968) is a doubtful species
characterised by its unusual conidia which are about 20–30
× 8.5–11 µm and 1–2-septate. A re-examination of type
material or examinations of new collections are necessary to
reassess this species. Prathigada terminaliae (Sutton 1994)
is better considered a species of Pseudocercospora (P.
terminaliae). Prathigada terminaliae-bellericae (Kamal 2010)
is tentatively reduced to synonymy with P. terminaliae. The
signicance of differences in the length of conidiophores and
conidia between P. terminaliae and P. terminaliae-bellericae
is unclear and doubtful as the values for these structures of
the latter species are intermediate between P. terminaliae
and P. kenemensis, which has been reduced to synonym with
the former species as well (Crous & Braun 2003). Prathigada
ziziphi (Rao & Ramakrishnan 1965) is tentatively reduced
to synonymy with Pseudocercospora jujubae. Type material
was not available, but the shape and size of the conidia agree
well.
There are several other morphological characteristics
which were proposed for a further splitting of
Pseudocercospora into smaller generic units, but all of
them failed due to a morphological continuum between the
particular morphological types and numerous intermediate
taxa. Furthermore, there is no molecular (phylogenetic)
evidence for and support of genera to be segregated based
on morphology (Crous et al. 2000, 2001, 2013). Braun (1998)
tried to use morphological characters for a separation of
Pseudocercospora into morphological, non-phylogenetic
sections, which are, however, little useful. Pallidocercospora
Braun et al.
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274 ima fUNGUS
(Crous et al. 2013), recently introduced for the “Mycosphaerella
heimii” complex (Crous et al. 2004a), is a segregation from
Pseudocercospora mainly based on its distinct phylogenetic
position and culture characteristics (formation of red crystals
on MEA, PDA, SNA and WA), whereas the morphology of
the asexual and sexual morphs is pseudocercospora- and
mycosphaerella-like, respectively.
Cercoseptoria was recognised and reintroduced in
Deighton (1976) for pseudocercosporoid species with acicular
conidia. Most of the species assigned to Cercoseptoria are
distinctly sporodochial. Deighton (1987) and Braun (1988)
questioned Cercoseptoria as properly distinguished genus
and reduced it to synonymy with Pseudocercospora. Absence
or presence of stromata and conidial shape are too variable
in Pseudocercospora to be used for the discrimination of
smaller units on generic rank. Braun’s (1993) introduction of
Cercostigmina for pseudocercospora-like former Stigmina
species (conidiophores and conidia thin-walled, conidia
scolecosporous) with consistently percurrently proliferating
conidiogenous cells, partly associated with mycosphaerella-
like sexual morphs, is another example. Sympodial and
percurrent proliferations of conidiogenous cells are often
mixed in particular Pseudocercospora species, and species
with consistently percurrent proliferation cluster within the big
Pseudocercospora clade as well (Crous et al. 2001, 2013;
Taylor et al. 2003).
The conidiogenous loci of Phaeoisariopsis griseola, type
species of Phaeoisariopsis, were previously considered
to be passalora-like, i.e. slightly thickened and darkened.
However, the loci in P. griseola range from being quite
inconspicuous (neither thickened nor darkened), as exhibited
in the type material of Pseudocercospora columnaris
(Cercospora columnaris), one of the synonyms of P. griseola,
to subconspicuous (i.e. non-protuberant, lying at around
the conidiogenous cells, unthickened, but slightly darkened-
refractive). Deighton (1990) maintained Phaeoisariopsis as
separate genus, but conned it to a few species with scars
similar to those of its type species, P. griseola. He reallocated
synnematous species with inconspicuous conidiogenous loci
to Pseudocercospora and those with distinctly sympodial
conidiogenous cells and conspicuously thickened and
darkened loci to Passalora, whereas Braun (1992) considered
Phaeoisariopsis a synnematous counterpart of Passalora.
However, the locus type of Phaeoisariopsis griseola, type
species of Phaeoisariopsis, falls within the range of loci in
Pseudocercospora, which has been proven by means of
results of molecular sequence analyses (Crous et al. 2006).
These comprehensive examinations of angular leaf spot of
bean (P. griseola) showed that this species clusters within the
Pseudocercospora clade and revealed that the formation of
synnematous conidiomata does not play any taxonomic role
on generic level within the Pseudocercospora complex (Crous
et al. 2006, 2013). Hence, this species was reallocated to the
latter genus and Phaeoisariopsis was reduced to synonym
with Pseudocercospora, which was proposed to be conserved
as the former genus name is older (Braun & Crous 2006),
i.e. former Phaeoisariopsis species with subconspicuous,
P. griseola-like scars as well as inconspicuous loci are now
part of Pseudocercospora and those with conspicuously
thickened loci belong to Passalora. Furthermore, Crous
et al. (2006) demonstrated the unexpected phylogenetic
position of Stigmina platani, type species of Stigmina,
within the Pseudocercospora clade. Although older than
Pseudocercospora, the latter genus name was conserved
over that of Stigmina (Braun & Crous 2006).
Paracercospora (Deighton 1979), with P. egenula as
type species, was introduced for former pseudocercosporoid
hyphomycetes characterised by having a special type of
conidiogenous loci, viz. the scars are only visible by a ring-
like structure formed by a slightly thickened and darkened
rim. P. jiensis was originally assigned to Pseudocercospora,
a position later conrmed in molecular sequence analyses
(Crous et al. 2000, 2001, Stewart et al. 1999, Arzanlou
et al. 2008). Therefore, Crous & Braun (2003) reduced
Paracercospora to synonymy with Pseudocercospora, but
the situation is more complicated as P. jiensis is, indeed,
a species of the latter genus, but P. egenula, the type
species of Paracercospora, does not cluster within the
Pseudocercospora clade and is tentatively maintained as
separate genus (Crous et al. 2013). The problem is that the
scar structure (Paracercospora type) is not informative and
distinctive in this case, i.e. Paracercospora is conned to its
type species and only phylogenetically distinguished from
Pseudocercospora. Phaeocercospora (Crous et al. 2013)
is another pseudocercospora-like genus just based on its
separate phylogenetic position.
Based on detailed multilocal phylogenetic examinations,
Crous et al. (2013) revealed a large monophyletic
Pseudocercospora (s. str.) clade, including the type species
of this genus, representing Pseudocercospora emend.,
which is now a holomorph genus in its own right. Beside
species of the “Mycosphaerella heimii” complex, forming
a separate clade that is now treated as Pallidocercospora,
and Phaeocercospora, there are some additional species
morphologically indistinguishable from Pseudocercospora
which cluster, however, apart from Pseudocercospora (s.
str.) in other clades. Such phylogenetically unresolved
species, as for instance P. colombiensis on Eucalyptus
(Crous 1998), P. thailandica on Acacia (Crous et al. 2004b),
and P. tibouchinigena on Tibuchina (Crous et al. 2013), are
tentatively retained in Pseudocercospora. Pseudocercospora
s. str. is characterised as follows:
Pseudocercospora Speg., Anales Mus. Nac. Buenos
Aires 20: 438, 1910 (nom. cons.).
Type species: Pseudocercospora vitis (Lév.) Speg. 1910.
Synonyms: Stigmina Sacc., Michelia 2: 22 (1880), nom. rej.
[type species: S. platani (Fuckel) Sacc. 1880].
Phaeoisariopsis Ferraris, Ann. Mycol. 7: 280 (1909), nom rej.
[type species: Ph. griseola (Sacc.) Ferraris 1909].
Cercosporiopsis Miura, Flora of Manchuria and East Mongolia,
3, Cryptogams: 527 (1928), nom. illeg. (Art. 53.1).
Septoriopsis F. Stevens & Dalbey, Mycologia 11: 4 (1918),
nom. illeg. (Art. 53.1).
Cercoseptoria Petr., Ann. Mycol. 23: 69 (1925) [type species:
C. chamaesyceae (F. Stevens & Dalbey) Petr. 1925].
Ancyclospora Sawada, Rep. Govt. Agric. Res. Inst. Taiwan
87: 78 (1944), nom. inval. (Art. 39.1).
Helicomina L.S. Olive, Mycologia 40: 17 (1948) [type species:
H. caperoniae L.S. Olive 1948].
Cercosporoid fungi 1
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Macraea Subram., Proc. Proc. Indian Acad. Sci, Section B,
Biol. Sci., 36: 164 "1952" (1953), nom. illeg. (Art. 53.1).
Prathigada Subram., in Subramanian & Ramakrishnan, J.
Madras Univ., B, 26: 366 (1956).
Cercocladospora G.P. Agarwal & S.M. Singh, Proc. Natl.
Acad. Sci. India, B, 42: 439 “1972” (1974) [type species:
C. andinae G.P. Agarwal & S.M. Singh 1974].
Cercostigmina U. Braun, Cryptog. Bot. 4: 107 (1993) [type
species: C. concentrica (Cooke & Ellis) U. Braun 1993].
Pseudophaeoramularia U. Braun, Trudy Bot. Inst. Im. V.L.
Komarova 20: 18 (1997) [type species: P. geranii (W.B.
Cooke & C.G. Shaw) U. Braun 1997].
Paracercospora p.p. [see Crous et al. (2013)].
Literature: Deighton (1976), Yen & Lim (1980: 168–190),
Pons & Sutton (1988), Hsieh & Goh (1990), Braun (1995a:
42), Guo & Hsieh (1995), Braun & Mel’nik (1997: 18), Guo et
al. (1998), Shin & Kim (2001: 158), Crous & Braun (2003: 25),
Kamal (2010: 143), Seifert et al. (2011: 364–367).
Description [emend. Crous et al. (2013)]: Hyphomycetes
(asexual morphs or asexual holomorphs) or
Pseudocercospora with mycosphaerella-like sexual
morphs; Mycosphaerellaceae. Foliicolous, occasionally also
caulicolous or on fruits, usually phytopathogenic, causing
leaf spots or other lesions, occasionally symptomless or
endophytic. Mycelium in vivo internal and/or external,
supercial, sometimes climbing leaf hairs; hyphae
branched, septate, hyaline to pigmented, mostly thin-
walled, smooth or only faintly rough-walled, mostly about
1–6 µm wide, external hyphae emerging through stomata,
rarely erumpent. Stromata lacking to well-developed, small
to large, substomatal, intraepidermal to deeply immersed,
mostly between 10 and 120 µm diam, pale olivaceous to
dark brown. Colonies on leaves, amphigenous, sometimes
also on fruits and stems, almost colourless to dark brown,
scattered to dense. Conidiophores semimacronematous
to mosty macronematous, in small to large, loose to dense
fascicles, arising from internal hyphae or stromata, emerging
through stomata or erumpent, and/or conidiophores solitary,
arising from supercial hyphae, lateral or occasionally
terminal, sometimes forming sporodochial conidiomata
(numerous conidiogenous cells or short conidiophores
arising from well-developed stromata) or distinct synnemata,
conical, cylindrical, liform to strongly geniculate-sinuous,
unbranched to branched, aseptate (conidiophores reduced
to conidiogenous cells) to plurieuseptate, subhyaline to dark
brown, wall thin to moderately thick, smooth to somewhat
rough-walled, occasionally with annellations caused by
rejuvenation; conidiogenous cells integrated, terminal,
occasionally intercalary, or conidiophores often reduced to
conidiogenous cells, mono- to polyblastic, sympodially or
percurrently proliferating, sometimes both types of proliferation
mixed, conidiogenous loci inconspicuous or conspicuous
by being more or less denticle-like, but locus wall always
unthickened or almost so, usually neither darkened nor
refractive, rarely more rigid and more conspicuous by being
somewhat darkened-refractive or slightly thickened along the
rim, but not thickened throughout. Conidia usually formed
singly, rarely disarticulating or in short acropetal chains,
usually scolecosporous, i.e. obclavate, cylindrical, liform,
acicular and pluriseptate, rarely amero- to phragmosporous,
i.e. ellipsoid-ovoid, short cylindrical, fusiform, aseptate or
only with few septa, euseptate, rarely eusepta mixed with
few distosepta, but not consistently distoseptate, hyaline or
subhyaline, olivaceous, olivaceous-brown to medium dark
brown, wall thin to somewhat thickened, smooth to faintly
verruculose, ends rounded, truncate to obconically truncate,
hila unthickened or almost so, planate, rarely slightly bulging,
without marginal frill or rarely with minute frill, conidial
secession schizolytic.
Pallidocercospora Crous, Stud. Mycol. 75: 73 (2012)
[2013].
Type species: Pallidocercospora heimii (Crous) Crous 2013.
Pseudocercospora p.p. [see Crous et al. (2013)].
Description: Hyphomycetes (asexual morphs or asexual
holomorphs) or Pallidocercospora with mycosphaerella-like
sexual morphs; Mycosphaerellaceae. Phylogenetically distinct
from Pseudocercospora, forming a separate clade. In vivo
morphologically indistinguishable from Pseudocercospora,
but in vitro forming red crystals on MEA, PDA, SNA and WA.
Phaeocercospora Crous, Persoonia 28: 171 (2012)
[Fungal Planet 122, 2012]
Type species: Phaeocercospora colophospermi Crous 2012.
Description: Hyphomycetes (asexual morphs or asexual
holomorphs); Mycosphaerellaceae. Phylogenetically distinct
from Pseudocercospora, belonging to the “Dothistroma
clade”. Morphologically close to and barely distinguishable
from former Cercostigmina species (Cercostigmina-like
Pseudocercospora species), i.e. with unilocal, determinate
to percurrent conidiogenous cells. Hitherto monotypic
(the type species occurs in South Africa on the legume
Colophospermum mopane).
STENELLA AND ZASMIDIUM
Stenella was introduced by Sydow (1930) as monotypic genus
for S. araguata, a species with conspicuously thickened,
darkened conidiogenous loci and hila. However, this genus
was little applied until Ellis (1976) and Deighton (1979)
widened its concept and added numerous morphologically
similar species, previously mostly assigned to Cercospora,
characterised by forming distinctly verruculose supercial
mycelium in vivo and verruculose conidia, formed singly or in
chains. The number of new and reallocated species increased
rapidly, encompassing a wide range of morphological types,
i.e. species with exclusively solitary conidiophores arising from
supercial hyphae to taxa with solitary as well as fasciculate
conidiophores, with or without stromata, with catenate or
solitary amero- to scolecosporous conidia or both types mixed
in particular species. Similar species with stenella-like conidia,
but without any verruculose supercial hyphae in vivo have
previously been placed in Stenellopsis, but Braun & Crous
(2005) reduced the latter genus to synonym with Stenella,
a conclusion supported by results of molecular sequence
Braun et al.
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276 ima fUNGUS
analyses (Shivas et al. 2009). The presence of supercial
mycelium in Stenella is not essential on generic rank, which
is comparable to the situation in other cercosporoid genera
like Passalora and Pseudocercospora. Verrucisporota,
introduced by Shaw & Alcorn (1993) as replacement for
the illegitimate Verrucospora, with V. proteacearum as type
species, is morphologically very close to Zasmidium and
barely distinguishable. Previous attempts to distinguish this
genus from Stenellopsis on the base of differences in the
stroma structure, broad loci and rugose conidia (Ellis 1971,
David 1997) are neither practicable nor tenable. New species
of this genus described by Beilharz & Pascoe (2002) are
rather zasmidium-like. One of these species, V. davisiae, has
a mycosphaerella-like sexual morph. Phylogenetic analyses
of Verrucisporota species carried out by Crous et al. (2009c)
showed that they cluster within the Mycosphaerellaceae clade
together with Zasmidium species. Therefore, Verrucisporota
is tentatively considered a synonym of the non-monophyletic
genus Zasmidium until the exact phylogenetic position of its
type species will be known.
Arzanlou et al. (2007) carried out comprehensive molecular
examinations of Ramichloridium and morphologically similar
genera, including Stenella and Zasmidium. Stenella araguata,
the type species of Stenella, surprisingly clustered in the
Teratosphaeriaceae, whereas all other examined Stenella
species belong to the Mycosphaerellaceae phylogenetically.
These results led to the conclusion that Stenella has to be
conned to its type species, whereas the other species in the
Mycosphaerellaceae need to be assigned to another genus.
Zasmidium proved to be an available name, as the type
species of this genus clusters in the Mycosphaerellaceae
and coincides morphologically with Stenella (Arzanlou
et al. 2007). Therefore, Braun et al. (2010a) introduced
Zasmidium as genus for stenella-like fungi belonging to
the Mycosphaerellaceae and reallocate numerous species,
although Zasmidium s. lat. does not form a single monophyletic
clade within the Mycosphaerellaceae (Crous et al. 2009a, b).
The conidiogenous loci and hila of the conidia in Zasmidium
are cercosporoid, i.e. planate and somewhat thickened and
darkened. Stenella s. str. is currently monotypic, i.e. conned
to its type species, and the conidiogenous loci are according
to David (1993) pileate. Otherwise, Stenella and Zasmidium
species are morphologically barely distinguished. Braun et al.
(2010b) and Kamal (2010) reallocated numerous additional
Stenella species described from Brazil, New Zealand,
Venezuela and known from India to Zasmidium.
Zasmidium Fr., Summa Veg. Scand. 2: 407 (1848).
Type species: Zasmidium cellare (Pers. : Fr.) Fr. 1848.
Synonyms: Biharia Thirum. & Mishra, Sydowia 7: 79 (1953)
[type species: B. vangueriae Thirum. & Mishra 1953].
Stenellopsis B. Huguenin, Bull. Trimestriel Soc. Mycol.
France 81: 695 (1966) [type species: St. fagraeae B.
Huguenin 1966].
Verrucispora D.E. Shaw & Alcorn, Proc. Linn. Soc. New
South Wales 92: 171 (1967), nom. illeg. (Art. 53.1).
Verrucisporota D.E. Shaw & Alcorn, Austral. Syst. Bot. 6:
273 (1993) [type species: V. proteacearum D.E. Shaw &
Alcorn 1993].
Stenella p.p. [see Braun et al. (2010a, b)].
Literature: Deighton (1979: 52–54, as Stenella), Ellis (1976:
307–314, as Stenella), Braun & Mel’nik (1997: 21), Braun et
al. (2010a, b), Shivas et al. (2009), Seifert et al. (2011: 478).
Description [emend. Braun et al. (2010a)]: Hyphomycetes
(asexual morphs or asexual holomorphs) or Zasmidium with
mycosphaerella-like sexual morphs; Mycosphaerellaceae.
Saprobic or mostly biotrophic, usually foliicolous, symptomless
or causing various lesions, ranging from yellowish
discolorations to distinct leaf spots. In plant pathogenic
species, mycelium mostly immersed as well as supercial,
rarely only immersed; hyphae branched, septate, colourless
or almost so to pigmented, pale olivaceous to brown, wall thin
to somewhat thickened, immersed hyphae smooth or almost
so to faintly rough, external hyphae distinctly verruculose to
verrucose (in culture immersed hyphae usually smooth or
almost so, aerial hyphae verruculose). Stromata lacking to
well-developed, pigmented. Conidiophores solitary, arising
from supercial hyphae, lateral, occasionally terminal, in vivo
(in plant pathogenic taxa) sometimes also fasciculate, arising
from internal hyphae or stromata, semimacronematous to
macronematous, in culture occasionally micronematous,
cylindrical, liform, subuliform, straight to strongly geniculate-
sinuous, mostly unbranched, aseptate, i.e. reduced to
conidiogenous cells, to pluriseptate, subhyaline to pigmented,
pale olivaceous to medium dark brown, wall thin to
somewhat thickened, smooth to verruculose; conidiogenous
cells integrated, terminal, occasionally intercalary, rarely
pleurogenous, or conidiophores reduced to conidiogenous
cells, mostly polyblastic, sympodial, conidiogenous loci
conspicuous, somewhat thickened and darkened-refractive,
planate. Conidia solitary or catenate, in simple or branched
acropetal chains, shape and size variable, ranging from amero-
to scolecosporous, aseptate to transversely plurieuseptate,
subhyaline to pigmented, pale olivaceous to brown, wall thin to
somewhat thickened, smooth or almost so to usually distinctly
verruculose (in plant pathogenic species without supercial
mycelium always verruculose), hila somewhat thickened and
darkened-refractive, planate, conidial secession schizolytic.
Stenella Syd., Ann. Mycol. 28: 205 (1930).
Type species: Stenella araguata Syd. 1930.
Description: Morphologically agreeing with plant pathogenic
Zasmidium species, except for pileate conidiogenous loci.
Phylogenetically belonging in the Teratosphaeriaceae (in its
current circumscription monotypic). Stenella araguata was
redescribed in Ellis (1971).
TENTATIVELY RECOGNISED CERCOSPOROID
GENERA
Asperisporium Maubl., Lavoura 16: 212, “1912”
(1913) and Bull. Trimestriel Soc. Mycol. France 29: 357
(1913).
Type species: Asperisporium caricae (Speg.) Maubl. 1913.
Literature: Ellis (1971: 273–274; 1976: 240–243), Sutton
(1975: 182–185), von Arx (1983), Braun (1995a: 40), Braun
Cercosporoid fungi 1
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& Mel’nik (1997: 14), Crous & Braun (2003: 13), Minnis et al.
(2011), Seifert et al. (2011: 95).
Description: Usually foliicolous, leaf-spotting hyphomycetes,
Mycosphaerellaceae. Mycelium in vivo internal; hyphae
branched, septate, colourless to pigmented, thin-walled,
smooth or almost so. Stromata usually well-developed,
substomatal to intraepidermal, often somewhat erumpent,
pigmented. Conidiophores macronematous, usually densely
fasciculate, forming sporodochial conidiomata, continuous
to septate, pigmented, wall thin to slightly thickened,
smooth or almost so; conidiogenous cells integrated,
terminal or conidiophores reduced to conidiogenous cells,
usually polyblastic, sympodial, but mostly not strongly
geniculate, conidiogenous loci conspicuous, thickened and
darkened. Conidia solitary, amero- to phragmosporous (non-
scolecosporous), mostly ellipsoid-ovoid, obovoid, fusiform
to short cylindrical or obclavate, mostly with 0–3 eusepta,
sometimes with a single or several oblique or longitudinal
septa, pigmented, distinctly verruculose to coarsely
verrucose, basal hilum thickened and darkened, conidial
secession schizolytic.
Notes: Asperisporium is morphologically very close to
Passalora, as already emphasized by Crous & Braun (2003),
except for verrucose conidia. The phylogenetic position of
A. caricae, its type species, has recently been elucidated in
Minnis et al. (2011), based on DNA sequences from the ITS
region and nLSU, showing that this species clusters within
the Mycosphaerellaceae clade close to several Passalora
species, including P. brachycarpa. According to this result,
Asperisporium is undoubtedly a potential synonym of
Passalora. However, as Passalora is still para- or even
polyphyletic and the phylogenetic position of its type species
is still unknown, a formal reassessment of Asperisporium was
postponed by Minnis et al. (2011) until the whole Passalora
complex will be phylogenetically examined and reassessed
in detail. However, the postponed reassessment of the genus
name Asperisporium does only concern its type species. Other
species assigned to Asperisporum have to be individually
reassessed, if possible based on molecular sequence data.
Asperisporium sequoiae and A. juniperinum, two species on
Gymnosperms with relatively delicately rough-walled conidia,
are typical examples. Sequences of the two species are rather
similar, cluster with other species of Passalora s. lat., but they
are only distantly related to the type species of Asperisporium
(Solheim 2013). Therefore, these species are better assigned
to the polyphyletic Passalora in its current circumscription.
Comprehensive phylogenetic examinations of the whole
complex of asperisporium- and passalora-like hyphomycetes
are necessary to reach a better more natural genus concept
in this group of cercosporoid hyphomycetes. The surface
sculpture of the conidia as character for the discrimination of
Asperisporium from Passalora is undoubtedly unreliable, at
least species with delicately verruculose conidia may belong
to Passalora, which is supported by phylogenetic data.
Cladosporiella Deighton, Mycol. Pap. 101: 34 (1965).
Type species: Cladosporiella cercosporicola Deighton 1965.
Literature: Deighton (1969: 33–39), Ellis (1971: 303), Braun
(1995a: 39), Seifert et al. (2011: 145).
Notes: Morphologically close to Cladosporium and
mycovellosiella-like Passalora species (with supercial
hyphae, conidiophores fasciculate or solitary, arising
from supercial hyphae, conidiogenous loci conspicuous,
thickened and darkened, conidia catenate, pigmented), but
the loci and hila are not cladosporium-like (not coronate) and
all species assigned to this genus are hyperparasitic. The
hyperparasitic habit as single character is barely acceptable
and usable to discriminate this genus from Passalora, but
since the phylogenetic position of Cladosporiella and its
relation to the Mycosphaerellaceae are still unclear and
unproven, we tentatively prefer to maintain it as separate
genus.
Denticularia Deighton, Trans. Brit. Mycol. Soc. 59: 421
(1972).
Type species: Denticularia modesta (Syd.) Deighton 1972.
Literature: Ellis (1976: 182–183), Braun (1995a: 42), Crous &
Braun (2003: 23), Seifert et al. (2011: 176).
Notes: Morphologically close to Pseudocercospora (leaf
spotting hyphomycetes with unthickened, not darkened
conidiogenous loci and hila), but the conidiogenous loci are
distinctly denticle-like, and the catenate conidia are non-
scolecosporous, only with 0–1(–3) septa.
Cultures of the type species of this genus and results
of molecular sequence analyses are necessary to
resolve its phylogenetic position and clarify its relation to
Pseudocercospora. It is still unclear and unproven if this
genus belongs in the Mycosphaerellaceae. Therefore,
Denticularia is tentatively retained as genus on its own.
Elletevera Deighton, Mycol. Pap. 118: 17 (1969).
Type species: Elletevera parasitica (Ellis & Everh.) Deighton
1969.
Literature: Braun (1995a: 37), Crous & Braun (2003: 17),
Seifert et al. (2011: 193).
Description: Fungicolous, hyperparasitic dematiaceous
hyphomycetes. Mycelium immersed; hyphae hyaline to faintly
pigmented; stromata lacking; conidiophores macronematous,
erect, laxly fasciculate, frequently branched, septate,
faintly pigmented, thin-walled, smooth; conidiogenous
cells integrated, terminal or formed as lateral branchlets,
conidiogenous loci neither thickened nor darkened, but
distinct by being denticle-like and somewhat refractive;
conidia solitary, rarely catenate, phragmo- to scolecosporous,
pigmented, with slightly refractive basal hilum.
Notes: This genus was introduced for hyperparasitic
cercosporoid hyphomycetes with “distinct” conidiogenous
loci, morphologically resembling species of the former genus
Mycovellosiella. However, the re-examination of several
collections of the type species of Elletevera showed that
Deighton’s (1969) original description of the conidiogenous
Braun et al.
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278 ima fUNGUS
loci (scars) is misleading (Crous & Braun 2003). Due to
denticle-like loci with unthickened, not darkened walls,
Elletevera is possibly related to Pseudocercospora or perhaps
not cercosporoid at all. But this question cannot be answered
without knowledge of the phylogenetic afnity of this genus,
which is still unproven and unclear. The hyperparasitic habit
alone is insufcent to uphold Elletevera as separate genus,
but until its phyologenetic position is proven the latter genus
is tentatively maintained.
Eriocercospora Deighton, Mycol. Pap. 118: 5 (1969).
Type species: Eriocercospora balladynae (Hansf.) Deighton
1969.
Literature: Ellis (1971: 248–249), Rao et al. (1982: 1155),
Braun (1995a: 39), Seifert et al. (2011: 199).
Description: Fungicolous, hyperparasitic dematiaceous
hyphomycetes. Mycelium supercial; hyphae branched,
septate, pigmented, thin-walled, smooth. Stromata lacking.
Conidiophores macronematous, mononematous, solitary, in
vivo arising from supercial hyphae, lateral, erect, usually
unbranched, continuous to septate, pigmented, thin-
walled, smooth; conidiogenous cells integrated, terminal
and intercalary, sympodially proliferating, but usually not
distinctly geniculate, conidiogenous loci subconspicuous, i.e.
unthickened, not or barely protuberant, not or only slightly
darkened-refractive. Conidia solitary, scolecosporous,
transversely euseptate, pigmented, thin-walled, smooth or
almost so, hila unthickened, not or barely darkened-refractive.
Notes: Crous & Braun (2003) re-examined type material and
numerous other collections of E. balladynae and found that
Deighton’s (1969) original description of the conidiogenous
loci (scars), which were compared with those of the former
genus Mycovellosiella, are misleading. The loci are neither
thickened nor conspicuously darkened (at most they
are only slightly darkened-refractive), i.e. they are rather
Pseudocercospora-like. In E. olivacea, another hyperparasitic
species, the loci and hila are quite unthickened and also
not darkened-refractive. Eriocercospora websteri and E.
moghaniae are two excluded plant pathogenic species that
were reallocated to Pseudocercospora (Braun 2002, Crous
& Braun 2003). Eriocercospora is possibly a synonym of
Pseudocercospora, but without proof based on molecular
DNA sequence analyses a nal decision has to be postponed.
Stenospora (type species: Stenospora uredinicola, Deighton
1969) is very close to Eriocercospora but mucedinaceous
(colourless).
Eriocercosporella R. Kumar, A.N. Rai & Kamal ex
U. Braun, Monogr. Cercosporella, Ramularia 2: 398
(1998).
Type species: Eriocercosporella indica R. Kumar, A.N. Rai &
Kamal ex U. Braun 1998.
Synonym: Eriocercosporella R. Kumar, A.N. Rai & Kamal,
Indian Phytopathol. 47: 127 (1994), nom. inval.
Literature: Seifert et al. (2011: 199).
Description: Foliicolous hyphomycetes, associated with leaf
spots. Mycelium internal and external, supercial hyphae
emerging through stomata, branched, pigmented, septate,
thin-walled, smooth. Stromata lacking. Conidiophores
macronematous, mononematous, in vivo solitary, arising from
supercial hyphae, lateral, simple, occasionally branched,
pigmented, septate, thick-walled, smooth; conidiogenous
cells integrated, terminal, uni- to multilocal, sympodially or
occasionally percurrently proliferating, loci truncate, at,
broad, neither thickened nor darkened, conidiogenesis
thalloblastic, i.e. at rst blastic, then thallic (base of
conidia ± agreeing in width with the diameter of the broad
conidiogenous loci). Conidia solitary, cylindrical to subclavate,
occasionally disarticulating, plurieuseptate, occasionally with
1–2 additional distosepta, thick-walled, brown, smooth, not
attenuated at the base, hila truncate, broad, width ± agreeing
with the diameter of the conidiogenous loci, neither thickened
nor darkened, conidial secession schizolytic.
Notes: Due to the conidiogenesis, the structure of the
conidiogenous cells and the thick-walled conidia, this
genus seems to be allied to the Sporidesmium complex,
but an afnity to Pseudocercospora can also not be
excluded with certainty. However, the phylogenetic afnity of
Eriocercosporella and its relation to the Mycosphaerellaceae
are currently quite unclear. Molecular examinations of the
phylogenetic position of the type species of this genus are
necessary to decide if Eriocercosporella belongs to the
cercosporoid or to the sporidesmioid complex. Therefore,
this genus is only tentatively treated in the present context of
cercosporoid genera.
Pantospora Cif., Ann. Mycol. 36: 242 (1938).
Type species: Pantospora guazumae Cif. 1938.
Dictyocephala A.G. Medeiros, Publ. Univ. Recife Inst. Micol.
373: 13 (1962) [type species: D. ulmifoliae (Obreg.-Bot.)
A.G. Medeiros 1962].
Literature: Deighton (1976: 156–159), Crous & Braun (2003:
23), Minnis et al. (2011), Seifert et al. (2011: 325).
Description: Foliicolous hyphomycetes, associated with
leaf spots, Mycosphaerellaceae. Mycelium internal;
hyphae colourless or almost so. Stromata developed,
pigmented. Conidiophores macronematous, in dense
coremioid fascicles or synnemata, septate, pigmented, thin-
walled, smooth; conidiogenous cells integrated, terminal,
proliferation sympodial and percurrent, conidiogenous loci
planate to slightly convex, neither thickened nor darkened
(pseudocercospora-like). Conidia formed singly, shape
variable, ellipsoid-ovoid, fusiform, clavate to obclavate,
didymo- to scolecosporous, with 1–11 transverse eusepta
and often a single or few oblique to longitudinal septa, hila
neither thickened nor darkened.
Notes: The genus Pantospora was described by Ciferri
(1938). Pantospora guazumae, the type species, is a leaf-
spotting hyphomycete on Guazuma ulmifolia. Dictyocephala,
based on Cercospora ulmifolia described from Colombia on
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G. ulmifolia, is undoubtedly a synonym of Pantospora as
already pointed out by Deighton (1976), who recognised the
latter genus although its type species is very similar to the
type species of Pseudocercospora by forming conidiophores
in dense coremioid fascicles or distinct synnemata and
sympodially to percurrently proliferating conidiogenous cells
with unthickened, not darkened conidiogenous loci. The
regular formation of oblique to longitudinal conidial septa was
considered to be the only difference, although a few non-
horizontal septa may also occur in some Pseudocercospora
species, including P. vitis, its type species. As there is no
rational base for a morphological discrimination between
the two genera, Crous & Braun (2003) reduced Pantospora
to synonymy with Pseudocercospora. However, sequence
analyses based on data from the rDNA ITS and nLSU
region, recently carried out by Minnis et al. (2011), showed
that Pantospora guazumae did neither cluster within the
Pseudocercospora s. str. clade [= clade 16, including the
type species P. vitis, according to Crous et al. (2009b)] nor
in clade 14 with other pseudocercospora-like species. The
closest relatives were rather passalora-like. These results
suggest that Pantospora cannot be considered a synonym
of Pseudocercospora. This genus is, at least tentatively,
maintained as separate genus until a better resolution of the
assemblage of species around P. guazumae will be available.
Paracercospora Deighton, Mycol. Pap. 144: 47 (1979).
Type species: Paracercospora egenula (Syd.) Deighton
1979.
Literature: Crous & Braun (2003: 22), Crous et al. (2012).
Description: Dematiaceous hyphomycete genus morphologi-
cally barely distinguishable from Pseudocercospora, but phy-
logenetically distinct; Mycosphaerellaceae. Mycelium in vivo
internal. Conidiophores macronematous, fasciculate, pig-
mented; conidiogenous cells integrated, terminal or conidi-
ophores reduced to conidiogenous cells, conidiogenous loci
subconspicous by being circular with very slightly thickened
and darkened-refractive rim. Conidia solitary, scolecospor-
ous, subhyaline to very pale olivaceous, hila very slightly
thickened and darkened-refractive along the rim.
Notes: The phylogenetic position and taxonomy
of Paracercospora is complicated and not yet fully
understood. Based on early phylogenetic analyses, Stewart
et al. (1999) considered Paracercospora a synonym of
Pseudocercospora and concluded that conidiogenous loci
(scars) only slightly thickened and darkened around the rim
are not sufciently different from quite unthickened loci to
be used as distinctive character on generic level, which was
supported by additional rDNA ITS examinations (Crous et
al. 2000, 2001). Therefore, Crous & Braun (2003) followed
this conclusion and treated Paracercospora as synonym
of Pseudocercospora. However, the reassessment of
Paracercospora was based on data derived from non-type
species, mainly P. jiensis and P. basiramifera (Crous 1998,
Arzanlou et al. 2008), which indeed cluster within the big
Pseudocercospora clade. Paracercospora egenula, the
type species of Paracercospora, has recently been included
in molecular sequence analyses, but clustered apart of
the Pseudocercospora clade in a not yet fully understood
clade together with Passalora brachycarpa, a species
with catenate conidia, Pseudocercospora tibuchinigena,
which is morphologically indistinguishable from genuine
Pseudocercospora spp., and several Mycosphaerella
spp. (Crous et al. 2012). Hence, Paracercospora is
currently conned to its type species, characterised
by circular conidiogenous loci with slightly thickened,
darkened-refractive rim and very pale conidia. Since such
paracercosporoid loci as well as subhyaline or pale conidia
are also known in several genuine Pseudocercospora
species, there is no rational base for a morphological
discrimination of the two genera, i.e. Paracercospora s. str.
is currently just a phylogenetically differentiated monotypic
genus.
Parastenella J.C. David, Mycol. Res. 95: 124 (1991).
Type species: Parastenella magnoliae (Weedon) J.C. David
1991.
Synonym: Stenellopsis Morgan-Jones, Mycotaxon 10: 405
(1980), nom illeg. (Art. 53.1).
Literature: Braun (1995a: 41), Seifert et al. (2011: 330).
Description: Dematiaceous hyphomycete genus resembling
Zasmidium (in vivo with supercial mycelium, hyphae,
conidiophores and solitary conidia pigmented, distinctly
verruculose to verrucose), but the conidiogenous cells are
terminal and intercalary, denticulate, with lateral short peg-
like protuberances, conidiogenous loci inconspicuous,
neither thickened nor darkened.
Notes: The phylogenetic position of this genus is unknown.
It is quite unclear if it is part of the Capnodiales and
Mycosphaerellaceae at all.
Pseudoasperisporium U. Braun, Schlechtendalia 5:
72 (2000).
Type species: Pseudoasperisporium tupae (Speg.) U. Braun
2000.
Literature: Seifert et al. (2011: 363).
Notes: Morphologically close to Asperisporium, but
the conidiogenous loci and hila at the base of conidia
are unthickened and not darkened. Species of
Pseudoaspersporium are distinguished from supercially
similar Fusicladium species (Venturiaceae) by having
coarsely verruculose conidia. The phylogenetic afnity of this
genus is, however, unclear and unproven.
Pseudocercosporidium Deighton, Mycol. Pap. 133:
55 (1973).
Type species: Pseudocercosporidium venezuelanum (Syd.)
Deighton 1973.
Literature: Ellis (1971: 297–299), Braun (1995a: 37), Crous &
Braun (2003: 14), Seifert et al. (2011: 367).
Braun et al.
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Description: Foliicolous, plant pathogenic, leaf spotting
hyphomycetes, teleomorph unknown. Mycelium internal.
Stromata lacking. Conidiophores in vivo solitary or in small
loose fascicles (groups) emerging through stomata, laxly erect,
macronematous, frequently branched, septate, pigmented
(very pale brown), thin-walled, smooth; conidiogenous cells
integrated, terminal, intercalary or pleurogenous (as lateral
branchlets), sympodial, polyblastic, conidiogenous loci
conspicuous, protruding, convex (papilla-like), but wall of the
loci neither thickened nor darkened, only somewhat refractive.
Conidia solitary, didymo- to scolecosporous, pigmented
(deeper in pigmentation than the conidiophores), thin-walled,
smooth or almost so, hila neither thickened nor darkened.
Notes: The phylogenetic afnity of P. venezuelanum and
its relation to the Mycosphaerellaceae are unknown.
Pseudocercosporidium resembles Passalora, but the
structure of the conidiogenous loci is quite distinct and closer
to scars of genera like Neoovularia and Pseudodidymaria
(Braun 1998).
Quasiphloeospora B. Sutton, Crous & Shamoun,
Mycol. Res. 100: 979 (1996).
Type species: Quasiphloeospora saximontanensis (Deighton)
B. Sutton, Crous & Shamoun 1996.
Literature: Deighton (1983: 7–8), Braun (1998: 400–401),
Crous & Braun (2003: 14), Seifert et al. (2011: 376–377).
Description: Cercosporoid hyphomycetes characterised by
forming large immersed sporodochium-like conidiomata
with liform, somewhat pigmented, irregularly verruculose
conidiophores, aseptate, i.e. reduced to conidiogenous cells,
monoblastic, determinate or sympodially to percurrently
proliferating, with slightly thickened and darkened
conidiogenous loci, and very pale to somewhat pigmented
scolecosporous conidia formed singly.
Notes: Quasiphloeospora is a cercosporoid genus with intricate
morphology and complex morphological relations to several other
genera, including Cercospora, Passalora and Pseudocercospora
(Crous & Braun 2003), but due to very pale, almost colourless
structures also to genera like Pseudocercosporella. Sutton et al.
(1996) classied the conidiomata as acervuli, although they may
better be referred to as sporodochia. The particular characters of
Q. saximontanensis, above all the structure of the conidiogenous
loci, are intermediate between the three similar genera cited
above. A clear afliation to one of these genera, just based on
morphology, is not possible. It is also possible that this species is
unrelated to any of the cercosporoid genera. Afnity and position
of Quasiphloeospora can only be proven by means of results
of molecular sequence analyses, which are, however, not yet
available. This genus might belong in the Mycosphaerellaceae,
but this is unproven. Thus, Quasiphloeospora is only tentatively
maintained as separate cercosporoid genus.
Scolecostigmina U. Braun, New Zealand J. Bot. 37:
323 (1999).
Type species: Scolecostigmina mangiferae (Koord.) U. Braun
& Mouch. 1999.
Stigmina p.p. [see Braun (1999), Crous et al. (2013)].
Literature: Crous & Braun (2003: 24), Crous et al. (2013: 74–
75), Seifert et al. (2011: 396).
Description: Scolecostigmina is morphologically close to
Pseudocercospora, above all to former Cercostigmina spe-
cies (leaf spotting dematiaceous hyphomycetes with sporo-
dochial conidiomata, macronematous densely fasciculate
conidiophores, percurrently proliferating conidiogenous cells,
neither thickened nor darkened applanate loci, and scole-
cosporous, plurieuseptate, pigmented conidia formed singly),
but the wall of the conidiophores is somewhat thickened and
mostly verruculose, possesses conspicuous, coarse annella-
tions and the conidia are transversely and occasionally also
obliquely or longitudinally septate.
Notes: S. mangiferae, the type species, which belongs
in the Mycosphaerellaceae, does not cluster within the
Pseudocercospora clade, i.e. it is phylogenetically distinct,
supporting Scolecostigmina as separate genus. The correct
placement of numerous additional species in Scolecostigmina
is, however, phylogenetically unproven and quite unclear.
Sequence data derived from cultures of Scolecostigmina
chibaensis on Pinus spp. in Japan (Nakashima et al. 2007)
have been analysed and showed that this species is not allied
to the type species of Scolecostigmina and does not belong
to the Mycosphaerellaceae at all. It takes an isolated position
with unclear afnity on family level, and does not belong to
any other families containing cercosporoid hyphomycetes
as for instance Cladosporiaceae, Dissoconiaceae or
Teratosphaericeae (Nakashima, unpubl. data). Due to
the phylogenetic position of Cercospora cryptomeriicola
within the Pseudocercospora clade, this species has been
reallocated to the latter genus, although its morphology
is rather scolecostigmina-like. The conidiomata are
sporodochial, conidiogenous cells proliferate percurrently
with conspicuous annellations, and the walls of the conidia
are thickened (Nakashima et al. 2007). These data indicate
that the typical “Scolecostigmina” morphology is unreliable,
i.e. all species assigned to Scolecostigmina have to be
re-examined and reassessed once molecular data are
available.
Semipseudocercospora J.M. Yen, Mycotaxon 17:
361 (1983).
Type species: Semipseudocercospora peristrophes-
acuminatae (J.M. Yen) J.M. Yen 1983.
Description: Morphologically close to Pseudocercospora
(leaf spotting hyphomycetes with unthickened, not darkened
conidiogenous loci and hila), but the conidiogenous cells are
not geniculate, i.e. not distinctly sympodially proliferating,
the conidiogenous loci are distinctly denticle-like, and the
solitary conidia are didymo- to phragmosporous, i.e. not
scolecosporous.
Notes: The phylogenetic position of the type species of
this genus and its relation to the Mycosphaerellaceae as
well as to the genus Pseudocercospora are still unknown
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and unproven. Therefore, Semipseudocercospora is only
tentatively maintained as separate cercosporoid genus.
Sirosporium Bubák & Serebrian., Hedwigia 52: 272
(1912).
Type species: Sirosporium antenniforme (Berk. & M.A. Curtis)
Bubák & Serebrian. 1912.
Literature: Ellis (1963: 2–11; 1971: 288–290; 1976: 299–303),
Braun (1995a: 39), Mel’nik (2000: 284–288), Crous & Braun
(2003: 18), Seifert et al. (2011: 404).
Description: Morphologically close to Passalora, i.e.
above all mycovellosiella-like (leaf spotting dematiaceous
hyphomycetes with internal and external mycelium,
supercial hyphae giving rise to solitary conidiophores,
lateral and terminal, conidiophores may also be formed in
fascicles, conspicuous conidiogenous loci and hila, thickened
and darkened, conidia solitary, size, shape and septation
variable), but the conidia are relatively thick-walled and at
least partly dictyosporous.
Notes: The morphological differentiation between
Sirosporium and mycovellosiella-like Passalora species is
vague. The phylogenetic meaning and value of thick conidial
walls and oblique to longitudinal septa as distinguishing
characters between Sirosporium and Passalora is unclear.
The phylogenetic position of the type species of Sirosporium
is still unknown. As already proposed in Braun (1995a) as
well as Crous & Braun (2003), Sirosporium is tentatively
treated as separate genus and should be conned to species
with dictyosporous conidia with thick walls.
DOUBTFUL AND EXCLUDED GENERA
Several hyphomycete genera have previously been
considered to be and treated as cercosporoid genera, but
based on modern phylogenetic examinations they are not
part of the family Mycosphaerellaceae and the corresponding
clade, i.e. they belong elsewhere and are not cercosporoid s.
str. Species of such genera are not treated here:
Miuraea Hara, Byochugai-Hoten (Manual of Pests and
Diseases): (260), 779 (1948).
Type species: Miuraea degenerans (Syd. & P. Syd.) Hara
1948.
Literature: von Arx (1983: 39), Braun (1995a: 218–223),
Seifert et al. (2011: 293), Crous et al. (2013: 69).
Notes: Statements that Miuraea is associated with
mycosphaerella-like teleomorphs and belongs to the
Mycosphaerellaceae were based on Miuraea persicae
(sexual morph: Mycosphaerella pruni-persicae). This species
clusters together with Pseudocercosporella species within
the Mycosphaerellaceae (Crous et al. 2013) and should
rather be assigned to the latter genus, which is in its current
circumscription non-monophyletic. It is quite unclear and
phylogenetically unproven if M. persicae is congeneric with
M. degenerans, the type species of Miuraea. The latter
species differs from M. persicae in having thin-walled,
hyaline hyphae and conidia, which become thick-walled
and pigmented with age. Longitudinal and oblique septa are
rather common. Miuraea s. str. does possibly not belong to
the Mycosphaerellaceae at all, but this is not yet proven.
Miuraea asiminae, another species treated under Miuraea in
Braun (1995a), was later reallocated to Pseudocercospora
(Braun & Crous 2008).
Phaeomycocentrospora Crous, H.D. Shin & U. Braun,
Stud. Mycol. 75: 61 (2012) [2013] [Pleosporales]
Type species: Phaeomycocentrospora cantuariensis (E.S.
Salmon & Wormald) Crous, H.D. Shin & U. Braun 2013.
Notes: The type species, originally described as species
of Cercospora, was later reallocated to Pseudocercospora
(Braun 1993). However, based on molecular sequence
analyses, recently carried out by Crous et al. (2013), it
was demonstrated that it represents an undescribed genus
belonging to the Pleosporales.
Thedgonia B. Sutton, Trans. Brit. Mycol. Soc. 61: 426
(1973) [Helotiales]
Type species: Thedgonia ligustrina (Boerema) B. Sutton
1973.
Literature: von Arx (1981, 1983), Braun (1990: 71; 1995a:
211–215), Crous & Braun (2003: 22), Seifert et al. (2011:
437), Crous et al. (2013: 61).
Notes: Thedgonia was previously considered a cercosporoid
genus with a type species originally assigned to Cercospora
s. lat. (von Arx 1983, Braun 1995a), which was seemingly
supported by the nding of the sexual morph of T. lupini,
described as Mycosphaerella lupini (Kaiser & Crous
1998), and the phylogenetic position of this species in the
Mycosphaerellaceae (Crous & Braun 2003). However, T.
ligustrina, the type species of Thedgonia, and T. lupini are not
congeneric and phylogenetically distinct since T. ligustrina
clusters within the Helotiales (Crous et al. 2009a, 2013).
Thedgonia lupini has to be reassessed and reallocated to
a genus within the Mycosphaerellaceae, maybe to the non-
monophyletic genus Pseudocercosporella to which this
species roughly ts. The afnity of other species assigned to
Thedgonia is unclear and unproven.
Xenostigmina Crous, Mycol. Mem. 21: 154 (1998)
[Pleosporales, Phaeosphaeriaceae]
Type species: Xenostigmina zilleri (A. Funk) Crous 1998.
Notes: Xenostigmina and its synasexual morphs in
Mycopappus belong to the Pleosporales (Crous et al. 2013),
i.e. they are not part of the Mycosphaerellaceae, in contrast
to Stigmina s. str. which has been reduced to synonym with
Pseudocercospora (Crous et al. 2006).
Braun et al.
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282 ima fUNGUS
KEY TO CURRENTLY RECOGNISED CERCOSPOROID GENERA
Expanded keys to cercosporoid genera and morphologically similar and confusable genera have been published in Braun
(1995a: 23–36), Crous & Braun (2003: 28–32) and Braun in Seifert et al. (2011: 887–893).
1 Saprobic or biotrophic, plant pathogenic hyphomycetes, causing various lesions, mostly leaf-spotting ........................ 2
Hyperparasitic (pathogenic on other fungi) or strictly fungicolous ................................................................................. 23
2 (1) Very large immersed sporodochium-like conidiomata, about 40–130 µm diam, with liform,
somewhat pigmented, irregularly verruculose conidiophores, aseptate, i.e. reduced to conidiogenous cells,
monoblastic, determinate or sympodially to percurrently proliferating, with slightly thickened and darkened
conidiogenous loci, and pale (subhyaline) to somewhat pigmented scolecosporous conidia formed singly;
on Ribes saximontanensis in North America .............................................................................. Quasiphloeospora
With other characters or combinations of characters ...................................................................................................... 3
3 (2) Stromata lacking; conidiophores in vivo solitary or in small loose fascicles (groups) emerging through stomata,
laxly erect, frequently branched, very pale brown; conidiogenous cells integrated, terminal, intercalary or
pleurogenous (as lateral branchlets), conidiogenous loci conspicuous, protruding, convex (papilla-like),
but wall of the loci neither distinctly thickened nor darkened, only somewhat refractive; conidia solitary,
didymo- to scolecosporous, pigmented (deeper in pigmentation than the conidiophores), hila neither
thickened nor darkened ..................................................................................................... Pseudocercosporidium
With other characters or combinations of characters; conidiogenous loci different, not papilla-like, but
truncate, either inconspicuous or more denticle-like, but always unthickened and not darkened, or
with conspicuously thickened and darkened-refractive loci ..................................................................................... 4
4 (3) Conidiogenous loci inconspicuous, neither thickened nor darkened or subconspicuous by being more
rigid or denticle-like, but wall of the loci always unthickend and not darkened, at most somewhat
refractive, or only slightly thickened and darkened around the rim (formed as minute somewhat darker
rim visible as darker circle) ...................................................................................................................................... 5
Conidiogenous loci conspicuous, thickened and darkened throughout, except for a very minute centre pore
(in front view visible as minute dark circle) ............................................................................................................ 14
5 (4) Forming supercial mycelium in vivo, hyphae verruculose-verrucose; conidiogenous cells terminal
and intercalary, denticulate, with lateral short peg-like protuberances, conidiogenous loci
inconspicuous, neither thickened nor darkened .................................................................................... Parastenella
Supercial hyphae in vivo lacking or, if present, smooth or almost so; with genuine conidiophores,
at least not with consistently peg-like protuberances ............................................................................................... 6
6 (5) Conidiophores in vivo in synnematous conidiomata ....................................................................................................... 7
Conidiophores in vivo solitary, fasciculate or in sporodochia .......................................................................................... 8
7 (6) Conidia often dictyosporous; on Guazuma ulmifolia, South America ............................................................. Pantospora
Conidia usually consistenty transversely septate; on other hosts ............... Pseudocercospora (synnematous species)
8 (6) Conidiophores in sporodochial conidiomata; wall of the densely arranged conidiophores somewhat
thickened and mostly distinctly verruculose, forming conspicuous, coarse annellations;
conidia transversely and occasionally also obliquely or longitudinally septate, wall often
somewhat thickened ...................................................................................................................... Scolecostigmina
Conidiophores thin-walled, rarely somewhat thickened, but then always smooth, annellations lacking
or if present ne and rather inconspicuous; conidia usually thin-walled and transversely septate .......................... 9
9 (8) Conidiophores in dense fascicles or in sporodochial conidiomata, distinctly verruculose; conidia solitary,
didymo- to phragmosporous, distinctly verruculose-verrucose ............................................. Pseudoasperisporium
Conidiophores and conidia smooth or almost so, at most faintly rough-walled ............................................................ 10
10 (9) Conidiogenous cells with distinct denticles; conidia amero- to phragmosporous, i.e. not scolecosporous ................... 11
Conidiogenous cells not denticulate or only subdenticulate (conidiogenous loci on shoulders caused
by sympodial proliferation) and then scolecosporous ............................................................................................ 12
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11 (10) Conidiogenous cells not geniculate, i.e. not distinctly sympodially proliferating, conidiogenous loci
formed as distinct terminal to lateral denticles; conidia solitary, didymo- to phragmosporous;
on Peristrophe, Asia ........................................................................................................... Semipseudocercospora
Conidiogenous cells sympodially proliferating; conidia catenate, 0–1(–3)-septate ....................................... Denticularia
12 (10) Conidiophores fasciculate and solitary, arising from supercial hyphae; conidiogenesis thalloblastic;
conidiogenous loci truncate, at, broad, loci and conidium initials of similar width, conidial
base not or barely constricted; conidia thick-walled; on Marsdenia in India ................................ Eriocercosporella
Conidiogenesis holoblastic; width of loci and conidium initials different, narrowed at the
attachment point between conidiogenous cells and conidium initial ...................................................................... 13
13 (12) Conidiophores solitary, arising from supercial hyphae, fasciculate, in sporodochia or even synnemata;
conidiogenous loci inconspicuous or conspicuous by being more or less denticle-like,
but locus wall always unthickened or almost so, usually neither darkened nor refractive,
rarely more rigid and more conspicuous by being somewhat darkened-refractive or slightly
thickened and darkened along the rim [loci ring-like, slightly thickened and darkened along the rim,
on Solanum, see Paracercospora s. str.]; conidia solitary, rarely in short chains, usually
scolecosporous, rarely amero- to phragmosporous, subhyaline to usually pigmented
[in vitro without red crystals] ....................................................................................................... Pseudocercospora
[On Colophospermum copane, South Africa; conidiogenous cells unilocal, determinate to percurrent,
Cercostigmina-like; phylogenetically distinct from Pseudocercospora s. str., see Phaeocercospora]
Conidiophores solitary or fasciculate; morphologically barely distinguishable from Pseudocercospora,
but phylogenetically distinct and with red crystals in vitro (P. heimii complex) ............................ Pallidocercospora
14 (4) In vivo with distinctly verruculose-verrucose supercial hyphae (aerial mycelium in vitro also verruculose);
conidia smooth or almost so to mostly verruculose-verrucose as well .................................................................. 15
Supercial hyphae in vivo lacking or, if present, smooth or almost so .......................................................................... 16
15 (14) Conidiogenous loci pileate; conidiophores in vivo solitary as well as fasciculate; conidia catenate;
on Pithecellobium, South America [Teratosphaeriaceae] ............................................................................. Stenella
Conidiogenous loci planate (Cercospora-like) [Mycosphaerellaceae] ........................................................... Zasmidium
16 (14) Conidiophores pigmented, rarely hyaline or subhyaline [subgen. Hyalocercospora]; conidia always
colourless, usually scolecosporous, acicular, liform, obclavate-cylindrical, and pluriseptate,
rarely amero- to phragmosporous .......................................................................................................... Cercospora
Conidiophores and conidia pigmented, at least faintly olivaceous ................................................................................ 17
17 (16) Conidia consistently distoseptate; conidiophores mostly frequently branched ..................................... Distocercospora
Conidia aseptate to euseptate or at most few distosepta mixed with eusepta ............................................................. 18
18 (17) Conidia distinctly (usually coarsely) verruculose-verrucose .......................................................................................... 19
Conidia smooth or almost so ......................................................................................................................................... 20
19 (18) Conidiophores mostly numerous in dense sporodochial conidiomata; conidia solitary, usually amero-
to phragmosporous, occasionally with longitudinal or oblique septa ................................................ Asperisporium
Conidiophores fasciculate; conidia solitary, but scolecosporous, only transversely septate
.................................................................................... Zasmidium (p.p., species without supercial hyphae in vivo)
20 (18) Conidia solitary, ± scolecosporous, fully developed conidia usually attenuated towards the tip,
somewhat rostrate, often somewhat curved, wall thickened; conidiogenous loci subconspicuous,
ranging from inconspicuous to conspicuous by being denticle-like or somewhat darkened-refractive,
but always unthickened; in vivo without supercial mycelium ......... see Pseudocercospora p.p. (incl. Prathigada)
Conidia solitary to catenate, amero- to scolecosporous, not rostrate, usually thin-walled; in vivo
with or without supercial mycelium ....................................................................................................................... 21
21 (20) In vivo with solitary conidiophores arising from supercial hyphae; conidia scolecosporous,
fairly thick-walled, with transverse and occasionally also longitudinal and oblique septa .................... Sirosporium
In vivo with internal or internal and external hyphae; conidia amero- to scolecosporous,
thin-walled, transversely septate ............................................................................................................................ 22
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22 (21) Conidia solitary or catenate; conidiogenous loci and conidial hila always distinctly
thickened and darkened ............................................................................................................................ Passalora
Conidia catenate; conidiogenous loci subconspicuous, ranging from inconspicuous to conspicuous
by being somewhat darkened-refractive, but always unthickened
....................................................................................... see Pseudocercospora p.p. (incl. Pseudophaeoramularia)
23 (1) Conidiophores laxly fasciculate, often branched; conidiogenous cells denticulate, but conidiogenous
loci not thickened and not darkened; hyperparasitic on ascomycetes (Phyllachora) ............................... Elletevera
In vivo with solitary conidiophores arising from supercial hyphae; conidiogenous loci not denticle-like ..................... 24
24 (23) Conidia catenate; conidiogenous loci somewhat thickened and darkened; on cercosporoid
hyphomycetes and rusts .................................................................................................................... Cladosporiella
Conidia solitary; conidiogenous loci subconspicuous, unthickened, not or only slightly
darkened-refractive; on sooty moulds .............................................................................................. Eriocercospora
TAXONOMIC TREATMENT
Fungicolous cercosporoid species
The genera Cladosporiella, Elletevera and Eriocercospora,
which may have afnity to cercosporoid genera, are treated
in this chapter. However, the phylogenetic positions of
these genera are quite unclear. Molecular data are not yet
known. Elletevera and Eriocercospora species are rather
pseudocercospora-like, but might also be related to similar
denticulate hyphomycete genera or to the Sporidesmium
complex. Afnity and position of Cladosporiella,
morphologically supercially similar to the Cladosporium
complex, is also quite unclear. This genus is treated here due
to rough similarity to phaeoramularia-like Passalora species.
All hyperparasitic taxa are only tentatively included in this
work on cercosporoids. The name Cercospora uredinophila
(Deighton 1969) suggests a hyperparasitic habit, but proved
to be a common phytopathogenic fungus on various Scirpus
spp. A comprehensive treatment of this species with full
synonymy was published in Braun (1995a). Cercospora
uromycestri and C. acori, reallocated to Passalora in Crous &
Braun (2003), seems to be the only true cercosporoid species
hyperparasitic or at least fungicolous on rust fungi.
CLADOSPORIELLA
Key to the species of Cladosporiella
1 Conidiophores very long, about 25–500 µm; on rust fungi ............................................................................................. 2
Conidiophores much shorter, about 8–40 µm; on cercosporoid hyphomycetes ............................................................. 3
2 (1) Stromata lacking; conidia (7–)15–40(–50) × 3.5–4.5 µm, (0–)1–2(–5)-septate ......................................... C. uredinicola
Stromata well-developed and large, 20–90(–150) µm diam; conidia broader, 4–6.5 µm,
(0–)1(–3)-septate .................................................................................................................................... C. uredines
3 (1) Conidia 20–135 × 2–5 µm, 0–9-septate .............................................................................................. C. cercosporicola
Conidia much shorter, 9–15 × 3–4 µm, 0–3-septate .................................................................................... C. deightonii
List of Cladosporiella species
Cladosporiella cercosporicola Deighton, Mycol. Pap.
101: 35 (1965).
(Fig. 2)
Illustrations: Deighton (1965: 36, g. 14), Ellis (1971: 303, g.
209), Seifert et al. (2011: 709, g. 227B).
Literature: Ellis (1971: 303).
Description: Colonies hyperparasitc on Passalora koepkei,
loosely occose, pale. Mycelium supercial, some pale hyphae
tightly coiled around conidiophores and conidia, branched,
septate, thin-walled, smooth. Stromata none. Conidiophores
arising from supercial hyphae, erect, straight to exuous,
unbranched, 8–40 × 3–4 µm, continuous or sparingly
septate, pale olivaceous, thin-walled, smooth; conidiogenous
cells integrated, terminal or conidiophores reduced to
conidiogenous cells, about 5–20 µm long, conidiogenous
loci conspicuous, slightly thickened and darkened, small,
about 0.8–1 µm diam. Conidia catenate, in branched chains,
subcylindrical to liform or somewhat obclavate, 20–135 ×
2–4.5(–5) µm, 0–9-septate, pale olivaceous, smooth, thin-
walled, ends short obconical, occasionally slightly swollen,
hila slightly thickened and darkened.
Holotype: Sabah: on Passalora koepkei, Mycosphaerellaceae,
on Saccharum ofcinarum, Poaceae, 1964, J. Solomon (K(M)
IMI 107538b).
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Host range and distribution: Only known from the type
collection.
Cladosporiella deightonii R.F. Castañeda & U. Braun,
Cryptog. Bot. 1: 43 (1989).
(Fig. 3)
Illustration: Castañeda & Braun (1989: 45, g. 9).
Description: Colonies on leaf spots of and associated
with Cercospora coffeicola, hypophyllous, pale. Mycelium
supercial; hyphae exuous, richly branched, 1–4 µm
wide, pale olivaceous-brown, septate, thin-walled, smooth.
Stromata lacking. Conidiophores arising from supercial
hyphae, lateral, erect, almost straight to geniculate-sinuous,
narrowed towards the apex, unbranched, 8–30 × 2–3 µm,
aseptate, rarely with 1–2 septa, olivaceous, thin-walled,
smooth; conidiophores usually reduced to conidiogenous
cells, rarely with integrated, terminal conidiogenous cells,
conidiogenous loci conspicuous, minute, about 1 µm diam,
slightly thickened and darkened, often situated on short
peg-like protuberances or shoulders caused by sympodial
proliferation near the apex. Conidia formed singly or in short
chains, ellipsoid, subcylindrical, subclavate, 9–15 × 3–4
µm, 0–3-septate, pale olivaceous to brownish, thin-walled,
smooth, ends rounded to attenuated, hila slightly thickened
and darkened, about 1–1.5 µm wide.
Types: Cuba: Los Corrales de Guisa, Granma, on
Cercospora coffeicola, Mycosphaerellaceae, on Coffea
arabica, Rubiaceae, 24 June 1987, R. F. Castañeda (INIFAT
C87/171 - holotype; HAL 1649 F – isotype).
Host range and distribution: Only known from the type
collection.
Cladosporiella uredinicola Deighton, Mycol. Pap.
118: 33 (1969).
(Fig. 4)
Illustration: Deighton (1969: 34–35, gs 19–20).
Description: Colonies on uredosori, effuse, subvelutinous,
grey-brown to brown. Mycelium immersed in the sori
as well as supercial; hyphae 2–3 µm wide, sparingly
branched, olivaceous, thin-walled, smooth. Stromata lacking.
Conidiophores arising from supercial hyphae, lateral or
sometimes terminal, erect, straight to curved or somewhat
Figs 2–5. 2. Cladosporiella cercosporoides (K(M) IMI 107538b). 3. C. deightonii (HAL 1649 F). 4. C. uredinicola (K(M) IMI 43280b). 5. C. uredinis
(K(M) IMI 16432b). A. Solitary conidiophores arising from supercial hyphae. B. Supercial hyphae. C. Conidia. D. Conidiophore fascicles. E.
Conidiophore tips. Bar = 10 µm.
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geniculate-sinuous, unbranched or branched, 25–500 ×
2.5–4 µm, pluriseptate, moderately olivaceous, paler towards
the tip, thin-walled, smooth; conidiogenous cells integrated,
terminal to intercalary, about 10–30 µm long, proliferation
sympodial; conidiogenous loci conspicuous, mostly slightly
prominent or on peg-like protuberances, slightly thickened
and darkened. Conidia in simple or occasionally branched
chains, ellipsoid-ovoid, subcylindrical, fusiform, obclavate,
(7–)15–40(–50) × (3.5–)4–4.5 µm, (0–)1–2(–5)-septate, pale
olivaceous, thin-walled, smooth, ends rounded to narrowed,
hila slightly thickened and darkened.
Holotype: Sierra Leone: Dodo, on uredosori of Puccinia
eucomi, Pucciniaceae, on Andropogon auriculatus, Poaceae,
15 Apr. 1940, F. C. Deighton (K(M) IMI 43280b).
Host range and distribution: On uredo- and teleutosori of
rust fungi, on Puccinia eucomi, Pucciniaceae, and Ravenelia
zygiae, Raveneliaceae, Africa (Sierra Leone).
Cladosporiella uredinis Deighton, Mycol. Pap. 118:
36 (1969).
(Fig. 5)
Illustrations: Deighton (1969: 37–38, gs 21–22).
Exsiccatae: Sydow, Fungi Exot. Exs. 444.
Description: Colonies on uredo- and teleutosori, pale to deep
grey-green, short and densely velutinous-occose to densely
occose, sometimes forming dense round to elliptic masses
up to 4 mm diam. Mycelium immersed within the sori and
extending into the surrounding leaf-tissue; hyphae branched,
1.5–2 µm diam, septate, pale olivaceous, thin-walled,
smooth, sometimes completely suppressing the formation of
uredospores. Stromata small to well-developed, immediately
above the sori hyphae aggregated, forming dense stromata,
often 25–80 µm wide and 20–90 µm deep, occasionally up
to 150 µm diam, composed of densely packed, ascending,
sparingly branched hyphae developing to conidiophores.
Conidiophores in smaller to mostly large or very large, dense
fascicles, arising from stromata, laxly erect, straight or almost
so to exuous, liform, simple or occasionally with short
branchlets, 60–350 × 2.5–4 µm, pluriseptate, pale yellowish
olivaceous to medium olivaceous, paler towards the tip, thin-
walled, smooth; conidiogenous cells integrated, terminal
to intercalary, conidiogenous loci truncate, occasionally
subdenticulate, 1–1.5(–2) µm diam, very slightly thickened
and darkened-refractive. Conidia in simple or branched
acropetal chains, ellipsoid-ovoid, fusiform, subcylindrical,
straight to somewhat curved, (12–)15–35(–45) × 4–6.5 µm,
(0–)1(–3)-septate, pale olivaceous, thin-walled, smooth,
primary conidia with broadly rounded apex, otherwise both
ends short obconically truncate, hila 1–1.5 µm wide, very
slightly thickened, darkened-refractive.
Types: Philippines: on uredosori (Uredo sp.) on Scirpus
grossus, Cyperaceae, 20 Mar. 1913, P. W. Graff, Sydow,
Fungi Exot. Exs. 444 (p.p.) (K(M) IMI 164332b – holotype;
Sydow, Fungi Exot. Exs. 444 (e.g. BPI 420980) – isotypes).
Host range and distribution: On uredo- and teleutosori of
rust fungi, on Puccinia (polygoni-amphibi, scleriae, solmsii,
thaliae) and Uredo sp., Pucciniaceae, Asia (India, Malaysia,
Philippines), West Indies (Trinidad).
Excluded Cladosporiella species
Cladosporiella foliicola R.F. Castañeda, Fungi
Cubenses 3: 3 (1988).
Illustration: Castañeda (1988: g. 2).
Holotype: Cuba: Prov. Pinar del Rio: Soroa, on living leaves
of Blechnum occidentale L., Blechnaceae, 11 Mar. 1987, R.
F. Castañeda (INIFAT C87/82).
Notes: This is a foliicolous species with very long, simple or
branched, pigmented, 8–15-septate conidiophores, 180–300
× 6–7 µm, ramoconidia and extremely long, brown conidia,
100–310 × 4–5 µm, with 15–30(–35) septa. The generic
afnity of this unusual species is quite unclear.
ELLETEVERA
Key to the species of Elletevera
1 Conidiophores very long, 100–250 µm, frequently branched; conidia 12–60 × (4–)5–7.5(–10) µm,
(0–)2–4(–6)-septate, very pale olivaceous; on Phyllachora parasitica and Phyllachora sp., Africa,
North and South America ...................................................................................................................... E. parasitica
Conidiophores shorter, 85–120 µm, unbranched; conidia 17–23 × 7–9 µm, (1–)2(–3)-septate,
pale olivaceous-ochraceous; on Phyllachora shiraiana, Asia, Japan .................................................... E. ochracea
List of Elletevera species
Elletevera ochracea Katum., Bull. Fac. Agric.
Yamaguchi Univ. 35: 110 (1988).
(Fig. 6)
Illustration: Katumoto (1988: 110, g. 2).
Description: Colonies hypophyllous, on stromata of
Phyllachora shiraiana, ochraceous to brownish, elliptical to
oblong in shape, 1.5–7 × 1.5–3.5 mm. Mycelium immersed;
hyphae 2.5–3 µm wide, branched, septate, subhyaline to
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very pale olivaceous. Conidiophores numerous, arising
from supercial stromata, at, pseudoparenchymatous,
35–40 µm diam, olivaceous-ochraceous, erect, divergent,
cylindrical, straight to sinuous, sometimes geniculate in the
upper portion, unbranched, 85–120 × 4.5–6.5 µm, attenuated
towards the tip (to about 2.5 µm), 1–2-septate in the lower
portion, pale olivaceous, thin-walled, smooth; conidiogenous
cells integrated, terminal, long, conidiogenous loci visible,
subdenticulate. Conidia solitary, short cylindrical-obovoid
to oblong, straight to slightly curved, 17–23 × 7–9 µm,
(1–)2(–3)-septate, pale olivaceous-ochraceous, thin-walled,
smooth, apex broadly rounded, base short obconically
truncate, hila 0.8–1 µm wide.
Holotype: Japan: Yamaguchi Pref.: Nishiki-cho, Mt. Jakuchi,
on stromata of Phyllachora shiraiana, Phyllachoraceae, on
leaves of Sasa palmata, Poaceae, 6 May 1985, K. Katumoto
(YAM 24294).
Host range and distribution: Only known from the type
collection.
Elletevera parasitica (Ellis & Everh.) Deighton, Mycol.
Pap. 118: 19 (1969).
(Fig. 7)
Basionym: Pyricularia parasitica Ellis & Everh., Proc. Acad.
Nat. Sci. Philadelphia 45: 462 “1893” (1894).
Synonym: Pyricularia grisea var. parasitica Ellis & Everh., in
Sumstine, Mycologia 41: 13 (1949), nom. nud.
Illustrations: Deighton (1969: 19, g. 11 and pl. 1), Seifert et
al. (2011: 701, g. 219C).
Exsiccatae: Davis, Fungi Wiscon. Exs. 152.
Description: Colonies amphigenous, mainly hypophyllous,
on and around Phyllachora spp., pale greenish, occose,
often contiguous. Mycelium immersed in ascomata of the
host fungus; hyphae 1.5–3 µm wide, branched, septate,
subhyaline to faintly olivaceous, thin-walled, smooth.
Fig. 6. Elletevera ochracea (YAM 24294). A. Conidiophores arising
from stroma cells. B. Conidiophores. C. Conidiophore tip. D. Conidia.
Bar = 10 µm.
Fig. 7. Elletevera parasitica (NY, holotype). A. Conidiophores. B.
Conidia. Bar = 10 µm.
Braun et al.
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288 ima fUNGUS
Stromata formed by aggregated pale olivaceous, ascending
hyphae, about 3–4 µm wide, in ostioles of the ascomata,
40–50 µm diam. Conidiophores numerous, in divergent
occose fascicles, arising from stromata, through ostioles,
also erumpent through the wall of ascomata and from repent
external hyphae, erect, cylindrical-liform, 100–250 × 4–5
µm, sometimes narrowed towards the tip and 2.5–4 µm
wide, simple or branched, with 1–2 main branches and short
lateral branchlets, about 20–25 µm long, pluriseptate, very
pale olivaceous, thin-walled, smooth; conidiogenous cells
integrated, terminal, conidiogenous loci neither thickened nor
darkened, but distinct by being denticle-like and somewhat
refractive, about 1–1.5 µm diam. Conidia solitary, broadly
ellipsoid-ovoid, obovoid, fusiform, obclavate, cylindrical-
clavate, occasionally rostrate, straight to somewhat curved,
12–60 × (4–)5–7.5(–10) µm, (0–)2–4(–6)-septate, very pale
olivaceous, thin-walled, smooth, apex broadly rounded,
more attenuated in conidia with rostrate apex, base short
obconically truncate, hila about 1–2 µm wide.
Holotype: USA: Wisconsin: Kenosha Co., on Phyllachora
graminis, on leaves of Elymus virginicus, 13 Aug. 1883, J. J.
Davis 9311 (NY).
Host range and distribution: On Phyllachora (graminis,
vulgata, Phyllachora sp.), Phyllachoraceae, on Beckeropsis
uniseta, Elymus (canadensis, hystrix, virginicus),
Muhlenbergia sp. and Paspalum virgatum, Poaceae, Africa
(Zambia), North America (USA, Delaware, Indiana, Iowa,
Oregon, Wisconsin), South America (Venezuela).
Notes: Most collections have been made in the USA on
Phyllachora graminis on Elymus ssp. and Muhlenbergia
sp. Other collections have been made on Phyllachora sp.
in Zambia on Beckeropsis uniseta (Deighton 1969) and
Venezuela on Paspalum virgatum (Urtiaga 1986). A collection
of this species on Musa paradisiaca from Sierra Leone (K(M)
IMI 45910) is unclear and unproven.
ERIOCERCOSPORA
Key to the species of Eriocercospora
1 Conidiophores up to 600 µm long and (3.5–)5–9(–13) µm wide; conidia (3.5–)5–9(–13) µm wide;
on various hosts belonging to the Asterinaceae, Englerulaceae and Parodiopsidaceae ................... E. balladynae
Conidiophores shorter, up to 120 µm long, and only 2.5–4 µm wide; conidia narrower,
only up to 4.5 µm; on Meliolina molle, Meliolinacea ................................................................................ E. olivacea
List of Eriocercospora species
Eriocercospora balladynae (Hansf.) Deighton, Mycol.
Pap. 118: 6 (1969).
(Fig. 8)
Basionym: Helminthosporium balladynae Hansf., Proc. Linn.
Soc. London 157: 39 (1945).
Synonyms: Cercospora schiffnerulae Hansf., Mycol. Pap. 15:
216 (1946).
Cercospora balladynae Hansf., Proc. Linn. Soc. London 158:
50 (1947).
Literature: Ellis (1971: 248–249).
Illustrations: Deighton (1969: 7–15, gs 2–9), Ellis (1971:
249, g. 170), Seifert et al. (2011: 693, g. 211D).
Description: Colonies overgrowing those of the host fungi,
olivaceous, velutinous to wooly. Mycelium supercial;
hyphae growing between host hyphae, branched, 2–3.5 µm
diam, occasionally swollen, up to 5 µm wide, septate, pale
olivaceous, darker around conidiophores. Conidiophores
arising from supercial hyphae, erect or almost so, straight
to curved, exuous, sinuous or somewhat geniculate,
simple or branched, length very variable, 15–600 ×
4–7.5(–10.5) µm, width often variable throughout their
length, pluriseptate, pale to medium olivaceous, thin-
walled, smooth; conidiogenous cells integrated, terminal
or intercalary, about 10–30 µm long, conidiogenous loci
subconspicuous, i.e. unthickened, not or barely protuberant,
not or only slightly darkened-refractive, about 1.5–2.5 µm
diam. Conidia solitary, shape rather variable, ellipsoid-
ovoid, obovoid, fusiform, subclavate to obclavate, (15–)
25–70(–135) × (3.5–)5–9(–13) µm, (1–)3–6(–9)-septate,
occasionally slighty constricted at the septa, very pale
olivaceous to medium pale olivaceous, thin-walled, smooth,
apex obtuse, broadly rounded to somewhat attenuated,
base obconically truncate, hila about 1–2.5 µm wide, not to
slightly darkened-refractive, barely thickened.
Holotype: Uganda: Entebbe road, on Balladynopsis
entebbeensis, on leaves of Grumilea succulenta, Nov. 1943,
C. G. Hansford 3264 (K(M) IMI 562c).
Host range and distribution: On Asterina sp., Asterinaceae;
Balladyna (tenuis, velutina, Balladyna sp.), Balladynocallia
glabra, Balladynopsis entebbeensis, Parodiopsidaceae;
Clypeolella (camelliae, Clypeolella sp.), Schiffnerula
(hippocrateae, mirabilis, periplocae, toddaliae, Schiffnerula
sp.), Englerulaceae, Africa (Ghana, Sierra Leone, South
Africa, Uganda), Asia (Malaysia, Myanmar, Papua New
Guinea), Central America (Costa Rica, Panama).
Eriocercospora olivacea Piroz., Kavaka 2: 38 (1974).
(Fig. 9)
Illustration: Pirozynski (1974: 39, g. 3).
Description: Colonies conned to colonies of Meliolina
molle, olivaceous, felty, 1–2 mm thick, completely obscuring
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the host fungus. Mycelium supercial; hyphae appressed
to host hyphae but not penetrating them, often toruloid,
2.5–5 µm wide, septate, pale olivaceous, thin-walled,
smooth. Conidiophores arising from supercial hyphae,
lateral, occasionally terminal, erect, geniculate-sinuous,
simple or branched, up to 120 µm long and 2.5–4 µm
wide, pluriseptate, pale olivaceous, thin-walled, smooth;
conidiogenous cells integrated, terminal and intercalary, 10–
20 µm long; conidiogenous loci truncate, about 1.8–2.5 µm
diam, neither thickened nor darkened or at most refractive
or very slightly darkened. Conidia solitary, long, cylindrical-
vermiform or liform to somewhat obclavate, up to 150
µm long and 4.5 µm wide, pluriseptate (up 22 septa), pale
olivaceous to olivaceous, thin-walled, smooth, apex obtuse,
rounded, base truncate to short obconically truncate, about
2–2.5 µm, hila neither thickened nor darkened.
Types: India: Tamil Nadu: Nilgiris, Gudalpur, on Meliolina
molle, Meliolinacea, on Syzygium cf. grande (S. cf.
montanum), Myrtaceae, 22 Feb. 1966, K. A. Pirozynski (K(M)
120229b – holotypes; DAOM 145763 – isotype).
Host range and distribution: Only known from the type
collection.
Excluded Eriocercospora species
Eriocercospora moghaniae Suj. Singh, Indian
Phytopathol. 33: 610 “1980” (1981).
Synonym:Stenella moghaniae (Suj. Singh) Kamal & Narayan,
Indian Phytopathol. 39: 201 (1987).
= Pseudocercospora moghaniae (Suj. Singh) U. Braun
& Crous, in Crous & Braun, Mycosphaerella and
Anam. 1: 18 (2003).
Eriocercospora palustris R.F. Castañeda & W.B.
Kendr., Univ. Waterloo Biol. Ser. 35: 52 (1991).
Notes: This is a saprobic mucedinaceous hyphomycete
of unknown generic afnity, in any case not belonging to
Eriocercospora. It is supercially somewhat reminiscent of
plant pathogenic species of Pseudocercosporella.
Eriocercospora websteri P. Rag. Rao, Manohar. & P.
Ram Rao, Curr. Sci. 51: 1155 (1982).
= Pseudocercospora websteri (P. Rag. Rao, Manohar.
& P. Ram Rao) U. Braun, Schlechtendalia 8: 36 (2002).
Fig. 8. Eriocercospora balladynae (K(M) IMI 562c). A. Conidiophores.
B. Conidia. Bar = 10 µm.
Fig. 9. Eriocercospora olivacea (K(M) IMI 120229b). A. Supercial
hyphae. B. Conidiophores. C. Conidia. Bar = 10 µm.
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FUNGIICOLOUS PASSALORA SPECIES
Key to fungiicolous Passalora species
1 Mycelium internal and external, supercial; stromata lacking; conidiophores consistently solitary,
arising from supercial hyphae; on Uromyces sparganii, Asia ....................................................................... P. acori
Mycelium immersed; stromata present, dark brown, columnar, globose to linear, often conuent
to form continuous stromatic areas over and within aecia; conidiophores arising from
stromatic structures, in small to moderately large fascicles; on aecia of Uromyces cestri, North
America .............................................................................................................................................. P. uromycestri
Fig. 10. Passalora acori (PC, holotype). A. Hyphae in uredo spores.
B. Solitary conidiophores arising from a supercial hypha. C. Conidia.
Bar = 10 µm.
Fig. 11. Passalora uromycesti (BPI 71801). A. Stroma. B.
Conidiophore fascicle. C. Conidiogenous cells. D. Conidia. Bar = 10
µm.
List of fungiicolous Passalora species
Passalora acori (J.M. Yen) U. Braun & Crous, in Crous
& Braun, Mycosphaerella and Anam. 1: 43 (2003).
(Fig. 10)
Basionym: Cercospora acori J.M. Yen, Rev. Mycol. 29: 209
(1964).
Synonym: Mycovellosiella acori (J.M. Yen) J.M. Yen, Bull.
Trimestriel Soc. Mycol. France 97: 131 (1981).
Description: Yen & Lim (1980: 154).
Illustration: Yen & Lim (1980: 205, g. 8).
Description: Leaf spots indistinct, sometimes forming dark
grey specks. Colonies on uredospores, amphigenous,
effuse, dark brown. Mycelium internal and external, hyphae
branched, septate, 2.5–5 µm wide, yellowish brown to
brown, thin-walled, smooth. Conidiophores consistently
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solitary, arising from supercial hyphae, erect, cylindrical, not
geniculate, unbranched, 20–190 × 4–5 µm, 2–6(–8)-septate,
brown to dark brown, tips rounded to conical; conidiogenous
cells integrated, terminal, with several minute brown
conidiogenous loci. Conidia solitary, obclavate to obclavate-
subacicular, 40–120 × 2–2.5 µm, 4–9(–12)-septate, pale
olivaceous-brown, thin-walled, smooth, apex pointed, base
obconically truncate, hila slightly thickened and darkened.
Holotype: Singapore: Botanic Gardens, on uredospores of
Uromyces sparganii [probably U. sparganii subsp. acori ≡
U. acori], Pucciniaceae, on Acorus calamus, Acoraceae, 26
Jan. 1964, S. H. Yen, No. 7 (PC).
Host range and distribution: Only known from the type
collection.
Passalora uromycestri (Pollack) U. Braun & Crous,
in Crous & Braun, Mycosphaerella and Anam. 1: 417
(2003).
(Fig. 11)
Basionym: Cercospora uromycestri Pollack, Mycologia 63:
691 (1971).
Illustrations: Pollack (1971: 690, gs 1–8).
Description: Colonies formed as dark brown to blackish felt-like
growth on aecial pustules of Uromyces cestri, which formed
small, swollen punctiform spots, 1–3 mm diam, surrounded
by a yellowish halo. Mycelium immersed. Stromata dark
brown, columnar, globose to linear, often conuent to form
continuous stromatic areas over and within the aecia.
Conidiophores arising from stromatic structures, in small
to moderately large fascicles, loose to rather dense, erect,
straight to slightly geniculate-sinuous, mostly unbranched,
about (15–)20–110(–130) × 4–6 µm, pluriseptate, light
to dark brown, wall thin to slightly thickened, smooth or
almost so; conidiogenous cells integrated, terminal, about
10–30 µm long, conidiogenous loci conspicuous, thickened
and darkened, slightly protuberant or often lateral and at,
appressed, 1–2 µm diam. Conidia solitary, obclavate, 30–100
× 2.5–4 µm, (0–)3–6(–7)-septate, subhyaline to usually pale
olivaceous, thin-walled, smooth, apex obtuse, base short
obconically truncate, hila 1.5–2 µm broad, slightly thickened
and darkened.
Types: USA: Florida: Miami, on aecia of Uromyces cestri, on
leaves of Cestrum diurnum, 6 Nov. 1969, F. D. Matthews (BPI
71801 – holotype; BPI 71802–71804 and K(M) IMI 16083 –
paratypes).
Host range and distribution: On Uromyces cestri,
Pucciniaceae, on leaves of Cestrum diurnum, Solanaceae,
North America (USA, Florida).
CERCOSPOROID SPECIES ON FERNS (PTERIDOPHYTA)
Cercospora
Tabular key to Cercospora species on ferns
Most species on ferns belong to the Cercospora apii complex (sensu Crous & Braun 2003), i.e. they are morphologically very
similar and morphologically little differentiated, so that it is barely possible to key out the species concerned just based on
morphology. Phylogenetic data are not yet available. Therefore, we prefer to list these species in form of a tabular key based on
host genera and families.
Adiantaceae = Pteridaceae subfam. Vittarioideae
Aspleniaceae
1 Conidia acicular, base truncate; on Asplenium spp., Europe, North America ................................................. C. asplenii
Conidia narrowly obclavate-liform, base obconically truncate; on Asplenium radicans .......... Cercospora camptosori
Cheilanthaceae = Pteridaceae subfam. Cheilanthoideae
Cyatheaceae
On Alsophila, North America ....................................................................................................................................... C. asplenii
Dryopteridaceae
1 Stromata 20–50 µm diam; conidia 30–125 × 1.5–3.5 µm, base obconically truncate, 1–2 µm wide;
on Dryopteris sp., North America ....................................................................................................... C. camptosori
Stromata lacking or very small; conidia 95–250 × 3.5–5 µm, base truncate, wider, about 3–4 µm wide;
on Dryopteris polypodioides, China .................................................................................................. C. dryopteridis
Lygodiaceae
On Lygodium japonicum, India ............................................................................................................................... C. lygodiicola
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Marsileaceae
On Marsilea quadrifolia, India ................................................................................................................................... C. marsileae
Polypodiaceae
1 Stromata lacking or almost so; on Platycerium spp., North America ............................................................. P. platycerii
Stromata 20–45 µm diam; on Microsorum membranaceum (Pleopeltis membranacea), Asia (India) ...... P. pleopeltidis
Pteridaceae subfam. Cheilanthoideae (= Cheilanthaceae)
On Aleuritopteris farinosa (Cheilanthes farinosa), India ......................................................................................... C. cheilanthis
Pteridaceae subfam. Vittarioideae (= Adiantaceae)
1 Conidia broadly obclavate-cylindrical, short conidia sometimes fusiform, 40–90 × (4–)5–8 µm; on Adiantum, Tanzania
............................................................................................................................................................. C. adiantigena
Conidia acicular, only 2–5 µm wide ................................................................................................................................. 2
2 (1) Conidia 145.5–455.5 × 3.5–5 µm; on Adiantum, India .............................................................................. C. pteridigena
Conidia much shorter and narrower, up to 120 µm long and 2–4 µm wide ..................................................................... 3
3 (2) Stromata lacking or very small; on Adiantum, Asia .................................................................................... C. adianticola
Stromata well-developed, up to 75 µm diam; on Adiantum, North America .................................................... C. asplenii
Selaginellaceae
On Selaginella spp., Europe, North America .................................................................................................... C. selaginellarum
Thelypteridaceae
1 Conidia long, 60–400 × 2–4 µm, 1–35-septate; on Abacopteris, Singapore ......................................... C. abacopteridis
Conidia shorter, 50–110 × 3–4 µm, usually 4–10-septate; on Cyclosorus, India, Taiwan ............................. C. cyclosori
Woodsiaceae
1 Conidia acicular, 20–90 µm long, base truncate; on Diplazium esculentum, Philippines .................................. C. athyrii
Conidia acicular to narrowly obclavate, 45–155(–230) µm, base truncate to short obconically truncate;
on Diplazium esculentum, India ......................................................................................................... C. diplaziicola
List of Cercospora species on ferns
Cercospora abacopteridis J.M. Yen & Lim, Cah.
Pacique 17: 96 (1973).
(Fig. 12)
Literature: Yen & Lim (1980: 154), Crous & Braun (2003: 37).
Illustrations: Yen & Lim (1980: 204, g. 7C–E).
Description: Leaf spots amphigenous, vein-limited, brown,
later conuent. Caespituli hypophyllous, inconspicuous.
Mycelium internal. Stromata lacking. Conidiophores in
relatively small fascicles, 2–18, divergent or somewhat
denser, arising from internal hyphae, emerging through
stomata, erect, straight, subcylindrical to somewhat
sinuous, 0–2 times geniculate above, unbranched, 15–120
× 4–5 µm, 1–5-septate, dark brown, paler towards the tip,
wall thin to slightly thickened, smooth; conidiogenous cells
integrated, terminal, occasionally intercalary, conidiogenous
loci conspicuous, 2–2.5 µm wide, darkened and thickened.
Conidia solitary, acicular to liform, straight to somewhat
curved, 60–400 × 2–4 µm, 1–35-septate, hyaline, thin-walled,
smooth, apex acute, base truncate, hilum about 2–3 µm wide,
thickened and darkened.
Holotype: Singapore: on Abacopteris urophylla,
Thelypteridaceae, 3 Mar. 1972, G. Lim (PC).
Host range and distribution: Only known from the type
collection.
Cercospora adianticola R.K. Srivast., A.K. Srivast.
& Kamal, Kavaka 20/21: 38 “1992/1993” (1995); as
“adiantiicola“.
(Fig. 13)
Literature: Crous & Braun (2003: 44), Kamal (2010: 13).
Illustration: Srivastava et al. (1995: 38, g. 1).
Description: Leaf spots amphigenous, 0.5–3 mm diam,
irregular, streak-like, sometimes conuent, spread
over the whole leaf surface. Mycelium internal; hyphae
septate, branched. Stromata lacking or poorly developed.
Conidiophores in small fascicles, erect, straight,
subcylindrical to 1–3 times geniculate, unbranched, 70–
140 × 3–6 µm, septate, olivaceous; conidiogenous cells
integrated, terminal and intercalary, conidiogenous loci
conspicuous, 2–4.5 µm diam, thickened and darkened.
Conidia solitary, acicular, 50–100 × 2–4 µm, 3–11-septate,
hyaline, thin-walled, smooth, apex acute, base truncate or
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slightly atenuated, hila about 1.5–2.5 µm wide, thickened
and darkened.
Holotype: India: U.P.: Maharaj Ganj, Nepal, Pakari Forest,
Narayan Ghat, on Adiantum venustum, Pteridaceae subfam.
Vittarioideae (= Adiantaceae), Nov. 1989, V. P. Pandey (K(M)
IMI 243785).
Host range and distribution: only known from the type
collection.
Notes: This species belongs to the C. apii s. lat. complex.
Thaung (1984) recorded “Cercospora asplenii” on Adiantum
sp. from Myanmar, but this collection seems to belong to C.
adianticola.
Cercospora adiantigena U. Braun & Crous, in Crous
& Braun, Mycosphaerella and Anam. 1: 44 (2003).
(Fig. 14)
Literature: To-anun et al. (2011: 32), Begum et al. (2012).
Illustrations: Crous & Braun (2003: 45, g. 2), To-anun et al.
(2011: 32, g. 14).
Description: Leaf spots amphigenous, oblong, linear, spread
between veins, 2–15 × 0.5–1.5 mm or conuent, forming
larger patches, brown. Caespituli amphigenous, punctiform
to subeffuse, greyish brown to dingy greyish white.
Mycelium internal. Stromata well-developed, immersed,
10–50 µm diam, subhyaline to pale olivaceous-brown.
Conidiophores in small to moderately large fascicles, loose
to fairly dense, arising from stromata, erumpent, erect,
straight, subcylindrical to moderately geniculate-sinuous,
unbranched, 25–150 × 4–10 µm, pluriseptate, subhyaline,
occasionally with a faintly greenish to olivaceous tinge, thin-
walled, smooth; conidiogenous cells integrated, terminal or
occasionally intercalary, 10–35 µm long, conidiogenous loci
conspicuous, planate, thickened and darkened throughout,
2.5–4 µm diam. Conidia solitary, broadly obclavate-cylindrical,
short conidia sometimes fusiform, 40–90 × (4–)5–8 µm,
(1–)3–6(–8)-septate, colourless, thin-walled, smooth, apex
Fig. 12. Cercospora abacopteridis (PC, holotype). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm. Fig. 13. Cercospora adianticola (K(M) IMI 243785). A. Conidiophore
fascicle. B. Conidiogenous cells. C. Conidia. Bar = 10 µm.
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obtuse, base short obconically truncate, 3–4 µm wide, hila
thickened and darkened.
Holotype: Tanzania: Kigoma, Kakombe, on Adiantum
philippense, 21 Feb. 1964, K. A. Pirozynski (K(M) IMI
106709b).
Host range and distribution: On Adiantum philippense
and Doryopteris sp., Pteridaceae subfam. Vittarioideae
(Adiantaceae), Africa (Tanzania), Asia (Philippines, Thailand).
Notes: This species is reminiscent of Cercosporella, but the
conidiogenous loci are planate, thickened and darkened
throughout, i.e. they are cercosporoid. Thus this species
rather pertains to Cercospora subgen. Hyalocercospora.
Cercospora asplenii Jaap, Ann. Mycol. 14: 43 (1916).
(Fig. 15)
Synonym: Mycosphaerella asplenii Thirum. & Govindu,
Sydowia 8: 345 (1954), nom. illeg., (Art. 53.1).
Literature: Chupp (1954: 454), Vassiljevsky & Karakulin
(1937: 219), Vasudeva (1963: 43–44), Crous & Braun (2003:
69), Kamal (2010: 19), Motohashi et al. (2010).
Exsiccatae: Jaap, Fungi Sel. Exs. 616. Petrak, Mycoth. Gen.
327.
Description: Leaf spots variable in shape and size, small to
relatively large, pale to dark brown or almost blackish, centre
later paler, pale brown to dingy grey. Caespituli amphigenous.
Stromata subglobose, small to large, up to 75 µm diam, brown
to almost blackish brown. Conidiophores in small to moderately
large fascicles, 2–20, occasionally solitary, divergent, straight,
subcylindrical to geniculate-sinuous, 0–5 times geniculate,
usually unbranched, 20–120 × 4–5.5 µm, pluriseptate, medium
olivaceous-brown throughout, wall slightly thickened, smooth;
conidiogenous cells integrated, mostly terminal, conidiogenous
loci conspicuous, thickened and darkened. Conidia solitary,
acicular, 20–120 × 2–4 µm, indistinctly pluriseptate, hyaline,
thin-walled, smooth, apex subacute, base truncate, hilum
about 2–2.5 µm wide, thickened and darkened.
Fig. 14. Cercospora adiantigena (K(M) IMI 106709b). A. Conidiophore
fascicle. B. Conidiogenous cells. C. Conidia. Bar = 10 µm. Fig. 15. Cercospora asplenii (HBG, holotype). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
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Holotype: Croatia: Dalmatia, Lapad near Ragusa, on
Asplenium trichomanes, 18 Mar. 1914, O. Jaap (HBG).
Host range and distribution: On ?Adiantum sp., Pteridaceae
subfam. Vittarioideae (Adiantaceae); ?Alsophila australis,
Cyatheaceae; Asplenium (nidus [antiquum], trichomanes,
Asplenium sp.), Aspleniaceae; North America (Canada,
USA, Florida), Asia (India, Japan, Myanmar), Europe (Czech
Republic, Germany, Croatia, Switzerland).
Cercospora athyrii Mendoza, Philipp. J. Sci. 75: 165
(1941).
Literature: Chupp (1954: 454), Crous & Braun (2003: 70).
Description: Leaf spots subcircular, 1–8 mm diam, brown,
somewhat sunken on the upper surface. Mycelium internal.
Stromata small, only lling the substomatal cavity, brown.
Conidiophores in dense fascicles, arising from stromata,
emerging through stomata, erect, subcylindrical or somewhat
attenuated towards the tip, or only 1–2 times geniculate,
unbranched, 30–105 × 3.5–5 µm, pluriseptate, medium to
dark olivaceous-brown, paler towards the tip; conidiogenous
loci conspicuous, thickened and darkened. Conidia solitary,
acicular, straight to curved, 20–90 × 3–5 µm, distinctly
pluriseptate, hyaline, thin-walled, smooth, apex acute, base
truncate.
Holotype: Philippines: Luzon, Manila, on Diplazium
esculentum (Athyrium esculentum), Woodsiaceae, Mendoza,
No. 55483 (not seen).
Host range and distribution: Only known from the type
collections.
Cercospora camptosori Davis, Trans. Wisconsin
Acad. Sci. 18: 267 (1915).
(Fig. 16)
Literature: Chupp (1954: 455), Crous & Braun (2003: 98).
Description: Leaf spots amphigenous, circular to angular-
irregular, small to large, sometimes covering large
leaf segments, brown, margin indenite. Caespituli
amphigenous, punctiform. Mycelium internal. Stromata
20–50 µm diam, dark brown. Conidiophores in dense
fascicles, arising from stromata, erect, subcylindrical or
somewhat attenuated towards the tip, straight to somewhat
geniculate near the apex, unbranched, 20–60 × 4–5 µm,
septate, pale olivaceous-brown, paler towards the tip, thin-
walled, smooth; conidiogenous cells integrated, terminal,
conidiogenous loci conspicuous, thickened and darkened,
1–2 µm diam. Conidia solitary, narrowly obclavate-liform,
30–125 × 1.5–3.5 µm, indistinctly pluriseptate, hyaline, thin-
walled, smooth, apex acute or subacute, base obconically
truncate, 1–2 µm wide, hila somewhat thickened and
darkened.
Lectotype (designated here, MycoBank, MBT176146):
USA: Wisconsin: Grant County, Marquette State Park, on
Asplenium radicans (Camptosorus rhizophyllus), 1 Aug.
1914, J. J. Davis (WIS). Isolectotype: BPI 433994.
Host range and distribution: On Asplenium radicans,
Aspleniaceae; ?Dryopteris sp., Dryopteridaceae; North
America (USA, ?Florida, Wisconsin).
Notes: This species differs from fern-inhabiting species
belonging to the C. apii complex in having narrowly obclavate-
liform conidia with obconically truncate base and narrow
conidiogenous loci and hila, only 1–2 µm diam.
Cercospora cheilanthis Chowdhry, D. Gupta & Padhi,
Indian Phytopathol. 36: 624 (1983); as “cheilanthei”.
(Fig. 17)
Fig. 16. Cercospora camptosori (BPI 433994). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
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Literature: Chupp & Braun (2003: 118), Kamal (2010: 31).
Illustration: Chowdhry et al. (1983: 625, g. 1).
Description: Leaf spots amphigenous, circular to elongated,
1–3 mm diam, blackish brown, Caespituli amphigenous,
mainly epiphyllous. Mycelium internal. Stromata well-
developed, 15–45 µm diam, globular, dark brown.
Conidiophores in large, dense fascicles, arising from
stromata, about 10–40, erect, straight to somewhat curved,
cylindrical, 1–3 times geniculate above, unbranched, 30–
165 × 3–5.5 µm, 2–6-septate, deep olivaceous-brown;
conidiogenous cells integrated, terminal and intercalary,
conidiogenous loci conspicuous, thickened and darkened.
Conidia solitary, acicular, about 10–130 × 1.5–3 µm,
1–19-septate, hyaline, thin-walled, smooth, apex acute,
base truncate, hila thickened and darkened.
Holotype: India: Odisha (Orissa): Bhubaneswar, on leaves
of Aleuritopteris farinosa (Cheilanthes farinosa), Pteridaceae
subfam. Cheilanthoideae (= Cheilanthaceae), 30 Nov. 1977,
D. Gupta (HCIO 32848).
Host range and distribution: Only known from the type
collection.
Note: This species belongs to Cercospora apii s. lat. (sensu
Crous & Braun 2003).
Cercospora cyclosori Goh & W.H. Hsieh, Trans.
Mycol. Soc. Republ. China 4: 26 (1989).
(Fig. 18)
Synonym: Cercospora cyclosori Sarbajna & Chattopadh, J.
Mycopathol. Res. 28: 14 (1990), nom. illeg. (Art. 53.1)
[type: on Cyclosorus sp., India, West Bengal, Hooghly,
Fig. 17. Cercospora cheilanthis (based on the original illustration). A.
Conidiophore fascicle. B. Conidia. Bar = 10 µm. Fig. 18. Cercospora cyclosori (K(M) IMI 312082). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
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Mankundu, 28 Aug. 1986, K. K. Sarbajna (K(M) IMI
311128)].
Literature: Hsieh & Goh (1990: 327).
Illustration: Hsieh & Goh (1990: 329, g. 249).
Description: Leaf spots irregularly shaped, 1–4 mm diam, often
conuent and larger, dark brown, margin indenite. Caespituli
amphigenous. Mycelium internal. Stromata lacking or small,
only formed as small aggregation of swollen hyphal cells,
substomatal or intraepidermal, dark brown. Conidiophores
solitary or in small, loose fascicles, 2–8, arising from stromata,
erect, straight or almost so, subcylindrical or somewhat
geniculate, above all in the upper half, unbranched, uniform
in width or somewhat attenuated towards the apex, 25–160
× 4–5 µm, 0–4-septate, brown, paler towards the apex;
conidiogenous cells integrated, terminal, conidiogenous loci
conspicuous, about 2–2.5 µm diam, thickened and darkened.
Conidia formed singly, acicular, 50–110 × 3–4 µm, indistinctly
pluriseptate (about 4–10), hyaline, thin-walled, smooth,
apex acute or subacute, base truncate, 2.5–3 µm wide, hila
somewhat thickened and darkened.
Type: Taiwan: Hwalien Hsien, Fengpin, on Cyclosorus
acuminatus, 30 Aug. 1984, T. K. Goh (NCHUPP-37a –
holotype; K(M) IMI 312082 – isotype)..
Host range and distribution: On Cyclosorus (acuminatus,
Cyclosorus sp.), Thelypteridaceae, Asia, (India, Taiwan).
Notes: Type material of C. cyclosori (Sarbajna &
Chattopadhyay 1990) has been examined and proved to be
identical with C. cyclosori Goh & W.H. Hsieh. To-anun et al.
(2011: 75, g. 58) described and illustrated a Cercospora
collection on Pteris biaurita from Thailand and referred it to C.
cyclosori. This identication is, however, doubtful. Cyclosorus
is part of the Thelypteridaceae whereas Pteris belongs in the
Pteridaceae. Several Cercospora spp. have been described
from other hosts of the Pteridaceae. Furthermore, C. cyclosori,
to which this collection was assigned, is morphologically C.
apii-like.
Cercospora diplaziicola A.K. Das, Indian J. Mycol.
Res. 27: 37 (1989).
(Fig. 19)
Literature: Crous & Braun (2003: 163), Kamal (2010: 41),
Phengsintham et al. (2010b, 2013a).
Illustrations: Das (1989: 38, g. 1), Phengsintham et al.
(2013a: 38, gs 20–21).
Description: Leaf spots subcircular to irregular, 1–5 mm diam,
grey-brown to dark brown in the centre, margin indenite to
dark brown. Caespituli amphigenous, scattered, dark brown.
Mycelium internal; hyphae branched, 2–3 µm wide, septate,
constricted at the septa, distance between septa 5–10 µm,
brownish or green-hyaline, wall thin, smooth, forming plate-like
plectenchymatous stromatic hyphal aggregations. Stromata
small to well-developed, oval to ellipsoidal, 15–25 µm diam,
brown, substomatal or intraepidermal, composed of swollen
hyphal cells, subglobose, rounded to angular in outline, 6–17
µm wide, brown to dark brown, wall thin. Conidiophores
formed singly or fasciculate, arising from stromata (2–12 per
fascicle), emerging through stomata, unbranched, straight to
curved, cylindrical, 40–190 × 4–6.5 µm, 2–8-septate, distance
between septa 8–28 µm long, medium brown or olivaceous-
brown, paler at the apex, wall 0.5–0.8 µm wide, smooth,
slightly to strongly geniculate, width ± uniform; conidiogenous
cells integrated, terminal or intercalary, cylindrical, 10–20
× 3–5 µm, pale brown; conidiogenous loci conspicuous,
subcircular, 2–3 µm wide, brown. Conidia solitary, acicular to
narrowly obclavate, straight to curved, 45–155(–230) × 3–5(–
6) µm, 3–20-septate, hyaline, thin-walled, smooth, tip acute,
base truncate to short obconically truncate, 2–3 µm wide, hila
thickened and darkened.
Fig. 19. Cercospora diplaziicola (K(M) IMI 292584). A. Conidiophore
fascicle. B. Conidiogenous cells. C. Conidia. Bar = 10 µm.
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Holotype: India: West Bengal: Jalpaiguri, Baradighi, on
Diplazium esculentum, 10 Oct. 1983, A. K. Das PCC 3669
(K(M) IMI 292584).
Host range and distribution: On Diplazium esculentum,
Woodsiaceae, Asia (India, Laos, Thailand).
Notes: Cercospora diplaziicola with acicular to obclavate
conidia is very close to C. athyrii, which was described to have
acicular conidia. However, type material of the latter species
could not be traced, is probably not maintained and thus
cannot be re-examined to prove the variation of the conidial
shape. It is possible that the two species described from
Diplazium esculentum in Asia are conspecic. Cercospora
diplaziicola is only tentatively maintained as separate species.
Cercospora dryopteridis Y.L. Guo, Mycosystema 16:
1 (1997).
(Fig. 20)
Literature: Crous & Braun (2003: 166), Zhuang (2005).
Illustration: Guo (1997: 2, g. 1).
Description: Leaf spots amphigenous, subcircular to angular-
irregular, 1–3 mm diam, often conuent, greyish white to
pale yellowish brown, with dark brown halo on the upper
side, paler on the lower side. Caespituli mainly hypophyllous.
Mycelium internal. Stromata lacking or only with a few
aggregated swollen hyphal cells. Conidiophores solitary or in
small fascicles, 2–12, divergent, erect, straight, subcylindrical
to 1–4 times geniculate, usually unbranched, 40–150(–210)
× 4–6 µm, 1–5-septate, olivaceous-brown to medium brown,
paler towards the tip; conidiogenous cells integrated, terminal
and intercalary, conidiogenous loci conspicuous, thickened
and darkened, 2.5–3.5 µm diam. Conidia solitary, acicular,
95–250 × 3.5–5 µm, pluriseptate, hyaline, thin-walled,
smooth, apex acute, base truncate, about 3–4 µm wide, hila
somewhat thickened and darkened.
Holotype: China: Prov. Shaanxi: Foping, on Dryopteris
polypodioides, Dryopteridaceae, 27 Nov. 1991, Y. L. Guo
3139 (HMAS 73548).
Host range and distribution: Only known from the type
collection.
Note: This species is part of the Cercospora apii complex.
Cercospora lygodiicola Lall, J.N. Kapoor & Munjal,
Indian Phytopathol. 17: 181 (1964).
Literature: Crous & Braun (2003: 258), Kamal (2010: 61).
Illustration: Lall et al. (1964: 182, g. 1).
Description: Leaf spots amphigenous, irregularly shaped, 2–15
mm diam, fuliginous, margin paler, sometimes covering the
whole surface of leaets. Caespituli amphigenous. Stromata
lacking or small, composed of a few swollen hyphal cells.
Conidiophores solitary or few to many in loose fascicles, arising
from internal hyhae or small stromatic hyphal aggregations, erect,
subcylindrical to somewhat geniculate, unbranched, about 15–
50 × 4–5 µm, continuous to sparingly septate, olivaceous-brown
below, paler towards the apex; conidiogenous cells integrated,
terminal or conidiophores reduced to conidiogenous cells,
conidiogenous loci thickened and darkened. Conidia solitary,
narrowly obclavate, 25–130 × 3–4 µm, indistinctly pluriseptate,
subhyaline, thin-walled, smooth, apex subacute or acute, base
short obconically truncate.
Holotype: India: Punjab: Kulu, Manali, on Lygodium
japonicum, Lygodiaceae, 9 Nov. 1962, V. S. Sharma (HCIO
28172).
Host range and distribution: Only known from the type
collection.
Fig. 20. Cercospora dryopteridis (HMAS 73548). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
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Cercospora marsileae Ragunathan, Prasad &
Purushot., Curr. Sci. 39: 305 (1970).
(Fig. 21)
Literature: Thaung (1984: 441), Crous & Braun (2003: 268),
Kamal (2010: 63).
Illustration: Ragunanthan et al. (1970: 305, gs 1–2).
Description: Leaf spots brown to rusty brown, with chlorotic
halo, 2–14 mm in length, sometimes spreading over
the whole leaet. Caespituli amphigenous. Mycelium
internal. Stromata well-developed, consisting of brown
cells. Conidiophores in fascicles, 2–10, erect, straight,
subcylindrical, unbranched, apical part geniculate, about 50–
110 × 3.5–5 µm, 1–3-septate, brown, paler towards the tip,
thin-walled, smooth; conidiogenous cells integrated, terminal,
conidiogenous loci thickened and darkened. Conidia formed
singly, acicular, about 45–225 × 3–5.5 µm, 4–26-septate,
hyaline, thin-walled, smooth, apex pointed, base truncate.
Holotype: India: Tamil Nadu: Annamalainagar, on living
leaves of Marsilea quadrifolia, 5 Nov. 1965, V. Ragunathan
(HCIO 29330).
Host range and distribution: On Marsilea quadrifolia,
Marsileaceae, Asia (India, Myanmar).
Note: This species belongs to the Cercospora apii complex.
Cercospora platycerii Chupp, A monograph of the
fungus genus Cercospora: 456 (1954).
(Fig. 22)
Fig. 21. Cercospora marsileae (based on the original illustration). A.
Conidiophore fascicle. B. Conidia. Bar = 10 µm. Fig. 22. Cercospora platycerii (CUP, holotype). A. Conidiophore
fascicle. B. Conidiogenous cells. C. Conidia. Bar = 10 µm.
Braun et al.
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300 ima fUNGUS
Literature: Raabe et al. (1981), Crous & Braun (2003: 327),
Phengsintham et al. (2010b), To-anun et al. (2011: 73).
Illustration: To-anun et al. (2011: 74, g. 57).
Description: Leaf spots amphigenous, subcircular, 10–30 mm
diam, dark brown to blackish, zonate, margin slightly raised
and centre depressed or vice versa. Caespituli hypophyllous.
Mycelium internal. Stromata lacking or small, only with small
aggregations of a few swollen hyphal cells, substomatal,
brown. Conidiophores solitary or in small fascicles, 2–9,
divergent, arising from internal hyphae or hyphal aggregations,
through stomata, erect, straight, subcylindrical, geniculate
above, unbranched, 30–200 × 3–5 µm, pluriseptate, pale to
medium olivaceous-brown, wall slightly thickened, smooth;
conidiogenous cells integrated, terminal, conidiogenous
loci conspicuous, thickened and darkened, 2–3.5 µm diam.
Conidia solitary, acicular, straight to somewhat curved, 40–
150 × 2.5–5 µm, pluriseptate, hyaline, thin-walled, smooth,
apex subacute, base truncate, 2–4 µm wide, hila somewhat
thickened and darkened.
Holotype: USA: New Jersey: East Orange, on Platycerium
sp., 21 Sep. 1933, D. L. Gill (CUP).
Host range and distribution: on Platycerium (bifurcacum,
grande, Platycerium sp.), Polypodiaceae, Asia (Thailand),
North America (USA, Florida, New Jersey), Hawaii.
Cercospora pleopeltidis Chidd., Mycopathol. Mycol.
Appl. 17: 73 (1962).
(Fig. 23)
Synonym: Cercospora microsori P. Kumar & Kamal, Curr.
Sci. 50: 509 (1981); as “microsorii” [type: on Microsorum
membranaceum, India, U.P., Gorakhpur, Feb. 1978, P.
Kumar 96 (K(M) IMI 230606b)].
Literature: Crous & Braun (2003: 275, 328), Kamal (2010:
65, 76).
Illustrations: Chiddarwar (1962: pl. I, gs 7–9), Kumar &
Kamal (1981: 509, g. 2).
Description: Leaf spots amphigenous, circular to oval,
conuent, forming irregular patches, pale brown to brown.
Caespituli amphigenous. Mycelium internal. Stromata
substomatal, subglobose, 20–45 µm diam, brown.
Conidiophores 5–16, in loose fascicles, arising from
stromata, through stomata, erect, straight, subcylindrical to
distinctly geniculate-sinuous, unbranched, 20–110(–150) ×
3–4.5(–5) µm, 2–7-septate, pale to brown, wall somewhat
thickened, smooth; conidiogenous cells integrated, terminal
or intercalary, with conspicuous conidiogenous loci, about
2–2.5 µm wide, thickened and darkened. Conidia solitary,
acicular to somewhat obclavate, straight to curved, 15–150
× 3–4 µm, 2–16-septate, hyaline, thin-walled, smooth, apex
acute or subobtuse, base truncate or slightly attenuated at
the very base, 2–2.5 µm wide, hila somewhat thickened and
darkened.
Types: India: Bombay, Mahabaleshwar, on leaves of
Microsorum membranaceum, 14 Oct. 1955, P. P. Chiddarwar
(K (M) IMI 83185 – holotype; BPI 439783 – isotype).
Host range and distribution: on Microsorum membranaceum
(Pleopeltis membranacea), Polypodiaceae, Asia (India, M.S.,
U.P.).
Notes: A morphologically similar Cercospora was found
on Nephrolepis cordifolia (Polypodium cordifolium),
Davalliaceae, in Venezuela (Lara, La Cuchilla, Dec. 2006,
R. Urtiaga, HAL 2601 F, deposited as Cercospora sp.).
The sample concerned is, however, too meager for a nal
determination and description.
Cercospora pteridigena M.K. Khan, R.K. Verma &
Kamal, Indian Phytopathol. 45: 26 (1994).
(Fig. 24)
Fig. 23. Cercospora pleopeltidis (K(M) IMI 83185). A. Conidiophore
fascicle. B. Conidia. Bar = 10 µm.
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Literature: Crous & Braun (2003: 340), Kamal (2010: 78).
Illustration: Khan et al. (1992: 27, g. 1).
Description: Lesions amphigenous, mainly on young leaves
and leaets, 2.5–15 mm diam, scattered, sometimes covering
the whole leaf surface, light yellow to dark brown. Mycelium
internal; hyphae up to 2.5 µm wide, septate, smooth. Stromata
lacking or only poorly developed, small aggregations of swollen
hyphal cells, about 10–20 µm diam, olivaceous. Conidiophores
solitary or in small fascicles, arising from internal hyphae
or small stromata, erect, straight to somewhat geniculate,
unbranched, about 80–205 × 6.5–7 µm, pluriseptate, pale
brown, thin-walled, smooth; conidiogenous cells integrated,
terminal or intercalary, with conspicuously thickened and
darkened conidiogenous loci, 4–5.5 µm diam. Conidia solitary,
acicular, very rarely branched, 145.5–495.5 × 3.5–5 µm,
10–25-septate, hyaline, base truncate, about 3–4 µm wide,
apex pointed, hila thickened and darkened.
Types: India: U.P.: Baharaich, on leaves of Adiantum
venustum, 14 Nov. 1987, M. K. Khan, (GPU, KS 239 –
holotype; K(M) IMI 3212623 – isotype).
Host range and distribution: On Adiantum (philippense
[lunulatum], venustum), Pteridaceae subfam. Vittarioideae
(Adiantaceae), Asia (India).
Note: An additional collection of this species from India on
Adiantum philippense is deposted at K (IMI 136020).
Cercospora selaginellarum P. Joly, Rev. Mycol. 30:
228 “1965” (1966).
Literature: Farr & Horner (1968).
Illustration: Cour & Joly (1965: 227, pl. 1–4, 229, g. 1).
Description: Lesions formed as small leaf spots. Mycelium
internal; hyphae 4–7 µm wide, brown. Stromata composed
of aggregated swollen hyphal cells, 15–40 µm diam, brown.
Conidiophores loosely fasciculate, 2–20, arising from
stromata, erect, unbranched, straight to geniculate-sinuous,
length variable, about 150–400 µm long and 4.5–6 µm wide,
pluriseptate, brown; conidiogenous cells integrated, terminal
and intercalary, conidiogenous loci conspicuous, thickened
and darkened. Conidia solitary, acicular, up to 300 µm
long, pluriseptate, hyaline, thin-walled, apex pointed, base
truncate, hila thickened and darkened.
Syntypes: France: on leaves of Selaginella denticulata and
S. helvetica (details not indicated).
Host range and distribution: on leaves of Selaginella (apoda,
helvetica, kraussiana [denticulata]), Selaginellaceae, Europe
(France), North America (USA, Minnisota).
Notes: This species belongs to the C. apii complex. Cour
& Joly (1965) compared C. selaginellarum with the latter
species and discussed the morphological similarity of the
two species. Type material was not indicated in the original
publication, and two different host species were cited,
Selaginella helvetica and S. denticulata. Original material
could not be traced. Jaap, Fungi Sel. Exs. 619, issued as
Leptosphaeria helvetica var. major, on Selaginella helvetica,
Italy, Liguria, Portono, 26 Mar. 1913, O. Jaap (material
deposited at HAL examined) contains conidiophores and
conidia of Cercospora sp., but the latter material does not
belong to C. selaginellarum. The conidia are much shorter,
narrower and obclavate, and the conidiogenous loci are
rather small, 1.5–2 µm diam (stromata immersed, about
10–40 µm diam, brown; conidiophores fasciculate, straight,
subcylindrical, 80–150 × 3–6 µm, pluriseptate, medium to
darker brown, tips much paler, conidiogenous loci 1.5–2
µm wide, slightly thickened and darkened; conidia solitary,
obclavate, about 40–90 × 4–5 µm, 3–8-septate, hyaline or
almost so, smooth, apex subobtuse, base rounded to short
obconically truncate, 2 µm wide, hila slighty thickened and
darkened). However, the material from Italy is not sufcient
for a nal taxonomic conclusion.
Fig. 24. Cercospora pteridigena (K(M) IMI 321263). A. Conidiophore
arising from a stroma. B. Conidiogenous cells. C. Conidia. Bar = 10 µm.
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Passalora
Key to the species of Passalora on ferns
1 Conidiophores 30–80 × 3–5 µm; conidia obclavate, long and narrow, 40–210 × 2–4 µm,
subhyaline to pale olivaceous; on Pteridium aquilinum, widespread ........................................................ P. pteridis
Conidiophores longer and broader, about 25–350 × 4–7.5 µm; conidia obclavate-cylindrical,
shorter and broader, 25–100 × 4–6 µm, olivaceous-brown or brown; on Pyrrosia lingua, Japan
................................................................................................................................................................. P. pyrrosiae
List of Passalora species on ferns
Passalora pteridis (Siemaszko) U. Braun & Crous,
in Crous & Braun, Mycosphaerella and Anam. 1: 340
(2003).
(Fig. 25)
Basionym: Cercospora pteridis Siemaszko, Arch. Nauk. Biol.
Towarz. Nauk. Warszaw. 1: 57 (1923).
Synonyms: Pseudocercospora pteridis (Siemaszko) Y.L. Guo
& X.J. Liu, Acta Mycol. Sin. 11: 298 (1992) [misapplied].
Mycovellosiella pteridis (Siemaszko) U. Braun, Trudy Bot.
Inst. Im. V.L. Komarova 20: 83 (1997).
Cercospora pteridicola Gonz.Frag., Mem. Real. Acad.
Ci. Exact. Madrid, Ser. 2, 6: 227 (1927) [holotype: on
Pteridium aquilinum, Spain, near San Filipe de Ballaróls,
Fuente de la Salud, Gerona, 16 Nov. 1920, Sennen (MA)].
Literature: Chupp (1954: 457), Vassiljevsky & Karakulin
(1937: 219), Lall et al. (1964: 182–183), Greene (1960),
Saccardo (1972: 1382), Crous & Braun (2003: 340), Kamal
(2010: 133).
Illustrations: Lall et al. (1964: 182, g. 3), Braun & Mel’nik
(1997: 121, g. 49).
Description: Leaf spots amphigenous, formed as irregular
discolorations, yellowish brown, brown, later darker, margin
indenite. Caespituli hypophyllous, subeffuse, not very
conspicuous. Mycelium internal and external; supercial
hyphae branched, septate, 2–6 µm wide, smooth, thin-walled,
subhyaline to olivaceous. Stromata lacking or only formed as
small substomatal hyphal aggregations. Conidiophores in
small, loose fascicles, arising from internal hyphae or hyphal
aggregations, or solitary, arising from supercial hyphae, erect
to decumbent, exuous, simple or occasionally branched,
subcylindrical to somewhat clavate, to geniculate-sinuous,
20–100 × 5–10 µm, continuous to pluriseptate, olivaceous to
brownish, thin-walled, smooth; conidiogenous cells integrated,
terminal, often somewhat swollen, subclavate, about 20–40 µm
long, conidiogenous loci conspicuous, somewhat thickened and
darkened. Conidia solitary, broadly obclavate-subcylindrical,
(30–)40–120(–140) × 5–8 µm, pluriseptate, hyaline, subhyaline
to pale olivaceous, thin-walled, smooth, apex obtuse, base
obconically truncate, hila slightly thickened and darkened.
Holotype: Russia: Pskhu, Caucasus Mts., on Pteridium
aquilinum, 1917, W. Siemaszko (not seen).
Host range and distribution: On Pteridium (aquilinum,
Pteridium sp.), Dennstaedtiaceae, Asia (India, Japan),
Caucasus (Georgia), Europe (Russia, Spain), North America
(USA, Wisconsin).
Notes: Type material of C. pteridis could not be traced in
Siemaszko’s herbarium at WA. The reallocation of this
species to Mycovellosiella and Passalora, respectively, was
based on the examination of type material of Cercospora
pteridicola. The identity of a Japanese collection on
Pteridium aquilinum subsp. japonicum (P. aquilinum var.
latiusculum), Iwate, 31 Aug. 1947, K. Sawada (IUM-FS79),
recorded by Sawada (1958) as Cercospora pteridis, is
unclear and remains unproven as this sample is devoid of
any fructication, which has been conrmed by a recently
made re-examination. Records of this species from India and
Myanmar on Pteridium sp. (Lall et al. 1964, Thaung 1984)
are also unclear and unproven. Material was not available
for a re-examination. A previous record from Colombia on
Fig. 25. Passalora pteridis (MA, holotype of Cercospora pteridicola).
A. Supercial hypha. B. Solitary conidiophore arising from a
supercial hypha. C. Conidiophore fascicle. D. Conidiogenous cells.
E. Conidia. Bar = 10 µm.
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Pteris sp. (Chupp 1954) was based on a misidentication and
belongs to a Pseudocercospora described in this paper as P.
pteridigena.
Passalora pyrrosiae (Togashi & Katsuki) C. Nakash.
& I. Araki, Mycoscience 49: 144 (2008).
(Fig. 26)
Basionym: Pseudocercospora pyrrosiae Togashi & Katsuki,
J. Jap. Bot. 28: 287 (1953).
Literature: Katsuki (1965: 76).
Illustration: Motohashi et al. (2008: 143, g. 6).
Description: Lesions indistinct or with distinct discolorations,
3–8 mm diam, greyish brown to reddish brown above,
with darker border line and greyish brown or brown
below, sometimes covering the whole surface. Caespituli
hypophyllous, fuliginous. Mycelium internal. Stromata
intraepidermal, small to large, up to 135 µm diam, olivaceous-
brown to dark brown. Conidiophores solitary or in moderately
large fascicles, 2–18, loose to dense, arising from stromata,
erumpent, erect, straight to slightly curved, subcylindrical
to geniculate-sinuous, unbranched, about 25–350 × 4–7.5
µm, 4–10-septate, pale olivaceous-brown to brown, thick-
walled, asperulate; conidiogenous cells integrated, terminal,
sympodially or percurrently proliferating, conidiogenous loci
thickened and darkened, 2.5–5 µm diam. Conidia solitary,
obclavate-cylindrical, straight to curved, 25–100 × 4–6 µm,
1–11-euseptate, rarely distoseptate, occasionally constricted
at the septa, olivaceous-brown or brown, wall thickened,
asperulate, base obconically truncate, apex obtuse, hila
somewhat thickened and darkened.
Types: Japan: Pref. Kagoshima, Kosugidani, Yaku I., on
Pyrrosia lingua, 7 Aug. 1951, K. Togashi & S. Katsuki
(holotype not preserved); neotype (designated in Motohashi
et al. 2008), Aichi Pref., Nagoya, Chikusa, Higashiyama
Botanical Garden, on Pyrrosia lingua, 6 June 2005, I. Araki
(TFM : FPH-7852, ex-neotype culture: MAFF 240280).
Host range and distribution: On Pyrrosia lingua,
Polypodiaceae; Asia (Japan).
Fig. 26. Passalora pyrrosiae (TFM : FPH-7852). a. Drawing (A. Conidiophore fascile, B. Conidia). b. Micrograph of conidiophores (SEM).
c. Micrograph of conidiophores (light microscopy).
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Pseudocercospora
Key to the species of Pseudocercospora on ferns
1 Mycelium internal and external; supercial hyphae usually with solitary conidiophores ................................................. 2
Mycelium internal; supercial hyphae with solitary conidiophores lacking .................................................................... 14
2 (1) Stromata lacking; conidiophores only solitary, arising from supercial hyphae .............................................................. 3
Stromata mostly present, small to large; conidiophores fasciculate, arising from internal hyphae or
stromata as well as solitary, arising from supercial hyphae ................................................................................... 6
3 (2) Conidiophores very long, 100–300 µm, pluriseptate (10–30 septa); conidia subcylindrical,
ellipsoid to short obclavate-obovoid, 12–25 × 4.5–6 µm, (2–)3–4(–5)-septate;
on Dicranopteris linearis, Africa, Gabon .............................................................................................. P. gleicheniae
Conidiophores much shorter, 2–40 µm, 0–2-septate; conidia acicular-liform or narrowly
obclavate-cylindrical, much longer and narrower; on other hosts ............................................................................ 4
4 (3) Conidia narrowly cylindrical to obclavate-cylindrical, 30–80 × 2–3 µm; on Abacopteris urophylla,
Singapore ........................................................................................................................................ P. abacopteridis
Conidia acicular-liform, 60–120 × 2–3.5 µm; on other hosts ......................................................................................... 5
5 (4) Conidia 3–3.5 µm wide, hila 2.5–3 µm wide; on Rumohra adiantiformis, Taiwan ......................................... P. rumohrae
Conidia narrower, 2–3 µm, hila 1.5–2 µm wide; on Ptisana salicina, New Zealand ........................................ P. ptisanae
6 (2) Stromata lacking; forming fasciculate conidiophores arising from internal hyphae, emerging through stomata;
conidia short, only 10–35 × 1.5–4 µm, 1–3-septate; on Lygodium japonicum, Asia ................................... P. lygonii
Stromata developed, giving rise to fasciculate conidiophores; conidia longer, pluriseptate ........................................... 7
7 (6) Stromata very large, up to 150 µm diam; on Nephrolepis falcata, Australia ..................................... P. nephrolepidicola
Stromata much smaller, 10–80 µm diam ........................................................................................................................ 8
8 (7) Supercial hyphae present, but without solitary conidiophores; on Davallia fejeensis, Brazil ................... P. davalliicola
Supercial hyphae with solitary conidiophores present and usually abundant ............................................................... 9
9 (8) Conidia narrow, only 2–4 µm wide; on Lygodium, Nephrolepis or Pteris spp. .............................................................. 10
Conidia wider, 3–5.5 µm; on other hosts ....................................................................................................................... 13
10 (9) Conidiophores very long, 30–150 µm; conidia acicular to narrowly obclavate, very long, 40–250 µm;
on Pteris sp., Colombia ....................................................................................................................... P. pteridigena
Conidiophores much shorter, 5–70 µm; conidia much shorter, up to 125 µm ................................................................ 11
11 (10) Conidia acicular to narrowly cylindrical, base truncate; on Nephrolepis spp., Asia ......................... P. nephrolepidigena
Conidia obclavate-cylindrical, base obconically truncate; on Lygodium or Pteris spp. ................................................. 12
12 (11) Hyphae subhyaline; conidiophores thin-walled; on Pteris spp., Asia .......................................................... P. pteridicola
Hyphae subhyaline, but distinctly pigmented where conidiophores arise; wall of longer
conidiophores somewhat thickened; on Lygodium sp., Asia .............................................................. P. lygodiigena
13 (9) Leaf spots lacking or indistinct; colonies hypophyllous; stromata small, 10–20 µm diam;
conidiophores up to 150 × 7 µm; on Adiantum, Asia, South America ......................................................... P. adianti
Leaf spots developed; colonies amphigenous; with large epiphyllous stromata, 20–60 µm diam;
conidiophores up to 100 × 5 µm; on Lonchitis hirsuta, Venezuela ....................................................... P. lonchitidis
14 (1) Stromata lacking or almost so ....................................................................................................................................... 15
Stromata developed, 10–110 µm diam ......................................................................................................................... 16
15 (14) Conidiophores 60–180 × 4–7 µm; conidia 3–5 µm wide; on Plagiogyria euphlebia, Taiwan
............................................................................................................................................................ P. plagiogyriae
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Conidiophores much shorter and narrower, 7–35 × 2.5–3 µm; conidia only 2–3 µm wide;
on Nephrolepis auriculata, Taiwan .................................................................................................. P. nephrolepidis
16 (14) Stromata epiphyllous, immersed, very large, 60–110 µm diam; on Cyathea sp., Japan ............................... P. cyatheae
Stromata much smaller, 10–75 µm diam ...................................................................................................................... 17
17 (16) Conidia acicular-liform .................................................................................................................................................. 18
Conidia obclavate-cylindrical ......................................................................................................................................... 22
18 (17) Conidia very narrow, 1–1.5 µm wide; on Cyclosorus acuminatus, Taiwan ...................................... P. pteridophytophila
Conidia wider, 1.5–4 µm; on other hosts ....................................................................................................................... 19
19 (18) Conidiophores relatively short, 5–30 µm; on Arachniodes or Microsorum spp. ............................................................ 20
Conidiophores longer, up to 70 µm; on other hosts ...................................................................................................... 21
20 (19) Colonies hypophyllous; conidiophores usually aseptate; on Arachniodes sp., China .............................. P. arachniodis
Colonies amphigenous; conidiophores 1–5-septate; on Microsorum ........................................................... P. microsori
21 (19) Conidiophores usually aseptate; conidia 50–120 µm long; on Metathelypteris laxa, Taiwan .................. P. thelypteridis
Conidiophores 0–2-septate; conidia 60–80 µm long; on Deparia japonica, Taiwan ........................................... P. athyrii
22 (17) Conidiophores very short, 9–14 × 2–4 µm; on Christella parasitica, Thailand ............................................. P. christellae
Conidiophores longer, (5–)10–280 µm; on other hosts ................................................................................................. 23
23 (22) Conidiophores very long, 160–280 µm; on Lygodium sp., India ...................................................... P. polypodiacearum
Conidiophores shorter, 5–150 µm; on other hosts ........................................................................................................ 24
24 (23) Conidiophores long, up to 150 µm; on Angiopteris or Pityrogramma ........................................................................... 25
Conidiophores shorter, 10–75 µm; on other hosts ........................................................................................................ 26
25 (24) Conidiophores 2–6 µm wide; conidia 30–60 × 3–5 µm; on Angiopteris sp., Asia ................................... P. angiopteridis
Conidiophores 5–7 µm wide; condia 25–200 × 5–7.5 µm; on Pityrogramma, Ecuador
.............................................. (?) Cercospora trismeriae (see “Doubtful, excluded and insufciently known species”)
26 (24) Supercial hyphae present, but without solitary conidiophores; on Davallia fejeensis, Brazil ................... P. davalliicola
Superal hyphae lacking ............................................................................................................................................... 27
27 (26) Conidia narrow, 2–4 µm ................................................................................................................................................ 28
Conidia wider, 3–6 µm .................................................................................................................................................. 30
28 (27) Conidia 20–80 µm long, only 3–5-septate; on diverse ferns of different families;
North and South America and Europe (introduced) ............................................................................... P. phyllitidis
Conidia 20.5–110 µm long, 2–12-septate ..................................................................................................................... 29
29 (28) Conidia never acicular, base short obconically truncate; on Odontosoria chinensis, India ............................ P. davalliae
Conidia acicular to somewhat obclavate-cylindrical, base mostly truncate or only slightly obconically
truncate; on Microsorum pustulatum, Australia ...................................................................................... P. microsori
30(27) Stromata 20–75 µm diam; conidiophores 10–70 µm long, 0–3-septate; conidia 4–7 µm wide;
on Tectaria zeylanica, Asia ................................................................................................... P. helminthostachydis
Stromata smaller, 10–40 µm diam; conidiophores 15–40 µm long, 0–1-septate; conidia 3–5 µm wide;
on Lygodium japonicum, China ............................................................................................................ P. lygodiicola
Tabular key to Pseudocercospora species on ferns
Adiantaceae (Pteridaceae subfam. Vittarioideae)
Athyriaceae
On Deparia japonica (Athyrium japonicum), Taiwan ........................................................................................................ P. athyrii
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Cyatheaceae
1 Distinct leaf spots formed; colonies epiphyllous; stromata very large, 60–110 µm diam;
conidiophores 25–50 × 3–5 µm; conidia cylindrical-obclavate, 30–50 × 3.5–5.5 µm,
base rounded to distinctly obconically truncate; on Cyathea sp., Japan ................................................ P. cyatheae
On dead fronds; colonies amphigenous; stromata smaller, up to 60 µm diam; conidiophores
longer and narrower, 30–70 × 2–3 µm; conidia subcylindrical or subacicular, width often irregular,
base truncate to somewhat obconically truncate; on Cyathea australis, Australia ............................. P. cyatheicola
Davalliaceae
1 Supercial hyphae with solitary conidiophores present .................................................................................................. 2
Supercial hyphae lacking, if present without solitary conidiophores ............................................................................. 3
2 (1) Stromata large, up to 130 µm diam; on Nephrolepis falcata, Australia ............................................. P. nephrolepidicola
Stromata smaller, 10–80 µm diam; on Nephrolepis biserrata and Nephrolepis sp., Brunei, Thailand
.................................................................................................................................................. P. nephrolepidigena
3 (1) Stromata lacking or small, up to 35 µm diam; conidia subcylindrical-acicular, usually 65–100 µm long,
base truncate; on Nephrolepis auriculata, Taiwan .......................................................................... P. nephrolepidis
Stromata larger, up to 75 µm diam; conidia obclavate-subcylindrical, 12–80 µm long,
base obconically truncate ........................................................................................................................................ 4
4 (3) Mycelium internal and external, but supercial hyphae without solitary conidiophores;
conidia 3–5 µm wide, hila (1.5–)2(– 2.5) µm diam; on leaves of Davallia fejeensis, Brazil ................ P. davalliicola
Mycelium only internal; conidia 2–3.5 µm wide, hila 1–1.5 µm diam; on Davallia trichomanoides and
Nephrolepis exaltata, North America ..................................................................................................... P. phyllitidis
Dennstadtiaceae
On Lonchitis hirsuta, Venezuela .............................................................................................................................. P. lonchitidis
Dryopteridaceae
1 Stromata lacking; with solitary conidiophores arising from supercial hyphae; conidia acicular,
60–120 µm long; on Rumohra adiantiformis, Taiwan ............................................................................. P. rumohrae
Stromata small to well-developed; solitary conidiophores lacking .................................................................................. 2
2 (1) Conidiophores 6.5–30 µm long; fully developed conidia acicular, short conidia sometimes obclavate;
on Arachniodes sp., China ................................................................................................................. P. arachniodis
Conidiophores 10–75 µm long; conidia always obclavate; on Rumohra adiantiformis, North America ........ P. phyllitidis
Gleicheniaceae
On Dicranopteris linearis, Africa, Gabon .............................................................................................................. P. gleichneniae
Lindsaeaceae
On leaves of Odontosoria chinensis [Davallia tenuifolia], India .................................................................................. P. davalliae
Lygodiaceae
1 Supercial hyphae present .............................................................................................................................................. 2
Mycelium only internal, supercial hyphae lacking ......................................................................................................... 3
2 (1) Stromata lacking; conidia small, 10–35 × 1.5–4 µm, 1–3-septate; on Lygodium spp. ....................................... P. lygodii
Stromata present; conidia 40–115.5 µm long, pluriseptate; on Lygodium sp. ........................................... P. lygodiigena
3 (1) Conidiophores short, 15–40 µm, 0–1-septate; on leaves of Lygodium japonicum ..................................... P. lygodiicola
Conidiophores much longer, 160–280 µm, pluriseptate; on Lygodium sp., India ............................ P. polypodiacearum
Marattiaceae
1 Supercial mycelium with solitary conidiophores developed; stromata lacking;
conidia acicular-liform; on Ptisana salicina, New Zealand ..................................................................... P. ptisanae
Supercial mycelium with solitary conidiophores lacking; stromata well-developed; conidia obclavate-cylindrical ........ 2
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2 (1) Conidia 3–5 µm wide, hila (1.5–)2(–2.5) µm wide; on Angiopteris spp., Asia ......................................... P. angiopteridis
Conidia narrower, 2–3.5 µm, hila 1–1.5 µm wide; on Angiopteris sp., North America .................................. P. phyllitidis
Osmundaceae
On Osmunda regalis, North America ......................................................................................................................... P. phyllitidis
Plagiogyriaceae
On Plagiogyria euphlebia, Taiwan ........................................................................................................................ P. plagiogyriae
Polypodiaceae
1 Conidia distinctly obclavate, base distinctly obconically truncate, 20–80 × 2–3.5 µm, 3–5-septate;
on Campyloneurum phyllitidis and Polypodium polypodioides, North America ..................................... P. phyllitidis
Conidia acicular to somewhat obclavate-cylindrical, base usually truncate or only slightly
obconically truncate, longer, 50–110 µm, with 2–12 septa; on Microsorum pustulatum, Australia ........ P. microsori
Pteridaceae subfam. Pteridoideae
1 Mycelium internal; supercial hyphae with solitary conidiophores lacking; conidiophores only in fascicles ................... 2
Mycelium internal and external; supercial hyphae with solitary conidiophores present ................................................ 3
2 (1) Conidiophores up to 150 × 7 µm; conidia up to 200 µm long and 5–7.5 µm wide; on Pityrogramma, Ecuador
.............................................. (?) Cercospora trismeriae (see “Doubtful, excluded and insufciently known species”)
Conidiophores up to 100 × 3–5 µm; conidia 20–80 × 2–5 µm; on Pteris spp., North America ..................... P. phyllitidis
3 Conidiophores 10–70 µm long, pale olivaceous, olivaceous-brown to medium brown, thin-walled;
conidia 40–125 µm long; on Pteris spp., Asia ...................................................................................... P. pteridicola
Conidiophores longer, 30–150 µm, medium dark brown, wall somewhat thickened; conidia longer, 40–250 µm;
on Pteris sp., Colombia ....................................................................................................................... P. pteridigena
Pteridaceae subfam. Vittarioideae (= Adiantaceae)
On Adiantum spp., Asia (India), South America (Venezuela) ......................................................................................... P. adianti
Tectariaceae
On Tectaria zeylanica (Helminthostachys zeylanica), Asia ...................................................................... P. helminthostachydis
Thelypteridaceae
1 Stroma lacking; supercial hyphae developed, giving rise to solitary conidiophores, fasciculate
conidiophores not formed; on Abacopteris urophylla, Singapore .............................................. P. abacopteridicola
Stroma developed; supercial hyphae and solitary conidiophores lacking; conidiophores fasciculate .......................... 2
2 (1) Conidia acicular-liform, base ± truncate ........................................................................................................................ 3
Conidia obclavate, base distinctly obconically truncate .................................................................................................. 4
3 (2) Conidia narrow, 30–70 × 1–1.5 µm, hyaline; on Cyclosorus acuminatus, Taiwan .......................... P. pteridophytophila
Conidia longer and wider, 50–120 × 2.5–3 µm, subhyaline to pale olivaceous;
on Metathelypteris laxa (Thelypteris laxa), Taiwan ........................................................................... P. thelypteridis
4 (2) Caespituli epiphyllous; conidiophores very short, 9–15 µm, 0–1-septate; conidia long, 50–105 µm;
on leaves of Christella parasitica, Thailand .......................................................................................... P. christellae
Caespituli hypophyllous; conidiophores longer, 10–75 µm, aseptate or with several septa;
conidia shorter, 20–80 µm; on Thelypteris tetragona, North America .................................................... P. phyllitidis
List of Pseudocercospora species on ferns
Pseudocercospora abacopteridicola (J.M. Yen &
Lim) J.M. Yen, in Yen & Lim, Gard. Bull., Singapore 33:
168 (1980).
(Fig. 27)
Basionym: Cercospora abacopteridicola J.M. Yen & Lim,
Cah. Pacique 17: 97 (1973).
Literature: Crous & Braun (2003: 37).
Illustration: Yen & Lim (1980: 204, g. 7 A–B).
Description: Leaf spots amphigenous, vein-limited, brown,
later conuent. Caespituli hypophyllous, vein-limited, brown,
later conuent. Mycelium internal and external, supercial
hyphae branched, septate, pale olivaceous-brown, 1.5–
2.5 µm wide, thin-walled, smooth. Stromata lacking.
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Conidiophores solitary, arising from supercial hyphae,
lateral, occasionally terminal, erect, straight, subcylindrical or
slightly attenuated towards the tip, unbranched, short, about
5–15 × 2.5–3 µm, 0–1-septate, concolorous with the hyphae,
thin-walled, smooth; conidiophores reduced to conidiogenous
cells, conidiogenous loci inconspicuous, neither thickened
nor darkened. Conidia solitary, narrowly cylindrical to
obclavate-cylindrical, 30–80 × 2–3 µm, 1–7(–9)-septate,
pale olivaceous-brown, thin-walled, smooth, tips obtuse
or subobtuse, base truncate, hilum neither thickened nor
darkened.
Holotype: Singapore: on Abacopteris urophylla, Thelypteri-
daceae, 3 Mar. 1972, G. Lim (PC).
Host range and distribution: Only known from the type
collection.
Pseudocercospora adianti (Syd.) Deighton, Mycol.
Pap. 140: 135 (1976).
(Fig. 28)
Basionym: Cercospora adianti Syd., Ann. Mycol. 28: 206
(1930).
Literature: Chupp (1954: 453), Das (1989), Crous & Braun
(2003: 44), Kamal (2010: 145).
Exsiccatae: Syd., Fungi Exot. Exs. 884.
Description: Leaf spots lacking or indistinct, diffuse brown
discolorations on the upper side. Colonies hypophyllous,
effuse, often covering the whole lower surface, dark olivaceous
to blackish. Mycelium internal and external, supercial
hyphae branched, septate, 1–3.5 µm wide, subhyaline to
pale olivaceous-brown, thin-walled, smooth. Stromata lacking
or only formed as small aggregations of swollen hyphal
cells, about 10–20 µm diam, substomatal to intraepidermal.
Conidiophores in small to moderately large fascicles, 2–20,
loose to rather dense, erect to decumbent, arising from internal
hyphae or small stromata, through stomata or erumpent, or
conidiophores solitary, arising from supercial hyphae, lateral,
straight, subcylindrical to usually distinctly to strongly geniculate-
Fig. 27. Pseudocercospora abacopteridicola (PC, holotype). A.
Solitary conidiophores arising from supercial hyphae emerging
through a stoma. B. Conidia. Bar = 10 µm.
Fig. 28. Pseudocercospora adianti (HBG, lectotype). A. Supercial
hyphae. B. Solitary conidiophore arising from a supercial hypha.
C. Conidiophore fascicle. D. Conidiogenous cells. E. Conidia. Bar
= 10 µm.
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sinuous, sometimes tortuous, usually unbranched, 20–150
× 4–7 µm, continuous to pluriseptate, mostly 2–8-septate,
pale to medium brown, wall thin to slightly thickened, smooth;
conidiogenous cells integrated, terminal to intercalary, 10–
35 µm long, conidiogenous loci inconspicuous to distinctly
denticle-like, but wall always unthickened and not darkened.
Conidia solitary, obclavate to cylindrical-obclavate, occasionally
broadly subacicular, straight to curved, occasionally sinuous-
sigmoid, 40–120 × 4–5.5 µm, 5–15-septate, subhyaline to pale
olivaceous, thin-walled, smooth, apex obtuse to subacute, base
short obconically truncate, about 2–2.5 µm wide, hila neither
thickened nor darkened.
Lectotype (designated here, MycoBank, MBT176147):
Venezuela: El Limon, near Puerto La Cruz, on Adiantum
tetraphyllum, 15 Jan. 1928, H. Sydow, Syd., Fungi Exot. Exs.
884 (HBG). Isolectotypes: Syd., Fungi Exot. Exs. 884 (e.g. B,
BPI 432406, CUP, MICH 15252, S, UPS, etc.).
Host range and distribution: On Adiantum (latifolium,
lunulatum, tetraphyllum, Adiantum sp.), Pteridaceae subfam.
Vittarioideae (= Adiantaceae), Asia (India, Maharashtra, West
Bengal), South America (Venezuela).
Pseudocercospora angiopteridis Goh & W.H. Hsieh,
Trans. Mycol. Soc. Republ. China 4(2): 27 (1989).
(Fig. 29)
Synonym: Cercospora angiopteridis Boedijn, Nova Hedwigia
3: 424 (1961) [types: on Angiopteris evecta, Indonesia,
Java, Bogor, Hortus Bogoriensis, 27 July 1953, Boedijn (L
53845 – holotype; K(M) IMI 91584 – isotype)].
Literature: Hsieh & Goh (1990: 222), Guo & Hsieh (1995: 15),
Guo et al. (1998: 26–27), Braun (2001: 420), Crous & Braun
(2003: 57).
Illustration: Boedijn (1961: Tab. 109), Hsieh & Goh (1990:
223, g. 172), Guo & Hsieh (1995: 16, g. 15), Guo et al.
(1998: 27, g. 15).
Description: Leaf spots amphigenous, subcircular to angular-
irregular, often conuent, forming larger patches, up to 10
mm diam, brown, margin indenite. Caespituli epiphyllous,
punctiform, dark brown to blackish. Mycelium internal.
Stromata immersed, well-developed, 20–70 µm diam, dark
brown. Conidiophores in larger fascicles, loose to dense, erect,
straight, subcylindrical to sinuous or somewhat geniculate,
unbranched, 10–140 × 2–6 µm, continuous to pluriseptate,
brown throughout or paler towards the tip, thin-walled, smooth;
conidiogenous cells integrated, terminal, about 10–30 µm
long, conidiogenous loci inconspicuous, neither thickened nor
darkened. Conidia solitary, obclavate-cylindrical, straight to
somewhat curved, 30–60 × 3–5 µm, 3–6-septate, subhyaline
to pale olivaceous or olivaceous-brown, thin-walled, smooth,
apex obtuse to subacute, base obconically truncate, (1.5–)2(–
2.5) µm wide, hila neither thickened nor darkened.
Types: Taiwan: Hwalien Hsieh, Taluke, on Angiopteris
lygodiifolia, 1 Feb. 1985, T. K. Goh (NCHUPP-148a –
holotype; K(M) IMI 312070 – isotype).
Host range and distribution: On Angiopteris (evecta,
lygodiifolia), Marattiaceae, Asia (Indonesia, Taiwan).
Notes: Braun (2001) re-examined type material of Cercospora
angiopteridis and reduced this species to synonym with
Pseudocercospora angiopteridis.
Pseudocercospora arachniodis Y.L. Guo,
Mycosystema 6: 103 (1993).
(Fig. 30)
Literature: Guo & Hsieh (1995: 89), Guo et al. (1998: 103–
104).
Illustrations: Guo (1993: 104, g. 1), Guo & Hsieh (1995: 88,
g. 80), Guo et al. (1998: 104, g. 84).
Description: Leaf spots amphigenous, at rst elliptical to
angular, 0.5–1 mm diam, later oblong, 1–4 × 0.5–1 mm, often
Fig. 29. Pseudocercospora angiopteridis (L 53845). A. Conidiophore
fascicles. B. Conidia. Bar = 10 µm.
Braun et al.
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310 ima fUNGUS
conuent, reddish brown, with dark brown halo on the upper
side, paler brown below. Caespituli hypophyllous. Mycelium
internal. Stromata substomatal, small, composed of a few
swollen, olivaceous-brown cells or globose, 10–30 µm diam.
Conidiophores in small to moderately large fascicles, usually
dense, arising from stromata, emerging throught stomata,
straight to curved, subcylindrical, rarely geniculate, unbranched,
6.5–30 × 2.5–4 µm, 0(–1)-septate, uniformly pale olivaceous
to olivaceous, thin-walled, smooth, apex conic to conically
truncate; conidiophores usually reduced to conidiogenous
cells, conidiogenous loci inconspicuous, unthickened and
not darkened, sometimes visible as truncate tip. Conidia
solitary, acicular, shorter conidia obclavate, 30–110 × 1.5–3
µm, 3–11-septate, pale olivaceous, thin-walled, smooth, apex
obtuse to subacute, base truncate to short obconically truncate,
about 1–2 µm wide, hila unthickened, not darkened.
Holotype: China: Shaanxi Province: Xian, on leaves of
Arachniodes sp., Dryopteridaceae, 17 Jul. 1973, J. Y. Li & T.
Y. Zhang 5046 (HMAS 62750).
Host range and distribution: Only known from the type
collection.
Pseudocercospora athyrii Goh & W.H. Hsieh, Trans.
Mycol. Soc. Republ. China 4(2): 27 (1989).
(Fig. 31)
Literature: Hsieh & Goh (1990: 38), Guo & Hsieh (1995: 32),
Guo et al. (1998: 45).
Illustrations: Hsieh & Goh (1990: 38, g. 23), Guo & Hsieh
(1995: 34, g. 32), Guo et al. (1998: 45, g. 32).
Description: Leaf spots amphigenous, angular-irregular,
up to 3 mm diam, brown, margin indenite. Caespituli
Fig. 30. Pseudocercospora arachniodis (HMAS 62750). A. Conidio-
phore fascicles. B. Conidia. Bar = 10 µm.
Fig. 31. Pseudocercospora athyrii (NCHUPP-228). A. Conidiophore
fascicle. B. Conidiogenous cells. C. Conidia. Bar = 10 µm.
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epiphyllous, punctiform, dark brown to blackish. Mycelium
internal. Stromata epiphyllous, immersed to somewhat
erumpent, subcircular to irregularly shaped, 20–50 µm
diam, dark brown. Conidiophores numerous, in dense
fascicles arising from stromata, erect, subcylindrical-liform,
geniculate-sinuous, usually simple, rarely branched, 10–70
× 2–3 µm, 0–2-septate, subhyaline to pale olivaceous, thin-
walled, smooth; conidiogenous cells integrated, terminal or
conidiophores reduced to conidiogenous cells, 10–25 µm
long, conidiogenous loci inconspicuous, neither thickened nor
darkened. Conidia solitary, acicular, almost straight to usually
curved-sinuous, 60–80 × 2–3 µm, indistinctly pluriseptate,
subhyaline, thin-walled, smooth, apex acute, base truncate,
2–2.5 µm wide, hila neither thickened nor darkened.
Holotype: Taiwan: Taipei, Yangmingshan, on Deparia
japonica (Athyrium japonicum), Athyriaceae, 30 Aug. 1986,
T. K. Goh (NCHUPP-228).
Host range and distribution: Only known from the type
collection.
Pseudocercospora christellae Phengsintham,
McKenzie, K.D. Hyde & U. Braun, Mycosphere 1: 207
(2010).
(Fig. 32)
Illustrations: Phengsintham et al. (2010a: 208–209, gs 6–7;
2013a: 61–62, gs 60–61).
Literature: Phengsintham et al. (2013a: 60–61).
Description: Leaf spots subcircular to irregular, 3–9 mm diam,
at rst brownish, later becoming brown to dark yellowish brown
at the margin. Caespituli epiphyllous, conspicuous. Mycelium
internal; hyphae branched, 2–3 µm wide, septate, constricted
at the septa, distance between septa 6–8 µm, subhyaline to
brownish, wall 0.3–0.5 µm wide, smooth, forming plate-like
plectenchymatous stromatic hyphal aggregations. Stromata
oval to ellipsoidal, 20–40 µm diam, brown to dark brown, stroma
cells oval, ellipsoidal and angular, 3–10 µm wide, dark brown,
wall 0.5–0.8 µm wide, smooth. Conidiophores fasciculate,
arising from stromata (3–16 per fascicle), geniculate,
unbranched, 9–15 × 2–4 µm, 0–1-septate, slightly constricted
at the septa, distance between septa 3–12 µm, uniformly
pale to medium brown, paler and narrower towards the tip,
wall 0.3–0.5 µm wide, smooth. Conidiogenous cells terminal,
8–12 × 2–4 µm, obtuse; conidiogenous loci inconspicuous,
unthickened, not darkened. Conidia solitary, obclavate,
straight to slightly curved, 50–105 × 2–4 µm, 3–9-septate, pale
olivaceous-brown, wall 0.3–0.5 µm wide, smooth, tip subacute,
base obconically truncate, hila 1–2 µm wide.
In vitro: Colonies on PDA after 3 wk at 25 C dark grey,
reaching 10–15 mm diam, hyphae 2–7 µm wide, septate,
constricted at the septa, distance between septa 9–19 µm,
brownish or subhyaline, wall 0.3–0.8 µm wide, smooth.
Colonies sterile.
Holotype: Thailand: Chiang Rai Province: Muang District,
Mae Chan Village, on leaves of Christella parasitica,
Thelypteridaceae, 18 July 2010, P. Phengsintham (MFLU10-
0405).
Host range and distribution: Only known from the type
collection.
Pseudocercospora cyatheae C. Nakash. & S. Inaba,
Mycoscience 47: 48 (2006).
(Fig. 33)
Illustration: Nakashima et al. (2006: 49, g. 1).
Description: Leaf spots distinct, irregularly shaped, brown
to dark brown, 2–5 mm diam, margin indistinct or distinct.
Caespituli epiphyllous, punctiform, dark. Mycelium internal.
Stromata well-developed, immersed, globose, 60–110
Fig. 32. Pseudocercospora christellae (MFLU10-0405). A.
Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10
µm.
Braun et al.
ARTICLE
312 ima fUNGUS
µm diam, pale brown to brown. Conidiophores in large,
dense fascicles, arising from stromata, erumpent, erect,
straight to somewhat sinuous, subcylindrical or attenuated
towards the tip, unbranched, 25–50 × 3–5 µm, 0–3-septate,
pale brown to brown, paler towards the tip, thin-walled,
smooth; conidiophores reduced to conidiogenous cells
or conidiogenous cells integrated, terminal, proliferation
sympodial or sometimes percurrent, conidiogenous loci
inconspicuous, unthickened, not darkened or rim-like. Conidia
solitary, cylindrical to obclavate, mildly curved to sigmoid,
30–50 × 3.5–5.5 µm, 3–8-septate, pale brown, thin-walled,
smooth, apex attenuated, obtuse to subacute, base rounded
to obconically truncate, hila unthickened, not darkened.
Holotype: Japan: Okinawa Pref.: Ishigaki, Tomigusuku, on
leaves of Cyathea sp., Cyatheaceae, 28 Mar. 2003, S. Inaba
(NBRC H-12398, culture ex-type: NBRC 101080).
Host range and distribution: Only known from the type
collection.
Pseudocercospora cyatheicola Crous & R.G. Shivas,
Persoonia 26: 121 (2011); as “cyathicola”.
(Fig. 34)
Illustration: Crous et al. (2011: 120).
Description: On dead fronds, associated with a
mycosphaerella-like teleomorph. Mycelium internal; hyphae
septate, branched, 2.5–3 µm wide, pale brown, smooth,
thin-walled. Stromata amphigenous on fronds, well-
developed, erumpent, up to 60 µm diam and 40 µm high,
brown. Conidiophores arising from stromata, in loose to
Fig. 33. Pseudocercospora cyatheae (NBRC-H-12398). a. drawing
(A. Conidiophore fascicle, B. Conidia, U. Braun del.). b. Conidium
(light microscopy). c. Conidiophore fascicle (light microscopy). Bar
= 10 µm.
Fig. 34. Pseudocercospora cyatheicola (CBS H-20580). A.
Conidiophore fascicles. B. Conidiophore tip. C. Conidia. Bar = 10
µm.
ab
c
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rather dense well-developed fascicles, erect, subcylindrical,
straight to geniculate-sinuous, unbranched, 30–70 × 2–3
µm, 1–3-septate, pale to medium brown, smooth, thin-
walled; conidiogenous cells integrated, terminal, pale brown,
proliferation sympodial and percurrent, conidiogenous loci
visible as truncate ends, unthickened and not darkened,
1.5–2 µm wide. Conidia solitary, subcylindrical to subacicular,
somewhat irregular in width, straight to irregularly curved,
(35–)60–80(–90) × (2–)3(–3.5) µm, 3–9-septate, pale brown,
smooth, thin-walled, guttulate, apex obtuse to subacute,
base truncate to somewhat obconically truncate, hila about 2
µm wide, unthickened, not darkened.
In vitro (in the dark, 25 °C after 1 mo): Colonies spreading,
somewhat erumpent, with moderate aerial mycelium and
smooth, lobate margins, reaching 35–45 mm diam. On malt
extract agar surface olivaceous-grey, with patches of smoke-
grey, reverse iron-grey; on potato-dextrose agar surface
pale olivaceous-grey, margin olivaceous-grey, reverse iron-
grey; on oatmeal agar surface pale olivaceous-grey, margin
olivaceous-grey.
Holotype: Australia: Queensland: Brisbane, Botanical
Garden, on fronds of Cyathea australis, Cyatheaceae, 14 Jul.
2009, P. W. Crous & R. G. Shivas (CBS H-20580, culture ex-
type: CBS 129520).
Host range and distribution: Only known from the type
collection.
Pseudocercospora davalliae (A.K. Kar & M. Mandal)
U. Braun & Crous, in Crous & Braun, Mycosphaerella
and Anam. 1: 154 (2003).
(Fig. 35)
Basionym: Cercospora davalliae A.K. Kar & M. Mandal,
Trans. Brit. Mycol. Soc. 53: 355 (1969).
Literature: Crous & Braun (2003: 154), Kamal (2010: 170).
Illustration: Kar & Mandal (1969: 355, g. 17).
Description: Leaf spots amphigenous, irregular in shape, 1–5
mm diam, dingy brown to darker brown, margin indenite.
Mycelium internal. Stromata substomatal, 15–50 µm diam,
dark olivaceous, composed of swollen hyphal cells, about 2–5
µm diam. Conidiophores fasciculate, in small to moderately
large, loose to dense fascicles, arising from stromata, emerging
through stomata, erect, straight, subcylindrical to attenuated
towards the tip, mostly geniculate-sinuous, unbranched,
about 5–65 × 2.5–5.5 µm, 0–6-septate, uniformly very
pale olivaceous-brown, thin-walled, smooth; conidiophores
reduced to conidiogenous cells or conidiogenous cells
integrated, terminal, occasionally intercalary, about 5–25 µm
long, conidiogenous loci inconspicuous or visible by being
truncate or rarely subdenticulate. Conidia solitary, narrowly
obclavate-subcylindrical or linear, 20.5–105.5 × 2–3.5 µm,
2–11-septate, pale olivaceous, thin-walled, smooth, apex
obtuse to subacute, base short obconically truncate, about
1–1.5 µm wide, hila unthickened, not darkened.
Types: India: West Bengal: Darjeeling, Kalimpong, 1372 m
alt., on leaves of Odontosoria chinensis (Davallia tenuifolia),
Lindsaeaceae, 8 May 1967, A. K. Kar & M. Mandal (K(M) IMI
135178 – holotype; BPI 435482 – isotype).
Host range and distribution: Only known from the type
collection.
Notes: Braun & Freire (2004: 229–230) recorded this species
on Davallia fejeensis from Brazil, which represents, however,
a distinct species described below as Pseudocercospora
davalliicola.
Pseudocercospora davalliicola U. Braun, sp. nov.
MycoBank MB805524
(Fig. 36)
Etymology: Derived from its host genus (inhabitant of
Davallia).
Fig. 35. Pseudocercospora davalliae (K(M) IMI 135178). A.
Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
Braun et al.
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314 ima fUNGUS
Exsiccatae: U. Braun, Fungi Sel. Exs. 14.
Diagnosis: Pseudocercosporae davalliae similis, sed hyphis
supercialibus cum conidiophoris solitariis formantibus,
conidiophoris saepe 0–1-septatis et conidiis brevioribus, 12–
70 µm, solitariis vel brevicatenatis.
Description: Leaf spots absent or irregular in shape and
size, formed as brown discolorations, margin indenite, later
large leaf segments or entire leaves becoming necrotic.
Caespituli amphigenous, mostly hypophyllous, punctiform,
scattered, dark greyish brown. Mycelium internal and
external; supercial hyphae emerging through stomata,
sparingly branched, septate, 1.5–3 µm wide, subhyaline to
pale olivaceous, thin-walled, smooth. Stromata substomatal,
10–50 µm diam, olivaceous-brown, composed of swollen
hyphal cells, about 2–5 µm diam. Conidiophores in small to
moderately large fascicles, loose to mostly dense, arising from
stromata, through stomata, erect, straight, subcylindrical to
distinctly geniculate-sinuous, unbranched, 5–40 × 2–4.5 µm,
0–1(–2)-septate, subhyaline to pale olivaceous or olivaceous-
brown, thin-walled, smooth; conidiophores reduced to
conidiogenous cells or conidiogenous cells integrated,
terminal, 5–25 µm long, conidiogenous loci inconspicuous.
Conidia solitary or in short chains, occasionally even in
branched chains, short conidia subcylindrical-fusoid, longer
conidia narrowly obclavate-cylindrical, 12–70 × 2–4 µm,
1–7-septate, subhyaline to pale olivaceous, thin-walled,
smooth, apex obtuse to pointed, base obconically truncate,
1–1.5 µm wide, hila unthickened, not darkened.
Types: Brazil: Ceará: Ubajara city, on leaves of Davallia
fejeensis, Davalliaceae, 10 Oct. 2002, F. Freire (HAL 2589 F –
holotype; U. Braun, Fungi Sel. Exs. 14 (as “Pseudocercospora
davalliae”), BPI, GZU, HMAS, K(M) IMI, KR, KUS, LE, PDD,
VPIR – isotypes).
Host range and distribution: Only known from the type
collection.
Notes: Braun & Freire (2004) recorded this collection as
Pseudocercospora davalliae. Davallia fejeensis and the type
host of the latter species, Odontosoria chinensis [Davallia
tenuifolia] are, however, not allied and belong in different
families. Furthermore, there are various morphological
differences between collections on the two hosts. The conidia
in P. davalliae are consistently formed singly and much longer,
up to about 100 µm, and pluriseptate. The conidiophores are
also longer and up to 6-septate, and supercial hyphae with
solitary conidiophores are lacking.
Pseudocercospora gleicheniae (J.M. Yen) U. Braun,
comb. nov.
MycoBank MB805525
(Fig. 37)
Basionym: Cercospora gleicheniae J.M. Yen, Bull. Trimestriel
Soc. Mycol. France 90: 41 (1974).
Literature: Crous & Braun (2003: 198).
Illustration: Yen (1974: 42, g. 1).
Description: Lesions lacking. Caespituli hypophyllous.
Mycelium internal and external; hyphae emerging through
stomata, supercial, branched, septate, pale olivaceous-
brown, 1.5–2 µm wide, thin-walled, smooth. Stromata lacking
or small, substomatal, somewhat pigmented. Conidiophores
solitary, arising from supercial hyphae, lateral or occasionally
terminal, erect, very long and liform, straight to somewhat
sinuous, unbranched or branched, about 100–300 µm long
and 3.5–5 µm wide, pluriseptate throughout (about 10–30
septa), dark brown, wall thickened, smooth; conidiogenous
cells integrated, terminal or intercalary, 10–30 µm long,
conidiogenous loci subconspicuous to conspicuous by being
± denticle-like, but wall neither thickened nor darkened, about
1–2 µm broad. Conidia solitary, subcylindrical, ellipsoid, short
subclavate-obovoid, 12–25 × 4.5–6 µm, (2–)3–4(–5)-septate,
Fig. 36. Pseudocercospora davalliicola (HAL 2589 F). A. Solitary
conidiophores arising from supercial hyphae. B. Conidiophore tips.
C. Conidiophores. D. Conidia. Bar = 10 µm.
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subhyaline, pale olivaceous to olivaceous-brown, thin-walled,
smooth or almost so, apex rounded, base short obconically
truncate to somewhat peg-like, hila neither thickened nor
darkened.
Holotype: Gabon: Libreville, 16.3 km sur route de Kongo,
on Dicranopteris linearis, Gleicheniaceae, 23 Feb. 1969, G.
Gilles 139 (PC).
Host range and distribution: Only known from the type
collection.
Notes: This is a very unusual species. The generic afnity is
not quite clear. Crous & Braun (2003) excluded this species.
Denticulate conidiogenous cells and phragmosporous
conidia resemble species of the Dactylaria complex.
However, the very long thick-walled conidiophores are not
dactylarioid. The type of loci (neither thickened nor darkened,
subdenticulate loci caused by geniculation due to sympodial
proliferation to distinctly denticle-like) are within the range
of Pseudocercospora loci. Phragmosporous conidia are
unusual for the latter genus but may occur. Therefore, this
species is tentatively assigned to Pseudocercospora.
Pseudocercospora helminthostachydis (Henn.)
Deighton, Mycol. Pap. 140: 145 (1976).
(Fig. 38)
Basionym: Cercospora helminthostachydis Henn., Hedwigia
47: 265 (1908).
Literature: Saccardo (1913: 1424), Chupp (1954: 424),
Bhurgava & Misra (1961), Thaung (1984), Crous & Braun
(2003: 215), Kamal (2010: 182).
Description: Leaf spots amphigenous, small indistinct or
irregular spots on the upper side, 2–5 mm diam, on the
lower side dark, blackish by dense colonies. Caespituli
hypophyllous, pustulate, dark brown to black. Mycelium
internal. Stromata small to well-developed, substomatal to
intraepidermal, 20–75 µm diam, brown, composed of swollen
hyphal cells, 2–7 µm diam. Conidiophores in moderately large
to large fascicles, arising from stromata, emerging through
stomata or erumpent, dense, sometimes very dense, erect,
unbranched, straight, subcylindrical to somewhat attenuated
towards the tip, slightly geniculate-sinuous, 10–70 × 3–6 µm,
0–3-septate, pale to medium olivaceous-brown, thin-walled,
smooth; conidiogenous cells integrated, terminal, 10–30 µm
long, conidiogenous loci unthickened, not darkened. Conidia
solitary, obclavate-cylindrical, straight to somewhat curved,
Fig. 37. Pseudocercospora gleicheniae (PC, holotype). A. Hyphae
emerging through a stoma. B. Conidiophore base. C. Conidiophore
tips. D. Conidia. Bar = 10 µm.
Fig. 38. Pseudocercospora helminthostachydis (B, holotype).
A. Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar =
10 µm.
Braun et al.
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316 ima fUNGUS
40–100 × 4–6 µm, 3–7-septate, olivaceous to olivaceous-
brown, thin-walled, smooth, apex obtuse, rounded, base
short obconically truncate, hila 1.5–2 µm wide, neither
thickened nor darkened.
Types: Philippines: Mindanao, Davao, on Tectaria zeylanica,
15 Mar. 1904, E. B. Copeland 543 (B – holotype; BPI 437080
– isotype [date given as "15 Mar. 1906"]).
Host range and distribution: On Tectaria zeylanica
(Helminthostachys zeylanica), Tectariaceae, Asia (India,
Philippines, Myanmar).
Pseudocercospora lonchitidis (Chupp) U. Braun &
Crous, in Crous & Braun, Mycosphaerella and Anam.
1: 255 (2003).
(Fig. 39)
Basionym: Cercospora lonchitidis Chupp, A monograph of
the fungus genus Cercospora: 455 (1954).
Literature: Crous & Braun (2003: 255).
Description: Leaf spots amphigenous, irregularly shaped,
3–8 mm diam, dull red above, below often only with a
narrow marginal line surrounding a greenish area. Caespituli
amphigenous, punctiform on the upper side, effuse below,
grey to dark brown. Mycelium internal and external, supercial
hyphae only formed on the lower side, branched, septate,
subhyaline to pale olivaceous, 1–4 µm diam, thin-walled,
smooth. Stromata only epiphyllous, immersed, subglobose,
20–60 µm diam, brown to dark brown. Conidiophores on
the upper side in larger fascicles, loose to mostly dense,
arising from stromata, erumpent, on the lower side in
small, loose fascicles and solitary, arising from supercial
hyphae, lateral or occasionally terminal, conidiophores erect,
straight, subcylindrical to geniculate-sinuous, unbranched
or occasionally branched, 10–100 × 3–5 µm, continuous
to pluriseptate throughout, subhyaline to pale olivaceous-
brown throughout, thin-walled, smooth; conidiogenous cells
integrated, terminal, about 10–30 µm long, conidiogenous loci
inconspicuous to subdenticulate, but always unthickened and
not darkened. Conidia solitary, acicular to obclavate, 30–160
× 3–5 µm, 3- to pluriseptate, subhyaline to pale olivaceous,
thin-walled, smooth, apex obtuse to subacute, base truncate
to slightly obconically truncate, about 2–3.5 µm wide, hila
neither thickened nor darkened.
Holotype: Venezuela: Aragua, road Maracaya Choroni,
on Lonchitis hirsuta, Dennstadtiaceae, 9 Apr. 1939, H. H.
Whetzel & A. S. Muller (CUP-VZ-3385).
Host range and distribution: Only known from the type
collection.
Pseudocercospora lygodii Goh & W.H. Hsieh, Trans.
Mycol. Soc. Republ. China 2(2): 131 (1987).
(Fig. 40)
Synonym: Cercospora lygodii Sawada, Rep. Taiwan Agric.
Res. Inst. 87: 83, 1944, nom. inval. (ICN, Art. 39.1).
Literature: Hsieh & Goh (1990: 305–306), Guo & Hsieh
(1995: 187), Guo et al. (1998: 199), Crous & Braun (2003:
258), Phengsintham et al. (2013a: 66–67).
Illustrations: Hsieh & Goh (1990: 306, g. 232), Guo &
Hsieh (1995: 187, g. 160), Guo et al. (1998: 199, g. 165),
Phengsintham et al. (2013a: 67, gs 70–71).
Description: Leaf spots amphigenous, circular to elliptical,
scattered, 2–30 mm diam, greyish to dark brown, centre
becoming paler, margin indenite. Colonies amphigenous,
mainly hypophyllous. Mycelium internal and external;
supercial hyphae creeping and climbing setae of the
leaf surface, branched, septate, 2–5 µm wide, olivaceous
to pale brown, thin-walled, smooth. Stromata lacking.
Conidiophores in small fascicles (2–10), divergent, arising
from internal hyphae, emerging through stomata, or solitary,
arising from supercial hyphae, lateral, erect, straight,
subcylindrical to slightly geniculate-sinuous, unbranched,
20–90 × 3–4 µm, 2–7-septate, yellowish ochraceous
to medium brown, thin-walled, smooth; conidiogenous
cells integrated, terminal, conidiophores occasionally
reduced to conidiogenous cells, about 20–35 µm long,
conidiogenous loci inconspicuous to subdenticulate, but
always unthickened and not darkened. Conidia solitary,
obclavate-cylindrical, straight to somewhat curved, 10–35
× 1.5–4 µm, 1–3-septate, colourless, thin-walled, smooth,
apex acute to subobtuse, base obconically truncate, about
1–2 µm wide, hila unthickened, not darkened.
Fig. 39. Pseudocercospora lonchitidis (CUP-VZ-3385). A. Supercial
hyphae. B. Solitary conidiophores arising from supercial hyphae. C.
Conidiophore fascicle. D. Conidia. Bar = 10 µm.
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Holotype: Taiwan: Hsinchu Hsien, Hsinpu, on Lygodium
japonicum, 2 May 1920, K. Sawada (NTU-PPE).
Host range and distribution: On Lygodium (exuosum,
japonicum [microstachyum]), Lygodiaceae, Asia (Taiwan,
Thailand).
Pseudocercospora lygodiicola Y.L. Guo & U. Braun,
sp. nov.
MycoBank, MB805526
(Fig. 41)
Pseudonym: Pseudocercospora polypodiacearum sensu
Guo & Liu (1992: 298), Guo & Hsieh (1995: 188) and Guo
et al. (1998: 199–200).
Etymology: Epithet derived from its host genus (inhabitant of
Lygodium).
Illustrations: Guo & Hsieh (1995: 189, g. 161), Guo et al.
(1998: 200, g. 166).
Diagnosis: Pseudocercosporae polypodiacearum similis, sed
conidiophoris brevioribus, 15–40 µm, 0–1-septatis, apice non
inatis.
Description: Leaf spots amphigenous, irregularly shaped,
1–6 mm diam, often conuent and larger, covering up to
half of a leaf, greyish brown above, greyish black below.
Caespituli amphigenous, mainly hypophyllous. Mycelium
immersed. Stromata substomatal, subglobose, 10–40
µm diam, dark brown. Conidiophores in dense fascicles,
arising from stromata, through stomata, erect, straight,
subcylindrical to geniculate-sinuous, 15–40 × 2.5–5 µm,
0–1-septate, uniformly olivaceous to olivaceous-brown,
thin-walled, smooth, apex obtuse to conic; conidiophores
reduced to conidiogenous cells or conidiogenous cells
integrated, terminal, about 10–25 µm long, conidiogenous
loci inconspicuous or subdenticulate, but always unthickened
and not darkened. Conidia solitary, obclavate, occasionally
subcylindrical, 40–115 × 3–5 µm, 3–13-septate, olivaceous-
brown, thin-walled, smooth, apex obtuse to subacute, base
subtruncate to usually short obconically truncate, about 1.5–
2 µm wide, hila unthickened, not darkened.
Types: China: Guangdong Province: Guangzhou, on leaves
of Lygodium japonicum, Lygodiaceae, 8 VI 1962, Q. M. Ma
& X. J. Liu 1439 (HMAS 59148 – holotype; BPI 1109717 –
isotype).
Host range and distribution: Only known from the type
collection.
Notes: This species was previously recorded, described and
illustrated from China as Pseudocercospora polypodiacearum
(Guo & Hsieh 1995, Guo et al. 1998), but differs from the latter
species in having much shorter, 0–1-septate, apically usually
not inated conidiophores, and is thus better considered a
separate species.
Pseudocercospora lygodiigena U. Braun, nom. nov.
MycoBank MB806067
(Fig. 42)
Basionym: Stenella lygodii Sarbajna, J. Econ. Taxon. Bot. 14:
489 (1990), non Pseudocercospora lygodii Goh & W.H.
Hsieh, 1987.
Synonym: Zasmidium lygodii (Sarbajna) Kamal, Cercosporoid
fungi India: 245 (2010).
Illustration: Sarbajna (1990: 488, g. 1).
Description: Leaf spots amphigenous, subcircular to angular-
irregular, 1–4 mm diam., sometimes vein-limited, centre
greyish, margin darker, brown. Caespituli amphigenous,
mostly hypophyllous, punctiform, scattered to aggregated,
brown or greyish by abundant sporulation. Mycelium
internal and external; supercial hyphae emerging through
Fig. 40. Pseudocercospora lygodii (NTU-PPE, holotype). A. Small conidiophore fascicle. B. Solidary conidiophores arising from supercial
hyphae. C. Conidia. Bar = 10 µm.
Braun et al.
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318 ima fUNGUS
stomata, branched, 1–4 µm wide, septate, pigmented near
conidiophores, otherwise hyaline, subhyaline or very pale
olivaceous, thin-walled, smooth or almost so, occasionally
faintly rough-walled. Stromata lacking to well-developed,
mostly substomatal, 10–30 µm diam., pigmented.
Conidiophores in small to moderately large, loose fascicles,
3–30, arising from internal hyphae or stromata and solitary,
arising from supercial hyphae, lateral, erect, straight,
subcylindrical to somewhat sinuous or subgeniculate,
unbranched, 5–55 × 2–4 µm, 0–3(–5)-septate, pale to
medium brown or olivaceous-brown, wall thin to somewhat
thickened, up to 0.8 µm, smooth; conidiogenous cells
integrated, terminal, 5–25 µm long, conidiogenous loci
inconspicuous, neither thickened nor darkened. Conidia
solitary, narrowly obclavate-subcylindrical, occasionally
subacicular, straight to curved, 30–115.5 × 2–4 µm,
2–14-septate, hyaline to faintly pigmented, thin-walled,
smooth or almost so to somewhat rough, apex subobtuse
to pointed, base short obconically truncate, 1–1.5 µm wide,
hila neither thickened nor darkened.
Neotype (designated here, MycoBank, MBT176675): India:
West Bengal: Baruipur, on Lygodium sp., Lygodiaceae, 15
Jul. 1984, A.K. Das 3623 (K(M) IMI 288612).
Host range and distribution: On Lygodium sp., Lygodiaceae,
Asia (India).
Notes: This species does neither belong to Stenella nor to
Zasmidium. Due to inconspicuous conidiogenous loci and
unthickened, non-pigmented hila of the conidia, it has to be
reallocated to Pseudocercospora. Sarbajna (1990) cited IMI
288612 as number of the holotype, but the data (locality,
collector and date) specied in the original description
for the holotype do not agree with the data on the label
of the material deposited at Kew (K) under this number.
The search for another collection of this species (under
another IMI number) corresponding to the data cited in
Sarbajna (1990) failed. Thus, type material of this species
is undoubtedly not preserved. The material deposited as
Stenella lygodii under IMI 288612 agrees with Sarbajna’s
(1990) original description, but it cannot be considered the
Fig. 41. Pseudocercospora lygodiicola (HMAS 59148). A.
Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10
µm.
Fig. 42. Pseudocercospora lygodiigena (K(M) IMI 288612). A.
Conidiophores arising from supercial hyphae. B. Fasciculate
conidiophores. C. Conidia. Bar = 10 µm.
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holotype. Therefore, we propose to designate this material
as neotype of S. lygodii.
Pseudocercospora microsori R.G. Shivas, A.J.
Young & B.C. McNeil, Persoonia 25: 157 (2010).
(Fig. 43)
Illustration: Shivas et al. (2010: 156).
Description: Leaf spots on fonds, amphigenous, scattered to
conuent, often covering much of the fond surface, circular
to irregular, 5–15 mm diam, dark reddish brown, centre
becoming grey, margin conspicuous, uneven, with chlorotic
halo, vein-limited. Caespituli amphigenous. Mycelium internal.
Stromata well-developed, 20–60 µm diam, substomatal,
brown. Conidiophores in loose to dense fascicles, 5–30,
arising from stromata, through stomata, forming sporodochial
conidiomata, erect, unbranched, geniculate-sinuous, 30–65 ×
3–5 µm, reddish brown, paler towards the apex, 1–5-septate,
thin-walled, smooth; conidiogenous cells integrated, terminal,
subcylindrical, 10–35 × 2.5–4 µm, conidiogenous loci
inconspicuous or visible as truncate tip, unthickened, not
darkened. Conidia solitary, acicular to narrowly obclavate-
subcylindrical, curved to exuous, 50–110 × 2.5–4 µm,
2–12-septate, pale brown, thin-walled, smooth, apex rounded
or subacute, base truncate to slightly obconically truncate,
hila neither thickened nor darkened.
Holotype: Australia: Queensland: Brisbane, West End, Doris
Street, on Microsorum pustulatum Copel., Polypodiaceae, 6
Aug. 2010, B. C. McNeil (BRISP 53617).
Host range and distribution: Only known from the type
collection.
Pseudocercospora nephrolepidicola Crous & R.G.
Shivas, Persoonia 25: 139 (2010).
(Fig. 44)
Illustration: Crous et al. (2010: 138).
Description: Leaf spots amphigenous, shape and size
irregular, 2–12 mm diam, medium brown, margin indenite.
Mycelium internal and external; hyphae branched, 2–3 µm
wide, septate, brown, thin-walled, smooth. Stromata well-
developed, subepidermal, up to 150 µm diam and 50 µm high,
Fig. 43. Pseudocercospora microsori (BRISP 53617). A. Conidiophore
fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm. Fig. 44. Pseudocercospora nephrolepidicola (CBS H-20492). A.
Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
Braun et al.
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320 ima fUNGUS
medium brown. Conidiophores arising from stromata in loose
fascicles, or solitary, arising from supercial hyphae, erect,
subcylindrical, irregularly geniculate-sinuous, unbranched or
branched below, 25–45(–90) × 2.5–3(–3.5) µm, 1–4-septate,
medium brown, thin-walled, smooth; conidiogenous cells
integrated, terminal, subcylindrical, 15–25(–40) × (2–)2.5(–3)
µm, sympodially and 1–2 times percurrently proliferating,
conidiogenous loci neither thickened nor darkened.
Conidia solitary, subcylindrical-liform, occasionally slightly
aciculare (somewhat attenuated towards the tip), straight to
irregularly exuous, (40–)50–60(–95) × (2.5–)3.5(–4) µm,
3–6(–9)-septate, pale brown, thin-walled, smooth, apex
obtuse, rounded, base truncate, hila neither thickened nor
darkened. Ascomata globose, erumpent, up to 80 µm diam,
brown, with a central ostiole; asci subcylindrical to narrowly
obovoid, 35–50 × 8–10 µm. Ascospores fusiform-ellipsoid,
widest in the middle of the apical cell, tapering towards both
ends, 9–11 × 2.5–3.5 µm, colourless, apex acutely rounded,
constricted at the septum.
In vitro (in the dark at 25 °C, after 2 wk): Colonies
spreading, erumpent, with folded surface and even, lobate
margins, up to 15 mm diam. On potato-dextrose agar surface
smoke-grey with patches of grey-olivaceous, iron-grey in
reverse; on oatmeal agar olivaceous-grey with patches of
pale olivaceous-grey.
Holotype: Australia: Queensland: Brisbane Botanical
Garden, on fronds of Nephrolepis falcata, Davalliaceae, 14
Jul. 2009, P. W. Crous & R. G. Shivas (CBS H-20492); ex-
type culture: CBS 128211.
Host range and distribution: Only known from the type
collection.
Notes: Pseudocercospora nephrolepidis, described
from Taiwan on Nephrolepis auriculata, differs from P.
nephrolepidicola in lacking or much smaller stromata, lacking
supercial hyphae, shorter, only 0–1-septate conidiophores
and shorter conidia.
Pseudocercospora nephrolepidigena U. Braun,
Meeboon & C. Nakash., sp. nov.
MycoBank MB805527
(Fig. 45)
Etymology: Derived from its host genus, Nephrolepis.
Literature: Braun & Sivapalan (1999: 21, as “P. thelypteridis“),
Nakashima et al. (2007: 265, as “P. phyllitidis”), Meeboon et
al. (2007, as “P. phyllitidis”).
Illustration: Braun & Sivapalan (1999: 20, g. 12, as “P.
thelypteridis“).
Fig. 45. Pseudocercospora nephrolepidigena (HAL 2602 F). a. Based on type material. b. Based on material from Brunei at HAL. A. Solitary
conidiophores arising from supercial hyphae. B. Supercial hyphae. C. Conidiophore fascicles. D. Conidiophores. E. Conidia. Bar = 10 µm.
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Diagnosis: Pseudocercosporae nephrolepidis et P. phyllitidis
similis, sed hyphis supercialibus cum conidiophoris solitariis
crescentibus.
Description: Leaf spots amphigenous, subcircular to an-
gular-irregular, 1–15 mm diam, or oblong, up to 20 mm,
sometimes conuent, forming large blotches or entire leaf
segments becoming necrotic, pale to dark brown, margin
indenite or somewhat darker, sometimes with yellowish
halo, occasionally somewhat zonate. Caespituli amphig-
enous, conspicuous on the upper side, punctiform, scat-
tered, blackish or greyish by abundant conidial formation,
less conspicuous on the lower side. Mycelium internal and
external; supercial hyphae amphigenous, abundant on
the lower side, less developed but also present on the up-
per side, branched, 1.5–4.5 µm wide, septate, subhyaline
to pale olivaceous, thin-walled, smooth. Stromata small to
well-developed, immersed, 10–80 µm diam, pale to dark
olivaceous-brown, cells 2.5–8 µm diam. Conidiophores in
small to rather large, loose to mostly dense fascicles, arising
from stromata, erumpent, or solitary, arising from supercial
hyphae, lateral, erect, straight, subcylindrical or somewhat
attenuated towards the tip, unbranched, occasionally some-
what geniculate, 5–50 × 1.5–4 µm (with attached persistent
conidia up to about 80 µm long), 0–3-septate, subhyaline
to pale olivaceous or brownish, thin-walled, smooth; conidi-
ophores often aseptate, i.e. reduced to conidiogenous cells,
or conidiogenous cells integrated, terminal, 5–25 µm long,
often monoblastic, determinate, sometimes polyblastic,
sympodially proliferating, conidiogenous loci inconspicuous
or visible as truncate tips, about 1.5–2.5 µm wide, neither
thickened nor darkened. Conidia solitary, persistent, i.e. at-
tached to the conidiogenous cells for a longer time, cylindri-
cal, long conidia liform to subacicular, i.e. somewhat atten-
uated towards the tip, straight to somewhat curved, 25–115
× (1.5–)2–3.5(–4) µm, 2–10-septate, septa not very distinct,
subhyaline to very pale olivaceous, thin-walled, smooth,
apex obtuse to subacute, base truncate, about 1.5–2.5 µm
wide, hila neither thickened nor darkened.
Types: Thailand: Chiang Mai Province: Suthep-Pui National
Park, on Nephrolepis biserrata, 31 May 2013, J. Meeboon
(HAL 2602 F – holotype; U. Braun, Fungi Sel Exs. 208, BPI,
BRIP, GZU, HAL, HMAS, K (M), KR, KUS, LE, M, PDD –
isotypes). Paratypes: Thailand: Chiang Mai Province:
Suthep-Pui National Park, 21 Nov. 2004, C. Nakashima &
J. Meeboon (CMU 27962) and 10 Dec. 2006, I. Araki & J.
Meeboon (CMU 28058).
Host range and distribution: On Nephrolepis (biserrata,
cordifolia, Nephrolepis sp.), Davalliaceae, Asia (Brunei,
Thailand).
Notes: Nakashima et al. (2007) recorded collections on this
host from Thailand as Pseudocercospora phyllitidis, but the
latter American species differs in lacking supercial hyphae
and in having much longer conidiophores. Furthermore, the
type host of P. phyllitidis, Campyloneurum phyllitidis, belongs
to the Polypodiaceae. Pseudocercospora phyllitidis, record-
ed on various ferns belonging to different families, is prob-
ably heterogeneous. “Pseudocercospora thelypteridis” on
Nephrolepis from Brunei, described and illustrated in Braun
& Sivapalan (1999), agrees well with P. nephrolepidigena.
Pseudocercospora nephrolepidis R. Kirschner,
Fungal Diversity 26: 223 (2007).
(Fig. 46)
Illustration: Kirschner & Chen (2007: 224, g. 2).
Description: Leaf spots amphigenous, on yellowing leaets
with green margin, 1–10 mm long and 3–4 mm wide, not
passing the midrib of leaets, shape irregular, medium
brown. Caespituli hypophyllous. Mycelium internal; hyphae
intercellular, 1–4 µm diam, pale brown, smooth. Stromata
absent or small, within and just below stomata, shape
irregular, rarely up to 35 µm diam, pale brown. Conidiophores
Fig. 46. Pseudocercospora nephrolepidis (TNM, holotype). A.
Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
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322 ima fUNGUS
fasciculate, up to 22, arising from internal hyphae or stromata,
emerging through stomata, straight, subcylindrical to slightly
geniculate, unbranched (7–)13–26(–35) × 2.5–3 µm,
0–1-septate, pale brown, thin-walled, smooth; conidiogenous
cells integrated, terminal or conidiophores mostly aseptate,
i.e. reduced to conidiogenous cells, conidiogenous loci
broadly truncate, neither thickened nor darkened. Conidia
solitary, subcylindrical to acicular, i.e. somewhat tapering
from base to top, straight to curved or sinuous, about (30–)
65–100(–115) × 2–3 µm, inconspicuously 2–9-septate,
subhyaline to pale brown (paler than the conidiophores), thin-
walled, smooth, apex subobtuse to subacute, base truncate,
hila neither thickened nor darkened.
Holotype: Taiwan: Taipei County, Yangmingshan, ca. 800 m
alt., on living leaves of Nephrolepis auriculata, 2 Mar. 2006,
R. Kirschner 2555 (TNM).
Host range and distribution: on Nephrolepis auriculata,
Davalliaceae, Asia (Taiwan).
Note: Differences between this species and the Australian P.
nephrolepidicola are discussed under the latter species.
Pseudocercospora phyllitidis (H.H. Hume) U. Braun
& Crous, in Crous & Braun, Mycosphaerella and Anam.
1: 321 (2003).
(Fig. 47)
Basionym: Cercospora phyllitidis H.H. Hume, Bull. Torrey
Bot. Club 27: 577 (1900).
Literature: Saccardo (1902: 1074), Chupp (1954: 456), Crous
& Braun (2003: 321).
Description: Leaf spots amphigenous, angular-irregular,
2–18 mm diam, brown, margin usually indenite. Caespituli
hypophyllous. Mycelium internal; rarely with a few supercial
hyphae, but without conidiophores. Stromata small to large,
only lling the substomatal cavity or larger, 10–80 µm diam,
subglobose, brown. Conidiophores in small to rather large
fascicles, arising from stromata, emerging through stomata,
erect, straight to somewhat curved or sinuous, subcylindrical,
barely geniculate, unbranched, 10–75(–100) × 3–5 µm,
continuous to pluriseptate, uniformly subhyaline, pale to
medium olivaceous-brown or brown, wall thin to somewhat
thickened, smooth; conidiogenous cells integrated, terminal or
conidiophores reduced to conidiogenous cells, conidiogenous
loci inconspicuous, neither thickened nor darkened. Conidia
solitary, obclavate, 20–80 × 2–5 µm, 3–5-septate, subhyaline
to pale olivaceous, thin-walled, smooth, apex subacute or
obtuse, base obconically truncate, about 1.5–2.5 µm wide,
hila neither thickened nor darkened.
Lectotype (designated here, MycoBank, MBT176148):
USA: Florida: east coast, Hobe Sound, on Campyloneurum
phyllitidis (Polypodium phyllitidis), 14 Mar. 1900, H. H. Hume
(NY). Isolectotype: BPI 439635.
Host range and distribution: On Angyopteris sp., Marattiaceae;
Campyloneurum phyllitidis, Polypodium polypodioides,
Polypodiaceae; Davallia trichomanoides, Nephrolepis
exaltata, Davalliaceae; Osmunda regalis, Osmundaceae;
Pteris (P. biaurita, P. ensifolia), Pteridaceae; Rumohra
adiantiformis, Dryopteridaceae; Thelypteris tetragona,
Thelypteridaceae; non-identied fern, North America
(Canada; USA, Florida, Indiana), Jamaica, Panama, Puerto
Rico, Europe (UK, in a botanical garden).
Notes: Records of this species from Asia, India (Lall et al.
1964, Kamal 2010) are doubtful and seem to belong to P.
pteridicola. There are various records on ferns belonging to
different families, suggesting that P. phyllitidis is probably
heterogeneous.
Pseudocercospora plagiogyriae Goh & W.H. Hsieh,
Trans. Mycol. Soc. Republ. China 2(2): 134 (1987).
(Fig. 48)
Synonym: Cercospora plagiogyriae Sawada, Rep. Taiwan
Agric. Res. Inst. 85: 119 (1943), nom. inval. (Art. 39.1).
Literature: Hsieh & Goh (1990: 263), Guo & Hsieh (1995:
251), Guo et al. (1998: 262).
Illustrations: Hsieh & Goh (1990: 265, g. 204), Guo & Hsieh
(1995: 251, g. 212), Guo et al. (1998: 262, g. 218).
Fig. 47. Pseudocercospora phyllitidis (NY, lectotype). A. Conidiophore
fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.
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Description: Leaf spots amphigenous, subcircular, 7–10 mm
diam, dark brown, centre becoming grey, zonate. Caespituli
amphigenous. Mycelium internal. Stromata lacking or only few
swollen hyphal cells in the substomatal cavity. Conidiophores
fasciculate, 3–20, loose, arising from swollen substomatal
hyhal cells, emerging through stomata, erect, straight to
usually geniculate-sinuous, occasionally subnodulose,
unbranched, 60–180 × 4–7 µm, indistinctly 3–8-septate,
brown, paler towards the tip, almost hyaline at the apex, thin-
walled, smooth; conidiogenous cells integrated, terminal and
intercalary, conidiogenous loci inconspicuous to visible by
being truncate or subdenticulate. Conidia solitary obclavate,
straight to somewhat curved, 50–85 × 3–5 µm, 5–7-septate,
pale olivaceous, thin-walled, smooth, apex pointed, base
short obconically truncate, about 1.5–2 µm wide, hila neither
thickened nor darkened.
Types: Taiwan: Taipei Hsieh, Shihting, on Plagiogyria
euphlebia, Plagiogyriaceae, 6 Nov. 1927, K. Sawada (NTU-
PPE – holotype; HMAS 05180 – isotype).
Host range and distribution: Only known from the type
collection.
Pseudocercospora polypodiacearum D.N. Shukla,
A.K. Singh, P. Kumar & Kamal, Indian Phytopathol. 35:
86 (1982).
(Fig. 49)
Synonyms: Cercospora dharwarensis Rangasw., Seshadri,
Chann. & Raghun., Scheme Coll. Ident. Fungi S. India,
Final Rep: 13 (1969), nom. inval., Art. 36.1) [type: on
Lygodium exuosum, India, Karnataka, Dandeli, 25 Nov.
1968, V.S. Seshadri].
Phaeoisariopsis lygodii M.K. Khan & Kamal, Mycotaxon
54: 28 (1995) [holotype: on Lygodium exuosum, India,
South Gorakhpur Forest Division, Dec. 1990, Kamal
(HCIO 30794)].
Literature: Zhuang (2001), Kamal (2010: 210–211).
Illustration: Shukla et al. (1982: 85, g. 1).
Description: Leaf spots hypophyllous, irregularly shaped,
often effuse, olivaceous-brown. Caespituli hypophyllous.
Mycelium internal. Stromata distinct, substomatal to
subepidermal, subglobose, 10–45 µm diam, dark brown.
Conidiophores in small to rather large, loose to sometimes
dense, almost coremioid fascicles, arising from stromata,
Fig. 48. Pseudocercospora plagiogyriae (NTU-PPE, holotype). A.
Conidiophore fascicle. B. Conidia. Bar = 10 µm.
Fig. 49. Pseudocercospora polypodiacearum (K(M) IMI 242551). A.
Conidiophore fascicle. B. Conidia. Bar = 10 µm.
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324 ima fUNGUS
through stomata or erumpent, erect, straight to exuous,
geniculate-sinuous, unbranched, 60–280 × 3–7 µm,
sometimes swollen at the apex, pluriseptate thoughout,
brown, thin-walled, smooth; conidiogenous cells integrated,
terminal, 10–30 µm long, mono- to polyblastic, sympodially
proliferating, conidiogenous loci inconspicuous, unthickened,
not darkened. Conidia solitary, obclavate-cylindrical, 30–130
× 3–7.5 µm, (1–)4–8(–12)-septate, pale olivaceous to light
brown, thin-walled, smooth, apex obtuse to subacute, base
obconically truncate, occasionally somewhat peg-like, 1.5–
2.5 µm wide, hila unthickened, not darkened.
Types: India: U.P.: Gorakhpur, on leaves of Lygodium sp.,
Feb. 1980, D. N. Shukla (K(M) IMI 242551 – holotype; GPU
255 – isotype).
Host range and distribution: On Lygodium (exuosum, sp.),
Schizaeaceae, Asia (India, Karnataka, U.P.).
Notes: A Chinese collection on Lygodium japonicum was
described and illustrated by Guo & Hsieh (1995) and Guo et
al. (1998) as P. polypodiacearum. However, this collection is
morphologically quite distinct by its much shorter, 0–1-septate
and apically unswollen conidiophores, and thus better
separated and described as new species (see P. lygodiicola).
Pseudocercospora pteridicola U. Braun & Y.L. Guo,
in Braun & Mel’nik, Trudy Bot. Inst. Im. V.L. Komarova
20: 84 (1997).
(Fig. 50)
Pseudonym: Pseudocercospora pteridis sensu Guo & Hsieh
(1995: 258).
Literature: Guo & Hsieh (1995: 258), Guo et al. (1998: 268–
269).
Illustrations: Guo & Hsieh (1995: 257, g. 218), Guo et al.
(1989: 268, g. 223), Braun & Mel’nik (1997: 121, g. 50).
Description: Leaf spots amphigenous, subcircular, 1–6
mm diam, often conuent, yellowish brown, pale brown to
greyish brown, with purplish to dark brown margin. Caespituli
amphigenous, mostly hypophyllous. Mycelium internal and
external; supercial hyphae branched, septate, 1.5–2.5 µm,
subhyaline, thin-walled, smooth. Stromata lacking, small
or well-developed, 15–30 µm diam, substomatal, brown.
Conidiophores in small to large fascicles, loose to dense,
arising from stromata, emerging through stomata, or solitary,
arising from supercial hyphae, lateral, rarely terminal,
erect, straight, subcylindrical to geniculate-sinuous, usually
unbranched, rarely branched, 10–70 × 2.5–4 µm, 0–7-septate,
pale olivaceous, olivaceous-brown to medium brown, thin-
walled, smooth; conidiogenous cells integrated, terminal,
about 10–25 µm long, conidiogenous loci inconspicuous,
neither thickened nor darkened. Conidia solitary, narrowly
obclavate-subcylindrical, attenuated towards the tip, 40–125
× 2–4 µm, 2- to pluriseptate, subhyaline, thin-walled, smooth,
apex subacute, base obconically truncate, 1–2 µm wide, hila
neither thickened nor darkened.
Holotype: China: Sichuan Province: Chongqing, on Pteris
vittata, 31 Aug. 1962, Q. M. Ma & S. J. Han 1740 (HMAS
59158).
Host range and distribution: On Pteris (?biaurita, vittata),
Pteridaceae, Asia (China, ?India).
Notes: Braun & Mel’nik (1997) assigned a collection on
Pteris sp. from Colombia to P. pteridicola, which is, however,
morphologically deviating, geographically isolated, and
therefore described as P. pteridigena. “Cercospora phyllitidis”
on Pteris biaurita in India (Lall et al. 1964: 182, g. 2; Kamal
2010) and on Pteris sp. in India (Kakoti et al. 1998) belong
Fig. 50. Pseudocercospora pteridicola (HMAS 59158). A. Supercial
hypha. B. Solitary conidiophores arising from a supercial hypha.
C. Conidiophore fascicle. D. Conidiophores. E. Conidia. Bar = 10 µm.
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probably to P. pteridicola, but material was not available for a
re-examination.
Pseudocercospora pteridigena U. Braun, sp. nov.
MycoBank MB805528
(Fig. 51)
Etymology: Derived from its host genus, Pteris.
Diagnosis: Pseudocercosporae pteridicolae valde similis, sed
conidiophoris longioribus, ad 150 µm, parietibus atrioribus et
leviter incrassatis, conidiis longioribus, ad 250 µm longis.
Description: Leaf spots amphigenous, subcircular to
somewhat irregular, 3–6 mm diam, yellowish brown to
grey, with a wide dark reddish brown border. Caespituli
hypophyllous, scattered, ne, brownish. Mycelium internal
and external; supercial hyphae emerging through stomata,
sparingly branched, 1.5–3 µm wide, subhyaline to pale
olivaceous, septate, thin-walled, smooth. Stromata small,
substomatal, 10–35 µm diam, brown, composed of swollen
hyphal cells, 2–5 µm diam. Conidiophores in small fascicles,
loose to moderately dense, arising from stromata, emerging
through stomata or solitary, arising from supercial hyphae,
erect, straight, subcylindrical to somewhat sinuous or slightly
geniculate, unbranched, 30–150 × 3–5 µm, continuous to
pluriseptate, medium dark brown below, paler towards the
apex, wall somewhat thickened, smooth; conidiogenous cells
integrated, terminal, about 10–30 µm long, conidiogenous loci
unthickened, not darkened, but sometimes subdenticulate.
Conidia solitary, acicular to narrowly obclavate, 40–250 × 2–4
µm, 4-septate to pluriseptate, subhyaline to pale olivaceous,
thin-walled, smooth, apex pointed, base short obconically
truncate, 1–1.5 µm wide, hila neither thickened nor darkened.
Holotype: Colombia: Dept. Caldas, on Pteris sp., Pteridaceae,
14 Jul. 1929, C.E. Chardón (CUP).
Host range and distribution: Only known from the type
collection.
Notes: The Asian Pseudocercospora pteridicola resembles
P. pteridigena but differs in having much shorter, paler,
thin-walled conidiophores, 10–70 µm, and shorter conidia,
40–125 µm. The collection from Colombia was tentatively
assigned to P. pteridicola in Braun & Mel’nik (1987: 84), and
morpholological differences between this collection and the
Chinese type were discussed.
Pseudocercospora pteridophytophila Goh & W.H.
Hsieh, Trans. Mycol. Soc. Republ. China 4(2): 30
(1989).
(Fig. 52)
Fig. 51. Pseudocercospora pteridigena (CUP, holotype)). A. Solitary
conidiophores arising from supercial hyphae. B. Conidiophore
fascicles. C. Conidiogenous cells. D. Conidia. Bar = 10 µm.
Fig. 52. Pseudocercospora pteridophytophila (K(M) IMI 312078).
A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. A.
Conidiophore fascicle. B. Conidia. Bar = 10 µm.
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Literature: Hsieh & Goh (1990: 328), Guo & Hsieh (1995:
330), Guo et al. (1998: 344).
Illustrations: Hsieh & Goh (1990: 330, g. 250), Guo & Hsieh
(1995: 331, g. 278), Guo et al. (1998: 345, g. 283).
Description: Leaf spots amphigenous, angular-irregular, 2–4
mm diam, often conuent, dark brown, margin indenite.
Caespituli amphigenous. Mycelium internal. Stromata
substomatal, subglobose to oblong, up to 35 µm diam,
brown. Conidiophores numerous, in dense fascicles, arising
from stromata, emerging through stomata, erect, straight
to distinctly geniculate-sinuous, unbranched, 15–50 × 2–5
µm, 0–2-septate, subhyaline to very pale olivaceous, thin-
walled, smooth; conidiogenous cells integrated, terminal or
conidiophores reduced to conidiogenous cells, usually 10–25
µm long, conidiogenous loci inconspicuous to subdenticulate,
unthickened, not darkened. Conidia solitary, narrowly acicular-
liform, straight to usually curved or somewhat sinuous, 30–70
× 1–1.5 µm, 4–8-septate, colourless, thin-walled, smooth,
apex pointed, base truncate to somewhat obconically truncate,
about 1 µm wide, hila neither thickened nor darkened.
Types: Taiwan: Miaoli Hsien, Cholan, on Cyclosorus
acuminatus, Thelypteridaceae, 11 Aug. 1984, T. K. Goh
(NCHUPP-37 – holotype); K(M) IMI 312078 – isotype).
Host range and distribution: Only known from the type
collection.
Pseudocercospora ptisanae U. Braun, sp. nov.
MycoBank MB805529
(Fig. 53)
Literature: Braun & Hill (2002: 29, as Pseudocercospora cf.
rumohrae).
lllustration: Braun & Hill (2002: 27, g. 10).
Etymology: Epithet derived from its host genus, Ptisana.
Diagnosis: Pseudocercosporae rumohrae valde similis,
sed hospite divergenti (Ptisana, Marattiaceae) et hilis multo
angustioribus, 1.5–2 µm diam.
Description: Leaf spots amphigenous, shape and size variable,
brown, margin indenite. Caespituli hypophyllous, not very
conspicuous. Mycelium internal and external; supercial
hyphae emerging through stomata, branched, septate, 1–4
µm wide, thin-walled, subhyaline to pale olivaceous, smooth.
Stromata lacking. Conidiophores usually solitary, arising
from supercial hyphae, lateral, rarely terminal, occasionally
in small, loose fascicles, emerging through stomata, erect,
straight, subcylindrical to geniculate-sinuous, unbranched,
5–40 × 2–3 µm, 0–1-septate, subhyaline to pale olivaceous,
thin-walled, smooth; conidiophores usually reduced to
conidiogenous cells or conidiogenous cells sometimes
integrated, terminal, conidiogenous loci inconspicuous.
Conidia solitary, acicular to liform, 80–100 × 2–3 µm,
indistinctly pluriseptate, subhyaline to pale olivaceous, thin-
walled, smooth, apex subacute, base truncate, 1.5–2 µm
wide, hila unthickened, not darkened.
Holotype: New Zealand: Auckland, Auckland Domain,
FERNZ Fernery, on leaves of Ptisana salicina (Marattia
salicina), Marattiaceae, 6 Mar. 2005, C. F. Hill (PDD 82345),
ex-type culture: ICMP 15860.
Notes: This species was described and illustrated by Braun
& Hill (2002) as Pseudocercospora cf. rumohrae, based on
a collection from 2000 (New Zealand, Auckland, Auckland
University, 30 Apr. 2000, C.F. Hill 185). This collection,
originally deposited at HAL, is not maintained. It was lost
in June 2013 during a big ood that caused damage in the
herbarium HAL. Pseudocercospora on Ptisana salicina is,
indeed, morphologically very close to P. rumohrae, but it
occurs on a fern of another family, and the conidia and hila
are distinctly narrower.
Pseudocercospora rumohrae W.H. Hsieh & Goh,
Trans. Mycol. Soc. Republ. China 4(2): 29 (1989).
(Fig. 54)
Literature: Hsieh & Goh (1990: 37), Guo & Hsieh (1995: 31),
Guo et al. (1998: 44).
Illustrations: Hsieh & Goh (1990: 37, g. 22), Guo & Hsieh
(1995: 33, g. 31), Guo et al. (1998: 44, g. 31).
Fig. 53. Pseudocercospora ptisanae (HAL, CF Hill 185). A. Supercial
hyphae. B. Solitary conidiophores arising from supercial hyphae. C.
Conidiophore fascicle. D. Conidia. Bar = 10 µm.
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Description: Leaf spots amphigenous, angular-irregular, size
variable, centre brown, yellowish or paler brown towards the
periphery, margin indenite. Colonies hypophyllous, not very
conspicuous. Mycelium internal and external, supercial
hyphae emerging through stomata; branched, septate, very
pale olivaceous, thin-walled, smooth. Stromata lacking.
Conidiophores solitary, arising from supercial hyphae,
occasionally a few conidiophores emerging through stomata,
forming small, loose fascicles, erect, straight to slightly curved,
subcylindrical to slightly clavate, at most slightly geniculate-
sinuous, 2–25 × 2.5–4 µm, 0–2-septate, subhyaline to pale
brownish, thin-walled, smooth; conidiogenous cells integrated,
terminal or conidiophores reduced to conidiogenous cells, 2–20
µm long, conidiogenous loci inconspicuous, neither thickened
nor darkened. Conidia solitary, acicular, 60–120 × 3–3.5 µm,
indistinctly pluriseptate, hyaline to very pale olivaceous, thin-
walled, smooth, apex acute or subacute, base truncate, about
2.5–3 µm wide, hila neither thickened nor darkened.
Types: Taiwan: Yilan, on Rumohra adiantiformis,
Dryopteridaceae, 4 Dec. 1985, W. H. Hsieh (NCHUPP-228 –
holotype; K(M) IMI 312068 – isotype).
Host range and distribution: Only known from the type
collection.
Pseudocercospora thelypteridis Goh & W.H. Hsieh,
Trans. Mycol. Soc. Republ. China 4(2): 30 (1989).
(Fig. 55)
Literature: Hsieh & Goh (1990: 329), Guo & Hsieh (1995:
330), Guo et al. (1998: 345).
Illustrations: Hsieh & Goh (1990: 331, g. 251), Guo & Hsieh
(1995: 332, g. 279), Guo et al. (1998: 345, g. 284).
Fig. 54. Pseudocercospora rumohrae (K(M) IMI 312068). A. Solitary
conidiophores arising from supercial hyphae. B. Conidiophores and
hyphae emerging through a stoma. C. Conidia. Bar = 10 µm.
Fig. 55. Pseudocercospora thelypteridis (NCHUPP-227). A.
Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
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Description: Leaf spots amphigenous, angular-irregular, up to
3 mm diam, brown, margin indenite. Caespituli epiphyllous,
punctiform, dark brown to blackish. Mycelium internal.
Stromata immersed, irregularly shaped, 20–35 µm diam,
dark brown. Conidiophores numerous, densely fasciculate,
arising from stromata, erumpent, erect, straight to curved,
cylindrical-liform, not geniculate, simple or sometimes
branched, somewhat narrowed towards the apex, 10–60
× 2–3 µm, continuous, rarely septate, subhyaline to pale
olivaceous, paler towards the apex, tips rounded to truncate,
thin-walled, smooth; conidiogenous cells integrated,
terminal or conidiophores reduced to conidiogenous cells,
conidiogenous loci inconspicuous. Conidia solitary, acicular-
liform, 50–120 × 2.5–3 µm, 5–12-septate, subhyaline to pale
olivaceous, thin-walled, smooth, apex pointed, base truncate
or subtruncate, about 1.5–2 µm wide, hila neither thickened
nor darkened.
Holotype: Taiwan: Taipei, Yangmingshan, on Metathelypteris
laxa (Thelypteris laxa), Thelypteridaceae, 30 Aug. 1986, T. K.
Goh (NCHUPP-227).
Host range and distribution: Only known from the type
collection.
Zasmidium
A single species.
Zasmidium australiense (J.L. Mulder) U. Braun &
Crous, comb. nov.
MycoBank MB805530
(Fig. 56)
Basionym: Stenella australiense J.L. Mulder, Mycol. Res. 92:
119 (1989).
lllustration: Mulder (1989: 119–121, gs 1–6).
Description: Leaf spots lacking, attacked areas only
chlorotic corresponding to the areas delimited by veins,
similar chlorotic areas also on the upper surface. Colonies
hypophyllous, effuse, forming greyish brown patches.
Mycelium internal and external; supercial hyphae forming
mycelial mats or a loose reticulum on the leaf surface,
becoming darker brown with age, branched, septate, rarely
constricted at septa, about 4–5 µm wide, strongly echinulate,
rarely smooth. Stromata absent. Conidiophores solitary,
arising from supercial hyphae, lateral, rarely terminal, erect,
straight, subcylindrical, usually unbranched, at most apically
somewhat geniculate, up to 160 × 3.5–5.5 µm, septate, dark
brown, conspicuously echinulate with crested type projections,
denser than in conidia; conidiogenous cells integrated,
terminal, proliferation sympodial, occasionally percurrent;
conidiogenous loci distinct, non-protuberant, thickened and
darkened-refractive, about 1–1.5 µm diam. Conidia solitary,
cylindrical, straight to distinctly curved, about 30–70 ×
3–5.5 µm, (1–)3–4(–5)-septate, brown, wall thin, echinulate
(crested projections), less dense than in conidiophores and
lacking around hila, apex obtuse, rounded, base rounded to
truncate, hila slightly thickened and darkened.
Holotype: Australia: Peregian beach, on leaves of Blechnum
serrulatum (B. indicum), Blechnaceae, 3 Dec. 1982, J. L.
Alcorn 8240b (K(M) IMI 273944).
Host range and distribution: Only known from the type
collection.
Notes: Due to scolecosporous conidia formed singly and
truncate scars and hila, this species is reallocated to
Zasmidium.
Fig. 56. Zasmidium australiense (K(M) IMI 273944). A. Conidiophores
arising from supercial hyphae. B. Conidia. Bar = 10 µm.
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CERCOSPOROID SPECIES ON GYMNOSPERMAE
Passalora species on Gymnospermae
Key to the species of Passalora on Gymnospermae
1 Stromata 20–80 µm diam; conidiophores fasciculate, 20–90 µm long, 2–5-septate; conidia 30–65 × 4–6 µm,
1–6-septate; on hosts of various genera of the Cupressaceae, including Juniperus, Asia, North and
South America ......................................................................................................................................... P. sequoiae
Stromata larger, up to 150 µm wide and 120 µm deep; conidiophores in sporodochial conidiomata, shorter,
about 10–40 µm long, 0–1-septate; conidia about 15–45 × 4–5 µm, 0–4-septate; on Juniperus communis,
Europe, North America .......................................................................................................................... P. juniperina
List of Passalora species on Gymnospermae
Passalora juniperina (Georgescu & Badea) H.
Solheim, Agarica 33: 78 (2013).
(Fig. 57)
Basionym: Cercospora juniperina Georgescu & Badea,
Analele Inst. Cercet. Exp. Forest., Ser. 1, 2: 46 (1937).
Synonyms: Stigmina juniperina (Georgescu & Badea) M.B.
Ellis, Mycol. Pap. 72: 67 (1959).
Asperisporium juniperinum (Georgescu & Badea) B. Sutton &
Hodges, Mycologia 82: 317 (1990).
Literature: Jaczewski (1901), Kujala (1950), Chupp (1954:
440), Hodges (1962: 66), Ellis (1976: 119), Kurkela (1994),
Sutton & Hodges (1990), Crous & Braun (2003: 233),
Barklund (2006), Solheim (2013).
Illustrations: Ellis (1976: 120, g. 84B), Hodges (1962: 65,
g. 2C), Sutton & Hodges (1990: 318, g. 2), Solheim (2013:
gs 2–5).
Exsiccatae: Săvul., Herb. Mycol. Rom. 1200 (“Exosporium
deectens”). Scheuer, Mycoth. Graec. 249.
Description: Causing needle blight, attacked hosts only
partly with dead needles or needles almost completely
necrotic, needles at rst light brown, later greyish brown to
grey. Colonies amphigenous, mainly epiphyllous, scattered
or in lines, dark brown to blackish, punctiform to pustulate.
Mycelium internal, usually epidermal and subepidermal;
hyphae branched, septate, 2–6 µm wide, brown, thin-walled,
smooth. Stromata substomatal, intraepidermal to deeply
immersed, nally also erumpent, often expended and large,
up to 150 µm diam and 120 µm deep, medium to darker
brown, composed of dense, rounded to mostly angular cells,
about 2.5–8 µm diam, wall somewhat thickened. Conidiomata
sporodochial, composed of large stromata and numerous,
densely arranged conidiophores, emerging through stomata
or erumpent, erect, straight to slightly curved, subcylindrical,
unbranched, (5–)10–35(–40) × 3–7(–9) µm, 0–1-septate,
olivaceous-brown, yellowish brown, thin-walled, smooth or
almost so; conidiophores mostly reduced to conidiogenous
cells or conidiogenous cells integrated, terminal, proliferation
sympodial, occasionally percurrent, conidiogenous loci 1–3,
rather inconspicuous to conspicuous, slightly thickened
and somewhat refractive, little darkened, (1.5–)2(–2.2) µm
wide. Conidia solitary, cylindrical to obclavate-cylindrical of
fusiform, 15–45 × (3–)4–5(–6.5) µm, 0–3(–4)-septate, without
constrictions at septa, olivaceous to yellowish brown, thin-
walled, delicately verruculose, apex obtuse, base rounded to
short obconically truncate, 1.5–2.2 µm wide, hila barely or
only slightly thickened, refractive, barely darkened.
Types: Romania: on Juniperus communis, Georgescu
& Badea [holotype not seen, probably not maintained];
Romania: Sinaia, Distr. Prahova, Muntenia, on Juniperus
communis, 10 May 1937, T. Savulescu & C. Sandu, Herb.
Mycol. Rom. 1200, as “Exosporium deectens” (K(M) IMI
84601 – neotype designated here, MycoBank, MBT176149).
Host range and distribution: on Juniperus communis [var.
communis, var. depressa (J. intermedia), var. saxatilis (J.
nana)], Cupressaceae, Europe (Finland, Norway, Romania,
Sweden, Russia), North America (Canada, Ontario; USA,
Iowa, Michigan, Wisconsin).
Note: Material recently collected in Russia, Novgorod Oblast,
2002, by V.A. Mel’nik has been distributed in Mycotheca
Graecensis 249 (e.g. BPI 858449, GZU, HAL).
Passalora sequoiae (Ellis & Everh.) Y.L. Guo & W.H.
Hsieh, Fl. Fung. Sin. 20: 120 (2003).
(Fig. 58)
Basionym: Cercospora sequoiae Ellis & Everh., J. Mycol.
3(2): 13 (1887).
Synonyms: Asperisporium sequoiae (Ellis & Everh.) B. Sutton
& Hodges, Mycologia 82: 314 (1990).
Cercosporidium sequoiae (Ellis & Everh.) W.A. Baker &
Partr., Mycotaxon 76: 250 (2000).
Cercospora cryptomeriae Shirai, in Kitajima, Bot. Mag. Tokyo
30: 412 (1916).
Cercospora thujina Plakidas, Phytopathology 35: 181 (1945)
[syntypes: on Platycladus orientalis, USA, Louisiana,
Baton Rouge, 3 June 1943, A.G. Plakidas, BPI 441926A–
C, and 25 Jul. 1943, BPI 441925A–C].
Literature: Saccardo (1892: 653), Kitajima (1916: 412),
Chupp (1954: 439), Hodges (1962: 62–63), Katsuki (1965:
51), Mulder & Gibson (1973), Hsieh & Goh (1990: 326),
Sutton & Hodges (1990: 314–315), Baker et al. (2000: 250),
Crous & Braun 2003: 373).
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Illustrations: Hodges (1962: 63, gs. 1A, 2A), Hsieh & Goh
(1990: 326, g. 247), Sutton & Hodges (1990: 315, g. 1),
Baker et al. (2000: 251, g. 1), Guo et al. (2003: 121, g. 78).
Exsiccatae: Davis, Fungi Wiscon. Exs. 25. Ellis & Everh.,
North Amer. Fungi 1754.
Description: Leaves (needles, scale leaves) at rst with small,
subcircular lesions, yellowish brown to medium reddish or
dark brown, scattered, later enlarged and conuent, nally
entire leaves becoming necrotic, brown or greyish brown;
twig symptoms visible as pale brown, irregular lesions,
later enlarged and conuent, nally turning reddish brown
towards the tips. Caespituli punctiform-pustulate, scattered,
brown or visible as dark greyish masses of conidia. Mycelium
internal. Stromata substomatal to immersed, intraepidermal
to deeply immersed, erumpent, subglobose to hemispherical
or somewhat irregular, 25–100 µm diam, dark brown,
composed of swollen hyphal cells, rounded to isodiametric,
2.5–8 µm diam. Conidiophores in small to usually large
fascicles, loose to usually dense, arising from stromata,
through stomata or erumpent, erect, straight to curved or
frequently geniculate-sinuous, above all in the upper half,
unbranched, 20–90 × 2.5–8 µm, 2–5-septate, medium brown
to dark brown throughout or paler towards the tip, olivaceous-
brown, thin-walled, smooth or almost so, occasionally
somewhat verruculose; conidiogenous cells integrated,
terminal, about 10–40 µm long, polyblastic, proliferating
sympodially, rarely percurrently, with 1–6 distinctly thickened
and darkened conidiogenous loci, 2–2.5 µm diam, mostly on
shoulders formed by sympodial proliferation. Conidia solitary,
obclavate-cylindrical, straight to somewhat curved, 30–80 ×
4–9 µm, 1–15-septate, occasionaly somewhat constricted at
the septa, yellowish to pale olivaceous-brown or brown, thin-
walled, verruculose-echinulate, apex obtuse, rounded, base
short obconically truncate, about 2–2.5 µm wide, hila slightly
thickened and darkened.
Lectotype (designated here, MycoBank, MBT176150): USA,
Pennsylvania: Germantown, on Sequoiadendron giganteum,
Sep. 1886, Meehan Thomas H., Ellis & Everh., North Amer.
Fungi 1754 (BPI 441133). Isolectotypes: Ellis & Everh., North
Amer. Fungi 1754.
Fig. 57. Passalora juniperina (K(M) IMI 84601). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm. Fig. 58. Passalora sequoia (BPI 441133). A. Conidiophore fascicle.
B. Conidiophore tips. C. Conidia. Bar = 10 µm.
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Host range and distribution: On Cryptomeria japonica,
×Cupressocyparis leylandica, Cupressus (lusitanica,
macrocarpa, sempervirens), Glyptostrobus pensilis,
Hesperocyparis arizonica (Cupressus arizonica), Juniperus
(communis, chinensis, virginiana), Platycladus orientalis
(Thuja orientalis), Sequoiadendron giganteum, Sequoia
sempervirens, Taxodium (distichum, mucronatum),
Cupressaceae, Asia (China, Japan), North America (Canada;
USA, Alabama, North Carolina, Louisiana, Oklahoma,
Pennsylvania), South America (Brazil, Guatemala),
Caribbean (Jamaica), Hawaii.
Notes: A specimen on Cryptomeria japonica from Taiwan
(24 Sep. 1928, K. Sawada, TNS-F220395) and various
Japanese collections on C. japonica (IUM-FS58, IUM-
FS61, MUMH10801, TFM:FPH-41, TFM:FPH-67) and
Taxodium mucronatum (TFM:FPH-3395) have been re-
examined. Furthermore, cultures ex Cryptomeria japonica
(MAFF237184, MAFF410035, MAFF410800, MAFF 410802,
MUCC780 ex MUMH 10801) and ex Taxodium mucronatum
(MAFF410034) exist. ITS1-5.8S-ITS2 (V9G/ITS4) & ACT
(ACT512F/ACT783R) sequences obtained from these
isolates are identical, suggesting that this fungus, at least
in Japan, represents a uniform species. Sequences derived
from cultures of Cercospora juniperina cluster close to P.
sequoiae together with various other Passalora species,
which supports the position of the two species in Passalora,
at least tentatively as the latter genus is not monophyletic.
In Japan, the causal agent of needle blight of Cryptomeria
japonica was described by Shirai (in Kitajima 1916) as
Cercospora cryptomeriae, which was already reduced to
synonym with Cercospora sequoiae by Ito et al. (1958, 1967)
based on agreeing morphological characteristics.
Pseudocercospora species on Gymnospermae
Key to the species of Pseudocercospora on Gymnospermae
1 Conidia very broad, 5–8 µm ............................................................................................................................................ 2
Conidia narrower, 2–5 µm ............................................................................................................................................... 5
2 (1) Conidiophores very long, 50–120 µm; on Cryptomeria, Japan ........................................................... P. cryptomeriicola
Conidiophores much shorter, 10–60 µm; on other hosts ................................................................................................ 3
3 (2) Conidia very long and broad, 65–135 × 9–13 µm; on twigs of Pinus, Japan .................. [Scolecostigmina chibaensis]
Conidia shorter and narrower, 18–55 × 2–8 µm; on leaves ............................................................................................ 4
4 (3) Stromata well-developed, 150–250 µm diam and 70–130 µm deep or oblong, up to 500 µm in length;
conidia smooth; on Thuja plicata, North America, and Chaemaecyparis lawsoniana, Hawaii .................... P. thujina
Stromata lacking or small, up to 30 µm diam; conidia rough-walled; on Chamaecyparis obtusa, Japan
................................................................................................................................................... P. chamaecyparidis
5 (1) Stromata almost lacking or relatively small, 10–50 µm; conidiophores fasciculate ........................................................ 6
Stromata large, 20–150(–300) µm diam; conidiomata sporodochial .............................................................................. 9
6 (5) Stromata lacking or small, up to 30 µm; conidiophores usually percurrently proliferating,
with up to ve ne annellations; conidia 18–50 µm long and 5–10-septate;
on Chamaecyparis obtusa, Japan ............................................................................................ P. chamaecyparidis
Stromata 20–50 µm diam; conidiophores sympodially proliferating; conidia mostly longer,
exceeding 50 µm; on other hosts ............................................................................................................................. 7
7 (6) Conidiophores often branched; conidia pale brown; leaf lesions characteristically marginal;
on Ginkgo biloba, Taiwan ..................................................................................................................... P. ginkgoana
Conidiophores usually unbranched; conidia subhyaline to pale olivaceous; lesions not conned
to leaf margins; on other hosts ................................................................................................................................. 8
8 (7) Stromata 20–50 µm diam; on Sequoia sempervirens (in inoculation experiments also on Larix kaempferi),
Japan ..................................................................................................................................... P. paraexosporioides
Stromata 20–150 µm diam; on Pinus spp., Africa, Asia, Central and South America, Caribbean ...... P. pini-densiorae
9 (5) Conidia acicular or narrowly cylindrical, 35–93 × 2–3 µm, only 3–5-septate, base truncate;
on Sciadopitys verticillata, Japan ..................................................................................................... P. sciadopytios
Conidia cylindrical or obclavate-cylindrical, shorter, (10–)20–65(–80) µm long, at least partly short
obconically truncate at the base; on other hosts .................................................................................................... 10
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10 (9) Conidiogenous cells sympodially and often also percurrently proliferating, with ne annellations;
conidia more or less cylindrical, base truncate to short obconically truncate; on Cryptomeria,
Hesperocyparis and Juniperus, Cupressaceae, Asia and North America ................................................. P. juniperi
Conidiophores sympodially proliferating, without annellations; conidia short cylindrical or
obclavate-cylindrical, base short obconically truncate; on hosts of the Pinaceae .................................................. 11
11 (10) Conidia cylindrical, relatively short, (15–)20–40(–50) µm; on Larix decidua, Europe ........................... P. exosporioides
Conidia obclavate-cylindrical, usually (10–)20–60(–80) µm long; on Pinus spp., Africa, Asia,
Central and South America, Caribbean ....................................................................................... P. pini-densiorae
Tabular key to the species of Pseudocercospora on Gymnospermae
Cupressaceae
1 Conidia very broad, 5–8 µm ............................................................................................................................................ 2
Conidia narrower, 2–5 µm ............................................................................................................................................... 4
2 (1) Conidiophores very long, 50–120 µm; on Cryptomeria, Japan ............................................................ P. cryptomeriicola
Conidiophores much shorter, 10–60 µm; on other hosts ................................................................................................ 3
3 (2) Stromata well-developed, 150–230 µm diam and 70–120 µm deep; conidia smooth; on Thuja plicata,
North America ............................................................................................................................................. P. thujina
Stromata lacking or small, up to 30 µm diam; conidia rough-walled; on Chamaecyparis obtusa, Japan
.................................................................................................................................................... P. chamaecyparidis
4 (1) Stromata lacking or small, up to 30 µm; conidiophores fasciculate, usually percurrently proliferating,
with up to ve ne annellations; conidia cylindrical to obclavate, 5–10-septate;
on Chamaecyparis obtusa, Japan ............................................................................................ P. chamaecyparidis
Stromata large, 30–150 µm diam; conidiomata sporodochial; conidiogenous cells sympodially and
often also percurrently proliferating, with ne annellations; conidia more or less cylindrical,
3–5-septate, base truncate to short obconically truncate; on Cryptomeria, Hesperocyparis
and Juniperus, Cupressaceae, Asia and North America ........................................................................... P. juniperi
Ginkgoaceae
On Ginkgo biloba, Taiwan ........................................................................................................................................ P. ginkgoana
Pinaceae
1 Conidia very long and broad, 65–135 × 9–13 µm; on twigs of Pinus, Japan .................. [Scolecostigmina chibaensis]
Conidia narrower, 2–5 µm; on leaves ............................................................................................................................. 2
2 (1) Conidia cylindrical, relatively short, (15–)20–40(–50); on Larix decidua, Europe ................................. P. exosporioides
Conidia obclavate-cylindrical, longer, (10–)20–80 µm; on Pinus (or Larix but only in inoculation experiments .............. 3
3 (2) Stromata 20–150 µm diam; on Pinus spp., Africa, Asia, Central and South America, Caribbean
.......................................................................................................................................................P. pini-densiorae
Stromata 20–50 µm diam; usually on Sequoia sempervirens, Cupressaceae (in inoculation
experiments also on Larix kaempferi), Japan ........................................................................ P. paraexosporioides
Sciadopityaceae
On Sciadopitys verticillata, Japan ........................................................................................................................ P. sciadopytios
List of Pseudocercospora species on
Gymnospermae
Pseudocercospora chamaecyparidis (Sawada) C.
Nakash. & U. Braun, comb. nov.
MycoBank MB805531
(Fig. 59)
Basionym: Cercospora chamaecyparidis Sawada, Bull. Gov.
Forest Exp. Sta., Meguro, Tokyo 45: 122 (1950).
Description: Leaves turning pale brown to brown, nally
grey. Caespituli hypophyllous, formed as scattered, blackish
sporodochia, erumpent. Stromata lacking or small, up to 30
µm, brown to blackish brown. Conidiophores fasciculate,
arising from internal mycelium or stromata, straight,
subcylindrical, 10–50 × 2.5–5 μm, aseptate or septate, brown
to blackish brown, wall thin or only slightly thickened, smooth;
conidiogenous cells integrated, terminal, proliferating
percurently or occasionally sympodially, often with
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volume 4 · no. 2
annellations, loci thin, unthickened to somewhat protuberant.
Conidia solitary, cylindrical to obclavate, straight, curved or
sigmoid, 18–50 × 2–8 μm, 5–10-septate, brown to reddish
brown, smooth to faintly rough-walled (light microscopy),
verruculose under SEM, thin, apex obtuse, rounded, base
truncate and unthickened, 2–3.8 µm wide.
Holotype: Japan: Fukushima, on Chamaecyparis obtusa, 22
Dec. 1949, H. Ootomo (IUM-FS57).
Host range and distribution: On Chamaecyparis obtusa,
Cupressaceae, Asia (Japan).
Notes: Cercospora chamaecyparidis was described by
Sawada based on Japanese material on Chamaecyparis
obtusa. The type material, which has been re-examined,
is morphologically close to the North American Pseudo-
cercospora thujina (Stigmina thujina) on Thuja plicata, but
beside a different host it is distinguished by having much
smaller or even lacking stromata. Thus it is better to retain
this fungus as separate species.
Pseudocercospora cryptomeriicola (Sawada)
C. Nakash., Akashi & Akiba, in Nakashima et al.,
Mycoscience 48: 254 (2007).
(Fig. 60)
Basionym: Cercospora cryptomeriicola Sawada, Bull. Gov.
Forest, Exp. Sta. (Tokyo) 45: 53 (1950).
Synonym: Distocercospora cryptomeriicola (Sawada) Akashi,
C. Nakash. & Tak. Kobay., Abstr. Jap. Forest. Soc.: 117
(2006), nom. inal. (Art. 29.1).
Literature: Katsuki (1965: 51), Crous & Braun (2003: 146).
Illustrations: Nakashima et al. (2007: 252, g. 2; 253, gs
3–12).
Description: Causing black line needle blight of Cryptomeria,
leaves and stems discoloured, yellowish to brown, at rst with
smaller spots, nally entire needles affected. Colonies visible
as fungal masses consisting of olivaceous conidiophores
and conidia in lines from base to top on necrotic needles.
Mycelium internal. Stromata well-developed, immersed,
later erumpent, subglobose, about 90–275 µm diam, dark
olivaceous-brown to black, composed of swollen hyphal
cells, rounded to angular or oblong. Conidiophores in large,
loose to dense fascicles, arising from stromata, forming
sporodochial conidiomata, erumpent, simple or rarely
branched, subcylindrical, straight to slightly sinuous, 50–120
× 3–5 µm, 1–7-septate, dark brown to blackish, wall smooth,
Fig. 59. Pseudocercospora chamaecyparidis (IUM-F557). a.
Conidium (light microscopy). b. Conidium (SEM). c. Conidiophores
(SEM). Bar = 10 µm (a), 5 µm (b) and 2 µm (c).
Fig. 60. Pseudocercospora cryptomeriicola (TFM:FPH-1085). A.
Conidiophore fascicle. B. Conidiophores. C. Conidia. Bars = 10 µm.
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334 ima fUNGUS
slightly thickened; conidiogenous cells integrated, terminal,
percurently proliferating, with conspicuous annellations,
conidiogenous loci neither thickened nor darkened. Conidia
solitary, cylindrical-obclavate, straight to slightly curved, 50–
80 × 5–8 µm, 7–13-septate, occasionally distosepta mixed
with eusepta, pale brown to brown, wall somewhat thickened,
smooth or almost so, apex rounded, base truncate to
obconically truncate, 2.5–3.8 µm wide, hila neither thickened
nor darkened.
Holotype: Japan: Yamagata, Kamabuchi, on Cryptomeria
japonica, 8 Aug. 1949, Sato (Herbarium, Museum of Iwate
University, Japan).
Host range and distribution: On leaves and stems of
Cryptomeria japonica, Japan, Cupressaceae, Asia (Japan,
endemic).
Notes: A culture (NBRC 102150) and several collections
of this species, which is endemic in Japan, have been
examined (TFM:FPH-1085, 1088, 7850 and 7851). Due to
unthickened conidiogenous loci and conidial hila as well as
its proven phylogenetic position in the Pseudocercospora
clade, Nakashima et al. (2007) reallocated this species to
Pseudocercospora, although the formation of well-developed
sporodochia, percurrently proliferating conidiogenous cells
and large, above all broad, dark and thick-walled conidia are
rather Scolecostigmina-like.
Pseudocercospora exosporioides (Bubák) B. Sutton
& Hodges, Mycologia 82: 320 (1990).
(Fig. 61)
Basionym: Cercospora exosporioides Bubák, Ann. Mycol.
13: 33 (1915).
Literature: Saccardo (1931: 873), Vassiljevsky & Karakulin
(1937: 252), Chupp (1954: 439), Crous & Braun (2003: 181).
Illustration: Sutton & Hodges (1990: 321, g. 4).
Exsiccatae: Petrak, Fl. Bohem. Morav. Exs. 1351.
Description: Causing needle blight, at rst forming small
brown to reddish brown discolorations, later expanded, often
at tips, nally entire needles discoloured, necrotic, dead
and shed. Colonies amphigenous, punctiform to pustulate,
dark. Mycelium internal; hyphae branched, septate, 2–10
µm diam, brown, smooth or almost so. Stromata well-
developed, immersed, large, sometimes conuent, about
50–150 µm diam, sometimes up to 300 µm in length, dark
brown, sometimes with a reddish tinge, composed of swollen
hyphal cells, 2–8 µm diam, rounded in outline to angular.
Conidiophores numerous to very numerous, arising from
stromata, erumpent, forming large, dense sporodochial
conidiomata, erect, straight to somewhat curved-sinuous,
subcylindrical to somewhat conical (attenuated towards
the tip), not, barely or only slightly geniculate, unbranched,
5–30 × 2–4.5 µm, 0–1(–2)-septate, pale yellowish to
olivaceous-brown throughout or paler towards the tip, thin-
walled, smooth or almost so; conidiophores reduced to
conidiogenous cells or conidiogenous cells integrated,
terminal, 5–25 µm long, conidiogenous loci inconspicuous,
neither thickened nor darkened. Conidia solitary, cylindrical to
slightly obclavate-cylindrical, somewhat attenuated towards
the tip, straight to slightly curved, (15–)20–40(–50) × 2–3.5
µm, (1–)3–4(–5)-septate, subhyaline to pale olivaceous, thin-
walled, smooth or almost so to somewhat rough-walled, apex
obtuse, base short obconically truncate, 1.5–2 µm wide, hila
neither thickened nor darkened.
Lectotype (designated here, MycoBank, MBT176151):
Czech Republic: Moravia, Hranice (Mährisch Weisskirchen),
Ludinabach, on needles of Larix decidua, 8 Oct. 1914, F.
Petrak (BPI 436251); isolectotypes: K(M) IMI 24163, LEP;
Petr., Fl. Bohem. Morav. Exs. 1351, e.g. HBG.
Host range and distribution: On needles of Larix decidua,
Pinaceae, Europe (Czech Republic).
Fig. 61. Pseudocercospora exosporioides (LEP, isolectotype).
A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
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Notes: The material deposited at BPI is marked as “holotype”,
but there are several duplicates of the type collection
(syntypes) housed in several other herbaria. Therefore, it is
proposed to designate the BPI material as lectotype.
Pseudocercospora ginkgoana R. Kirschner, Mycol.
Progr. 12: 423 (2013).
(Fig. 62)
Illustration: Kirschner & Okuda (2013: 423, g. 1A–E).
Description: Lesions marginal, visible as necrotic, brown leaf
margins with yellowish border towards the healthy green leaf
blade. Caespituli amphigenous. Mycelium internal; hyphae
intercellular, 2–3 µm wide, hyaline to pale brown, smooth.
Stromata developed, amphigenous, about 30–40 µm diam,
immersed, erumpent through the epidermis, composed of
brown swollen hyphal cells, 3–11 × 3–5 µm. Conidiophores
numerous, in dense fascicles, arising from stromata, erumpent,
erect, straight, subcylindrical, somewat curved to distinctly
geniculate, simple or branched, (18–)26–57(–78) × 3–5(–7)
µm, continuous or 1- to pluriseptate, pale to medium brown,
thin-walled, smooth; conidiophores reduced to conidiogenous
cells or conidiogenous cells integrated, terminal, occasionally
pleurogenous, (8–)9.5–16.5(–21) × 2.5–4 µm, conidiogenous
loci inconspicuous to subdenticulate, truncate and 1–1.5 µm
wide, but always unthickened and not darkened. Conidia
solitary, subcylindrical to narrowly obclavate, i.e. narrowed
towards the apex and obconically truncate at the base,
straight or occasionally curved, (11–)33–65(–75) × 2–2.5(–
3) µm, 1–9-septate, pale brown, thin-walled, smooth, apex
obtuse to subacute, base short obconically truncate, 1.5 µm
wide, hila neither thickened nor darkened.
Holotype: Taiwan: Taoyuan County, Lasashan, New Century
Resort Farm with tea and fruit plantations, ca. 1,500 m alt.,
on leaves of Ginkgo biloba, 22 Oct. 2011, R. Kirschner et al.
3561 (TNM).
Host range and distribution: On leaves of Ginkgo biloba,
Ginkgoaceae, Asia (Taiwan).
Notes: This is the rst Pseudocercospora on Ginkgo. Kirschner
& Okuda (2013) cultivated this fungus, and derived ITS DNA
sequence data. A comparision showed a 99 % similarity with
sequences of Pseudocercospora chiangmaiensis.
Pseudocercospora juniperi (Ellis & Everh.) B. Sutton
& Hodges, Mycologia 82: 319 (1990).
(Fig. 63)
Basionym: Cercospora sequoiae var. juniperi Ellis & Everh.,
J. Mycol. 3: 14 (1987).
Synonym: Pseudocercospora ibusukii C. Nakash. & T. Kobay.
(in Kobayashi 2007: 775), nom. inval. (Art. 30.8).
Literature: Saccardo (1892: 653), Chupp (1954: 440),
Peterson & Wysong (1968), Peterson (1977), Crous & Braun
(2003: 234).
Illustration: Sutton & Hodges (1990: 320, g. 3).
Description: Causing needle blight, with discolorations and
leaf spots, yellowish to brown, nally entire leaves discoloured,
necrotic; often occurring in young plantations and windbreaks.
Colonies punctiform to pustulate, paler or dark by abundant
conidial formation. Mycelium internal; hyphae mainly
intraepidermal, but also in the mesophyll, branched, septate,
2–5 µm wide, brown. Stromata well-developed, substomatal
to deeply immersed, about 30–150 µm diam, medium to dark
brown, composed of swollen hyphal cells, about 3–6 µm
diam, wall somewhat thickened. Conidiophores in small to
usually larger, often almost sporodochial fascicles, loose to
moderately dense, arising from stromata, through stomata
or erumpent, erect, straight, subcylindrical to somewhat
geniculate-sinuous, unbranched or rarely branched, 15–
40 × 2.5–3.5 µm, 0–3-septate, olivaceous-brown, paler
towards the tip, thin-walled, smooth; conidiogenous cells
integrated, terminal, 10–25 µm long, proliferation sympodial
or percurrent, with ne annellations, conidiogenous loci
Fig. 62. Pseudocercospora ginkgoana (TNM, holotype). A.
Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
Braun et al.
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truncate, neither thickened nor darkened. Conidia solitary,
cylindrical, subcylindrical to slightly obclavate-cylindrical,
straight to somewhat curved, 30–60 × 2.5–3 µm, 3–5-septate,
not constricted at the septa, pale olivaceous to olivaceous-
brown in mass, thin-walled, almost smooth to rough-walled,
apex obtuse, base truncate or slightly obconically truncate,
about 2 µm wide, hila unthickened, not darkened.
Holotype: USA: Wisconsin: Kenosha Co., Power’s Lake, on
needles of Juniperus virginiana, Aug. 1886, J. J. Davis (NY).
Host range and distribution: On Cryptomeria japonica,
Hesperocyparis arizonica (Cupressus arizonica), Juniperus
(communis, virginiana, Juniperus sp.), Cupressaceae, Asia
(Japan), North America (Canada, Ontario; USA, Arizona,
Connecticut, Florida, Georgia, Kentucky, Missouri, North
Carolina, Nebraska, Oklahoma, South Carolina, Virginia,
Wisconsin).
Notes: This species was previously confused with Cercospora
juniperina (see Chupp 1954), but it is easily distinguishable
from the latter species by having unthickened, not darkened
conidiogenous loci as pointed out by Sutton & Hodges (1990).
Pseudocercospora paraexosporioides C. Nakash.
& U. Braun, sp. nov.
MycoBank MB805532
(Fig. 64)
Etymology: Named for its resemblance to Pseudocercospora
exosporioides.
Pseudonym: Cercospora exosporioides sensu auct.
Literature: Katsuki (1965: 51).
Diagnosis: Differt a P. exosporioides stromatibus minoribus,
20–50 µm diam, conidiophoris in fasciculis minoribus et
conidiis longioribus, 25–80 µm, pluriseptatis (3–10).
Description: Leaf spot scattered, brown, later enlarged and
conuent, yellowish brown to reddish brown or dark brown.
Caespituli hypophyllous, punctiform, dark olivaceous.
Mycelium internal. Stromata hypophyllous, substomatal to
immersed, pale to dark brown, 20–50 µm diam. Conidiophores
in small to moderately large, dense fascicles, arising from
stromata, through stomata or erumpent, straight, subcylindrical
or narrower towards the apex to geniculate, unbranched, 22–
48 × 2–3 µm, 0–2-septate, pale to pale brown, thin-walled,
smooth; conidiogenous cells integrated, terminal, proliferating
sympodially, conidiogenous loci truncate, unthickened, not
darkened. Conidia solitary, cylindrical to obclavate, straight
or mildly curved, 25–80 × 2–4 µm, 3–10-septate, pale to pale
olivaceous, thin-walled, smooth to rough-walled, apex obtuse
to subacute, base obconically truncate, about 2–2.5 µm wide,
hila unthickened, not darkened.
Holotype: Japan: Tokushima Pref.: on leaves of Sequoia
sempervirens, 4 Sept. 1959, K. Ito (TFM:FPH-551).
Fig. 63. Pseudocercospora juniperi (NY, holotype). A. Conidiophore
fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.
Fig. 64. Pseudocercospora paraexosporioides (TFM:FPH-551).
A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar =
10 µm.
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Host range and distribution: On Larix kaempferi (only in
inoculation tests), Pinaceae and Sequoia sempervirens,
Cupressaceae, Asia (Japan, endemic).
Notes: Additional material has been examined: on leaves
of Sequoia sempervirens, Japan, Ibaraki Pref., Tsukuba, 11
Sep. 1998, T. Kobayashi & C. Nakashima (CNS 448, herb.
Nakashima); Japan, Fukuoka Pref., Tanushimaru, 20 June
2000, T. Kobayashi & Y. Ono (CNS 970, herb. Nakashima);
culture ex Sequoia sempervirens (MAFF237788, ex CNS
448).
Hashimoto (1959) observed and collected a species of
Cercospora on Sequoia sempervirens in a nursery in Fukuoka
Prefecture, Japan. The specimen concerned was sent to C.
Chupp, who identied it as Cercospora exosporioides. To
conrm the pathogenicity, mycelial suspensions were used
to inoculate Larix kaempferi, Sequoia sempervirens, Pinus
densiora, Pinus thunbergii, and Cryptomeria japonica. Three
weeks later, symptoms and conidial masses were observed
on L. laempferi and S. sempervirens (Hashimoto 1959).
Natural infections are only known from S. sempervirens,
which seems to be the principal host of this species. The
identication of the Japanese species as P. exosporioides
is, however, not tenable since collections from Japan on
S. sempervirens and the type material of the latter species
from Europe are morphologically rather different. True P.
exosporioides differs from P. paraexosporioides in having
much larger stromata, up to 300 µm diam, large sporodochial
conidiomata, and much shorter, usually subcylindrical conidia
with few septa. The Japanese collections of “P. exosporioides”
represent undoubtedly a separate species described as P.
paraexosporioides. Reports of P. exosporioides from Japan
(e.g. Crous & Braun 2003) refer to the new species.
Pseudocercospora pini-densiorae (Hori & Nambu)
Deighton, Trans. Brit. Mycol. Soc. 88: 390 (1987).
var. pini-densiorae
(Fig. 65)
Basionym: Cercospora pini-densiorae Hori & Nambu, in
Nambu, J. Pl. Protec. Tokyo 4: 354 (1917).
Synonyms: Cercoseptoria pini-densiorae (Hori & Nambu)
Deighton, Mycol. Pap. 140: 167 (1976).
Mycosphaerella gibsonii H.C. Evans, Mycol. Pap. 153: 61
(1984).
Literature: Nambu (1917), Saccardo (1931: 873), Vassiljevsky
& Karakulin (1937: 252), Chupp (1954: 440), Katsuki (1965:
51), Mulder & Gibson (1972), Ellis (1976: 252), Suto (1979),
CMI Distribution Maps of Plant Diseases 481, 1982, Evans
(1984), Ivory & Wingeld (1986), Crous et al. (1990), Hsieh &
Goh (1990: 261), Ivory (1994), Guo & Hsieh (1995: 248–249),
Guo et al. (1998: 259), Crous & Braun (2004: 324), Kamal
(2010: 209), Quaedvlieg et al. (2012: 112).
Illustrations: Ellis (1976: 253, g. 190C), Hsieh & Goh (1990:
263, g. 202), Guo & Hsieh (1995: 250, g. 211), Guo et al.
(1998: 259).
Description: Causing needle blight, i.e. needles with
yellowish, yellowish brown to later greyish to blackish
brown lesions, nally becoming necrotic. Colonies
punctiform to pustulate, greyish to greyish brown. Mycelium
internal; hyphae branched, septate, pale olivaceous
to olivaceous-brown, thin-walled, smooth. Stromata
substomatal, subglobose, 20–150 µm diam, brown to
dark brown, composed of thick-walled, swollen hyphal
cells, 2–6 µm diam. Conidiophores in rich, dense to very
dense fascicles, often forming sporodochial conidiomata,
arising from stromata, through stomata or erumpent,
erect, straight, subcylindrical or narrowed towards the tip
to geniculate-sinuous, unbranched, 5–50 × 2.5–4 µm,
subhyaline to uniformly olivaceous or olivaceous-brown,
0–3-septate, thin-walled, smooth; conidiophores reduced
to conidiogenus cells or conidiogenous cells integrated,
terminal, 5–25 µm long, sympodially or occasionally
percurrently proliferating, with ne, not very conspicuous
annellations, conidiogenous loci inconspicuous or at least
neither thickened nor darkened. Conidia solitary, obclavate
or obclavate-cylindrical, straight to somewhat curved, (10–)
20–65(–80) × 2–5 µm, (1–)3–7(–10)-septate, subhyaline
to pale olivaceous, thin-walled, smooth, apex obtuse to
subacute, base obconically truncate, 1.5–2.5 µm wide, hila
Fig. 65. Pseudocercospora pini-densiorae (BPI 1109716). A.
Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar = 10
µm.
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338 ima fUNGUS
unthickened, not darkened; with Asteromella state. Sexual
morph forming variable ascostromata, dark brown to black,
discrete, immersed, subepidermal, globose, unilocular,
(50–)70–90(–120) µm diam to erumpent, linear, multilocal,
150–800(–1400) µm long, 70–125(–160) µm wide and
90–150 µm deep, occasionally laterally united in bands,
stromata pseudoparenchymatous, cells 3–8(–12) µm diam,
with thickened walls, locules globose to ask-shaped, (45–)
50–75(–95) × 55–75 µm, ostiolate, periphysate, often with
an apical stromatic shield, 70–90 µm diam, in longitudinal
series, asci bitunicate, clavate to cylindrical, (33–)35–38
× 5.5–7 µm, with a thickened bluntly rounded apex, rarely
saccate, 32–36 × 6–8 µm, 8-spored, ascospores ellipsoid,
cuneate, (7.5–)8.5–11(–12.5) × (1.8–)2.2–2.8 µm, 1-septate,
hyaline, guttulate.
In vitro: Colonies grey to greyish green or blackish,
attending 1.8–2.2 cm after 15 d on PCA-UV at 25 °C, with
compact, grey mycelium in the centre, becoming pulvinate,
black reverse (conidia in culture up to 90 µm long and 1.8–2.5
µm wide).
Type: Japan: Kagoshima, Magome, on needles of Pinus
densiora, 20 Sep. 1915 [not preserved]; Japan: Kagoshima,
Magome, on needles of Pinus densiora, 1 Oct. 1915,
identied by Hara (NIAES C-511 – neotype designated
here, MycoBank, MBT176152).
Host range and distribution: On needles of Pinus
(ayacahuite, canariensis, caribaea, cembra, clausa,
contorta, contorta var. murreyana, densiora, echinata,
elliottii, exilis, greggii, halepensis, hartwegii [rudis],
jeffreyi, kesiya, lambertiana, luchensis, massoniana,
merkusii, mugo, muricata, nigra, oocarpa, patula, parviora,
pentaphylla, pinaster, pinea, ponderosa, pseudostrobus,
radiata, resinosa, rigida, roxburghii, strobus, sylvestris,
taeda, taiwanensis, thunbergii, tubuliformis, and wallichiana
[grifthii]), Pinaceae. Africa (Kenya, Madagascar, Malawi,
South Africa, Swaziland, Tanzania, Zambia, Zimbabwe),
Asia (Bangladesh, China, India, Japan, Korea, Malaysia,
Nepal, Papua New Guinea, Philippines, Sri Lanka,
Thailand, Vietnam), Australia and New Zealand, Central
and South America (Brazil, Nicaragua), West Indies
(Jamaica, Netherlands Antilles). The following species
were susceptible in inoculation experiments carried out
by Suto (1979) in Japan: Abies veitchii, A. sachalinensis,
Cedrus deodara, Picea glehnii, P. jezoensis, Pseudotsuga
menziesii, and Larix kaempferi (L. leptolepsis).
var. montantiana (M.D. Mehrotra) U. Braun, comb. et
stat. nov.
MycoBank MB805533
Basionym: Pseudocercospora montantiana M.D. Mehrotra,
Trans. Brit. Mycol. Soc. 88: 577 (1987).
Illustration: Mehrotra (1987: 576, gs 1–6).
Diagnosis: Conidia narrowly obclavate-subcylindrical
to subacicular, only 1.5–2.5 µm wide, but otherwise
morphologically fully agreeing with var. pini-densiorae.
Holotype: India: Assam: on needles of Pinus kesiya,
Pinaceae, Apr. 1984, M. D. Mehrotra (K(M) IMI 286975).
Notes: Pseudocercospora pini-densiorae is the causal agent
of a common, widespread disease of Pinus spp. causing
needle blight. Type material is not preserved, but topotype
material has been found in the Herbarium of the National
Institute for Agro-Environmental Science, Japan, which is
proposed to serve as neotype.
Pseudocercospora montantiana was described from
India on Pinus kesiya and distinguished from P. pini-
densiorae by its narrower acicular conidia, only 1.8–2.5 µm
wide (Mehrotra 1987). However, type material of the latter
species was examined and proved to be very close to P.
pini-densiorae. The conidia are, indeed, narrower than in
common collections of P. pini-densiorae, but the conidia
are not consistently acicular. Shorter conidia are narrowly
obclavate-subcylindrical with short obconically truncate
base, which agrees with typical conidia of P. pini-densiorae.
Only some very long conidia may be subacicular. Otherwise
P. montantiana on Pinus kesiya, which has been recorded
as host species of P. pini-densiorae, is morphologically
indistinguishable from the latter species and barely more than
a morphological variety of the common pine needle blight.
Stromata are very variable, 10–80 µm diam or oblong and
up to 150 µm in length, the conidiophores are numerous, in
dense fascicles, 10–50 × 2–4 µm, and the conidia are mostly
25–70 µm long and 3–7-septate. Mehrotra (1987) described
conidiophores up to 100 µm long, which possibly refers to
conidiophores with attached conidia, and he described
conidia up to 105 µm in length.
Pseudocercospora sciadopityos C. Nakash. & Tak.
Kobay., sp. nov.
MycoBank MB805534
(Fig. 66)
Etymology: Name derived from its host genus, Sciadopitys.
Synonym: Pseudocercospora horiei C. Nakash. & Tak.
Kobay. (in Kobayashi 2007: 775), nom. inval. (Art.30.8).
Literature: Kobayashi (2007).
Diagnosis: Pseudocercosporae juniperi, P. exosporioides et P.
pini-densiorae similis, sed conidiis acicularibus vel anguste
cylindraceis, ad 83 µm longis, sed etiam 3–5-septatis, basi
truncata.
Description: Leaf symptoms visible as brown to yellowish
brown discolorations. Caespituli hypophyllous, punctiform,
formed as olivaceous masses of conidia and conidiophores.
Mycelium internal. Stromata immersed, intraepidermal to
substomatal, erumpent, brown to dark brown, 45–75 µm diam.
Conidiophores arising from stromata, densely fasciculate,
through stomata or erumpent, erect, straight to slightly curved,
35–85 × 2–3 μm (on average 55 × 2 μm), pluriseptate, pale
brown, thin-walled, smooth; conidiogenous cells integrated,
terminal, proliferating sympodially or percurrently, with
unthickend, not darkened conidiogenous loci. Conidia solitary,
mildly curved, acicular to narrowly cylindrical, 63–83 × 2–3
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μm (on average 71 × 3 μm), 3–5-septate, pale, smooth, base
truncate, 2–3.8 µm wide, hila neither thickened nor darkened.
Holotype: Japan: Tokyo, Chofu, on Sciadopitys verticillata,
16 June 1977, T. Kobayashi (TFM:FPH-4757).
Host range and distribution: On Sciadopitys verticillata,
Sciadopityaceae, Asia (Japan, Tochigi, Tokyo, Chiba).
Note: A rst description of this species, which caused early
defoliation of Sciadopitys several times in Japan, has been
published by Kobayashi (2007) as Cercospora sp.
Pseudocercospora thujina (Dearn.). U. Braun & C.
Nakash., comb. nov.
MycoBank MB805535
(Fig. 67)
Basionym: Coryneum thujinum Dearn., Mycologia 16: 171
(1924).
Synonyms: Sciniatosporium thujinum (Dearn.) Morgan-
Jones, Canad. J. Bot. 49: 1005 (1971).
Stigmina thujina (Dearn.) B. Sutton, Trans. Brit. Mycol. Soc.
58: 166 (1971).
Literature: Sutton (1971), Hodges (1982), Dick (1998), Cech
(2008), Cech & Diminić (2008).
Illustrations: Morgan-Jones (1971: 1006, g. 10), Hodges
(1982: 868, g. 1A–D).
Description: Leaves turning pale brown to brown, later
greyish brown or grey, nally they become necrotic and die, in
severe cases entire branches or twigs may die. Conidiomata
hypophyllous, scattered, blackish, sporodochial, erumpent.
Mycelium immersed; hyphae branched, septate, 2–4 µm
wide, brown, wall smooth. Stromata well-developed, at
rst immersed, later often rupturing the epidermis, dark
brown to black, 150–270 µm diam and 60–170 µm deep, or
oblong, up to 500 µm in length, composed of swollen hyphal
cells, rounded to angular in outline, about 2–6 µm diam.
Conidiophores numerous, in loose to usually dense fascicles,
arranged in palisade-like layers, arising from stromata,
forming sporodochial conidiomata, erumpent, erect, straight
to curved or somewhat exuous, subcylindrical or somewhat
attenuated towards the tip, unbranched, 10–80 × 3–7 µm,
aseptate to sparingly septate, light brown, yellowish brown
or pale olivaceous-brown throughout or usually somewhat
paler towards the tip, darker in mass, wall thin to slightly
thickened, smooth or somewhat rough; conidiophores
Fig. 67. Pseudocercospora thujina (BPI 404872). A. Conidiophore
fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.
Fig. 66. Pseudocercospora sciadopytios (TFM:FPI-4757). a.
Conidium (light microscopy). b. Conidiophore fascicle (light
microscopy). Bars = 5 µm (a) and 10 µm (b).
a
b
Braun et al.
ARTICLE
340 ima fUNGUS
reduced to conidiogenous cells or conidiophores septate
and conidiogenous cells integrated, terminal, monoblastic,
determinate or percurrently proliferating, with up to ve ne
annellations, sometimes also sympodially proliferating, with
two loci, conidiogenous loci unthickened, not darkened, about
2–4 µm wide, sometimes with minute frill. Conidia solitary,
straight to mostly curved or sigmoid, often abruptly curved at
the apex or sometimes at the base, subcylindrical-vermiform
to clavate, occasionally obclavate, 30–70 × (5–)6–9(–11)
µm, 4–16-septate, usually without constrictions at the septa
or only slightly constricted, cells often with distinct central
lumen, medium brown to medium dark brown, wall thin to
slightly thickened, up to 1 µm, smooth to faintly rough, apex
obtuse, rounded, base short obconically truncate, 2–5 µm
wide, hila unthickened, not darkened, often with minute frill.
Lectotype (designated here, MycoBank, MBT176153):
USA: Oregon: Lane Co., Cascade National Forest, 12.000
ft. alt., on Thuja plicata, 22 Oct. 1921, J. S. Boyce 831 (BPI
404872), isolectotypes: BPI 404883, 404884, syntypes: BPI
404871, 869772 (from “Hot Springs”, Boyce 541).
Host range and distribution: On Chamaecyparis lawsoniana,
Thuja plicata, Cupressaceae, North America (Canada, British
Columbia; USA, Idaho, North Carolina, Oregon, Washington),
introduced in Europe (Austria and Croatia), Hawaii and New
Zealand.
Notes: Based on the phylogenetic position of its type
species, Crous et al. (2006) reduced Stigmina to synonymy
with Pseudocercospora. Therefore, the generic position
of Stigmina thujina has to be reassessed. Due to its
sporodochial conidiomata with frequently percurrently
proliferating conidiogenous cells and relatively thin-walled
conidia, this species is rather Cercostigmina-like (Braun
1993), but the latter genus was reduced to synonymy with
Pseudocercospora. The type species of Cercostigmina and
additional species assigned to this genus cluster within the big
Pseudocercospora clade (Crous et al. 2001, 2013; Taylor et
al. 2003). Furthermore, the conidiogenous cells of S. thujina
are percurrent as well as sometimes sympodial. Therefore,
this species is better reallocated to Pseudocercospora.
Pseudocercospora thujina is also known on Chamaecyparis
lawsoniana and has been introduced on this host in Europe,
Hawaii and New Zealand (Hodges 1982, Dick 1998, Cech
2008, Cech & Diminić (2008). Two samples from Hawaii
have been re-examined (BPI 428035, 428036). The rst
European observation was from 2002 in Austria, Salzburg.
Cercospora chamaecyparidis, described from Japan, is
morphologically very close to P. thujina, but occurs on the Asian
Chamaecyparis obtusa and differs in having much smaller or
even lacking stromata and verruculose conidia. Therefore, C.
chamaecyparidis is maintained as a separate species and
reallocated to Pseudocercospora as well.
Doubtful, excluded and insufciently known
species
Cercospora licum Henn., Hedwigia 41: 310 (1902).
Literature: Saccardo (1906: 611), Chupp (1954: 455).
Holotype: Brazil: São Paulo, Botanical Garden, on
Nephrodium sp., Dryopteridaceae, A. Puttemans No. 258
[not preserved].
Notes: Due to rather broad, densely septate cylindrical conidia
with thick walls, this species is rather helminthosporioid, but
not cercosporoid (see Chupp 1954).
Cercospora equiseti Dobrozr., Bolez. Rast. 16: 202
(1927).
Synonyms: Didymaria equiseti (Dobrozr.) Chupp, Monogr.
Cercospora: 205 (1954).
Ramularia dobrozrakoviana U. Braun, Internat. J. Mycol.
Lichenol. 3: 281 (1988).
Monodidymaria equiseti (Dobrozr.) U. Braun, Mycotaxon
51: 44 (1994).
Literature: Vassiljevsky & Karakulin (1937: 219), Braun
(1998: 360).
Illustration: Braun (1998: 362, g. 618).
Holotype: Russia: Leningrad Oblast’, on Equisetum arvense,
Equisetaceae, 23 Jul. 1927, Dobrozrakova (LE 40397).
Cercospora trismeriae Petr., Sydowia 2: 379 (1948)
Literature: Chupp (1954: 457), Crous & Braun (2003: 411).
Description: Leaf spots narrow, oblong between secondary
veins and often extending from the midrib to the leaf margin, in
severe infections covering large portions of the leaf blade, dull
yellow to greyish brown on both surfaces. Caespituli mainly
hypophyllous. Mycelium internal. Stromata substomatal,
globose, dark olivaceous. Conidiophores straight, curved
to sinuous, 40–150 × 5–7 µm, pale olivaceous, sparingly
septate. Conidia obclavate, straight to curved, rarely
sigmoid, 25–200 × 5–7.5 µm, 2–9-septate, subhyaline to
pale olivaceous-brown, base subtruncate, apex subobtuse to
somewhat attenuated.
Holotype: Ecuador: Pichinicha Province: Mindo, on living
leaves of Pityrogramma sp. (Trismeria sp.), 5 Nov. 1937, H.
Sydow, No. 1937.
Host range and distribution: Only known from the type
collection.
Notes: The generic afnity of this species is unclear and could
not be proven since it was not possible to trace type material,
neither in Petrak’s herbarium at W nor at GZU. Based on the
original description, C. trismeriae is probably a species of
Pseudocercospora.
Scolecostigmina chibaensis C. Nakash., Tak. Kobay. &
Tosh. Yamada, in Nakashima et al. Mycoscience 48:
25 (2007).
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341
volume 4 · no. 2
Illustration: Nakashina et al. (2007: 251, g. 1).
Description: Twig symptoms visible as swellings, nally
cracking. Stromata lacking or small, blackish brown to black,
erumpent, small aggregations of swollen hyphal cells, 2–13 μm
diam. Conidiophores solitary to densely fasciculate, straight,
olivaceous-brown to blackish brown, subcylindrical, 30–90
× 4–10 μm, pluriseptate, wall thickened, up to about 2 µm,
smooth or almost so, occasionally rough; conidiogenous cells
integrated, terminal, determinate or percurrently proliferating,
with truncate, unthickened, not darkened terminal locus with
distinct, relatively large porus, up to 2 µm diam (conidiogenesis
probably tretic). Conidia solitary, occasionally in short chains,
phragmo- to scolecosporous, broadly obclavate-subcylindrical,
65–135 × 9–13 μm, 5–13-euseptate or eu- and distosepta
mixed, often somewhat constricted at septa, brown to blackish
brown, wall thick, up to 2 µm, smooth to somewhat verruculose,
apex sometimes somewhat rostrate, tips obtuse, base
obconically truncate, 4–5.5 µm wide, hila neither thickened nor
darkened or entire base somewhat darker.
Holotype: Japan: Chiba, Kamogawa, on twigs of Pinus
parviora, 19 May 2006, T. Yamada (TFM:FPH-7858), isotype
MUMH 10314 (ex-holotype culture: NBRC 102148).
Host range and distribution: On Pinus (ayacahuite, parviora
var. parviora, parviora var. pentaphylla, peuce, strobiformis,
wallichiana, strobus, Pinus sp.), Pinaceae, Asia (Japan).
Notes: In preliminary phylogenetic analyses using
rDNA ITS data, S. chibaensis did neither cluster with
Pseudocercospora species (including Stigmina s. str. [type
species: S. platani]) nor Scolecostigmina mangiferae, the
type species of Scolecostigmina. This species does not
belong to the Mycosphaerellaceae at all. The conidiogenous
cells possess a conspicuous, relatively large porus, and
the conidiogenesis seems to be tretic. Scolecostigmina
chibaensis is morphologically rather corynesporoid. It has to
be excluded from Scolecostigmina in any case, but a nal
conclusion must be postponed as its phylogenetic afnity to a
specic order and family is not yet clear.
ACKNOWLEDGEMENTS
We are very grateful to the directors and curators of BPI, CUM, CUP,
HMAS, IUM, K, PC, TFM, and TNS for loaning type material and
other collections in their keeping during the course of monographic
studies of cercosporoids fungi on other fungi, gymnosperms and
ferns.
REFERENCES
Arx JA von (1981) The Genera of Fungi sporulating in Pure Culture.
3rd edn. Vaduz: J. Cramer.
Arx JA von (1983) Mycosphaerella and its anamorphs. Proceedings
of the Koninklijke Nederlandse Akademie van Wettenschappen,
Series C, 86: 15–54.
Arzanlou M, Groenewald JZ, Fullerton RA, Abeln ECA, Carlier J, et
al. (2008) Multiple gene genealogies and phenotypic characters
differentiate several novel species of Mycosphaerella and related
anamorphs on banana. Persoonia 20: 19–37.
Arzanlou M, Groenewald JZ, Gams W, Braun U, Crous PW (2007)
Phylogenetic and morphotaxonomic revision of Ramichloridium
and allied genera. Studies in Mycology 58: 57–93.
Beilharz V, Pascoe IG, Parbery DG (2002) Three new
Pseudocercospora species, one with a Mycosphaerella
teleomorph, from Kennedia in Australia. Mycotaxon 82: 397–407.
Baker WA, Partridge EC, Morgan-Jones G (2000) Notes on
Hyphomycetes. LXXVII. Asperisporium sequoiae, the causal
organism of conifer needle blight, reclassied in Cercosporidium,
with comments on the status of the genus. Mycotaxon 76: 247–
256.
Barklund P (2006) Recent disease problems in Swedish forests.
Aktuelt fra Skogforskningen 1(6): 84.
Begum MM, Dalisay TU, Cumagun CJR (2012) Taxonomic review
of and development of a lucida key for Philippine cercosporoids
and related fungi. In: Plant Pathology (CJR Cumagun, ed.):
1–40. Rijeka: Intech.
Beilharz V, Pascoe I (2002) Two additional species of Verrucisporota,
one with a Mycosphaerella teleomorph, from Australia.
Mycotaxon 87: 357–365.
Boedijn KB (1961) The genus Cercospora in Indonesia. Nova
Hedwigia 3: 411–436.
Braun U (1988) Studies on Ramularia and allied genera (II). Nova
Hedwigia 47: 335–349.
Braun U (1990) Taxonomic problems of the Ramularia/Cercosporella
complex. Studies in Mycology 32: 65–75.
Braun U (1992) Taxonomic notes on some species of the Cercospora
complex. Nova Hedwigia 55: 211–221.
Braun U (1993) New genera of phytopathogenic Hyphomycetes.
Cryptogamic Botany 4: 107–114.
Braun U (1995a) A monograph of Cercosporella, Ramularia and
allied genera (phytopathogenic hyphomycetes). Vol. 1. Eching:
IHW Verlag.
Braun U (1995b) Miscellaneous notes on phytopathogenic
hyphomycetes (II). Mycotaxon 55: 223–241.
Braun U (1998) A monograph of Cercosporella, Ramularia and allied
genera (phytopathogenic hyphomycetes). Vol. 2. Eching: IHW
Verlag.
Braun U (2001) Revision of Cercospora species described by K.B.
Boedijn. Nova Hedwigia 73: 419–436.
Braun U (2002) Miscellaneous notes on some micromycetes (II).
Schlechtendalia 8: 33–38.
Braun U (2012) The impact of the discontinuation of dual
nomenclature of pleomorphic fungi: the trivial facts, problems,
and strategies. IMA Fungus 3: 81–86.
Braun U, Crous PW (2005) Additions and corrections to names
published in Cercospora and Passalora. Mycotaxon 92: 395–
416.
Braun U, Crous PW (2006) (1732) Proposal to conserve the name
Pseudocercospora against Stigmina and Phaeoisariopsis
(Hyphomycetes). Taxon 55: 803.
Braun U, Crous PW (2008) Cercosporoid hyphomycetes on hosts
of the Annonaceae: Cercospora annonaceae and Isariopsis
annonarum revisited. Mycotaxon 105: 207–224.
Braun U, Freire F (2004) Some cercosporoid hyphomycetes from
Brazil – III. Cryptogamie Mycologie 25: 221–244.
Braun U, Melnik VA (1997) Cercosporoid fungi from Russia and
adjacent countries. Trudy Botanicheskogo Instituta Imeni V.L.
Braun et al.
ARTICLE
342 ima fUNGUS
Komarova, Rossijskaya Akademiya Nauk St. Petersburg 20:
1–130.
Braun U, Sivapalan A (1999) Cercosporoid hyphomycetes from
Brunei. Fungal Diversity 3: 1–27.
Braun U, Crous PW, Schubert K, Shin HD (2010a) Some reallocations
of Stenella species to Zasmidium. Schlechtendalia 20: 99–104.
Braun U, Freire F, Urtiaga R (2010b) New species and new records
of cercosporoid hyphomycetes from Brazil, New Zealand and
Venezuela. Polish Botanical Journal 55: 281–291.
Bhurgava KS, Misra PC (1961) Cercospora helminthostachydis P.
Henn. in Yorakhpur, India. Sydowia 15: 231–232.
Castañeda RF (1988) Fungi Cubenses. Vol. 3. Habana: Instituto
de Investigaciones Fundamentales en Agricultura Tropical
"Alejandro de Humboldt".
Castañeda RF, Braun U (1989) Cercospora and allied genera of
Cuba (I). Cryptogamic Botany 1: 42–55.
Cech TL (2008) Phytopathologische Notizen 2008. Forstschutz
Aktuell 43: 21–23.
Cech TL, Diminić D (2008) New blight of Cupressaceae in Austria and
Croatia. In: Proceedings of the Conference of IUFRO working
party 7.02.02, 21–26 May 2007, Sopron, Hungary. Acta Silvatica
& Lignaria Hungarica, Special Edition (2007): 265.
Chiddarwar PP (1962) Contributions to our knowledge of the
Cercosporae of Bombay State III. Mycopathologia et Mycologia
Applicata 17: 71–81.
Chowdhry PN, Gupta D, Padhi B (1983) Some new species
of Cercosporae from ornamental plants in India. Indian
Phytopathology 36: 624–625.
Chupp C (1954) A monograph of the fungus genus Cercospora.
Ithaca: Published by the author.
Ciferri R (1938) Mycoora domingensis exsiccatae. Annales
Mycologici 36: 198–245.
Ciferri R (1954) Schedae Mycologicae XII–XXXIV. Sydowia 8: 245–
270.
Clements FE, Shear CL (1931) The Genera of Fungi. New York: H.
W. Wilson.
Cour P, Joly P (1965) Un parasite nouveau des Sélaginelles :
Cercospora sellaginellarum. Revue de Mycologie 30: 225–
230.
Crous PW (1998) Mycosphaerella spp. and their anamorphs
associated with leaf spot diseases of Eucalyptus. Mycologia
Memoir 21: 1–170.
Crous PW, Braun U (1996). Cercosporoid fungi from South Africa.
Mycotaxon 57: 233–321.
Crous PW, Braun U (2003) Mycosphaerella and its Anamorphs. 1.
Names published in Cercospora and Passalora. [CBS Biodiversity
Series no. 1.] Utrecht: CBS-KNAW Fungal Biodiversity Centre.
Crous PW, Wingeld MJ, Swart WJ (1990) Shoot and needle
diseases of Pinus spp. in South Africa. South African Forestry
Journal 154: 60–66.
Crous PW, Aptroot A, Kang JC, Braun U, Wingeld MJ (2000) The
genus Mycosphaerella and its anamorphs. Studies in Mycology
45: 107–121.
Crous PW, Kang JC, Braun U (2001) A phylogenetic redenition
of anamorph genera in Mycosphaerella based on ITS rDNA
sequences and morphology. Mycologia 93: 1081–1101.
Crous PW, Groenewald JZ, Mansilla JP, Hunter GC, Wingeld MJ
(2004a) Phylogenetic reassessment of Mycosphaerella spp. and
their anamorphs occurring on Eucalyptus. Studies in Mycology
50: 195–214.
Crous PW, Groenewald JZ, Pongpanich K, Himaman W, Arzanlou M,
Wingeld MJ (2004b) Cryptic speciation and host specicity among
Mycosphaerella spp. occurring on Australian Acacia species grown
as exotics in the tropics. Studies in Mycology 50: 457–469.
Crous PW, Liebenberg MM, Braun U, Groenewald JZ (2006) Re-
evaluationg the taxonomic status of Phaeoisariopsis griseola, the
causal agent of angular leaf spots of bean. Studies in Mycology
55: 163–173.
Crous PW, Braun U, Groenewald JZ (2007) Mycosphaerella is
polyphyletic. Studies in Mycology 58: 1–32.
Crous PW, Braun U, Hunter GC, Wingeld MJ, Verkley GJM, et al.
(2013) Phylogenetic lineages in Pseudocercospora. Studies in
Mycology 75: 37–114.
Crous PW, Braun U, Wingeld MJ, Wood AR, Shin HD, et al. (2009a)
Phylogeny and taxonomy of obscure genera of microfungi.
Persoonia 22: 139–161.
Crous PW, Groenewald JZ, Shivas RG (2010) Pseudocercospora
nephrolepidicola Crous & R.G. Shivas. Fungal Planet 59.
Persoonia 25: 138–139.
Crous PW, Groenewald JZ, Shivas RG, Edwards J, Seifert KA, et
al. (2011) Fungal Planet Description Sheets: 69–91. Persoonia
26: 108–156.
Crous PW, Schoch CL, Hyde KD, Wood AR, Gueidan C, et al. (2009b)
Phylogenetic lineages in the Capnodiales. Studies in Mycology
64: 17–47.
Crous PW, Summerell BA, Carnegie AJ, Wingeld MJ, Hunter GC,
et al. (2009c) Unravelling Mycosphaerella: do you believe in
genera? Persoonia 23: 99–118.
Crous PW, Summerell BA, Shivas RG, Burgess TI, Decock CA, et al.
(2012). Fungal Planet Description Sheets: 107–127. Persoonia
28: 138–182.
Curzi M (1932) De fungis et morbis africanis. I. De quibusdam
hyphomycetibus parasitis Somalie. Bollettino della Stazione di
Patologia Vegetale di Roma, Ser. 2, 12: 148–168.
Das AK (1989) Two new species of Cercospora from India. Indian
Journal of Mycological Research 27: 37–42.
David JC (1997) A contribution to the systematics of Cladosporium:
revision of the fungi previously referred to Heterosporium.
Mycological Papers 172: 1–157.
Deighton FC (1965) Various hyphomycetes, mainly tropical.
Mycological Papers 101: 28–43.
Deighton FC (1967) Studies on Cercospora and allied genera. II.
Passalora, Cercosporidium and some species of Fusicladium on
Euphorbia. Mycological Papers 112: 1–80.
Deighton FC (1969) Microfungi IV: Hyperparasitic hyphomycetes,
and a note on Cercosporella uredinophila Sacc. Mycological
Papers 118: 1–41.
Deighton FC (1971) Studies on Cercospora and allied genera. III.
Centrospora. Mycological Papers 124: 1–13.
Deighton FC (1973) Studies on Cercospora and allied genera.
IV. Cercosporella Sacc., Pseudocercosporella gen. nov. and
Pseudocercosporidium gen. nov. Mycological Papers 133: 1–62.
Deighton FC (1974) Studies on Cercospora and allied genera. V.
Mycovellosiella Rangel, and a new species of Ramulariopsis.
Mycological Papers 137: 1–75.
Deighton FC (1976) Studies on Cercospora and allied genera. VI.
Pseudocercospora Speg., Pantospora Cif. and Cercoseptoria
Petr. Mycological Papers 140: 1–168.
Deighton FC (1979) Studies on Cercospora and allied genera. VII.
New species and redispositions. Mycological Papers 144: 1–56.
Cercosporoid fungi 1
ARTICLE
343
volume 4 · no. 2
Deighton FC (1983) Studies on Cercospora and allied genera. VIII.
Further notes on Cercoseptoria and some new species and
redispositions. Mycological Papers 151: 1-13.
Deighton FC (1987) New species of Pseudocercospora and
Mycovellosiella, and new combinations into Pseudocercospora
and Phaeoramularia. Transactions of the British Mycological
Society 88: 365–391.
Deighton FC (1990) Observations on Phaeoisariopsis. Mycological
Research 94: 1096–1102.
Dick MA (1998) Stigmina thujina – a needle blight of Lawson’s
cypress. Forest Health News 70: 2–3.
Earle FS (1901) Some fungi from Puerto Rico. Muhlenbergia 1:
10–23.
Ellis MB (1959) Clasterosporium and some allied Dematiaceae –
Phragmosporae. II. Mycological Papers 72: 1–75.
Ellis MB (1963) Dematiaceous hyphomycetes IV. Mycological Papers
87: 1–42.
Ellis MB (1971) Dematiaceous Hyphomycetes. Kew: Commonwealth
Mycological Institute.
Ellis MB (1976) More Dematiaceous Hyphomycetes. Kew:
Commonwealth Mycological Institute.
Evans HC (1984) The genus Mycosphaerella and its anamorphs
Cercoseptoria, Dothistroma and Lecanosticta on pines.
Mycological Papers 153: 1–102.
Farr ML, Horner Jr HT (1968) Fungi on Selaginella. Nova Hedwigia
15: 239–287.
Ferraris T (1909) Osservazioni mycologische. Su specie del gruppo
Hyphales (Hyphomycetae), Ser. 1a, No. I-X. Annales Mycologici
7: 273–286.
Fresenius JGBW (1863) Beiträge zur Mykologie. Frankfurt a. M.:
H.L. Brönner.
Fries EM (1849) Summa vegetabilium Scandinaviae. Sectio
Posterior. Stockholm: A. Bonnier.
Fuckel KWGL (1863) Fungi rhenani exsiccati a L. Fuckel collecti.
Fasc. I–IV. 1863. Hedwigia 2: 132–136.
Goodwin SB, Dunkle LD, Zismann VL (2001) Phylogenetic analysis
of Cercospora and Mycosphaerella based on internal transcribed
spacer region of ribosomal DNA. Phytopathology 91: 648–658.
Groenewald JZ, Groenewald M, Braun U (2010) Cercospora
speciation and host range. In: Cercospora Leaf Spot of Sugar
Beet and related species (Lartey RT, Weiland JJ, Panella L,
Crous PW, Windels CE, eds): 21–37. St Paul, MN: APS Press.
Groenewald JZ, Nakashima C, Nishikawa J, Shin, HD, Park JH, et
al. (2013) Species concepts in Cercospora: spotting the weeds
among the roses. Studies in Mycology 75: 115–170.
Groenewald M, Groenewald JZ, Crous PW (2005) Distinct species
exist within the Cercospora apii morphotype. Phytopathology 95:
951–959.
Groenewald M, Groenewald JZ, Braun U, Crous PW (2006a) Host
range of Cercospora apii and C. beticola, and description of C.
apiicola, a novel species from celary. Mycologia 98: 275–285.
Groenewald M. Groenewald JZ, Harrington TC, Abeln ECA, Crous
PW (2006b) Mating type gene analysis in apparently asexual
Cercospora species is suggestive of cryptic sex. Fungal Genetics
and Biology 43: 813–825.
Guo YL (1993) Two species of Pseudocercospora. Mycosystema 6:
103–106.
Guo YL (1997) A new species of Cercospora. Mycosystema 16: 1–3.
Guo YL, Hsieh WH (1995) The genus Pseudocercospora in China.
Mycosystema, Monographicum Series 2: 1–388.
Guo YL, Liu XJ (1992) Studies on the genus Pseudocercospora in
China III. Acta Mycologica Sinica 11: 294–299.
Guo YL, Liu XJ, Hsieh WH (2003) Mycovellosiella, Passalora,
Phaeoramularia. [Flora Fungorum Sinicorum, vol. 20.] Beijing:
Science Press.
Guo YL, Liu XJ, Hsieh WH (2005) Cercospora. [Flora Fungorum
Sinicorum, vol. 24.] Beijing: Science Press.
Hashimoto H (1959) On the needle-blight of Sequoia sempervirens
Endl., caused by Cercospora exosporioides Bubák. The Bulletin
of Fukuoka-Ken Forest Experiment Station 11: 19–30.
Hawksworth D (2011) A new dawn for the naming of fungi: impacts
of decisions made in Melbourne in July 2011 on the future
publication and regulation of fungal names. IMA Fungus 2: 155–
162.
Hernández-Gutiérrez A, Dianese JC (2008) New cercosporoid fungi
from the Brazilian Cerrado 1. Species on hosts of the families
Anacardiaceae. Mycotaxon 106: 41–63.
Hernández-Gutiérrez A, Dianese JC (2009) New cercosporoid fungi
from the Brazilian Cerrado 2. Species on hosts of the subfamilies
Caesalpinioideae, Faboideae and Mimosoideae (Leguminosae
s. lat.). Mycotaxon 107: 1–24.
Hodges CS (1962) Comparison of four similar fungi from Juniperus
and related conifers. Mycologia 54: 62–69.
Hodges CS (1982) Needle blight of Port-Orford-Cedar caused by
Stigmina thujina in Hawaii. Plant Disease 66: 867–869.
Hsieh WH, Goh TK (1990) Cercospora and similar fungi from Taiwan.
Taipei: Maw Chang Book Co.
Ito K, Kobayashi T, Shibukawa K (1967) Etiological studies on
the needle blight of Cryptomeria japonica – III: A comparison
between Cercospora cryptomeriae Shirai and Cercospora
sequoiae Ellis et Everhart. Bulletin of the Government Forest
Experiment Station 204: 73–97.
Ito K, Shibukawa K, Kobayashi T (1958) Blight of Sequoia gigantea
seedlings caused by Cercospora cryptomeriae Shirai. Journal of
the Japanese Forestry Society 40: 407–410.
Ivory MH (1994) Records of foliage pathogens of Pinus species in
tropical countries. Plant Pathology 43: 511–518.
Ivory MH, Wingeld MJ (1986) First report of Mycosphaerella gibsonii
in South Africa. Phytophylactica 18: 51–53.
Jaczewski A v (1901) Über eine Pilzkrankheiten auf dem Wacholder.
Zeitschrift für Panzenkrankheiten 11: 203–207.
Kaiser W, Crous PW (1998) Mycosphaerella lupini sp. nov., a serious
leaf spot disease of perennial lupin in Southcentral Idaho, USA.
Mycologia 90: 726–731.
Kakoti RK, Saikia UN, Mamar D (1998) Phytopathogenic fungi of
north-east India – IV. Annals of Agri Bio Research 3: 207–210.
Kamal (2010) Cercosporoid Fungi of India. Dehra Dun: Bishen Singh
Mahendra Pal Singh.
Kar AK, Mandal M (1969) New Cercospora spp. from West Bengal.
Transactions of the British Mycological Society 53: 337–360.
Katsuki S (1965) Cercosporae of Japan. Transactions of the
Mycological Society of Japan, Extra Issue 1: 1–100.
Katumoto K (1988) Five hyperparasitic hyphomycetes from Japan.
Bulletin of the Faculty of Agriculture, Yamaguchi University 35:
107–119.
Khan MK, Verma RK, Kamal (1992) New species of Cercospora,
Mycovellosiella and Phaeoisariopsis. Indian Phytopathology 45:
26–34.
Kitashima K (1916) On the red-plague of “Sugi” seedlings [in
Japanese]. Botanical Magazine Tokyo 30: 411–414.
Braun et al.
ARTICLE
344 ima fUNGUS
Kirschner R, Chen CJ (2007) Foliicolous hyphomycetes from Taiwan.
Fungal Diversity 26: 219–239.
Kirschner R., Okada T (2013) A new species of Pseudocercospora
and new record of Bartheletia paradoxa on leaves of Ginkgo
biloba. Mycological Progress 12: 421–426.
Kobayashi T (2007) Index of Fungi Inhabiting Woody Plants in Japan:
host, distribution and literature. Tokyo: Zenkoku-Noson-Kyoiku
Kyokai Publishing.
Kujala V (1950) Über die Kleinpilze der Koniferen in Finland.
Ascomycetes, Fungi imperfecti, Uredinales. Metsätieteellinen
tutkimuslaitoksen julkisuja 38: 1–121.
Kumar P, Kamal (1981) Two new species of fungus genus Cercospora
Fres. Current Science 50: 505–510.
Kurkela T, 1994) Metsän taudit. Metsäpatologian perusteet. Tampere:
Otatieto OY.
Lall G, Kapoor JN, Munjal RL (1964) Some fern inhabiting Cercospora
species from India. Indian Phytopathology 17: 181–183.
Maublanc A (1913a [“1912”]) Sobre uma molestia do mamoeiro
(Carica papaya L.)/Sur une maladie des feuilles du papayer
“Carica papaya”. Lavoura 16: 204–212.
Maublanc A (1913b) Sur une maladie des feuilles du papayer (Carica
papaya). Bulletin Trimestriel de la Société Mycologique de
France 29: 353–358.
Meeboon J, Hidayat I, To-anun C (2007) An annotated list of
cercosporoid fungi in Northern Thailand. Journal of Agricultural
Technology 3: 51–63.
Mehrotra MD (1987) Pseudocercospora needle blight, a new disease
of Pinus kesiya from India. Transactions of the British Mycological
Society 88: 575–577.
Mel’nik VA (2000) Opredelitel’ gribov Rossii. Klass Hyphomycetes
Vol. 1. St.-Petersburg: Nauka.
Minnis AM, Kennedy AH, Grenier DB, Rehner SA, Bischoff JF
(2011) Asperisporium and Pantospora (Mycosphaerellaceae):
epitypications and phylogenetic placement. Persoonia 27: 1–8.
Miura M (1928) Flora of Manchuria and East Mongolia. Vol. 3.
Cryptogams, Fungi. Daliean: South Manchuria Railroad.
Montenegro-Calderón JG, Martinez-Álvarez JA, Vieyra-Hernández
MT, et al. (2011) Molecular identication of two strains of
Cercospora rodmanii isolated from water hyacinth present in
Yuriria lagoon, Guanajuato, Mexico and identication of new
hosts for several other strains. Fungal Biology 115: 1151–1162.
Morgan-Jones G (1971) Sciniatosporium Kalchbr., and its synonyms
Marcosia Syd., Stigmina Sacc., Thyrostroma Höhn., and
Thyrostromella Syd., non Höhn. Canadian Journal of Botany 49:
993–1009.
Motohashi K, Araki I, Nakashima C (2008) Four new species of
Phyllosticta, one new species of Pseudocercospora, and one
new combination in Passalora from Japan. Mycoscience 49:
138–146.
Motohashi K, Kobayashi T, Furukawa T, Ono Y (2010) Notes on
some plant-inhabiting fungi collected from the Nansei Islands (2).
Mycoscience 51: 93–97.
Mulder JL (1989) Stenella australiensis sp. nov. on Blechnum
indicum. Mycological Research 92: 118–122.
Mulder JL, Gibson IAS (1972) Cercospora pini-densiorae. CMI
Descriptions of Pathogenic Fungi and Bacteria 329: 1–2.
Farnham Royal: Commonwealth Agricultural Bureaux.
Mulder JL, Gibson IAS (1973) Cercospora sequoiae. CMI Descriptions
of Pathogenic Fungi and Bacteria 366: 1–2. Farnham Royal:
Commonwealth Agricultural Bureaux.
Muntañola M (1960) Algunos hyphomycetes criticos. Lilloa 30: 165–232.
Muthappa BN (1968) Fungi of Coorg, India. I. Mycopathologia et
Mycologia Applicata 34: 193–195.
Nakashima C, Akashi T, Takahashi Y, Yamada T, Akiba M, Kobayashi
T (2007) New species of the genus Scolecostigmina and revision
of Cercospora cryptomeriicola on conifers. Mycoscience 48:
250–254.
Nakashima C, Inabe S, Park J-Y, Ogawa Y (2006) Addition and
re-examination of Japanese species belonging to the genus
Cercospora and allied genera. IX. Newly recorded species from
Japan (4). Mycoscience 47: 48–52.
Nambu N (1917) Diseases of forest trees in nurseries. Journal of
Plant Protection, Tokyo 4: 353–354.
Norvell LL (2011) Fungal nomenclature. 1. Melbourne approves a
new Code. Mycotaxon 116: 481–490.
Penzes A (1927) Cercospora Hungarica. Folia Cryptogamica 1:
288–336.
Peterson GW (1977) Epidemiology and control of a blight of
Juniperus virginiana caused by Cercospora sequoiae var.
juniperi. Phytopathology 67: 234–238.
Peterson GW, Wysong DS (1968) Cercospora blight of juniper:
damage and control. Plant Disease Reporter 52: 361–362.
Petrak F (1951) Über die Gattungen Chaetotrichum Syd. und
Ragnhildiana Solh. Sydowia 5: 30–39.
Phengsintham P, Braun U, McKenzie EHC, Chukeatirote E, Cai
L, Hyde KD (2013a) Monograph of cercosporoid fungi from
Thailand. Plant Pathology & Quarantine 3: 19–90.
Phengsintham P, Chukeatirote E, Bahkali AH, Moslem MA, Hyde
KD, Braun U (2010b) Cercospora and allied genera from Laos 3.
Cryptogamie Mycologie 31: 305–322.
Phengsintham P, Chukeatirote E, McKenzie EHC, Hyde KD, Braun
U (2013b) Monograph of cercosporoid fungi from Laos. Current
Reseach in Environmental & Applied Mycology 3: 34–158.
Phengsintham P, Chukeatirote E, McKenzie EHC, Moslem MA, Hyde
KD, Braun U (2010a) Two new species and a new record of
cercosporoids from Thailand. Mycosphere 1: 205–212.
Pirozynski KA (1974) Meliolina mollis and two hyperparasites in
India. Kavaka 2: 33–41.
Pollack FG (1971) Cercospora uromycestri, hyperparasite of rust on
Cestrum diurnum. Mycologia 63: 689–693
Pollack FG (1987) An annotated compilation of Cercospora names.
Mycologia Memoirs 12: 1–212.
Pons N, Sutton BC (1988) Cercospora and similar fungi on yams
(Dioscorea species). Mycological Papers 160: 1–78.
Prasad SS, Verma RAB (1970) A new genus of Moniliales from India.
Indian Phytopathology 23: 111–113.
Pretorius MC, Crous PW, Groenewald JZ, Braun U (2003) Phylogeny
of some cercosporoid fungi from Citrus. Sydowia 55: 286–305.
Quaedvlieg W, Groenewald JZ, de Jesús Yáñez-Morales M, Crous
PW (2012) DNA barcoding of Mycosphaerella species of
quarantine importance to Europe. Persoonia 29: 101–115.
Raabe RD, Ibra L, Conners IL, Martinez AP (1981) Checklist of Plant
Diseases in Hawaii. [Information Text Series no. 022.] Honolulu:
Hawaii Institute of Agriculture and Human Resources, College
of Tropical Agriculture and Human Resources, University of
Hawaii. Ragunanthan V, Prasad NN, Purushothaman D (1970)
A new species of Cercospora causing leaf-blight of Marsilea
quadrifoliata. Current Science 39: 304–305.
Rangel E (1917) Algunos fungos nobas do Brasil. Archivos do Jardim
Botânico do Rio de Janeiro 2: 69–74.
Cercosporoid fungi 1
ARTICLE
345
volume 4 · no. 2
Rao AV, Ramakrishnan K (1965) Prathigada ziziphi sp. nov. Current
Science 34: 60–61.
Rao PR, Manoharachary C, Ramarao P (1982) Eriocercospora
websteri sp. nov. and related species. Current Science 51:
1155–1156.
Rao PR, Rao PR, Manoharachary C (1975) Notes on microfungi from
Andhra Pradesh. III. A new species of Prathigada on Bauhinia.
Nova Hedwigia 26: 437–440.
Saccardo PA (1880) Conspectus generum fungorum Italiae inferiorum,
nempe as Sphaeropsideas, Melanconicas et Hyphomycetas
pertinentium, systemate sporologico dispositorum. Michelia 2:
1–38.
Saccardo PA (1892) Sylloge fungorum omnium hucusque cognitum,
Vol. 10. Padova: P. A. Saccardo.
Saccardo PA (1902) Sylloge fungorum omnium hucusque cognitum,
Vol. 16 [Saccardo PA, Sydow P, eds]. Padova: P. A. Saccardo
Saccardo PA (1906) Sylloge fungorum omnium hucusque cognitum,
Vol. 18 (Supplementum universal, Pars VII) [Saccardo PA,
Saccardo D, eds]. Padova: P. A. Saccardo.
Saccardo PA (1913) Sylloge fungorum omnium hucusque cognitum,
Vol. 22 (Supplementum universal, Pars IX). Padova: P. A.
Saccardo.
Saccardo PA (1931) Sylloge fungorum omnium hucusque cognitum,
Vol. 25 (Trotter A, ed.). Avellino: sumptis coherendum Saccardo
Typis Pergola.
Saccardo PA (1972) Sylloge fungorum omnium hucusque cognitum,
Vol. 26 [Trotter A, ed., published by Cash K]. New York, London:
Johnson.
Sarbajna KK (1990) New foliicolous hyphomycetes from India.
Journal of Economic and Taxonomic Botany 14: 487–492.
Sarbajna KK, Chattopadhayay BK (1990) Studies on “Cercospora-
like fungi” from West Bengal. Journal of Mycopathological
Research 28: 13–18.
Sawada K (1958) Research on fungi in the Tohuku district of Japan
(IV). Fungi Imperfecti. Bulletin of the Government Forest
Experiment Station (Japan) 105: 35–140.
Seifert K, Morgan-Jones G, Gams W, Kendrick B (2011) The Genera
of Hyphomycetes. [CBS Biodiversity Series no. 9.] Utrecht: CBS-
KNAW Fungal Biodiversity Centre.
Shaw DE, Alcorn JL (1993) New name for Verrucispora and its
species. Australian Systematic Botany 6: 273–276.
Shin HD, Kim JD (2001) Cercospora and allied genera from Korea.
Plant Pathogens of Korea 7: 1–303.
Shivas RG, Young AJ, Braun U (2009) Zasmidium macluricola.
Fungal Planet 39. Persoonia 23: 190–191.
Shivas RG, Young AJ, McNeil BC (2010) Pseudocercospora
microsori. Fungal Planet 68. Persoonia 25: 156–157.
Shukla DN, Singh AK, Kumar P, Kamal (1982) Fungi of Gorakhpur-
XIV. Indian Phytopathology 35: 86–91.
Solheim H (2013) Passalora juniperina comb. nov. – new in Norway
or just overlooked? Agarica 33: 73–80.
Solheim WG (1930) Morphological studies of the genus Cercospora.
Illinois Biological Monographs 12(1): 1–85.
Spegazzini C (1910) Mycetes Argentinenses, Ser. V. Anales del
Museo Nacional de Buenos Aires, Ser. 3, 20: 329–467.
Srivastava RK, Srivastava AK, Kamal (1995) [“1992–1993”] – An
appraisal of Indian species of Cercospora Fres. sensu stricto:
some new species and new records. Kavaka 20/21: 37–47.
Stewart EL, Liu Z, Crous PW, Szabo L (1999) Phylogenetic
relationships among some cercosporoid anamorphs of
Mycosphaerella based on rDNA sequence analysis. Mycological
Research 103: 1491–1499.
Subramanian CV (1953) Fungi Imperfecti from Madras–II.
Proceedings, Indian Academy of Sciences, Section B, Biological
Sciences 36: 160–168.
Sutton BC (1971) Sciniatosporium. Transactions of the British
Mycological Society 58: 164–167.
Sutton BC (1975) Coelomycetes V. Coryneum. Mycological Papers
138: 1–224.
Sutton BC (1994) IMI Descriptions of Fungi and Bacteria, Set 119,
Nos 1181–1190. Mycopathologia 125: 43–64.
Sutton BC, Hodges CH (1990) Revision of Cercospora-like fungi on
Juniperus and allied conifers. Mycologia 82: 313–325.
Sutton BC, Shamoun SF, Crous PW (1996) Two leaf pathogens of
Ribes spp. in North America, Quasiphloeospora saximontanensis
(Deighton) comb. nov. and Phloeosporella ribis (J.J. Davis)
comb. nov. Mycological Research 100: 979–983.
Suto Y (1979) Pathogenicity of Cercospora pini-densiorae Hori et
Nambu to various coniferous seedlings. Journal of the Japanese
Forest Society 61(5): 180–183.
Sydow H (1930) Fungi venezuelani. Annales Mycologici 28: 29–224.
Taylor JE, Groenewald JZ, Crous PW (2003) A phylogenetic analysis
of Mycosphaerellaceae leaf spot pathogens of Proteaceae.
Mycological Research 107: 653–658.
Tessmann DJ, Charudattan R, Kistler HC, Rosskopf EN (2001) A
molecular characterization of Cercospora species pathogenic
to water hyacinth and emendation of P. piaropi. Mycologia 93:
323–334.
Thaung MM (1984) Some fungi of Cercospora from Burma.
Mycotaxon 19: 425–452.
Thomma BPHJ, van Esse HP, Crous PW, de Wit PJGM (2005)
Cladosporium fulvum (syn. Passalora fulva), a highly specialized
plant pathogen as a model for functional studies on plant
pathogenic Mycosphaerellaceae. Molecular Plant Pathology 6:
379–393.
To-anun C, Hidayat I, Meeboon J (2011) Genus Cercospora in
Thailand: taxonomy and phylogeny (with a dichotomous key to
species). Plant Pathology & Quarantine 1: 11–87.
Urtiaga R (1986) Indice de Enfermedades en Plantas de Venezuela
y Cuba. Barquisimeto: Published by the author.
Vassiljevsky NI, Karakulin BP (1937) Parazitnye Nesovershennye
Griby. Vol. 1. Gifomicety. Moskva, Leningrad: Izdatel'stvo
Akademii Nauk SSSR.
Vasudeva RS (1963) Indian Cercosporae. New Dehli: Indian Council
of Agricultural Research.
Verkley GJM, Crous PW, Groenewald JZ, Braun U, Aptroot A
(2004) Mycosphaerella punctiformis revisited: morphology,
phylogeny, and epitypication of the type species of the genus
Mycosphaerella (Dothideales, Ascomycota). Mycological
Research 108: 1271–1282.
Viégas AP (1945) Alguns fungos do Brasil, Cercosporae. Boletim da
Sociedade Brasileira de Agronomia 8: 1–160.
Yen JM (1974) Les Cercospora du Gabon. – IV. Bulletin Trimestriel
de la Société Mycologique de France 90: 41–47.
Yen JM, Lim G (1980) Cercospora and allied genera of Singapore
and the Malay Peninsula. Gardens’ Bulletin, Singapore 33: 151–
263.
Zhuang WY (2001) Higher Fungi of Tropical China. Ithaca:
Mycotaxon.
Zhuang WY (2005) Fungi of Northwestern China. Ithaca: Mycotaxon.
