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Studies in Mycology 75: 37–114. 

Phylogenetic lineages in Pseudocercospora 

P.W. Crous 1,2,3 , U. Braun 4 , G.C. Hunter 1,5,6 , M.J. Wingfield 5 , G.J.M. Verkley 1 , H.-D. Shin 7 , C. Nakashima 8 , J.Z. Groenewald 1 

1 

CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT, Utrecht, the Netherlands; 2 Microbiology, Department of Biology, Utrecht University, Padualaan 8, 3584 CH 

Utrecht, the Netherlands; 3 Wageningen University and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands; 

4 

Martin-Luther-Universität, FB. Biologie, Institut für Geobotanik und Botanischer Garten, Neuwerk 21, D-06099 Halle (Saale), Germany; 

5 

Forestry and Agricultural Biotechnology Institute (FABI), University of Pretoria, Pretoria 0002, South Africa; 6 Present address: Forest Research, Alice Holt Lodge, Farnham, 

Surrey GU10 4LH, UK; 7 Division of Environmental Science and Ecological Engineering, Korea University, Seoul 136-701, Korea; 8 Laboratory of Plant Pathology, Graduate 

School of Bioresources, Mie University, Kurima-Machiya 1577, Tsu 514-8507, Japan 

*Correspondence: P.W. Crous, p.crous@cbs.knaw.nl 

Abstract: Pseudocercospora is a large cosmopolitan genus of plant pathogenic fungi that are commonly associated with leaf and fruit spots as well as blights on a wide range 

of plant hosts. They occur in arid as well as wet environments and in a wide range of climates including cool temperate, sub-tropical and tropical regions. Pseudocercospora 

is now treated as a genus in its own right, although formerly recognised as either an anamorphic state of Mycosphaerella or having Mycosphaerella-like teleomorphs. The aim 

of this study was to sequence the partial 28S nuclear ribosomal RNA gene of a selected set of isolates to resolve phylogenetic generic limits within the Pseudocercospora 

complex. From these data, 14 clades are recognised, six of which cluster in Mycosphaerellaceae. Pseudocercospora s. str. represents a distinct clade, sister to Passalora 

eucalypti, and a clade representing the genera Scolecostigmina, Trochophora and Pallidocercospora gen. nov., taxa formerly accommodated in the Mycosphaerella heimii 

complex and characterised by smooth, pale brown conidia, as well as the formation of red crystals in agar media. Other clades in Mycosphaerellaceae include Sonderhenia, 

Microcyclosporella, and Paracercospora. Pseudocercosporella resides in a large clade along with Phloeospora, Miuraea, Cercospora and Septoria. Additional clades represent 

Dissoconiaceae, Teratosphaeriaceae, Cladosporiaceae, and the genera Xenostigmina, Strelitziana, Cyphellophora and Thedgonia. The genus Phaeomycocentrospora is 

introduced to accommodate Mycocentrospora cantuariensis, primarily distinguished from Pseudocercospora based on its hyaline hyphae, broad conidiogenous loci and hila. 

Host specificity was considered for 146 species of Pseudocercospora occurring on 115 host genera from 33 countries. Partial nucleotide sequence data for three gene loci, ITS, 

EF-1α, and ACT suggest that the majority of these species are host specific. Species identified on the basis of host, symptomatology and general morphology, within the same 

geographic region, frequently differed phylogenetically, indicating that the application of European and American names to Asian taxa, and vice versa, was often not warranted. 

Studies in Mycology 

Key words: Capnodiales, Cercospora, cercosporoid, Mycosphaerella, Mycosphaerellaceae, Paracercospora, Pseudocercosporella, Multi-Locus Sequence Typing (MLST), 

systematics. 

Taxonomic novelties: New genera - Pallidocercospora Crous, Phaeomycocentrospora Crous, H.D. Shin & U. Braun; New species - Cercospora eucommiae Crous, U. Braun 

& H.D. Shin, Microcyclospora quercina Crous & Verkley, Pseudocercospora ampelopsis Crous, U. Braun & H.D. Shin, Pseudocercospora cercidicola Crous, U. Braun & C. 

Nakash., Pseudocercospora crispans G.C. Hunter & Crous, Pseudocercospora crocea Crous, U. Braun, G.C. Hunter & H.D. Shin, Pseudocercospora haiweiensis Crous & X. 

Zhou, Pseudocercospora humulicola Crous, U. Braun & H.D. Shin, Pseudocercospora marginalis G.C. Hunter, Crous, U. Braun & H.D. Shin, Pseudocercospora ocimi-basilici 

Crous, M.E. Palm & U. Braun, Pseudocercospora plectranthi G.C. Hunter, Crous, U. Braun & H.D. Shin, Pseudocercospora proteae Crous, Pseudocercospora pseudostigminaplatani 

Crous, U. Braun & H.D. Shin, Pseudocercospora pyracanthigena Crous, U. Braun & H.D. Shin, Pseudocercospora ravenalicola G.C. Hunter & Crous, Pseudocercospora 

rhamnellae G.C. Hunter, H.D. Shin, U. Braun & Crous, Pseudocercospora rhododendri-indici Crous, U. Braun & H.D. Shin, Pseudocercospora tibouchinigena Crous & U. 

Braun, Pseudocercospora xanthocercidis Crous, U. Braun & A. Wood, Pseudocercosporella koreana Crous, U. Braun & H.D. Shin; New combinations - Pallidocercospora 

acaciigena (Crous & M.J. Wingf.) Crous & M.J. Wingf., Pallidocercospora crystallina (Crous & M.J. Wingf.) Crous & M.J. Wingf., Pallidocercospora heimii (Crous) Crous, 

Pallidocercospora heimioides (Crous & M.J. Wingf.) Crous & M.J. Wingf., Pallidocercospora holualoana (Crous, Joanne E. Taylor & M.E. Palm) Crous, Pallidocercospora 

konae (Crous, Joanne E. Taylor & M.E. Palm) Crous, Pallidoocercospora irregulariramosa (Crous & M.J. Wingf.) Crous & M.J. Wingf., Phaeomycocentrospora cantuariensis 

(E.S. Salmon & Wormald) Crous, H.D. Shin & U. Braun, Pseudocercospora hakeae (U. Braun & Crous) U. Braun & Crous, Pseudocercospora leucadendri (Cooke) U. Braun & 

Crous, Pseudocercospora snelliana (Reichert) U. Braun, H.D. Shin, C. Nakash. & Crous, Pseudocercosporella chaenomelis (Y. Suto) C. Nakash., Crous, U. Braun & H.D. Shin; 

Typifications: Epitypifications - Pseudocercospora angolensis (T. Carvalho & O. Mendes) Crous & U. Braun, Pseudocercospora araliae (Henn.) Deighton, Pseudocercospora 

cercidis-chinensis H.D. Shin & U. Braun, Pseudocercospora corylopsidis (Togashi & Katsuki) C. Nakash. & Tak. Kobay., Pseudocercospora dovyalidis (Chupp & Doidge) 

Deighton, Pseudocercospora fukuokaensis (Chupp) X.J. Liu & Y.L. Guo, Pseudocercospora humuli (Hori) Y.L. Guo & X.J. Liu, Pseudocercospora kiggelariae (Syd.) Crous & 

U. Braun, Pseudocercospora lyoniae (Katsuki & Tak. Kobay.) Deighton, Pseudocercospora lythri H.D. Shin & U. Braun, Pseudocercospora sambucigena U. Braun, Crous & 

K. Schub., Pseudocercospora stephanandrae (Tak. Kobay. & H. Horie) C. Nakash. & Tak. Kobay., Pseudocercospora viburnigena U. Braun & Crous, Pseudocercosporella 

chaenomelis (Y. Suto) C. Nakash., Crous, U. Braun & H.D. Shin, Xenostigmina zilleri (A. Funk) Crous; Lectotypification - Pseudocercospora ocimicola (Petr. & Cif.) Deighton; 

Neotypifications - Pseudocercospora kiggelariae (Syd.) Crous & U. Braun, Pseudocercospora lonicericola (W. Yamam.) Deighton, Pseudocercospora zelkovae (Hori) X.J. Liu 

& Y.L. Guo. 

Published online: 22 May 2012; doi:10.3114/sim0005. 

INTRODUCTION 

Until recently, Pseudocercospora was treated as an anamorphic 

genus linked to Mycosphaerella (Mycosphaerellaceae, 

Capnodiales), along with approximately 30 other anamorphic 

Copyright CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. 

genera (Crous 2009). The separation of the Mycosphaerella 

complex into families (Crous et al. 2007a, 2009b) and genera (Crous 

et al. 2009c) based on DNA sequence data and morphology had 

substantial implications for Pseudocercospora. Pseudocercospora 

is now recognised as a holomorphic genus in its own right, several 

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37

Crous et al. 

species of which have Mycosphaerella-like teleomorphs, for 

example, Pseudocercospora fijiensis and its Mycosphaerella-like 

teleomorph that cause black leaf streak of banana (Arzanlou et 

al. 2008). The name Mycosphaerella is restricted to species with 

Ramularia anamorphs (Verkley et al. 2004, Crous et al. 2009c, 

Koike et al. 2011), with Ramularia being an older name than 

Mycosphaerella. A single generic name is now used for species 

of Pseudocercospora (Hawksworth et al. 2011, Wingfield et al. 

2011), in compliance with the recently accepted changes to the 

International Code of Nomenclature for algae, fungi and plants 

(ICN) adoped during the Botanical Congress in Sydney in 2011, in 

particular, the abolishment of Article 59 dealing with pleomorphic 

fungi. 

Species of Pseudocercospora are well recognised as plant 

pathogens, endophytes or saprobes, with some used as biological 

control agents of weeds (Den Breeÿen et al. 2006). They occur on 

a large number of plants, many of which are important ornamentals 

or food crops including fruits, cereals and commercially propagated 

forest trees (Fig. 1). An early hypothesis was that the majority of 

Pseudocercospora species were strictly host specific. Later studies 

have reported that a few species occur on different hosts belonging 

to a single plant family (Deighton 1976, 1979), although DNA data 

or inoculation studies to support wider host ranges has often been 

lacking. 

The classic monograph of the hyphomycete genus Cercospora 

(Chupp 1954) considered morphological features, including the 

structure of conidiomata as well as conidial pigmentation, septation, 

wall thickness, length, width, and shape as valuable features 

to define species within the genus. Chupp’s circumscription of 

Cercospora was rather broadly defined, and the genus was later 

shown to be extremely heterogenous (Deighton 1976). Deighton 

(1976) distinguished different groups within Cercospora based on 

characters such as superficial mycelium (and the texture thereof), 

conidial scar type, conidiophore and conidium pigmentation, 

septation, and conidial catenulation. These additional features 

resulted in many Cercospora species being transferred to several 

alternative genera such as Cercosporella, Mycocentrospora, 

Mycovellosiella, Phaeoramularia, Paracercospora, Passalora, 

Pseudocercospora, Ramularia, Stenella and Stigmina (Deighton 

1971, 1976, 1979, 1987, Braun 1995, 1998). A subsequent 

morphological treatment of names published in Cercospora (Crous 

& Braun 2003) provided some rationalisation, with the following 

concepts proposed for the taxonomic treatment of cercosporoid 

fungi: structure of conidiogenous loci (scars) and hila, as either 

unthickened (or almost so, but slightly darkened or refractive) or 

unthickened; presence or absence of pigmentation in conidiophores 

and conidia. 

Pseudocercospora was originally introduced by Spegazzini 

(1910) based on the type species Pseudocercospora vitis, a 

foliar pathogen of grapevines. The majority of Pseudocercospora 

species known to date are regarded as pathogens on a wide variety 

of plants, predominantly in tropical and sub-tropical environments 

where they cause leaf spots, blights, fruit spot and fruit rot (Chupp 

1954, Deighton 1976, von Arx 1983, Pons & Sutton 1988). Some 

important plant pathogens include the species associated with 

Sigatoka disease on banana (Arzanlou et al. 2007, 2008, 2010, 

Churchill 2010), angular leaf spot of bean (Crous et al. 2006), 

husk spot of macadamia (Beilharz et al. 2003), Cercospora leaf 

spot of olive (Ávila et al. 2005), cactus (Ayala-Escobar et al. 2005), 

avocado (Deighton 1976), and eucalypts (Braun & Dick 2002). 

The importance of these diseases is also reflected in quarantine 

regulations, e.g. for Pseudocercospora angolensis the cause of 

fruit and leaf spot disease on citrus (Pretorius et al. 2003) (Fig. 

2), and P. pini-densiflorae the cause of brown needle blight of pine 

(Evans 1984, Crous et al. 1990). 

Pseudocercospora was established to accommodate 

synnematal analogues of Cercospora, as well as species that 

produce pigmented conidiogenous structures and conidia with 

neither thickened nor darkened conidial hila (Deighton 1976, Braun 

1995) (Fig. 3). It was proposed that Pseudocercospora be divided 

into several genera (Deighton 1976) based on morphological 

differences, a view later supported by several authors (Pons 

& Sutton 1988, Braun 1995, Crous & Braun 1996). Since the 

first study applied DNA phylogenetic analyisis to species in the 

Mycosphaerella complex (Stewart et al. 1999), Pseudocercospora 

has been shown to be heterogenous, accommodating hundreds of 

species (Crous et al. 2000, 2001, Crous & Braun 2003). 

There are very few morphological features that are informative 

at the generic level within the Pseudocercospora complex. 

Deighton (1983) found it difficult to distinguish Cercoseptoria from 

Pseudocercospora on the basis of conidial shape, with conidia 

in the former genus acicular and those in the latter obclavate to 

cylindrical. In delimiting Pseudocercospora as an anamorph of 

Mycosphaerella, von Arx (1983) considered Pseudocercospora 

together in a group of related genera characterised by hyaline or 

subhyaline conidiogenous structures and unthickened, truncate, 

flat and broad conidiogenous loci. Later, Braun (1992) and Crous 

et al. (2000) argued that the arrangement of the conidiophores 

did not distinguish between sections within Pseudocercospora 

due to transitions from solitary to fasciculate to subsynnematal 

conidiophores. Crous et al. (2001) also regarded the slight 

thickening of conidial scars as a taxonomically uninformative 

generic character. 

DNA sequence data for various gene regions have in recent 

years provided substantial information to support the generic 

circumscription of Pseudocercospora. Several studies have 

employed DNA sequence data from the Internal Transcribed 

Spacer (ITS) region of the rDNA operon for Pseudocercospora 

species from various hosts. Crous et al. (2000) examined isolates 

of Pseudocercospora from Eucalyptus and found that they could 

be separated into two clades within Mycosphaerella. Another clade 

of Pseudocercospora species occurred on banana, indicating that 

Pseudocercospora could be polyphyletic within the Mycosphaerella 

complex. Further evidence supporting this view emerged in 

subsequent studies that included many Pseudocercospora isolates 

(Crous et al. 2001). These phylogenetic studies have shown that 

several other genera are congeneric with Pseudocercospora 

and thus Cercostigmina, Paracercospora, Phaeoisariopsis 

and Pseudophaeoramularia were reduced to synonymy with 

Pseudocercospora (Stewart et al. 1999, Crous et al. 2001, Braun & 

Hill 2002, Crous et al. 2006). Based on these studies, the necessity 

arose to conserve Pseudocercospora over Stigmina, which 

represented an older generic name (Braun & Crous 2006). 

Extensive DNA-based phylogenetic research has in recent 

years been conducted on Mycosphaerella and many of its 

anamorphic genera. These studies have not provided substantial 

resolution of Pseudocercospora. The aims of this study were to 

define phylogenetic lineages (reflecting genera) within what is 

perceived to be Pseudocercospora. An additional aim was to use 

the molecular data to infer host range and thus to consider the 

importance of host specificity in this important genus. 

38


Fig. 1. Leaf spot symptoms associated with various species from the Pseudocercospora complex. A. P. fatouae on Fatoua villosa. B. P. clematidis on Clematis apiicola. C. P. 

griseola on Phaseolus vulgaris. D. P. rhododendron-indici on Rhododendron indicum. E. P. pyracanthae on Pyracantha angustifolia. F. P. lonicericola on Lonicera japonica. G. 

Scolecostigmina mangiferae on Mangifera indica. H. P. fraxinites on Fraxinus rhynchophylla. I. Pseudocercosporella potentillae on Potentilla kleiniana. J. Pseudocercospora 

udagawana on Hovenia dulcis. 

Fig. 2. Pseudocercospora species of quarantine importance. A. P. fijiensis on Musa (Black Leaf Streak or Black Sigatoka) (Photo G.H.J. Kema). B, C. P. angolensis on Citrus 

(Phaeoramularia Fruit and Leaf Spot). 

www.studiesinmycology.org 

39

Crous et al. 

Fig. 3. Morphological structures of Pseudocercospora spp. A. Synnematous conidiophore. B. Densely aggregated fascicle of conidiophores with well-developed brown stroma. 

C, D. Loosely branched fascicles of conidiophores with moderate (C) and poorly (D) developed brown stroma. E. Fascicle reduced to conidiogenous cells. F. Conidiophore 

fascicles arising from stomata. G, H. Solitary conidiogenous cells on superficial hyphae. I. Geniculate conidiophore (arrow) with truncate apical locus. J, K. Conidiophores 

branched below (arrows). L. Conidiogenous cells with percurrent proliferations (arrows). M, N. Conidiophores with sympodial proliferation. O. Conidiophores with conidiogenous 

cells (note minutely thickened scars, arrows). P. Subcylindrical conidium with subacute apex and truncate base. Q. Conidia with constrictions at septa. R. Conidium with guttules. 

S. Cylindrical conidium with obtuse apex, and truncate base. T. Undulate conidia. U. Curved conidium. Aseptate to 1-septate conidia. V. 1-septate conidia. W, X. Obclavate 

conidia with obconical base. Y. Obclavate conidium with short obconical base. Z. Dark brown, muriformly euseptate conidia (thick-walled, not distoseptate). 

40


MATERIALS AND METHODS 

Isolates 

Direct isolations were made from fascicles of conidiophores on 

leaves. Some leaves were incubated in moist chambers for up to 

1 wk to enhance sporulation before single conidial colonies were 

established on 2 % malt extract agar (MEA) (Crous 2002). Leaf 

spots bearing ascomata were soaked in water for approximately 

2 h, after which they were attached to the inner surface of Petri 

dish lids over plates containing MEA. Ascospore germination 

patterns were examined after 24 h, and single ascospore and 

conidial cultures established as described previously (Crous et 

al. 1991, Crous 1998). Colonies were sub-cultured onto synthetic 

nutrient-poor agar (SNA), potato-dextrose agar (PDA), oatmeal 

agar (OA), and MEA (Crous et al. 2009d), and incubated at 25 °C 

under continuous near-ultraviolet light to promote sporulation. 

Isolates were also sourced from the culture collections of the 

CBS-KNAW Fungal Biodiversity Centre (CBS), the working 

collection of Pedro Crous (CPC), Chiharu Nakashima (CNS) and 

the culture collection of the laboratory of plant pathology, Mie 

University, Japan (MUCC), and the mycological herbarium of Mie 

University (MUMH). Furthermore, isolates representing fungal 

species from genera allied to Pseudocercospora, e.g. Cercospora, 

Cercostigmina, Cyphellophora, Davidiella, Dissoconium, Miuraea, 

Mycocentrospora, Passalora, Phaeoisariopsis, Phleospora, 

Septoria, Strelitziana, Stigmina, Teratosphaeria, Thedgonia, 

Trochophora, and Xenostigmina, were included in this study (Table 

1). 

DNA isolation 

Mycelium from actively growing fungal cultures was scraped from 

the surface of MEA or PDA plates using a sterile scalpel blade. 

Harvested mycelium was ground to a fine powder using liquid 

nitrogen and DNA was isolated using the CTAB extraction protocol 

as outlined by Crous et al. (2009d) or the UltraClean TM Microbial 

DNA Isolation Kit (MoBio Laboratories, Inc., Solana Beach, CA, 

USA) following the manufacturers’ protocols. Isolated DNA was 

visualised by electrophoresis in 1 % agarose gels (w/v) stained 

with ethidium bromide and viewed under near ultra-violet light. DNA 

concentrations were determined by measuring electrophoresed 

DNA samples against a HyperLadder TM I molecular marker 

(BIOLINE) or alternatively by a NanoDrop quantification as outlined 

by the manufacturer. 

PCR amplification 

DNA isolated from fungal isolates was used as template for further 

Polymerase Chain Reaction (PCR) amplifications. Four nuclear 

gene regions were targeted for PCR amplification and subsequent 

sequencing. These regions included the Internal Transcribed 

Spacer regions ITS-1, ITS-2 and the 5.8S nrRNA gene regions 

(ITS), the first 900 bp of the Large Subunit (28S, LSU) (domains 

D1–D3) of the rDNA operon and partial gene regions of the 

translation elongation factor 1-alpha (EF-1α) and the actin (ACT) 

genes. 

The ITS region was amplified using primers ITS-1 or ITS-5 and 

ITS-4 (White et al. 1990) while primers used for amplification of 

the LSU region were LR0R (Rehner & Samuels 1994) or LSU1Fd 

(Crous et al. 2009b) and LR5 or LR7 (Vilgalys & Hester 1990). 

Primers employed for the amplification of EF-1α included EF1- 

728F and EF1-986R (Carbone & Kohn 1999) or EF-2 (O’Donnell et 

al. 1998) while ACT-512F and ACT-783R (Carbone & Kohn 1999) 

were used to amplify a portion of the ACT gene. All PCR reaction 

mixtures and conditions followed those outlined by Hunter et al. 

(2006b). Following PCR amplification, amplicons were visualized 

on 1.5 % agarose gels stained with ethidium bromide and viewed 

under ultra-violet light and sizes of amplicons were determined 

against a HyperLadder TM I molecular marker (BIOLINE). The PCR 

amplicons for the four loci were subsequently diluted 1 to 10 times 

in preparation for further DNA sequencing reactions. 

DNA sequencing and phylogenetic inference 

PCR amplicons of the four gene regions targeted in this study 

served as templates for DNA sequencing reactions with the 

BigDye® Terminator Cycle Sequencing Kit v. 3.1 (Applied 

Biosystems Life Technologies, Carlsbad, CA, USA) following the 

protocol of the manufacturer. DNA sequencing reactions used the 

same primers as those for the PCR reactions. However, additional 

internal primers LR3R (http://www.biology.duke.edu/fungi/mycolab/ 

primers.htm), LR16 (Moncalvo et al. 1993) and LR5 were used to 

sequence the LSU in order to obtain reliable sequences spanning 

the entire D1-D3 region. DNA sequencing amplicons were purified 

through Sephadex® G-50 Superfine columns (Sigma Aldrich, St. 

Louis, MO) in MultiScreen HV plates (Millipore, Billerica, MA). 

Purified sequence reactions were run on an ABI Prism 3730xl DNA 

Sequencer (Life Technologies, Carlsbad, CA, USA). 

Generated DNA sequence electropherograms were analysed 

using MEGA (Molecular Evolutionary Genetics Analysis) v. 4.0 

(Tamura et al. 2007), 4Peaks v. 1.7.2 (http://www.mekentosj.com/) 

and SeqMan v. 8.0.2. from the DNASTAR Lasergene® software 

package. Consensus sequences were generated and imported 

into MEGA for initial alignment and the construction of sequence 

datasets. DNA sequences representing isolates of closely allied 

genera, for which material could not be obtained were downloaded 

from the NCBI GenBank nucleotide database (www.ncbi.nlm.nih. 

gov) and added to the DNA sequence datasets generated in this 

study. Sequence datasets for the four genomic loci were aligned in 

MAFFT (“Multiple alignment program for amino acids or nucleotide 

sequences”) v. 6.0 (Katoh & Toh 2006, Katoh et al. 2005; http:// 

mafft.cbrc.jp/alignment/server/index.html) using the Auto alignment 

strategy with the 200PAM/ K=2 scoring matrix and a gap opening 

penalty of 1.53 with an offset value of 0.0. Resulting sequence 

alignments were manually evaluated and adjusted in MEGA, 

MacClade v.4.08 (Maddison & Maddison 2000) or Sequence 

Alignment Editor v. 2.0a11 (Rambaut 2002). 

A phylogenetic re-construction was conducted for the aligned 

LSU data set to determine generic relationships using MrBayes 

v. 3.1.2 (Ronquist & Huelsenbeck 2003). Subsequently, a species 

level phylogeny was derived from the combined ITS, ACT and EF- 

1α alignment of Pseudocercospora s. str. sequences using PAUP 

v. 4.0b10 (Swofford 2003). For the LSU alignment, MrModeltest 

v. 2.2 (Nylander 2004) was used to determine the best nucleotide 

substitution model settings for MrBayes. Based on the results of the 

MrModeltest, a phylogenetic analysis was performed with MrBayes 

v. 3.1.2 applying a general time-reversible (GTR) substitution 

model with inverse gamma rates and dirichlet base frequencies 

and a heating parameter set at 0.3. The Markov Chain Monte Carlo 

(MCMC) analysis of 4 chains started in parallel from a random tree 

topology and had 8 000 000 generations. Trees were saved each 


41

Crous et al. 

Table 1. Pseudocercospora and Pseudocercospora-like isolates included in the morphological and/or phylogenetic analyses. 

Species Culture accession numbers 1 Collector Host Family Country GenBank accession numbers 2 

LSU ITS EF-1α ACT 

Cercospora eucommiae CPC 10047 H.D. Shin Eucommia ulmoides Eucommiaceae South Korea GU253741 GU269702 GU384418 GU320406 

CPC 10802; CBS 131932 H.D. Shin Eucommia ulmoides Eucommiaceae South Korea GU214674 GU269851/ 

GU214674 

GU384563 GU320555 

CPC 11508; CBS 132026 H.D. Shin Eucommia ulmoides Eucommiaceae South Korea GU253742 GU269703 GU384419 GU320407 

Cercospora sojina CPC 12322; CBS 132018 H.D. Shin Glycine soja Fabaceae South Korea GU253861 GU214655 JQ324984 JQ325008 

Cyphellophora eucalypti CBS 124764; CPC 13412 P.W. Crous Eucalyptus sp. Myrtaceae Australia GQ303305 GQ303274 GU384510 JQ325009 

Dissoconium dekkeri CBS 110748; CPC 825; CMW 14906 G. Kemp Eucalyptus grandis Myrtaceae South Africa GU214422 AF173315 JQ324985 DQ147651 

Microcyclospora quercina CPC 10712; CBS 130827 G. Verkley Quercus sp. Fagaceae Netherlands GU214681 GU269789 GU384499 GU320490 

Miuraea persicae CPC 10069; CBS 132307 H.D. Shin Prunus persica Rosaceae South Korea GU253859 GU269843 GU384556 GU320546 

CPC 10828; CBS 131935 H.D. Shin Prunus armeniaca Rosaceae South Korea JQ324939 GU269844 GU384557 GU320547 

“Mycosphaerella” laricina CBS 326.52 E. Müller Larix decidua Pinaceae Switzerland GU253693 GU269643 GU384361 GU320353 

“Mycosphaerella” madeirae CBS 112895; CPC 3745 S. Denman Eucalyptus globulus Myrtaceae Portugal DQ204756 AY725553 DQ211672 DQ147641 

“Mycosphaerella” marksii CBS 110920; CPC 935; CMW 5150 A.J. Carnegie Eucalyptus botryoides Myrtaceae Australia DQ246250/ 

GU253694 

AF309588/ 

GU269644 

DQ235134 DQ147625 

Pallidocercospora acaciigena CBS 112516; CPC 3838 M.J. Wingfield Acacia mangium Fabaceae Venezuela GU214661/ 

GU253697 

GU269648 GU384366 GU320356 

CBS 120740; CPC 13290 B. Summerell Eucalyptus sp. Myrtaceae Australia GU253698 EF394822/ 

GU269649 

GU384367 GU320357 

Pallidocercospora crystallina CBS 681.95; CBS 116158; CPC 802; 

CMW 3033 

M.J. Wingfield Eucalyptus bicostata Myrtaceae South Africa DQ204747 AY490757 DQ147636/ 

DQ211662 

DQ147636 

Pallidocercospora heimii CBS 110682; CPC 760; CMW 4942 P.W. Crous Eucalyptus sp. Myrtaceae Madagascar DQ204751 AF309606 DQ211667 DQ147638 

Pallidocercospora heimioides CBS 111190; CPC 1312; CMW 3046 M.J. Wingfield Eucalyptus sp. Myrtaceae Indonesia DQ204753 AF309609 DQ211669 DQ147633 

Pallidocercospora irregulariramosa CBS 114774; CBS 114777; CPC 1360; 

CMW 4943 

M.J. Wingfield Eucalyptus saligna Myrtaceae South Africa DQ204754 AF309607 DQ211670 DQ147634 

Pallidocercospora konae CBS 120748; CPC 13469 W. Himaman Eucalyptus camaldulensis Myrtaceae Thailand GU253852 EF394842 GU384549 GU320538 

Paracercospora egenula CBS 485.81 N. Ponnapa Solanum melongena Solanaceae India JQ324940 GU269699 GU384415 GU320403 

CPC 12537; CBS 132030 H.D. Shin Solanum melongena Solanaceae South Korea GU253738 GU269698 GU384414 GU320402 

MUCC 883 T. Mikami Solanum melongena Solanaceae Japan GU253739 GU269700 GU384416 GU320404 

Passalora eucalypti CBS 111318; CPC 1457 P.W. Crous Eucalyptus saligna Myrtaceae Brazil GU253860 GU269845 GU384558 GU320548 

Phaeomycocentrospora cantuariensis CPC 10157 H.D. Shin Humulus scandens Cannabaceae South Korea GU253712 GU269664 GU384381 GU320370 

CPC 10762; CBS 131928 H.D. Shin Luffa cylindrica Cucurbitaceae South Korea GU253713 GU269665 GU384382 GU320371 

42


Table 1. (Continued). 



CPC 11646; CBS 132013 H.D. Shin Acalypha australis Euphorbiaceae South Korea GU253715 GU269667 GU384384 GU320373 

CPC 11694; CBS 132014 H.D. Shin Humulus scandens Cannabaceae South Korea GU253716 GU269668 GU384385 GU320374 

Phloeospora ulmi CBS 344.97 W. Gams Ulmus glabra Ulmaceae Austria GU253841 JQ324974 JQ324986 GU320528 

CBS 613.81 H.A. Van der Aa Ulmus sp. Ulmaceae Austria GU253842 GU269825 JQ324987 GU320529 

Pseudocercospora abelmoschi CPC 14478; CBS 132103 H.D. Shin Hibiscus syriacus Malvaceae South Korea GU253696 GU269647 GU384365 GU320355 

Pseudocercospora acericola CBS 122279 R. Kirschner Acer albopurpurascens Aceraceae Taiwan GU253699 GU269650 GU384368 GU320358 

Pseudocercospora ampelopsis CPC 11680; CBS 131583 H.D. Shin Ampelopsis brevipenduncula 

var. heterophylla 

Vitaceae South Korea GU253846 GU269830 GU384542 GU320534 

Pseudocercospora angolensis CBS 112933; CPC 4118 M.C. Pretorius Citrus sp. Rutaceae Zimbabwe GU214470 AY260063/ 

GU269836 

GU384548 JQ325010 

CBS 149.53 T. de Carvalho & O. Mendes Citrus sinensis Rutaceae Angola JQ324941 JQ324975 JQ324988 JQ325011 

Pseudocercospora araliae CPC 10154 H.D. Shin Aralia elata Araliaceae South Korea GU253701 GU269652 GU384370 GU320360 

MUCC 873 T. Kobayashi & C. Nakashima Aralia elata Araliaceae Japan GU253702 GU269653 GU384371 GU320361 

Pseudocercospora arecacearum CBS 118406 C.F. Hill Rhopalostylis sapidis Arecaceae New Zealand GU253704 GU269655 GU384373 GU320363 

CBS 118792 C.F. Hill Howea forsteriana Arecaceae New Zealand GU253703 GU269654 GU384372 GU320362 

Pseudocercospora assamensis CBS 122467 I. Buddenhagen Musa cultivar Musaceae India GU253705 GU269656 GU384374 GU320364 

Pseudocercospora atromarginalis CBS 114640 C.F. Hill Solanum sp. Solanaceae New Zealand GU253706 GU269658 GU384376 GU320365 

CPC 11372; CBS 132010 H.D. Shin Solanum nigrum Solanaceae South Korea GU214671 GU269657 GU384375 — 

Pseudocercospora balsaminae CPC 10044; CBS 131882 H.D. Shin Impatiens textori Balsaminaceae South Korea GU253708 GU269660 GU384379 GU320367 

Pseudocercospora basiramifera CBS 111072; CPC 1266 M.J. Wingfield Eucalyptus pellita Myrtaceae Thailand GU253709 GU269661 DQ211677 GU320368 

CBS 114757; CPC 1267 M.J. Wingfield Eucalyptus pellita Myrtaceae Thailand GU253802 GU269781 GU384492 GU320484 

Pseudocercospora basitruncata CBS 114664; CPC 1202 M.J. Wingfield Eucalyptus grandis Myrtaceae Colombia GU253710/ 

DQ204759 

DQ267600/ 

GU269662 

DQ211675 DQ147622 

Pseudocercospora callicarpae MUCC 888 T. Kobayashi Callicarpa japonica Verbenaceae Japan GU253711 GU269663 GU384380 GU320369 

Pseudocercospora catalpigena MUCC 743 C. Nakashima & I. Araki Catalpa ovata Bignoniaceae Japan GU253731 GU269690 GU384406 GU320395 

Pseudocercospora catappae MUCC 809 C. Nakashima & T. Akashi Terminalia catappa Combretaceae Japan GU253717 GU269669 GU384386 GU320375 

Pseudocercospora cercidicola MUCC 896 T. Kobayashi & Y. Kobayashi Cercis chinensis Fabaceae Japan GU253719 GU269671 GU384388 GU320377 

Pseudocercospora cercidis-chinensis CPC 14481; CBS 132109 H.D. Shin Cercis chinensis Fabaceae South Korea GU253718 GU269670 GU384387 GU320376 

Pseudocercospora cf. cruenta CBS 117232 R. Kirschner Phaseolus vulgaris Fabaceae Taiwan GU253730 GU269689 GU384405 GU320394 

Pseudocercospora cf. kaki CPC 10636; CBS 131921 H.D. Shin Diospyros lotus Ebenaceae South Korea GU214677 GU269728 GU384441 GU320430 


43

Crous et al. 




Pseudocercospora chengtuensis CPC 10696; CBS 131924 H.D. Shin Lycium chinense Solanaceae South Korea JQ324942 GU269673 GU384390 GU320379 

MUCC 828 I. Araki & M. Harada Lycium chinense Solanaceae Japan JQ324943 — — — 

Pseudocercospora chionanthi-retusi CPC 14683; CBS 132110 H.D. Shin Chionanthus retusus Oleaceae South Korea GU253721 GU269674 GU384391 GU320380 

Pseudocercospora chrysanthemicola CPC 10633; CBS 131888 H.D. Shin Chrysanthemum sp. Asteraceae South Korea GU253722 GU269675 GU384392 GU320381 

Pseudocercospora cladosporioides CBS 117482; CPC 10913 P.W. Crous Olea europaea Oleaceae Tunisia JQ324944 GU269678 GU384395 GU320383 

“Pseudocercospora” colombiensis CBS 110969; CPC 1106; CMW 4944 M.J. Wingfield Eucalyptus urophylla Myrtaceae Colombia DQ204744 AY752149 DQ211660 DQ147639 

Pseudocercospora contraria CPC 14714; CBS 132108 H.D. Shin Dioscorea quinqueloba Dioscoreaceae South Korea JQ324945 GU269677 GU384394 GU320385 

Pseudocercospora coprosmae CBS 114639 C. F. Hill Coprosma robusta Rubiaceae New Zealand JQ324946 GU269680 GU384397 GU320386 

Pseudocercospora cordiana CBS 114685; CPC 2552 P.W. Crous & R.L. Benchimol Cordia goeldiana Boraginaceae Brazil GU214472 AF362054/ 

GU269681 

GU384398 GU320387 

Pseudocercospora coriariae MUCC 840 I. Araki & M. Harada Coriaria japonica Coriariaceae Japan GU253725 GU269682 GU384399 GU320388 

Pseudocercospora cornicola MUCC 909 C. Nakashima & E. Imaizumi Cornus alba var. sibirica Cornaceae Japan GU253726 GU269683 GU384400 GU320389 

Pseudocercospora corylopsidis MUCC 874 T. Kobayashi & C. Nakashima Hamamelis japonica Hamamelidaceae Japan GU253757 GU269721 GU384437 GU320425 

MUCC 908 C. Nakashima & E. Imaizumi Corylopsis spicata Hamamelidaceae Japan GU253727 GU269684 GU384401 GU320390 

Pseudocercospora cotoneastri MUCC 876 T. Kobayashi & C. Nakashima Cotoneaster salicifolius Rosaceae Japan GU253728 GU269685 GU384402 GU320391 

Pseudocercospora crispans CPC 14883; CBS 125999 P.W.Crous Eucalyptus sp. Myrtaceae South Africa GU253825 GU269807 GU384518 GU320510 

Pseudocercospora crocea CPC 11668; CBS 126004 H.D. Shin Pilea hamaoi Urticaceae South Korea JQ324947 GU269792 GU384502 GU320493 

Pseudocercospora crousii CBS 119487 C.F. Hill Eucalyptus sp. Myrtaceae New Zealand GU253729 GU269686 GU384403 GU320392 

Pseudocercospora cruenta CPC 10846; CBS 132021 H. Booker Vigna sp. Fabaceae Trinidad GU214673 GU269688 GU384404 JQ325012 

Pseudocercospora cydoniae CPC 10678; CBS 131923 H.D. Shin Chaenomeles speciosa Rosaceae South Korea GU253732 GU269691 GU384407 GU320396 

Pseudocercospora cymbidiicola CBS 115132 C.F. Hill Cymbidium sp. Orchidaceae New Zealand GU253733 GU269692 GU384408 GU320397 

Pseudocercospora davidiicola MUCC 296 C. Nakashima & I. Araki Davidia involucrata Nyssaceae Japan GU253734 GU269693 GU384409 GU320398 

Pseudocercospora dendrobii MUCC 596 C. Nakashima & K. Motohashi Dendrobium sp. Orchidaceae Japan GU253737 GU269696 GU384412 GU320401 

Pseudocercospora destructiva MUCC 870 S. Uematsu & C. Nakashima Euonymus japonicus Celastraceae Japan GU253735 GU269694 GU384410 GU320399 

Pseudocercospora dianellae CBS 117746 C.F. Hill Dianella caerulae Liliaceae New Zealand GU253736 GU269695 GU384411 GU320400 

Pseudocercospora dodonaeae CBS 114647 C.F. Hill Dodonaea viscosa Sapindaceae New Zealand JQ324948 GU269697 GU384413 JQ325013 

Pseudocercospora dovyalidis CPC 13771; CBS 126002 P.W. Crous Dovyalis zeyheri Flacourtiaceae South Africa GU253818 GU269800 GU384513 GU320503 

Pseudocercospora elaeocarpi MUCC 925 C. Nakashima Elaeocarpus sp. Elaeocarpaceae Japan GU253740 GU269701 GU384417 GU320405 

“Pseudocercospora” epispermogonia CBS 110750; CPC 822 G. Kemp Eucalyptus grandis Myrtaceae South Africa DQ204757 DQ267596 DQ211673 DQ147629 

44





Pseudocercospora eucalyptorum CBS 110777; CPC 16; CMW 5228 P.W. Crous Eucalyptus nitens Myrtaceae South Africa DQ204762 AF309598 DQ211678 DQ147614 

CBS 114242; CPC 10390; CMW 14908 J.P. Mansilla Eucalyptus globulus Myrtaceae Spain GU214481 AY725526 DQ211681 DQ147613/ 

GU320465 

CBS 116359; CPC 3751 P.W. Crous Eucalyptus sp. Myrtaceae Madeira GU253829 GU269812 GU384524 GU320514 

CPC 10500; CBS 114243 P.W. Crous Eucalyptus nitens Myrtaceae New Zealand JQ324949 AY725527 GU384474 JQ325014 

CPC 10507; CBS 116371 P.W.Crous Eucalyptus nitens Myrtaceae New Zealand JQ324950 GU269687 JQ324989 GU320393 

CPC 10916 P.W. Crous Eucalyptus sp. Myrtaceae South Africa GU253788 GU269763 GU384475 GU320464 

CPC 11713; CBS 132015 P. Mansilla Eucalyptus globulus Myrtaceae Spain JQ324951 GU269811 GU384523 JQ325015 

CPC 12406; CBS 132029 I. Smith Eucalyptus globulus Myrtaceae Australia GU253811 GU269793 GU384503 GU320494 

CPC 12568; CBS 132309 C. Mohammed Eucalyptus nitens Myrtaceae Australia GU253814 GU269796 GU384506 GU320497 

CPC 12802; CBS 132032 A. Phillips Eucalyptus globulus Myrtaceae Portugal GU253789 JQ324976 JQ324990 GU320466 

CPC 12957; CBS 132033 B. Summerell Eucalyptus deanei Myrtaceae Australia GU253815 GU269797 JQ324991 JQ325016 

CPC 13455; CBS 132034 P.W. Crous Eucalyptus sp. Myrtaceae Portugal GU253816 GU269798 GU384511 GU320501 

CPC 13769; CBS 132035 P.W. Crous Eucalyptus punctata Myrtaceae South Africa GU253707 GU269659 GU384378 GU320366 

CPC 13816; CBS 132114 S. Denman Eucalyptus glaucescens Myrtaceae UK GU253819 GU269801 JQ324992 GU320504 

CPC 13926; CBS 132105 S. Denman Eucalyptus sp. Myrtaceae USA GU253820 GU269802 JQ324993 GU320505 

Pseudocercospora eupatoriella CBS 113372 M.J. Morris Chromolaena odorata Asteraceae Jamaica GU253743 GU269704 GU384420 GU320408 

Pseudocercospora eustomatis CBS 110822 G. Dal Bello Eustroma grandiflorum Gentianaceae Argentina GU253744 GU269705 GU384421 GU320409 

Pseudocercospora exosporioides MUCC 893 T. Kobayashi Sequoia sempervirens Taxodiaceae Japan GU253746 GU269707 GU384423 GU320411 

Pseudocercospora fijiensis CBS 120258; CIRAD 86 J. Carlier Musa sp. Musaceae Cameroon JQ324952 EU514248 Genome 3 Genome 3 

MUCC 792 T. Kobayashi & C. Nakashima Musa sp. Musaceae Japan GU253776 GU269748 JQ324994 GU320450 

Pseudocercospora flavomarginata CBS 118841; CMW 13586 M.J. Wingfield Eucalyptus camaldulensis Myrtaceae Thailand DQ153306 DQ155657 DQ156548 DQ166513 

CBS 124990; CPC 13492 W. Himaman Eucalyptus camaldulensis Myrtaceae Thailand GU253817 GU269799 GU384512 GU320502 

CPC 14142; CBS 126001 X. Zhou Eucalyptus sp. Myrtaceae China GU253822 GU269804 GU384515 GU320507 

Pseudocercospora fori CBS 113285; CMW 9095 G.C. Hunter Eucalyptus grandis Myrtaceae South Africa DQ204748 AF468869 DQ211664 DQ147618 

CPC 14880; CBS 132113 P.W. Crous Eucalyptus sp. Myrtaceae South Africa GU253824 GU269806 GU384517 GU320509 

Pseudocercospora fraxinites CPC 10743; CBS 131927 H.D. Shin Fontanesia phillyraeoides Oleaceae South Korea GU253720 GU269672 GU384389 GU320378 

MUCC 891 T. Kobayashi Fraxinus excelsior Oleaceae Japan GU253748 GU269710 GU384426 GU320414 

Pseudocercospora fukuokaensis CPC 14689; CBS 132111 H.D. Shin Styrax japonicus Styracaceae South Korea GU253750 GU269713 GU384429 GU320417 

MUCC 887 T. Kobayashi Styrax japonicus Styracaceae Japan GU253751 GU269714 GU384430 GU320418 


45

Crous et al. 




Pseudocercospora fuligena CPC 12296; CBS 132017 Z. Mersha Lycopersicon sp. Solanaceae Thailand JQ324953 GU269711 GU384427 GU320415 

MUCC 533 C. Nakashima Lycopersicon esculentum Solanaceae Japan GU253749 GU269712 GU384428 GU320416 

Pseudocercospora glauca CPC 10062; CBS 131884 H.D. Shin Albizzia julibrissin Fabaceae South Korea GU253752 GU269715 GU384431 GU320419 

Pseudocercospora gracilis CBS 243.94; CPC 730 P.W. Crous Eucalyptus urophylla Myrtaceae Indonesia DQ204750 DQ267582 DQ211666 DQ147616 

Pseudocercospora griseola f. griseola CBS 119112; CPC 10460 F.S. Ngulu & C. Mushi Phaseolus vulgaris Fabaceae Tanzania GU253753 GU269717 GU384433 GU320421 

CBS 194.47 — Phaseolus vulgaris Fabaceae Portugal JQ324954 DQ289801 JQ324995 DQ289868 

CBS 880.72 H.A. van Kesteren Phaseolus vulgaris Fabaceae Netherlands GU214476 GU269716 GU384432 GU320420 

CPC 10462 M.M. Liebenberg Phaseolus vulgaris Fabaceae South Africa GU253865 GU269849 GU384562 GU320553 

CPC 10480; CBS 131887 M.M. Liebenberg Phaseolus vulgaris Fabaceae South Africa GU253864 GU269848 GU384561 DQ289882 

CPC 10779; CBS 131929 H.D. Shin Phaseolus vulgaris Fabaceae South Korea GU253862 GU269846 GU384559 DQ289885 

CPC 12239 G. Mahuku Phaseolus vulgaris Fabaceae Colombia GU253863 GU269847 GU384560 DQ289887 

Pseudocercospora guianensis MUCC 855 C. Nakashima & T. Akashi Lantana camara Verbenaceae Japan GU253755 GU269719 GU384435 GU320423 

MUCC 879 C. Nakashima Lantana camara Verbenaceae Japan GU253756 GU269720 GU384436 GU320424 

Pseudocercospora haiweiensis CPC 14084; CBS 131584 X. Zhou Eucalyptus sp. Myrtaceae China GU253821 GU269803 GU384514 GU320506 

Pseudocercospora hakeae CBS 112226; CPC 3145 P.W. Crous & B. Summerell Grevillea sp. Proteaceae Australia GU253805 GU269784 GU384495 JQ325017 

Pseudocercospora humuli MUCC 742 C. Nakashima & I. Araki Humulus lupulus var. lupulus Cannabaceae Japan GU253758 GU269725 GU384439 GU320428 

Pseudocercospora humulicola CPC 10049; CBS 131883 H.D. Shin Humulus scandens Cannabaceae South Korea JQ324955 GU269724 JQ324996 JQ325018 

CPC 11358; CBS 131585 H.D. Shin Humulus scandens Cannabaceae South Korea JQ324956 GU269723 GU384438 GU320427 

Pseudocercospora indonesiana CBS 122473 I.W. Buddenhagen Musa sp. Musaceae Sumatra GU253765 GU269735 GU384448 GU320437/ 

EU514340 

CBS 122474 I.W. Buddenhagen Musa sp. Musaceae Indonesia JQ324957 EU514283 JQ324997 JQ325019 

Pseudocercospora ixorae CBS 118760 R. Kirschner Ixora sp. Rubiaceae Taiwan GU253759 GU269726 GU384440 GU320429 

Pseudocercospora jussiaeae CPC 14625; CBS 132117 H.D. Shin Ludwigia prostrata Onagraceae South Korea JQ324958 JQ324977 JQ324998 JQ325020 

Pseudocercospora kaki MUCC 900 S. Uematsu & C. Nakashima Diospyros kaki Ebenaceae Japan GU253761 GU269729 GU384442 GU320431 

Pseudocercospora kiggelariae CPC 11853; CBS 132016 W. Gams Kiggelaria africana Flacourtiaceae South Africa GU253762 GU269730 GU384443 GU320432 

Pseudocercospora latens MUCC 763 C. Nakashima & T. Akashi Lespedeza wilfordii Fabaceae Japan GU253763 GU269732 GU384445 GU320434 

Pseudocercospora leucadendri CPC 1869 S. Denman & P.W. Crous Leucadendron sp. Proteaceae South Africa GU214480 GU269842 GU384555 GU320545 

Pseudocercospora libertiae CBS 114643 C.F. Hill Libertia ixioides Iridaceae New Zealand JQ324959 GU269733 GU384446 GU320435 

Pseudocercospora lilacis CPC 12767; CBS 132031 C. Hodges Ligustrum japonicum Oleaceae USA GU253767 GU269737 GU384449 GU320439 

46





Pseudocercospora longispora CBS 122470 D.R. Jones Musa sp. Musaceae Malaysia GU253764 GU269734 GU384447 GU320436/ 

EU514342 

Pseudocercospora lonicericola MUCC 889 T. Kobayashi Lonicera gracilipes var. glabra Caprifoliaceae Japan GU253766 GU269736 JQ324999 GU320438 

Pseudocercospora luzardii CPC 2556 A.C. Alfenas Hancornia speciosa Apocynaceae Brazil GU214477 AF362057/ 

GU269738 

GU384450 GU320440 

Pseudocercospora lyoniae MUCC 910 C. Nakashima & E. Imaizumi Lyonia ovalifolia var. elliptica Ericaceae Japan GU253768 GU269739 GU384451 GU320441 

Pseudocercospora lythracearum CPC 10707; CBS 131925 H.D. Shin Lagerstroemia indica Lythraceae South Korea GU253769 GU269740 GU384452 GU320442 

MUCC 890 T. Kobayashi Lagerstroemia indica Lythraceae Japan GU253770 GU269741 GU384453 GU320443 

Pseudocercospora lythri CPC 14588; CBS 132115 H.D. Shin Lythrum salicaria Lythraceae South Korea GU253771 GU269742 GU384454 GU320444 

MUCC 865 I. Araki & M. Harada Lythrum salicaria Lythraceae Japan GU253772 GU269743 GU384455 GU320445 

Pseudocercospora macrospora CBS 114696; CPC 2553 P.W. Crous & R.L. Benchimol Bertholletia excelsa Lecythidaceae Brazil GU214478 AF362055/ 

GU269745 

GU384457 GU320447 

Pseudocercospora mali MUCC 886 T. Kobayashi Malus sieboldii Rosaceae Japan GU253773 GU269744 GU384456 GU320446 

Pseudocercospora marginalis CPC 12497; CBS 131582 H.D. Shin Fraxinus rhynchophylla Oleaceae South Korea GU253812 GU269794 GU384504 GU320495 

Pseudocercospora melicyti CBS 115023 M. Fletcher Melicytus macrophyllus Violaceae New Zealand JQ324968 GU269769 GU384481 GU320472 

Pseudocercospora metrosideri CBS 118795 C.F. Hill Metrosideros collina Myrtaceae New Zealand GU253774 GU269746 GU384458 GU320448 

Pseudocercospora musae CBS 116634 J. Carlier Musa sp. Musaceae Cuba GU253775 GU269747 GU384459 GU320449 

Pseudocercospora myrticola MUCC 632 C. Nakashima & K. Motohashi Myrtus communis Myrtaceae Japan GU253777 GU269749 GU384460 GU320451 

Pseudocercospora nandinae CBS 117745 C.F. Hill Nandina domestica Berberidaceae New Zealand GU253778 GU269750 GU384461 GU320452 

Pseudocercospora natalensis CBS 111069; CPC 1263 T. Coutinho Eucalyptus nitens Myrtaceae South Africa DQ267576 DQ303077 JQ325000 DQ147620 

CBS 111071; CPC 1265 T. Coutinho Eucalyptus nitens Myrtaceae South Africa GU253801 GU269780 GU384491 GU320483 

Pseudocercospora nephrolepidis CBS 119121 R. Kirschner Nephrolepis auriculata Oleandraceae Taiwan GU253779 GU269751 GU384462 GU320453 

Pseudocercospora nogalesii CBS 115022 C.F. Hill Chamaecytisus proliferus Fabaceae New Zealand JQ324960 GU269752 GU384463 GU320454 

Pseudocercospora norchiensis CBS 114641 C.F. Hill Rubus sp. Rosaceae New Zealand GU253794 GU269772 GU384484 GU320475 

CBS 120738; CPC 13049 W. Gams Eucalyptus sp. Myrtaceae Italy GU253780 EF394859/ 

GU269753 

GU384464 GU320455 

Pseudocercospora ocimi-basilici CPC 10283 M.E. Palm Ocimum basilicum Lamiaceae Mexico GU214678 GU269754 GU384465 GU320456 

Pseudocercospora oenotherae CPC 10290; CBS 131885 H.D. Shin Oenothera odorata Onagraceae South Korea JQ324961 GU269856 GU384567 GU320559 

CPC 10630; CBS 131920 H.D. Shin Oenothera odorata Onagraceae South Korea GU253781 GU269755 GU384466 GU320457 

Pseudocercospora paederiae CPC 10007 H.D. Shin Paederia foetida Rubiaceae South Korea GU253783 GU269757 GU384468 — 

Pseudocercospora palleobrunnea CBS 124771; CPC 13387 P.W. Crous Syzygium sp. Myrtaceae Australia GQ303319 GQ303288 GU384509 GU320500 


47

Crous et al. 




Pseudocercospora pallida CPC 10776; CBS 131889 H.D. Shin Campsis grandiflora Bignoniaceae South Korea GU214680 GU269758 GU384469 GU320459 

Pseudocercospora pancratii CBS 137.94 R.F. Castaneda — — Cuba GU253784 GU269759 GU384470 GU320460 

Pseudocercospora paraguayensis CBS 111286; CPC 1459 P.W. Crous Eucalyptus nitens Myrtaceae Brazil GU214479/ 

DQ204764 

DQ267602 DQ211680 DQ147606 

CBS 111317; CPC 1458 P.W. Crous Eucalyptus nitens Myrtaceae Brazil GQ852634 JQ324978 GU384522 JQ325021 

Pseudocercospora pini-densiflorae MUCC 534 Y. Tokushige Pinus thunbergii Pinaceae Japan GU253785 GU269760 GU384471 GU320461 

Pseudocercospora plecthranthi CPC 11462; CBS 131586 H.D. Shin Plectranthus sp. Lamiaceae South Korea JQ324962 GU269791 GU384501 GU320492 

Pseudocercospora pouzolziae CBS 122280 R. Kirschner Gonostegia hirta Urticaceae Taiwan GU253786 GU269761 GU384472 GU320462 

Pseudocercospora profusa CPC 10042 H.D. Shin Acalypha australis Euphorbiaceae South Korea GU253808 GU269787 GU384497 GU320488 

CPC 10055; CBS 132306 H.D. Shin Acalypha australis Euphorbiaceae South Korea GU253787 GU269762 GU384473 GU320463 

Pseudocercospora proteae CPC 15217; CBS 131587 F. Roets Protea mundii Proteaceae South Africa GU253826 GU269808 GU384519 GU320511 

Pseudocercospora prunicula CPC 14511; CBS 132107 H.D. Shin Prunus x yedoensis Rosaceae South Korea GU253723 GU269676 GU384393 GU320382 

Pseudocercospora pseudostigminaplatani 

CPC 11726; CBS 131588 H.D. Shin Platanus occidentalis Platanaceae South Korea JQ324963 GU269857 GU384568 GU320560 

Pseudocercospora puderi MUCC 906 S. Maruyama Rosa sp. Rosaceae Japan GU253790 GU269764 GU384476 GU320467 

Pseudocercospora punctata CPC 14734; CBS 132116 P.W. Crous Syzygium sp. Myrtaceae Madagascar GU253791 GU269765 GU384477 GU320468 

Pseudocercospora purpurea CBS 114163; CPC 1664 P.W. Crous Persea americana Lauraceae Mexico GU253804 GU269783 GU384494 GU320486 

Pseudocercospora pyracanthae MUCC 892 T. Kobayashi & C. Nakashima Pyracantha angustifolia Rosaceae Japan GU253792 GU269767 GU384479 GU320470 

Pseudocercospora pyracanthigena CPC 10808; CBS 131589 H.D. Shin Pyracantha angustifolia Rosaceae South Korea — GU269766 GU384478 GU320469 

Pseudocercospora ranjita CPC 11141; CBS 126005 M.J. Wingfield Gmelina sp. Verbenaceae Indonesia GU253810 GU269790 GU384500 GU320491 

Pseudocercospora ravenalicola CBS 122468 M. Arzanlou & W. Gams Ravenala madagascariensis Strelitziaceae India GU253828 GU269810 GU384521 GU320513 

Pseudocercospora rhabdothamni CBS 114872 M. Fletcher Rhabdothamnus solandri Gesneriaceae New Zealand JQ324964 GU269768 GU384480 GU320471 

Pseudocercospora rhamnellae CPC 12500; CBS 131590 H.D. Shin Rhamnella frangulioides Rhamnaceae South Korea GU253813 GU269795 GU384505 GU320496 

Pseudocercospora rhapisicola CBS 282.66 K. Tubaki Rhapis flabellifornis Arecaceae Japan GU253793 GU269770 GU384482 GU320473 

Pseudocercospora rhododendri-indici CPC 10822; CBS 131591 H.D. Shin Rhododendron indicum Ericaceae South Korea JQ324965 GU269722 — GU320426 

Pseudocercospora rhoina CPC 11464; CBS 131891 H.D. Shin Rhus chinensis Anacardiaceae South Korea JQ324966 GU269771 GU384483 GU320474 

Pseudocercospora robusta CBS 111175; CPC 1269; CMW 5151 M.J. Wingfield Eucalyptus robur Myrtaceae Malaysia DQ204767 AY309597 DQ211683 DQ147617 

Pseudocercospora rubi MUCC 875 T. Kobayashi & C. Nakashima Rubus allegheniensis Rosaceae Japan GU253795 GU269773 GU384485 GU320476 

Pseudocercospora rumohrae CBS 117747 C.F. Hill Marattia salicina Marattiaceae New Zealand GU253796 GU269774 GU384486 GU320477 

Pseudocercospora sambucigena CPC 10292; CBS 131886 H.D. Shin Sambucus williamsii Caprifoliaceae South Korea GU253809 GU269788 GU384498 GU320489 

48





CPC 14397; CBS 126000 P.W. Crous Sambucus nigra Caprifoliaceae Netherlands GU253823 GU269805 GU384516 GU320508 

Pseudocercospora sawadae CBS 115024 C.F. Hill Psidium guajava Myrtaceae New Zealand JQ324967 GU269775 — GU320478 

Pseudocercospora securinegae CPC 10793; CBS 131930 H.D. Shin Flueggea suffruticosa Euphorbiaceae South Korea GU253797 GU269776 GU384487 GU320479 

Pseudocercospora snelliana CPC 11654; CBS 131592 H.D. Shin Morus bombycis Moraceae South Korea — GU269731 GU384444 GU320433 

Pseudocercospora sordida MUCC 913 C. Nakashima & E. Imaizumi Campsis radicans Bignoniaceae Japan GU253798 GU269777 GU384488 GU320480 

Pseudocercospora sp. CBS 110993; CPC 1057 M.J. Wingfield Populus sp. Salicaceae South Africa GU253800 GU269779 GU384490 GU320482 

CBS 110998; CPC 1054 M.J. Wingfield Eucalyptus grandis Myrtaceae South Africa GU253799 GU269778 GU384489 GU320481 

CBS 111373; CPC 1493 M.J. Wingfield Eucalyptus globulus Myrtaceae Uruguay GU253803 GU269782 GU384493 GU320485 

CBS 112725; CPC 3961 K.A. Seifert Melilotus alba Fabaceae Canada GU253806 GU269785 — — 

CBS 113387 A. den Breeyen Lantana camara Verbenaceae Jamaica GU253754 GU269718 GU384434 GU320422 

CPC 10058 H.D. Shin Potentilla kleiniana Rosaceae South Korea — JQ324979 JQ325001 JQ325022 

CPC 10645; CBS 131922 P.W. Crous — — Brazil GU253700 GU269651 GU384369 GU320359 

CPC 14711; CBS 132102 H.D. Shin Pyracantha angustifolia Rosaceae South Korea — JQ324980 JQ325002 JQ325023 

CPC 15116; NC1 37A1a J. Batzer Malus sp. cv. Golden Delicious Rosaceae USA: North 

Carolina 

JQ324969 JQ324981 JQ325003 JQ325024 

Pseudocercospora stahlii CBS 117549 R. Kirschner Passiflora foetida Passifloraceae Taiwan GU253830 GU269813 GU384525 GU320515 

Pseudocercospora stephanandrae MUCC 914 C. Nakashima & E. Imaizumi Stephanandra incisa Rosaceae Japan GU253831 GU269814 GU384526 GU320516 

Pseudocercospora subsessilis CBS 136.94 R.F. Castaneda — — Cuba GU253832 GU269815 GU384527 GU320517 

Pseudocercospora subtorulosa CBS 117230 R. Kirschner Melicope sp. Rutaceae Taiwan GU253833 GU269816 GU384528 GU320518 

Pseudocercospora subulata CBS 118489; CPC 10849 M. Dick Eucalyptus botryoides Myrtaceae New Zealand JQ324970 DQ303090 JQ325004 GU320519 

Pseudocercospora tereticornis CBS 124996; CPC 12960 A.J. Carnegie Eucalyptus nitens Myrtaceae Australia GQ852647 JQ324982 GU384377 JQ325025 

CPC 13299; CBS 125214 P.W. Crous Eucalyptus tereticornis Myrtaceae Australia GQ852649 GQ852770 GU384508 GU320499 

“Pseudocercospora” thailandica CBS 116367; CPC 10547 K. Pongpanich Acacia mangium Fabaceae Thailand GU253837 — DQ835102/ 

GU384533 

GU320523/ 

AY752217 

CPC 10548; CBS 116367 K. Pongpanich Acacia mangium Fabaceae Thailand GU253853 AY752157 AY840477 GU320539 

Pseudocercospora theae CBS 128.30 M. Curzi Camelia sinensis Theaceae Italy GU253838 GU269821 GU384534 GU320524 

“Pseudocercospora” tibouchinigena CBS 116462 C.F. Hill Tibouchina sp. Melastomataceae New Zealand GU253839 GU269822 GU384535 GU320525 

Pseudocercospora timorensis MUCC 819 C. Nakashima & T. Akashi Ipomoea indica Convolvulaceae Japan GU253840 GU269823 GU384536 GU320526 

Pseudocercospora udagawana CPC 10799; CBS 131931 H.D. Shin Hovenia dulcis Rhamnaceae South Korea — GU269824 GU384537 GU320527 


49

Crous et al. 



Pseudocercospora variicolor MUCC 746 C. Nakashima & I. Araki Paeonia lactiflora var. 

trichocarpa 


Paeoniaceae Japan GU253843 GU269826 GU384538 GU320530 

Pseudocercospora viburnigena CPC 15249; CBS 125998 M.L. Crous Viburnum davidii Caprifoliaceae Netherlands GU253827 GU269809 GU384520 GU320512 

Pseudocercospora viticicola MUCC 777 C. Nakashima Vitex trifolia Verbenaceae Japan GU253845 GU269828 GU384540 GU320532 

Pseudocercospora vitis CPC 11595; CBS 132012 H.D. Shin Vitis vinifera Vitaceae South Korea GU214483 DQ289829/ 

GU269829 

GU384541 GU320533 

CPC 14661; CBS 132112 H.D. Shin Vitis vinifera Vitaceae South Korea GU253844 GU269827 GU384539 GU320531 

Pseudocercospora weigelae MUCC 899 T. Kobayashi & Y. Kobayashi Weigela coraeensis Caprifoliaceae Japan GU253847 GU269831 GU384543 GU320535 

Pseudocercospora xanthocercidis CPC 11665; CBS 131593 A.R. Wood Xanthocercis zambesiaca Fabaceae South Africa JQ324971 JQ324983 JQ325005 JQ325026 

Pseudocercospora xanthoxyli CPC 10065 H.D. Shin Xanthoxylum ailanthoides Rutaceae South Korea GU253848 GU269832 GU384544 GU320536 

Pseudocercospora zelkovae CPC 14484; CBS 132106 H.D. Shin Zelkova serrata Ulmaceae South Korea GU253849 GU269833 GU384545 JQ325027 

CPC 14717; CBS 132118 H.D. Shin Zelkova serrata Ulmaceae South Korea GU253850 GU269834 GU384546 JQ325028 

MUCC 872 T. Kobayashi & C. Nakashima Zelkova serrata Ulmaceae Japan GU253851 GU269835 GU384547 GU320537 

Pseudocercosporella arcuata CPC 10050 H.D. Shin Rubus oldhamii Rosaceae South Korea GU214685 GU269850 JQ325006 GU320554 

Pseudocercosporella capsellae CPC 14773; CBS 131896 H.D. Shin Raphanus sativus Brassicaceae South Korea GU253714 GU269666 GU384383 GU320372 

Pseudocercosporella chaenomelis CPC 14795; CBS 131897 H.D. Shin Chaenomeles speciosa Rosaceae South Korea GU253834 GU269817 GU384530 GU320520 

MUCC 1510; CBS 132131 C. Nakashima Chaenomeles sinensis Rosaceae Japan — JQ793663 — JQ793664 

Pseudocercosporella fraxini CPC 11509 H.D. Shin Fraxinus rhynchophylla Oleaceae South Korea GU214682 GU269709 GU384425 GU320413 

Pseudocercosporella koreana CPC 11414 H.D. Shin Vicia amurensis Fabaceae South Korea GU214683 GU269852 GU384564 GU320556 

Pseudocercosporella oxalidis CBS 118758 R. Kirschner Oxalis debilis Oxalidaceae Taiwan GU253782 GU269756 GU384467 GU320458 

Pseudocercosporella sp. CPC 10864; CBS 131890 H.D. Shin Trigonotis peduncularis Boraginaceae South Korea JQ324972 GU269858 GU384569 JQ325029 

Pseudocercosporella zelkovae CPC 11592; CBS 132011 H.D. Shin Zelkova serrata Ulmaceae South Korea GU214482 GU269853 — GU320557 

Scolecostigmina mangiferae CBS 125467; CPC 17351 P.W. Crous Mangifera indica Anacardiaceae Australia GU253877 GU269870 GU384578 GU320566 

CPC 17352; CBS 125467 P.W. Crous Mangifera indica Anacardiaceae Australia GU253878 GU269871 GU384579 GU320567 

Septoria cerastii CPC 12343; CBS 132028 H.D. Shin Cerastium holosteoides var. 

hallasanense 

Caryophyllaceae South Korea GU253869 GU269859 GU384570 JQ325030 

Septoria chelidonii CPC 12337; CBS 132027 H.D. Shin Chelidonium majus var. 

asiaticum 

Papaveraceae South Korea GU253870 GU269860 GU384571 GU320561 

Septoria crepidis CPC 12539; CBS 131895 H.D. Shin Crepis japonica Asteraceae South Korea GU253871 GU269861 GU384572 GU320562 

Septoria dysentericae CPC 12328; CBS 131892 H.D. Shin Inula britannica var. chinensis Asteraceae South Korea GU253866 GU269854 GU384565 GU320558 

Septoria erigerontis CPC 12340; CBS 131893 H.D. Shin Erigeron annuus Asteraceae South Korea GU253872 GU269862 GU384573 JQ325031 

50





Septoria eucalyptorum CPC 11282; CBS 118505 W. Gams Eucalyptus sp. Myrtaceae India GU253873 GU269863 GU384574 GU320563 

Septoria justiciae CPC 12509; CBS 131894 H.D. Shin Justicia procumbens Acanthaceae South Korea GU253874 GU269864 GU384575 GU320564 

Septoria quercicola CBS 663.94 H.A. van der Aa Quercus robur Fagaceae Netherlands GU253867 GU269855 GU384566 JQ325032 

Septoria rubi CPC 12331; CBS 132022 H.D. Shin Rubus crataegifolius Rosaceae South Korea GU253875 GU269865 GU384576 — 

Stigmina platani CBS 336.33 R.M. Nattrass Platanus orientalis Platanaceae India GU253868 — JQ325007 — 

Strelitziana australiensis CBS 124778; CPC 13421 P.W. Crous Eucalyptus sp. Myrtaceae Australia GQ303326 GQ303295 GU384362 — 

CPC 13556; CBS 132310 P.W. Crous Eucalyptus sp. Myrtaceae Australia GU253695 GU269645 GU384363 GU320354 

Teratosphaeria alcornii CBS 313.76; CPC 3632 J.L. Alcorn Eucalyptus tessellaris Myrtaceae Australia GU253876 GU269866 GU384577 GU320565 

Teratosphaeria dimorpha CPC 14132; CBS 124051 B.A. Summerell Eucalyptus caesia Myrtaceae Australia FJ493215 FJ023537 — — 

Teratosphaeria stellenboschiana CBS 124989; CPC 13767 P.W. Crous Eucalyptus punctata Myrtaceae South Africa GQ852715 GQ852823 — — 

Thedgonia ligustrina CPC 10019 H.D. Shin Ligustrum ovalifolium Oleaceae South Korea GU253854 GU269837 GU384550 GU320540 

CPC 10530; CBS 132130 P.W.Crous Ligustrum sp. Oleaceae Netherlands GU253855 GU269838 GU384551 GU320541 

CPC 10861; CBS 132025 H.D. Shin Ligustrum ovalifolium Oleaceae South Korea GU253856 GU269839 GU384552 GU320542 

Trochophora fasciculata CPC 10282 H.D. Shin Daphniphyllum macropodum Daphniphyllaceae South Korea FJ839668 FJ839632 — — 

Trochophora simplex CBS 124744 H.D. Shin Daphniphyllum macropodum Daphniphyllaceae South Korea GU253880 GU269872 GU384580 GU320568 

MUCC 952 C. Nakashima & I. Araki Daphniphyllum teijsmannii Daphniphyllaceae Japan GU253879 — — — 

Xenostigmina zilleri CBS 115685 K.A. Seifert Acer sp. Aceraceae Canada GU253857 GU269840 GU384553 GU320543 

CBS 115686 K.A. Seifert Acer sp. Aceraceae Canada FJ839676/ 

GU253858 

GU269841 GU384554 GU320544 

Zasmidium nabiacense CBS 125010; CPC 12748 A.J. Carnegie Eucalyptus sp. Myrtaceae Australia GQ852734 GQ852841 GU384507 GU320498 

1 CBS: CBS-KNAW Fungal Biodiversity Centre, Utrecht, The Netherlands; CIRAD: Centre de Coopération Internationale en Recherche Agronomique pour le Développement, UMR-BGPI, Montpellier, France; CMW: Culture Collection of the Forestry and 

Agricultural Biotechnology Institute (FABI) of the University of Pretoria, Pretoria, South Africa; CPC: Culture collection of Pedro Crous, housed at CBS; MUCC: Culture Collection, Laboratory of Plant Pathology, Mie University, Tsu, Mie Prefecture, Japan. 

2 LSU: partial 28S nrRNA gene; ITS: internal transcribed spacer regions 1 & 2 including 5.8S nrRNA gene; EF-1α: partial translation elongation factor 1-alpha gene; ACT: partial actin gene. 

3 Sequence for this locus obtained from: http://genome.jgi-psf.org/Mycfi1/Mycfi1.home.html 


51

Crous et al. 

1 000 generations, resulting in 8 001 saved trees in each of the 

two tree files. Burn-in was set at 2 000 000 generations after which 

the likelihood values were stationary. For parsimony analysis of the 

combined ITS, ACT and EF-1α alignment, alignment gaps were 

treated as a fifth character state and all characters were unordered 

and of equal weight. Maximum parsimony analysis was performed 

in PAUP using the heuristic search option with 100 random taxon 

additions and tree bisection and reconnection (TBR) as the branchswapping 

algorithm. Branches of zero length were collapsed 

and all multiple, equally most parsimonious trees were saved. 

The robustness of the trees was evaluated by 1 000 bootstrap 

replicates (Hillis & Bull 1993). Tree length (TL), consistency index 

(CI), retention index (RI) and rescaled consistency index (RC) were 

calculated and the resulting trees were printed with Geneious v. 

5.5.4 (Drummond et al. 2011). Sequences derived in this study 

were deposited in GenBank (Table 1), the alignments in TreeBASE 

(www.treebase.org/treebase/index.html), and taxonomic novelties 

in MycoBank (www.MycoBank.org; Crous et al. 2004b). 

Taxonomy 

All taxonomic descriptions were based on structures on 

herbarium material. Diseased leaf tissue was viewed under a 

Nikon® SMZ1500 stereoscopic zoom microscope and relevant 

morphological structures were lifted from lesions with a sterile 

dissecting needle and mounted on glass slides in clear lactic acid. 

For measurements, 30–50 replicates of all relevant morphological 

features were made at ×1 000 magnification using a Carl Zeiss® 

Axioskop 2 plus light microscope. High-resolution photographic 

images of diseased material, leaf lesions and microscopic fungal 

structures were captured with a Nikon® digital sight DS-fi1 high 

definition colour camera mounted on the light microscope or a 

Nikon® digital sight DS-5M camera mounted on a stereoscopic 

zoom microscope. Images of morphological structures were 

captured, and measurements taken, using the Nikon® software 

NIS-Elements v. 2.34 while Adobe Photoshop was used for the final 

editing of acquired images and photographic preparations. Novel 

Pseudocercospora taxa were plated onto MEA and incubated at 

24 °C for 2–4 wk in the dark in duplicate. The mycological colour 

charts of Rayner (1970) were used to define colours of the fungal 

colonies. 

RESULTS 

DNA sequencing and phylogenetic analyses 

Large Subunit (LSU) phylogeny: The final aligned LSU dataset 

contained 316 ingroup taxa with a total of 1305 characters and 

Saccharomyces cerevisiae (GenBank Accession: Z73326) served 

as the outgroup taxon. From this alignment 827 characters 

were used for the Bayesian analysis; the consensus trees and 

posterior probabilities were calculated (Fig. 4) from the 12 002 

trees left after discarding those used for burn-in. The resulting 

LSU phylogeny resolved several clades (Clades 1–14) grouping 

species of Pseudocercospora and allied genera (Fig. 4). Clade 1 

(Posterior Probability (PP) value of 1.0) including Cyphellophora 

and Strelitziana represented by one of the two basal lineages. 

Thedgonia ligustrina (100 %) represented the second basal clade 

(PP = 1.0). In the Pleosporales, Clade 3 included Xenostigmina 

zilleri (PP = 1.0) and Clade 4 Pseudocercospora cantuariensis (PP 

= 1.0), the latter being described below as Phaeomycocentrospora 

cantuariensis. Clade 5 contained Cladosporium species belonging 

to the teleomorph genus Davidiella (PP = 1.0). Clade 6 (PP = 1.0) 

represented species belonging to Teratosphaeria and including 

the recently established genus Microcyclospora. Clade 7 (PP = 

1.0) accommodated species of Dissoconium. Clade 8 (PP = 1.0) 

including species representing Mycosphaerella, Pseudocercospora 

and Zasmidium, as well as the recently established genus 

Microcyclosporella. Clade 9 (PP = 1.0) included Pseudocercospora 

tibouchinigena, Pseudocercospora egenula described below as 

Paracercospora egenula and the Mycosphaerella ellipsoidea 

complex. Clade 10 (PP = 1.0) accommodated species of other 

genera namely Pseudocercosporella, Mycosphaerella ulmi 

(Phleospora), Muiraea, Cercospora and Septoria. Clade 11 

(PP = 1.0) included Mycosphaerella species with Sonderhenia 

anamorphs. Clade 12 (PP = 1.0) is sister to Clade 11 and included 

species representing taxa of Mycosphaerella and their associated 

Pseudocercospora-like anamorphs, appeared to represent a 

novel genus. Other genera in this clade included Scolecostigmina 

and Trochophora. The isolates representing Trochophora are 

accommodated at a basal position in this clade with no PP support. 

The three isolates of Scolecostigmina mangiferae resided in a 

well-supported sub-clade (PP = 1.0) close to isolates regarded 

as part of the Mycosphaerella heimii complex (P. acaciigena, M. 

irregulariramosa, M. colombiensis, P. thailandica, M. heimii, M. 

heimioides, M. konae), described below in Pallidocercospora. 

Clade 13 (PP = 1.0) accommodated Passalora eucalypti. The 

remainder of the phylogeny encompassed Clade 14 (PP = 1.0), 

representing Pseudocercospora s. str., and accommodated the 

majority of Pseudocercospora species from many different hosts. 

The type species of Pseudocercospora, P. vitis was included in 

this clade. Interestingly, P. vitis was basal in this clade with the 

majority of Pseudocercospora species radiating out from the basal 

Pseudocercospora isolates. The LSU phylogeny provided a wellsupported 

sub-clade (PP = 1.0) representing the second half of 

the sensu stricto clade (Clade 14). Several isolates representing 

species from genera morphologically allied to Pseudocercospora 

were also grouped in Clade 14. These included Stigmina platani, 

Cercostigmina protearum var. leucadendri (as Pseudocercospora 

leucadendri, see below), Cercostigmina protearum var. hakeae 

(as Pseudocercospora hakea, see below), Phaeoisariopsis 

griseola f. griseola (as Pseudocercospora griseola f. griseola, 

see Crous et al. 2006) and Pseudophaeoramularia angolensis 

(as Pseudocercospora angolensis, see below), which supports 

previous proposals to include these genera in Pseudocercospora 

s. str. 

Pseudocercospora s. str. phylogeny: A further analysis was 

conducted on Clade 14 (Fig. 4), representing Pseudocercospora 

s. str. For this analysis, DNA sequence data from the ITS, ACT 

and EF-1α gene regions were combined in the parsimony analysis. 

For this dataset, there was a total of 194 taxa, each representing 

1 029 characters. Passalora eucalypti (CBS 111318) served as the 

outgroup taxon for this analysis. From the combined alignment of 

1 029 characters, 414 were constant, 124 were variable and 491 

characters were parsimony uninformative. Only the first 1 000 

equally most parsimonious trees were saved, the first of which is 

shown (Fig. 5) (TL = 4315, CI = 0.312, RI = 0.819, RC = 0.256). 

The phylogeny resulting from the combined sequence data 

was more structured towards the terminal nodes than the LSU 

phylogeny. Similar to the LSU phylogeny, a split was observed 

within Pseudocercospora s. str., with at least two main clades 

being evident. Although present in the strict consensus tree, this 

52


Saccharomyces cerevisiae Z73326 

Cyphellophora eucalypti CBS 124764 

Cyphellophora laciniata FJ358239 

Phialophora europaea FJ358248 

Fonsecaea pedrosoi AF356666 

Capronia peltigerae HQ613813 

Chaetothyriales 

Glyphium elatum AF346420 

Coniosporium perforans FJ358237 

Sarcinomyces petricola FJ358249 

Strelitziana australiensis CPC 13556 

Strelitziana australiensis GQ303326 

Thedgonia ligustrina EU040242 

Thedgonia ligustrina CPC 10019 


Helotiales 


Mycopappus aceris FJ839660 

Pleosporales, 

Xenostigmina zilleri GU253857 

Xenostigmina zilleri FJ839676 

Phaeosphaeriaceae 

Neottiosporina paspali EU754172 

Pleurophoma pleurospora EU754200 

Byssothecium circinans AY016357 

Corynespora olivacea GU301809 

Phoma carteri GQ387594 

Phoma violicola GU238156 

Ascochyta hordei var. hordei EU754134 

Phaeomycocentrospora cantuariensis CPC 10157 

Phaeomycocentrospora cantuariensis CPC 10762 Pleosporales 



Cladosporium cladosporioides EU019262 

Cladosporium herbarum DQ678074 

Cladosporium allicinum EU019263 

Cladosporium allicinum EU019261 

Teratosphaeria molleriana EU167583 

Teratosphaeria dimorpha FJ493215 

Teratosphaeria fibrillosa GU214506 

Teratosphaeria stellenboschiana CPC 13767 

Teratosphaeria alcornii CBS 313.76 

Teratosphaeria nubilosa DQ246228 

Penidiella columbiana EU019274 

Penidiella eucalypti EU882145 

Catenulostroma eucalyptorum JF951174 

Phaeothecoidea eucalypti EU019280 Capnodiales, 

Readeriella nontingens FJ493201 

Readeriella mirabilis EU019291 Teratosphaeriaceae 

Readeriella novaezelandiae DQ246239 

Teratosphaeria flexuosa FJ493216 

Teratosphaeria ohnowa EU019305 

Catenulostroma abietis EU019249 

Catenulostroma germanicum EU019253 

Pseudocercosporella fraxini GU214682 

Microcyclospora tardicrescens GU570552 

“Pseudocercospora” sp. GU214681 

Microcyclospora pomicola GU570551 

Microcyclospora malicola GU570549 

Microcyclospora malicola GU570550 

Dissoconium aciculare EU019266 

Dissoconium dekkeri GU214422 

Dissoconium dekkeri DQ204768 

Dissoconium commune DQ246262 

Dissoconium commune EU019267 

Posterior probability values 

= 1.0 

= 0.95 to 0.99 

= 0.90 to 0.94 

= 0.80 to 0.89 

= 0.70 to 0.79 

Capnodiales, 

Cladosporiaceae 

Capnodiales, 

Dissoconiaceae 

1 

2 

3 

4 

5 

6 

7 

0.5 

Fig. 4. Consensus phylogram (50 % majority rule) of 12 002 trees resulting from a Bayesian analysis of the LSU sequence alignment using MrBayes v. 3.1.2. Bayesian posterior 

probabilities are indicated with colour-coded branches (see legend) and the scale bar represents the expected changes per site. Important clades are indicated in coloured 

blocks and numbered. The tree was rooted to Saccharomyces cerevisiae (GenBank Z73326). 


53

Crous et al. 

Capnodiales, Mycosphaerellaceae 

Zasmidium nabiacense GQ852734 

Mycosphaerella marksii DQ246250 

Pseudocercospora epispermogonia DQ204757 

Mycosphaerella madeirae DQ204756 

Microcyclosporella mali GU570545 

Microcyclosporella mali GU570548 

Mycosphaerella laricina CBS 326.52 

Paracercospora egenula CPC 12537 

Paracercospora egenula MUCC 883 

Paracercospora egenula CBS 485.81 

Passalora brachycarpa GU214664 

“Pseudocercospora” tibouchinigena CBS 116462 

Mycosphaerella ellipsoidea GU214434 

Mycosphaerella africana DQ246257 

Mycosphaerella aurantia DQ246256 

Cercospora sojina CPC 12322 

Pseudocercosporella oxalidis CBS 118758 

Cercospora zebrinae GU214657 

Cercospora capsici GU214654 

Cercospora apii GU214653 

Cercospora rodmanii GQ884186 

Pseudocercosporella arcuata GU214685 

Pseudocercosporella sp. CPC 10864 

Septoria eucalyptorum CPC 11282 

Septoria justiciae CPC 12509 

Cercospora eucommiae CPC 10047 



Septoria cerastii CPC 12343 

Septoria chelidonii CPC 12337 

Septoria crepidis CPC 12539 

Septoria dysentericae CPC 12328 

Septoria erigerontis CPC 12340 

Septoria rubi CPC 12331 

Pseudocercosporella zelkovae GU214482 

Pseudocercosporella capsellae CPC 14773 

Phloeospora ulmi CBS 344.97 

Phloeospora ulmi CBS 613.81 

Pseudocercosporella koreana GU214683 

Septoria quercicola CBS 663.94 

Pseudocercosporella cydoniae CPC 14795 

Miuraea persicae CPC 10069 

Miuraea persicae CPC 10828 

Mycosphaerella swartii DQ923536 

Mycosphaerella walkeri DQ267574 

Trochophora fasciculata FJ839668 

Trochophora simplex MUCC 952 

Trochophora simplex CBS 124744 

Pallidocercospora holualoana GU214440 

Pallidocercospora holualoana JF770467 

Pallidocercospora acaciigena GU214661 

Pallidocercospora acaciigena CBS 120740 

Pallidocercospora crystallina DQ204747 

Pallidocercospora irregulariramosa DQ204754 

“Pseudocercospora” colombiensis DQ204744 

“Pseudocercospora” thailandica CPC 10548 

“Pseudocercospora” thailandica CBS 116367 

Pallidocercospora konae CBS 120748 

Pallidocercospora heimii DQ204751 

Pallidocercospora heimioides DQ204753 

Scolecostigmina mangiferae strain 3 

Scolecostigmina mangiferae CPC 17352 

Scolecostigmina mangiferae CBS 125467 

8 

9 

10 

11 

12 


= 1.0 

= 0.95 to 0.99 

= 0.90 to 0.94 

= 0.80 to 0.89 

= 0.70 to 0.79 

0.5 

Fig. 4. (Continued). 

54



Passalora eucalypti CBS 111318 

Passalora eucalypti DQ246244 

Pseudocercospora vitis CPC 14661 

Pseudocercospora vitis CPC 11595 

Pseudocercospora dianellae CBS 117746 

Pseudocercospora lyoniae MUCC 910 

Pseudocercospora metrosideri CBS 118795 

Pseudocercospora myrticola MUCC 632 

Pseudocercospora pouzolziae CBS 122280 

Pseudocercospora rumohrae CBS 117747 

Pseudocercospora luzardii CPC 2556 

Pseudocercospora nephrolepidis CBS 119121 

Pseudocercospora profusa CPC 10055 

Pseudocercospora proteae CPC 15217 

Pseudocercospora balsaminae CPC 10044 

Pseudocercospora crousii CBS 119487 

Pseudocercospora cymbidiicola CBS 115132 

Pseudocercospora viburnigena CPC 15249 

Pseudocercospora fori CBS 113285 

Pseudocercospora natalensis CBS 111069 

Pseudocercospora gracilis CBS 243.94 

Pseudocercospora robusta CBS 111175 

Pseudocercospora eucalyptorum CBS 110777 

Pseudocercospora eucalyptorum CPC 12406 










Pseudocercospora humulicola CPC 11358 

Pseudocercospora ixorae CBS 118760 

Pseudocercospora natalensis CBS 111071 


Pseudocercospora plecthranthi CPC 11462 

Pseudocercospora crocinus CPC 11668 


Pseudocercospora humulicola CPC 10049 

Pseudocercospora cladosporioides CBS 117482 

Pseudocercospora rhabdothamni CBS 114872 

Pseudocercospora sawadae CBS 115024 



Pseudocercospora angolensis CBS 112933 

Pseudocercospora angolensis CBS 149.53 

Pseudocercospora cf. kaki CPC 10636 

Pseudocercospora palleobrunnea CBS 124771 

Pseudocercospora sambucigena CPC 10292 

Pseudocercospora sambucigena CPC 14397 

Stigmina platani CBS 336.33 

Pseudocercospora pseudostigmina-platani CPC 11726 

Pseudocercospora arecacearum CBS 118406 

Pseudocercospora arecacearum CBS 118792 

Pseudocercospora sp. CBS 113387 

Pseudocercospora coprosmae CBS 114639 

Pseudocercospora griseola f. griseola CBS 194.47 

Pseudocercospora griseola f. griseola CPC 12239 

Pseudocercospora griseola f. griseola CBS 119112 




Pseudocercospora griseola f. griseola CBS 880.72 

13 

14 


= 1.0 

= 0.95 to 0.99 

= 0.90 to 0.94 

= 0.80 to 0.89 

= 0.70 to 0.79 

0.5 


55

Crous et al. 

Ps. macrospora CBS 114696 

Ps. norchiensis CBS 120738 

Ps. norchiensis CBS 114641 

Ps. sordida MUCC 913 


Ps. purpurea CBS 114163 

Ps. nogalesii CBS 115022 

Ps. ocimi-basilici CPC 10283 

Ps. rhododendri-indici CPC 10822 

Ps. punctata CPC 14734 

Ps. leucadendri CPC 1869 

Ps. fori CPC 14880 

Ps. theae CBS 128.30 

Ps. basitruncata CBS 114664 

Ps. subulata CBS 118489 

Ps. libertiae CBS 114643 

Ps. melicyti CBS 115023 

Ps. dendrobii MUCC 596 

Ps. jussiaeae CPC 14625 

Ps. lythri CPC 14588 

Ps. lythri MUCC 865 

Ps. araliae CPC 10154 

Ps. araliae MUCC 873 

Pseudocercospora sp. CPC 15116 

Ps. abelmoschi CPC 14478 


Ps. assamensis CBS 122467 

Ps. basiramifera CBS 111072 

Ps. callicarpae MUCC 888 

Ps. catappae MUCC 809 

Ps. cercidicola MUCC 896 

Ps. cercidis-chinensis CPC 14481 


Ps. chengtuensis MUCC 828 

Ps. chionanthi-retusi CPC 14683 

Ps. chrysanthemicola CPC 10633 

Ps. contraria CPC 14714 

Ps. coriariae MUCC 840 

Ps. cornicola MUCC 909 

Ps. corylopsidis MUCC 908 

Ps. cotoneastri MUCC 876 

Ps. cf. cruenta CBS 117232 

Ps. catalpigena MUCC 743 

Ps. davidiicola MUCC 296 

Ps. destructiva MUCC 870 

Ps. elaeocarpi MUCC 925 

Ps. eupatoriella CBS 113372 

Ps. exosporioides MUCC 893 

Ps. fraxinites CPC 10743 

Ps. fukuokaensis MUCC 887 

Ps. fuligena MUCC 533 

Ps. fukuokaensis CPC 14689 

Ps. glauca CPC 10062 

Ps. guianensis MUCC 855 

Ps. guianensis MUCC 879 

Ps. corylopsidis MUCC 874 

Ps. humuli MUCC 742 

Ps. kaki MUCC 900 

Ps. latens MUCC 763 

Ps. lilacis CPC 12767 

Ps. indonesiana CBS 122473 

Ps. lonicericola MUCC 889 

Ps. lythracearum CPC 10707 

Ps. lythracearum MUCC 890 


= 1.0 

= 0.95 to 0.99 

= 0.90 to 0.94 

= 0.80 to 0.89 

= 0.70 to 0.79 

Ps. = Pseudocercospora 

14 

cont. 

Ps. mali MUCC 886 

Ps. nandinae CBS 117745 

Ps. oenotherae CPC 10630 

Ps. pancratii CBS 137.94 

Ps. paraguayensis CBS 111317 

Ps. pini-densiflorae MUCC 534 

Ps. puderi MUCC 906 

Ps. pyracanthae MUCC 892 

Ps. rhapisicola CBS 282.66 

Ps. securinegae CPC 10793 



Ps. basiramifera CBS 114757 

Ps. ranjita CPC 11141 

Ps. dovyalidis CPC 13771 

Ps. haiweiensis CPC 14084 

Ps. stahlii CBS 117549 

Ps. stephanandrae MUCC 914 

Ps. timorensis MUCC 819 

Ps. variicolor MUCC 746 

Ps. viticicola MUCC 777 

Ps. weigelae MUCC 899 

Ps. zelkovae CPC 14484 

Ps. zelkovae CPC 14717 

Ps. acericola CBS 122279 

Ps. cydoniae CPC 10678 

Ps. eustomatis CBS 110822 

Ps. marginalis CPC 12497 

Ps. flavomarginata CBS 118841 

Ps. flavomarginata CBS 124990 

Ps. flavomarginata CPC 14142 

Ps. fraxinites MUCC 891 

Ps. ravenalicola CBS 122468 

Ps. subsessilis CBS 136.94 

Ps. crispans CPC 14883 

Ps. rubi MUCC 875 

Ps. rhamnellae CPC 12500 

Ps. zelkovae MUCC 872 

Ps. atromarginalis CPC 11372 

Ps. paederiae CPC 10007 

Ps. subtorulosa CBS 117230 

Ps. longispora CBS 122470 

Ps. xanthoxyli CPC 10065 

Ps. ampelopsis CPC 11680 

Ps. paraguayensis CBS 111286 

Ps. circumscissa CPC 14511 

Ps. atromarginalis CBS 114640 

Ps. cordiana CBS 114685 

Ps. cruenta CPC 10846 

Ps. pallida CPC 10776 

Ps. kiggelariae CPC 11853 

Ps. xanthocercidis CPC 11665 

Ps. chengtuensis CPC 10696 

Ps. indonesiana CBS 122474 

Ps. fuligena CPC 12296 

Ps. oenotherae CPC 10290 

Ps. tereticornis CPC 13299 

Ps. tereticornis CBS 124996 


Ps. rhoina CPC 11464 

Ps. fijiensis MUCC 792 

Ps. fijiensis CBS 120258 

Ps. hakeae CBS 112226 

Ps. musae CBS 116634 

Ps. dodonaeae CBS 114647 

0.5 

14 

cont. 

56


Passalora eucalypti CBS 111318 

100 Ps. vitis CPC 11595 Vitis, Vitaceae 

Ps. vitis CPC 14661 Vitis, Vitaceae 

Ps. luzardii CPC 2556 Hancornia, Apocynaceae 

100 66 Ps. purpurea CBS 114163 Persea, Lauraceae 

Ps. sordida MUCC 913 Campsis, Bignoniaceae 

95 

Ps. macrospora CBS 114696 Bertholletia, Lecythidaceae 

Ps. nogalesii CBS 115022 Chamaecytisus, Fabaceae 

57 

Pseudocercospora sp. CBS 111373 Eucalyptus, Myrtaceae 

100 

85 Ps. norchiensis CBS 120738 Eucalyptus, Myrtaceae 

100 Ps. norchiensis CBS 114641 Rubus, Rosaceae 

100 

Pseudocercospora sp. CBS 113387 Lantana, Verbenaceae 

100 Ps. sambucigena CPC 10292 Sambucus, Caprifoliaceae 

96 

Ps. sambucigena CPC 14397 Sambucus, Caprifoliaceae 

Ps. pseudostigmina-platani CPC 11726 Platanus, Platanaceae 

67 

Ps. griseola f. griseola CPC 10480 Phaseolus, Fabaceae 

100 Ps. griseola f. griseola CPC 10779 Phaseolus, Fabaceae 


83 

Ps. griseola f. griseola CBS 194.47 Phaseolus, Fabaceae 

Ps. griseola f. griseola CBS 880.72 Phaseolus, Fabaceae 

Ps. griseola f. griseola CBS 119112 Phaseolus, Fabaceae 


Ps. basitruncata CBS 114664 Eucalyptus, Myrtaceae 

100 Ps. angolensis CBS 149.53 Citrus, Rutaceae 

85 Ps. angolensis CPC 4118 Citrus, Rutaceae 

10 changes 

99 

Ps. ocimi-basilici CPC 10283 Ocimum, Lamiaceae 

84 

Ps. punctata CPC 14734 Syzygium, Myrtaceae 

Ps. udagawana CPC 10799 Hovenia, Rhamnaceae 

Ps. cladosporioides CBS 117482 Olea, Oleaceae 

81 

Pseudocercospora sp. CPC 10058 Potentilla, Rosaceae 

Ps. jussiaeae CPC 14625 Ludwigia, Onagraceae 

Ps. dendrobii MUCC 596 Dendrobium, Orchidaceae 

88 

62 Ps. araliae CPC 10154 Aralia, Araliaceae 

76 98 Ps. araliae MUCC 873 Aralia, Araliaceae 

100 Ps. lythri CPC 14588 Lythrum, Lythraceae 

68 

94 

99 

Ps. lythri MUCC 865 Lythrum, Lythraceae 

Ps. nephrolepidis CBS 119121 Nephrolepis, Oleandraceae 

100 

Ps. pouzolziae CBS 122280 Gonostegia, Urticaceae 

Ps. rumohrae CBS 117747 Marattia, Marattiaceae 

Ps. plecthranthi CPC 11462 Plectranthus, Lamiaceae 

99 Ps. balsaminae CPC 10044 Impatiens, Balsaminaceae 

81 Ps. dianellae CBS 117746 Dianella, Liliaceae 

Ps. rhabdothamni CBS 114872 Rhabdothamnus, Gesneriaceae 

95 

Ps. humulicola CPC 10049 Humulus, Cannabaceae 

Ps. humulicola CPC 11358 Humulus, Cannabaceae 

Ps. crocea CPC 11668 Pilea, Urticaceae 

95 57 

96 

Ps. profusa CPC 10055 Acalypha, Euphorbiaceae 

Ps. profusa CPC 10042 Acalypha, Euphorbiaceae 

Ps. robusta CBS 111175 Eucalyptus, Myrtaceae 

100 

100 

Ps. gracilis CBS 243.94 Eucalyptus, Myrtaceae 

Ps. eucalyptorum CPC 12957 Eucalyptus, Myrtaceae 



100 Ps. eucalyptorum CPC 13926 Eucalyptus, Myrtaceae 

64 




Ps. eucalyptorum CBS 116359 Eucalyptus, Myrtaceae 

Ps. eucalyptorum CBS 110777 Eucalyptus, Myrtaceae 


87 






Fig. 5. The first of 1 000 equally most parsimonious trees obtained from a heuristic search with 100 random taxon additions of the combined ITS, ACT and EF-1α sequence 

alignment using PAUP v. 4.0b10. The scale bar shows 10 changes, and bootstrap support values from 1 000 replicates are shown at the nodes. Thickened lines indicate those 

branches present in the strict consensus tree and the tree was rooted to Passalora eucalypti strain CBS 111318 (GenBank GU269845, GU320548 and GU384558, respectively). 


57

Crous et al. 

Ps. rhododendri-indici CPC 10822 Rhododendron, Ericaceae 

Ps. coprosmae CBS 114639 Coprosma, Rubiaceae 

83 100 

Ps. arecacearum CBS 118406 Rhopalostylis, Arecaceae 

Ps. arecacearum CBS 118792 Howea, Arecaceae 

Ps. cf. kaki CPC 10636 Diospyros, Ebenaceae 

55 

Ps. leucadendri CPC 1869 Leucadendron, Proteaceae 

Ps. myrticola MUCC 632 Myrtus, Myrtaceae 

83 

Ps. proteae CPC 15217 Protea, Proteaceae 

Ps. ixorae CBS 118760 Ixora, Rubiaceae 

Ps. cymbidiicola CBS 115132 Cymbidium, Orchidaceae 

76 Ps. lyoniae MUCC 910 Lyonia, Ericaceae 

Ps. theae CBS 128.30 Camelia, Theaceae 

Ps. fori CBS 113285 Eucalyptus, Myrtaceae 

Ps. fori CPC 14880 Eucalyptus, Myrtaceae 

89 

Ps. natalensis CBS 111069 Eucalyptus, Myrtaceae 

59 

Ps. natalensis CBS 111071 Eucalyptus, Myrtaceae 

Ps. libertiae CBS 114643 Libertia, Iridaceae 

Ps. metrosideri CBS 118795 Metrosideros, Myrtaceae 

Ps. melicyti CBS 115023 Melicytus, Violaceae 

Ps. palleobrunnea CPC 13387 Syzygium, Myrtaceae 

Ps. subulata CPC 10849 Eucalyptus, Myrtaceae 

76 

62 Ps. crousii CBS 119487 Eucalyptus, Myrtaceae 

Ps. sawadae CBS 115024 Psidium, Myrtaceae 

Ps. viburnigena CPC 15249 Viburnum, Caprifoliaceae 

Ps. rhoina CPC 11464 Rhus, Anacardiaceae 

100 Ps. xanthoxyli CPC 10065 Xanthoxylum, Rutaceae 

Ps. snelliana CPC 11654 Morus, Moraceae 

Ps. securinegae CPC 10793 Flueggea, Euphorbiaceae 

100 Ps. fijiensis MUCC 792 Musa, Musaceae 

Ps. fijiensis CBS 120258 Musa, Musaceae 

Ps. acericola CBS 122279 Acer, Aceraceae 

81 

Ps. subtorulosa CBS 117230 Melicope, Rutaceae 

60 Pseudocercospora sp. CPC 10645 

100 Ps. eustomatis CBS 110822 Eustroma, Gentianaceae 

94 Ps. oenotherae CPC 10630 Oenothera, Onagraceae 

65 Ps. oenotherae CPC 10290 Oenothera, Onagraceae 

Ps. cruenta CPC 10846 Vigna, Fabaceae 

100 Ps. atromarginalis CBS 114640 Solanum, Solanaceae 

Ps. atromarginalis CPC 11372 Solanum, Solanaceae 

100 Ps. chengtuensis CPC 10696 Lycium, Solanaceae 

Ps. fuligena CPC 12296 Lycopersicon, Solanaceae 

86 

Ps. fuligena MUCC 533 Solanum, Solanaceae 

91 Ps. nandinae CBS 117745 Nandina, Berberidaceae 

Ps. cordiana CPC 2552 Cordia, Boraginaceae 

82 

Pseudocercospora sp. CBS 110998 Eucalyptus, Myrtaceae 

51 Ps. paraguayensis CBS 111286 Eucalyptus, Myrtaceae 

Ps. paraguayensis CPC 1458 Eucalyptus, Myrtaceae 

89 

Ps. pini-densiflorae MUCC 534 Pinus, Pinaceae 

68 

Ps. pyracanthigena CPC 10808 Pyracantha, Rosaceae 

Ps. fukuokaensis CPC 14689 Styrax, Styracaceae 

94 

Ps. fukuokaensis MUCC 887 Styrax, Styracaceae 

64 

Ps. mali MUCC 886 Malus, Rosaceae 

Ps. cercidis-chinensis CPC 14481 Cercis, Fabaceae 

Ps. glauca CPC 10062 Albizzia, Fabaceae 

Ps. chrysanthemicola CPC 10633 Chrysanthemum, Asteraceae 

100 Ps. marginalis CPC 12497 Fraxinus, Oleaceae 

100 Ps. chionanthi-retusi CPC 14683 Chionanthus, Oleaceae 

10 changes 


split was not well-supported in the phylogeny. Deeper nodes of 

the backbone were poorly supported. There were high levels of 

support for several of the smaller sub-clades in this tree, which are 

discussed in the Taxonomy section below. 

Taxonomy 

Isolates representing 146 species of Pseudocercospora were 

subjected to DNA analysis and morphological comparison. 

Phylogenetic analyses based on the LSU gene resolved a total of 

14 clades in the Pseudocercospora complex. 

58


69 

69 

10 changes 

Ps. ampelopsis CPC 11680 Ampelopsis, Vitaceae 

Ps. hakeae CBS 112226 Grevillea, Proteaceae 

100 Ps. longispora CBS 122470 Musa, Musaceae 

Ps. musae CBS 116634 Musa, Musaceae 

99 Ps. assamensis CBS 122467 Musa, Musaceae 

Ps. indonesiana CBS 122474 Musa, Musaceae 

99 Ps. indonesiana CBS 122473 Musa, Musaceae 

Ps. cydoniae CPC 10678 Chaenomeles, Rosaceae 

Ps. cornicola MUCC 909 Cornus, Cornaceae 

99 

Ps. eupatoriella CBS 113372 Chromolaena, Asteraceae 

95 

66 Ps. lilacis CPC 12767 Ligustrum, Oleaceae 

97 Pseudocercospora sp. CPC 15116 Malus, Rosaceae 

100 Ps. crispans CPC 14883 Eucalyptus, Myrtaceae 

Pseudocercospora sp. CBS 110993 Populus, Salicaceae 

Ps. cf. cruenta CBS 117232 Phaseolus, Fabaceae 

55 

Ps. xanthocercidis CPC 11665 Xanthocercis, Fabaceae 

Ps. catappae MUCC 809 Terminalia, Combretaceae 

82 

Ps. destructiva MUCC 870 Euonymus, Celastraceae 

95 Ps. fraxinites CPC 10743 Fontanesia, Oleaceae 

100 Ps. fraxinites MUCC 891 Fraxinus, Oleaceae 

Ps. ravenalicola CBS 122468 Ravenala, Strelitziaceae 

Ps. dovyalidis CPC 13771 Dovyalis, Flacourtiaceae 

87 

Ps. stahlii CBS 117549 Passiflora, Passifloraceae 

Ps. kiggelariae CPC 11853 Kiggelaria, Flacourtiaceae 

Ps. basiramifera CBS 111072 Eucalyptus, Myrtaceae 

100 

Ps. basiramifera CBS 114757 Eucalyptus, Myrtaceae 

Ps. humuli MUCC 742 Humulus, Cannabaceae 

84 

Ps. puderi MUCC 906 Rosa, Rosaceae 

Ps. cercidicola MUCC 896 Cercis, Fabaceae 

Ps. abelmoschi CPC 14478 Hibiscus, Malvaceae 

60 

Ps. coriariae MUCC 840 Coriaria, Coriariaceae 

Ps. callicarpae MUCC 888 Callicarpa, Verbenaceae 

Ps. davidiicola MUCC 296 Davidia, Nyssaceae 

Ps. dodonaeae CBS 114647 Dodonaea, Sapindaceae 

Ps. ranjita CPC 11141 Gmelina, Verbenaceae 

Ps. timorensis MUCC 819 Ipomoea, Convolvulaceae 

100 Ps. tereticornis CBS 124996 Eucalyptus, Myrtaceae 

Ps. tereticornis CPC 13299 Eucalyptus, Myrtaceae 

100 Ps. flavomarginata CPC 13492 Eucalyptus, Myrtaceae 

Ps. flavomarginata CMW 13586 Eucalyptus, Myrtaceae 

Ps. flavomarginata CPC 14142 Eucalyptus, Myrtaceae 

Ps. pancratii CBS 137.94 Hippeastrum, Liliaceae 

100 

Ps. viticicola MUCC 777 Vitex, Verbenaceae 

Pseudocercospora sp. CPC 14711 Pyracantha, Rosaceae 

Ps. exosporioides MUCC 893 Sequoia, Taxodiaceae 

Ps. kaki MUCC 900 Diospyros, Ebenaceae 

Ps. lythracearum CPC 10707 Lagestroemia, Lythraceae 

78 Ps. lythracearum MUCC 890 Lagestroemia, Lythraceae 

Ps. lonicericola MUCC 889 Lonicera, Caprifoliaceae 

99 

Ps. variicolor MUCC 746 Paeonia, Paeoniaceae 

89 Ps. paederiae CPC 10007 Paederia, Rubiaceae 

Ps. subsessilis CBS 136.94 Melia, Meliaceae 

Ps. guianensis MUCC 855 Lantana, Verbenaceae 

100 

Ps. guianensis MUCC 879 Lantana, Verbenaceae 

100 Ps. pyracanthae MUCC 892 Pyracantha, Rosaceae 

61 Ps. weigelae MUCC 899 Weigela, Caprifoliaceae 

Ps. stephanandrae MUCC 914 Stephanandra, Rosaceae 

Ps. prunicola CPC 14511 Prunus, Rosaceae 

63 Ps. corylopsidis MUCC 908 Corylopsis, Hamamelidaceae 

Ps. corylopsidis MUCC 874 Hamamelis, Hamamelidaceae 

Ps. haiweiensis CPC 14084 Eucalyptus, Myrtaceae 

99 Ps. cotoneastri MUCC 876 Cotoneaster, Rosaceae 

Ps. elaeocarpi MUCC 925 Elaeocarpus, Elaeocarpaceae 

89 

Ps. latens MUCC 763 Lespedeza, Fabaceae 

72 

Ps. rhamnellae CPC 12500 Rhamnella, Rhamnaceae 

Ps. catalpigena MUCC 743 Catalpa, Bignoniaceae 

61 

Ps. pallida CPC 10776 Campsis, Bignoniaceae 

68 

Ps. rhapisicola CBS 282.66 Rhapis, Arecaceae 

58 Ps. contraria CPC 14714 Dioscorea, Dioscoreaceae 

Ps. rubi MUCC 875 Rubus, Rosaceae 

Ps. zelkovae MUCC 872 Zelkova, Ulmaceae 

88 

Ps. zelkovae CPC 14484 Zelkova, Ulmaceae 

95 

Ps. zelkovae CPC 14717 Zelkova, Ulmaceae 



59

Crous et al. 

Clade 1 represented Strelitziana (Pseudocercospora-like 

but with a separating cell between conidia and conidiogenous 

cells) and Cyphellophora (Pseudocercospora-like but phialides 

with flaring collarettes, situated directly on hyphae). Thedgonia 

ligustrina (Pseudocercosporella-like, but conidia in chains) 

represented Clade 2. Clade 3 included several isolates of 

Pseudocercospora cantuariensis, which represents a novel genus, 

distinguished from Pseudocercospora based on its broad conidial 

hila and scars, as well hyaline mycelium, and the presence of 

hyphopodia-like structures. Xenostigmina zilleri, characterised 

as being Stigmina-like, but also having sympodial proliferation 

of the conidiogenous cells, clustered in Clade 4, which was 

basal to Cladosporium (Cladosporiaceae; Clade 5). Clade 6 

represented several members of Teratosphaeriaceae, known 

to have a wide range of anamorphs, including Microcyclospora. 

Clade 7 represented species of Dissoconium (Dissoconiaceae), 

distinct due to their dimorphic conidia that are actively discharged. 

Clade 8 remains unresolved, and was represented by disjunct 

elements appearing Zasmidium- and Pseudocercosporalike 

in morphology, including Microcyclosporella. Clade 9 was 

represented by several Mycosphaerella species such as M. laricina 

(anamorph Pseudocercospora sp.), and Paracercospora egenula. 

Paracercospora was separated from Pseudocercospora based on 

a combination of characters, including pale olivaceous conidia, and 

a minute thickening along the rim of its conidial hila and scars. Clade 

10 included a diverse assemblage of genera. Two genera that differ 

mainly based on their conidiomatal structure, Pseudocercosporella 

and Septoria, clustered in this clade. Miuraea, a genus intermediate 

between Cercospora and Pseudocercospora, also resided within 

this clade. Clade 11 was represented by two coelomycetous species 

of Sonderhenia that clustered basal to Clade 12. The latter included 

a new genus with Pseudocercospora-like anamorphs, mostly 

distinguished from Pseudocercospora s. str. by having species 

with smooth, pale brown conidia, and the frequent production 

of red crystals in agar (previously referred to in literature as the 

Mycosphaerella heimii complex). Scolecostigmina (based on S. 

mangiferae), which is characterised by verruculose conidia and 

percurrently proliferating conidiogenous cells, clustered alongside 

to Trochophora, characterised by brown sickle-shaped conidia with 

three thick, dark septa. Passalora eucalypti formed a separate 

lineage in Clade 13 that was adjacent to Pseudocercospora s. str. 

in Clade 14. This clade included the type species, P. vitis that is 

basal in this cluster. Although there was structure within the clade, 

we regard it as representing a single genus, including Stigmina 

platani, the type of Stigmina, Phaeoisariopsis (P. griseola), and 

Pseudophaeoramularia (P. angolensis). Several isolates identified 

from different countries as representing the same species based on 

host, disease symptoms and general morphology, clustered apart 

from one another. These collections were found to represent novel 

cryptic species. 

DISCUSSION 

Several novel taxa were identified in this study on the basis 

of phylogenetic analyses of the various gene regions together 

with morphological examination of the specimens and isolates. 

Recognised clades, as well as novel species and genera, are 

described and discussed below. Where descriptions of known taxa 

are freely available online in MycoBank or journals, they are not 

repeated here, other than their generic circumscriptions. 

Clade 1: Strelitziana and Cyphellophora 

Strelitziana M. Arzanlou & Crous, Fungal Planet No. 8: 2006. 

Conidiophores erect, solitary, arising from aerial and submerged 

mycelium, subcylindrical, straight to geniculate-sinuous, pale 

brown. Conidiogenous cells terminal, integrated, rejuvenating 

percurrently, proliferating apically via several short, conspicuous 

denticles; conidiogenesis holoblastic with rhexolytic conidial 

secession. Conidia solitary, pale brown, smooth, long obclavate, 

multi-euseptate; microcyclic conidiation present in culture. 

Type species: Strelitziana africana M. Arzanlou & Crous, Fungal 

Planet No. 8. 2006. 

Notes: The genus Strelitziana presently accommodates four species 

that are primarily distinguished based on their conidial dimensions. 

These include S. africana, S. australiensis, S. eucalypti and S. mali 

(Arzanlou & Crous 2006, Cheewangkoon et al. 2009, Zhang et al. 

2009, Crous et al. 2010). 

Cyphellophora G.A. de Vries, Mycopathol. Mycol. Appl. 16: 

47. 1962. 

Colonies (on OA) with moderate to rapid growth, velvety to lanose, 

in various shades of grey; reverse black. Fertile hyphae pale brown, 

sometimes with constrictions at the septa. Conidiogenous cells 

phialidic, intercalary, sometimes on short side branches, each with 

a short, lateral or terminal collarette. Conidia sickle-shaped, brown, 

smooth-walled, transversely septate, adhering in small bundles 

(from de Vries 1962). 

Type species: Cyphellophora laciniata G.A. de Vries, Mycopathol. 

Mycol. Appl. 16: 47. 1962. 

Notes: The genus Cyphellophora, which is based on C. laciniata 

(isolated from human skin; De Vries et al. 1986), appears to be 

heterogeneous (Decock et al. 2003, Crous et al. 2007a, 2009a, 

Cheewangkoon et al. 2009) and requires further study. 

Clade 2: Thedgonia 

Thedgonia B. Sutton, Trans. Brit. Mycol. Soc. 61: 426. 1973. 

Foliicolous, phytopathogenic, causing discrete leaf spots. 

Conidiomata fasciculate, punctiform. Mycelium internal, hyphae 

subhyaline, septate, branched, forming substomatal stromata, 

hyaline to pale brown. Conidiophores fasciculate, arising from 

stromata, simple, rarely branched, subcylindrical, straight to 

geniculate-sinuous, continuous to septate, smooth, hyaline to 

pale yellowish green. Conidiogenous cells integrated, terminal, 

60


occasionally conidiophores reduced to conidiogenous cells, 

holoblastic-thalloblastic, sympodial, conidiogenous loci more or 

less planate, unthickened, non-pigmented. Conidia in disarticulating 

chains, rarely in branched chains, subcylindrical to obclavate, with 

one to several transverse eusepta, hyaline or almost so, apex 

rounded to truncate, base truncate, hila flat, unthickened, hyaline 

(Crous et al. 2009a). 

Type species: Thedgonia ligustrina B. Sutton, Trans. Brit. Mycol. 

Soc. 61: 426. 1973. 

Thedgonia ligustrina (Boerema) B. Sutton, Trans. Brit. 

Mycol. Soc. 61: 428. 1973. 

Basionym: Cercospora ligustrina Boerema, Tijdschr. Plantenziekten 

68: 117. 1962. 

≡ Cercoseptoria ligustrina (Boerema) Arx, Genera of Fungi Sporulating in 

Pure Culture, ed. 3: 306, Lehre 1981. 

Specimens examined: Asia, on Ligustrum sp., H. Evans, CPC 4296 = W2072, 

CPC 4297 = W 2073, CPC 4298 = W 1877. Netherlands, Eefde, on Ligustrum 

ovalifolium, 23 Mar. 1959, G.H. Boerema, holotype L, ex-type culture CBS 148.59; 

Bilthoven, on L. ovalifolium, 2003, P.W. Crous, CPC 10530 = CBS 124332, CPC 

10532, 10533. South Korea, Namyangju, on L. ovalifolium, 9 Oct. 2002, leg. H.D. 

Shin, isol. P.W. Crous, CBS H-20204, CPC 10019, 10861–10863; Suwon, on L. 

obtusifolium, 2 Oct. 2007, leg. H.D. Shin, isol. P.W. Crous, CBS H-20207, CPC 

14754–14756. 

Notes: Contrary to the earlier hypothesis that Thedgonia belonged 

to the Mycosphaerellaceae (Kaiser & Crous 1998), Crous et 

al. (2009a) showed that it resides in Helotiales. Consequently, 

Thedgonia-like anamorphs that occur in the Mycosphaerellaceae 

must be accommodated elsewhere. 

Clade 3: Xenostigmina 

Xenostigmina Crous, Mycol. Mem. 21: 154. 1998. 

Foliicolous, phytopathogenic, causing discrete leaf spots. Mycelium 

internal, consisting of hyaline to pale brown, septate, branched, 

smooth hyphae. Conidiomata sporodochial, brown to black. 

Conidiophores densely aggregated, arising from the upper cells 

of a pale brown stroma, finely verruculose, hyaline to pale brown, 

multiseptate, subcylindrical, straight to variously curved, branched. 

Conidiogenous cells terminal and intercalary, hyaline to pale brown, 

finely verruculose, doliiform to subcylindrical, tapering to flat tipped 

loci, mono- to polyblastic, proliferating sympodially and percurrent; 

loci not thickened or conspicuous. Conidia solitary, pale to medium 

brown, with pale brown apical and basal regions, finely verruculose, 

mostly straight, ellipsoidal, apex subobtuse, frequently extending 

into a beak; base truncate at dehiscence, inner part extending later 

to form a short, subobtuse basal appendage; septation muriform; 

basal marginal frill present (Crous et al. 2009a). 

Type species: Xenostigmina zilleri (A. Funk) Crous, Mycol. Mem. 

21: 155. 1998. 

Specimens examined: Canada, British Columbia, 15 km east of Sardis, on living 

leaves of Acer macrophyllum, 22 Oct. 1985, A. Funk & C.E. Dorworth, holotype 

DAVFP 23272; British Columbia, on living leaves of Acer sp., 2002, leg. K.A. Seifert, 

isol. P.W. Crous, CBS 115686 = CPC 4010, CBS 115685 = CPC 4011; Victoria BC, 

48°30’25.63”N, 123°30’46.99”W, 115 m, fallen leaves of A. macrophyllum, 6 Sep. 

2007, leg. B. Callan, isol. P.W. Crous, epitype designated here CBS H-20208, 

cultures ex-epitype CPC 14376 = CBS 124108, CPC 14377, 14378 (Xenostigmina 

zilleri), CPC 14379 = CBS 124109, CPC 14380, 14381 (Mycopappus aceris). 

Notes: Xenostigmina with its Mycopappus synanamorph is distinct 

from Stigmina s. str., which is a synonym of Pseudocercospora 

s. str. (Crous et al. 2006, Braun & Crous 2006, 2007). The genus 

Xenostigmina (Crous 1998) appears related to Seifertia (Seifert et 

al. 2007) in the Dothideomycetes (Crous et al. 2009b). 

Clade 4: Phaeomycocentrospora 

Phaeomycocentrospora Crous, H.D. Shin & U. Braun, gen. 

nov. MycoBank MB564813. 

Etymology: Name reflects the pale brown appearance of conidia 

and the superficial similarity to Mycocentrospora. 


Mycelium internal and external, consisting of hyaline, septate, 

branched, smooth, 3–5 μm diam hyphae; hyphopodium-like 

structures present. Caespituli amphigenous. Conidiophores in 

loose fascicles, arising from a poorly developed stroma, or from 

superficial hyphae emerging from stomata, or erumpent through 

the cuticle; erect on superficial hyphae, olivaceous-brown, straight 

to slightly curved, unbranched, not geniculate, obconically truncate 

at apex; conidiogenous cells integrated, terminal or conidiophores 

reduced to conidiogenous cells, mono- to polyblastic, proliferating 

sympodially, transversely septate; conidiogenous loci broad, more 

or less planate, neither thickened nor darkened. Conidia solitary, 

filiform to cylindrical, straight to moderately curved, subhyaline to 

pale olivaceous, transversely euseptate, usually not constricted 

at septa, tapering somewhat towards an obtuse apex, truncate at 

base; hilum unthickened, not darkened, broad. 

Type species: Phaeomycocentrospora cantuariensis (E.S. Salmon 

& Wormald) Crous, H.D. Shin & U. Braun, comb. nov. 

Notes: Phaeomycocentrospora is similar to Pseudocercospora in 

that its conidia and conidiophores appear to be pigmented and its 

conidiogenous loci are unthickened and not darkened. It is distinct 

from Pseudocercospora in that its mycelium is hyaline, hyphopodialike 

structures are present, and conidia are hyaline with a pale 

brown inner wall layer, giving the impression of pigmented conidia. 

This fungus also has extremely broad conidial loci and scars that 

are untypical of Pseudocercospora. Chupp (1954) commented that 

Cercospora cantuariensis represented an unusual species that should 

be transferred to a genus of its own. Based on its unique phylogenetic 

placement (Fig. 4) and morphology, Phaeomycocentrospora gen. 

nov. is established for this taxon. Deighton (1971, 1972) assigned 

this species to Mycocentrospora, but the type species M. acerina is 

phylogenetically distinct from other genera morphologically similar to 

it and differs in having conidia with filiform appendages and often with 

strongly swollen intercalary cells. 

Phaeomycocentrospora cantuariensis (E.S. Salmon 

& Wormald) Crous, H.D. Shin & U. Braun, comb. nov. 

MycoBank MB564814. Fig. 6. 

Basionym: Cercospora cantuariensis E.S. Salmon & Wormald, J. 

Bot. (London) 61: 134. 1923. 

≡ Centrospora cantuariensis (E.S. Salmon & Wormald) Deighton, Mycol. 

Pap. 124: 8. 1971. 

≡ Mycocentrospora cantuariensis (E.S. Salmon & Wormald) Deighton, 

Taxon 21: 716. 1972. 

≡ Pseudocercospora cantuariensis (E.S. Salmon & Wormald) U. Braun, 

Mycotaxon 48: 281. 1993. 


61

Crous et al. 

Fig. 6. Phaeomycocentrospora cantuariensis (CPC 11691–11693). A. Leaf spots on upper and lower leaf surface. B, C. Sporulation of leaf surface. D–I. Conidiophores and 

conidiogenous cells. J–M. Conidia. Scale bars = 10 μm. 

Leaf spots amphigenous, scattered, often confluent, subcircular 

to irregular, 1–5 mm diam, becoming up to 10 mm diam when 

confluent, greyish to white, centre reddish brown with yellowish 

brown zone on upper surface; greyish brown to grey on lower 

surface. Caespituli amphigenous, but predominantly hypophyllous. 

Mycelium internal and external; internal hyphae hyaline, septate, 

branched, smooth, 3–4 μm diam; external hyphae plagiotropous, 

branched, septate, smooth, hyaline, 3–5 μm diam. Conidiophores 

in loose fascicles, arising from a poorly developed stroma, or from 

superficial hyphae emerging from stomata, or erumpent through 

the cuticle; erect on superficial hyphae, olivaceous-brown, straight 

to slightly curved, unbranched, not geniculate, obconically truncate 

at apex, proliferating sympodially, 0–3-septate, 30–140 × 7–20 μm. 

Conidiogenous cells terminal, unbranched, pale brown, smooth, 

tapering to flat-tipped apical loci, with scars neither thickened nor 

darkened, 4–7 μm diam; at times proliferating percurrently, with 

1–3 percurrent proliferations at apex, 12–45 × 5–8 μm. Conidia 

solitary, filiform to cylindrical, straight to moderately curved, 

subhyaline to pale olivaceous, smooth, 3–15(–21)-septate, usually 

not constricted at septa, tapering somewhat towards obtuse apex, 

truncate at base, or long obconically subtruncate, (100–)140–200(– 

500) × (5–)7–12(–20) μm; hilum unthickened, not darkened, 4–7 

μm diam; conidia appear to have an inner wall layer that is pale 

brown when studied in culture (adapted from Shin & Kim 2001). 

Specimens examined: South Korea, Hoengseong, on Humulus scandens (= H. 

japonicus), 4 Sep. 2005, H.D. Shin, CBS H-20830; Suwon, Acalypha australis, 5 

Nov. 2004, H.D. Shin, cultures CPC 11691–11693; Suwon, H. scandens , 5 Nov. 

2004, H.D. Shin, CBS H-20831, cultures CPC 11694–11696; Hoengseong, on H. 

62


scandens, 11 Oct. 2004, H.D. Shin, CBS H-20832, cultures CPC 11646, 11647; 

Wonju, on H. scandens, 18 Oct. 2002, H.D. Shin, CBS H-20833, cultures 10157, 

10158; Namyangju, on Luffa aegyptica (= L. cylindrica), 22 Oct. 2003, H.D. Shin, 

CBS H-20834, cultures CPC 10762–10766. 

Clade 5: Cladosporium (Cladosporiaceae) 

Cladosporium Link, Ges. Naturf. Freunde Berlin Mag. 

Neuesten Entdeck. Gesammten Naturk. 7: 37. 1816. 

Teleomorph: Davidiella Crous & U. Braun, Mycol. Progr. 2: 8. 2003. 

Saprobic or phytopathogenic. Ascomata pseudothecial, black 

to red-brown, globose, inconspicuous and immersed beneath 

stomata to superficial, situated on a reduced stroma, with 1(–3) 

short, periphysate ostiolar necks; periphysoids frequently growing 

down into cavity; wall consisting of 3–6 layers of textura angularis. 

Asci fasciculate, short-stalked or not, bitunicate, subsessile, 

obovoid to broadly ellipsoid or subcylindrical, straight to slightly 

curved, 8-spored. Pseudoparaphyses frequently present in mature 

ascomata, hyaline, septate, subcylindrical. Ascospores bi- to 

multiseriate, hyaline, obovoid to ellipsoid-fusiform, with irregular 

luminar inclusions, mostly thick-walled, straight to slightly curved; 

frequently becoming brown and verruculose in asci; at times covered 

in mucoid sheath (from Schubert et al. 2007). Mycelium superficial, 

loosely branched, septate, sometimes constricted at septa, hyaline, 

subhyaline to pale brown, smooth or almost so to verruculose or 

irregularly rough-walled, sometimes appearing irregular in outline due 

to small swellings and constrictions, walls unthickened to somewhat 

thickened. Conidiophores both macro- and micronematous, arising 

laterally from plagiotropous hyphae or terminally from ascending 

hyphae. Macronematous conidiophores erect, straight to flexuous, 

somewhat geniculate-sinuous, nodulose or not, unbranched or 

occasionally branched, pluriseptate, pale to medium brown, older 

ones almost dark brown, walls thickened, sometimes even twolayered. 

Conidiogenous cells integrated, terminal or intercalary, 

mono- to usually polyblastic, nodulose to nodose or not, proliferation 

sympodial, with several conidiogenous loci, mostly situated on 

small lateral shoulders, more or less protuberant, characteristically 

coronate (SEM), i.e. with a convex central dome surrounded 

by a low to distinctly raised rim, appearing to be thickened and 

somewhat darkened-refractive. Micronematous conidiophores 

hardly distinguishable from hyphae, sometimes only as short lateral 

outgrowth with a single apical scar, short, conical to almost filiform 

or narrowly cylindrical, pluriseptate, usually short, subhyaline to pale 

brown, almost smooth to minutely verruculose or irregularly roughwalled, 

0–3-septate. Conidiogenous cells integrated, terminal or 

conidiophores reduced to conidiogenous cells, narrowly cylindrical or 

filiform, with a single or two loci. Conidia solitary (in Heterosporiumlike 

species) to usually catenate, in unbranched or loosely branched 

chains, straight to slightly curved; small terminal conidia without distal 

hilum, obovoid to ellipsoid to subcylindrical, aseptate, subhyaline to 

pale brown; intercalary conidia with a single or sometimes up to 

three distal hila, limoniform, ellipsoid to subcylindrical, 0–1-septate; 

secondary ramoconidia with up to four distal hila, ellipsoid to 

cylindrical-oblong, 0–1(–2)-septate, pale greyish brown or brown 

to medium brown, smooth to minutely verruculose to verrucose, 

walls slightly to distinctly thickened, apex obtuse or slightly truncate, 

towards the base sometimes distinctly attenuated with hila situated 

on short stalk-like prolongations, hila slightly to distinctly protuberant, 

coronate structure as in conidiogenous loci, somewhat thickened and 

darkened-refractive; microcyclic conidiogenesis occurring; primary 

ramoconidia similar to secondary ramoconidia, except base truncate, 

uniform with conidiogenous cell, and more subcylindrical in shape 

(adapted from Schubert et al. 2007). 

Type species: Cladosporium herbarum (Pers. : Fr.) Link, Ges. 

Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten 

Naturk. 7: 37. 1816. 

Notes: Cladosporium is well-defined by having Davidiella 

teleomorphs and conidiophores that give rise to conidial chains with 

unique coronate scars (David 1997, Braun et al. 2003a, Schubert et 

al. 2007, Bensch et al. 2010, 2012), which easily distinguish it from 

a range of other morphologically similar genera (Crous et al. 2007a, 

b; Braun & Crous, in Seifert et al. 2011). 

Clade 6: Teratosphaeriaceae 

Teratosphaeria Syd. & P. Syd., Ann. Mycol. 10: 39. 1912. 

Phytopathogenic, commonly associated with leaf spots, but also 

on fruit, or causing cankers on stems. Ascomata pseudothecial, 

superficial to immersed, frequently situated in a stroma of brown 

pseudoparenchymatal cells, globose, unilocular, papillate, 

ostiolate, canal periphysate, with periphysoids frequently present; 

wall consisting of several layers of brown textura angularis; inner 

layer of flattened, hyaline cells. Pseudoparaphyses frequently 

present, subcylindrical, branched, septate, anastomosing. Asci 

fasciculate, 8-spored, bitunicate, frequently with multi-layered 

endotunica. Ascospores ellipsoid-fusoid to obovoid, 1-septate, 

hyaline, but becoming pale brown and verruculose, frequently 

covered in mucoid sheath (from Crous et al. 2007a). 

Type species: Teratosphaeria fibrillosa Syd. & P. Syd., Ann. Mycol. 

10: 40. 1912. 

Notes: Teratosphaeria accommodates a group of plant pathogenic 

fungi that can cause serious leaf spot, blotch and canker diseases 

of a range of hosts (Crous 2009, Crous et al. 2007a, 2009b, Hunter 

et al. 2009, 2011). The Teratosphaeriaceae remains to be clearly 

resolved, and several different genera are presently recognised in 

the family. Some are plant-associated such as Batcheloromyces, 

Baudoinea, Capnobotryella, Catenulostroma, Davisoniella, 

Devriesia, Hortea, Penidiella, Phaeothecoidea, Pseudotaeniolina, 

Readeriella, Staninwardia, and Stenella s. str. (Crous et al. 2007a, 

2009a, 2011b), and others including Cystocoleus, Racodium, 

Friedmanniomyces, Elasticomyces, Recurvomyces (Selbmann et 

al. 2008) and Xanthoriicola (Ruibal et al. 2011) are lichenicolous or 

rock inhabiting. 

Microcyclospora Jana Frank, Schroers & Crous, Persoonia 

24: 99. 2010. 

Epiphytic and endophytic, occurring on leaves and fruit. Mycelium 

consisting of branched, septate, pale brown, smooth, 2–3 μm wide 

hyphae. Conidiophores reduced to conidiogenous cells, integrated 

in hyphae, giving rise to peg-like lateral protuberances, 1 µm 

wide, 1–2 µm tall, mono- to polyblastic. Conidia scolecosporous, 

cylindrical, straight to variously curved, flexuous, apex obtuse, 

base truncate, 1–multi-septate, somewhat constricted at septa, 

smooth, pale brown, guttulate, aggregated in mucoid masses; hila 

not thickened or darkened; microcyclic conidiation observed in 

culture. 


63

Crous et al. 

Type species: Microcyclospora pomicola Jana Frank, B. Oertel, 

Schroers & Crous, Persoonia 24: 100. 2010. 

Notes: Microcyclospora was recently introduced in 

Teratosphaeriaceae for three taxa associated with sooty blotch of 

apple (Frank et al. 2010). The species described here resembles 

others presently known in Microcyclospora by having pigmented 

structures and undergoing microcyclic condiation. Other than 

having distinct conidial dimensions, it differs from other genera in 

that its conidiogenous cells are annellidic (not mono- to polyblastic), 

and its conidia are darker brown and verruculose to warty, not pale 

brown and smooth. 

Microcyclospora quercina Crous & Verkley, sp. nov. 

MycoBank MB564815. Figs 7, 8. 

Etymology: Name reflects its host, Quercus. 

Foliicolous, endophytic. Mycelium consisting of branched, septate, 

brown, 1.5–3 μm diam hyphae, guttulate, smooth to verruculose 

or warty, with or without mucoid sheath. Conidiophores reduced to 

conidiogenous cells. Conidiogenous cells lateral on hyphae, brown, 

solitary, not aggregated, 1.5–2 μm diam, with 1–4 percurrent 

proliferations and flaring collarettes. Conidia solitary, subcylindrical 

(rarely obclavate), gently curved, apex obtuse (rarely subobtuse), 

base truncate or long obconically truncate, with slight basal taper 

to hilum that is 2 μm diam, unthickened, nor darkened, frequently 

with small marginal frill, brown, guttulate to granular, smooth, 

appearing warty or roughened due to external mucoid layer which 

is sometimes present, transversely (1–)3–4(–11)-euseptate, 

becoming constricted at septa with age, (12–)30–45(–70) × (2–) 

2.5–3 μm; microcyclic conidiation commonly observed. 

Culture characteristics: Colonies after 2 wk in the dark up to 15 mm 

diam, with sparse aerial mycelium, folded surface and uneven to 

somewhat feathery, lobate margins, exuding copious amounts of 

slime on PDA, but less so on MEA and OA; colonies olivaceousblack 

on all media. 

Specimen examined: Netherlands, endophytic in leaves of Quercus robur, Sep. 

2003, G.J.M. Verkley, holotype CBS H-20835, culture ex-type CPC 10712 = CBS 

130827. 

Clade 7: Dissoconium (Dissoconiaceae) 

Dissoconium de Hoog, Oorschot & Hijwegen, Proc. K. Ned. 

Akad. Wet., Ser. C, Biol. Med. Sci. 86(2): 198. 1983. 

Hyperparasitic, but also reported to be phytopathogenic. Ascomata 

pseudothecial, immersed, globose, unilocular, papillate, ostiolate, 

canal periphysate; wall consisting of 3–4 layers of brown textura 

angularis; inner layer of flattened, hyaline cells. Pseudoparaphyses 

absent. Asci fasciculate, 8-spored, bitunicate. Ascospores ellipsoidfusoid, 

1-septate, hyaline, with or without mucoid sheath. Mycelium 

internal and external, consisting of branched, septate, smooth, 

hyaline to pale brown hyphae. Conidiophores separate, arising 

from hyphae, subcylindrical, subulate or lageniform to cylindrical, 

tapering to a bluntly rounded or truncate apex, straight to once 

geniculate, smooth, medium brown, 0–multi-septate; conidiogenous 

cells polyblastic, with terminal and lateral conidiogenous loci, visible 

as slightly thickened, darkened scars on a rachis. Conidia solitary, 

pale olivaceous-brown, smooth, ellipsoid to obclavate or globose, 

Fig. 7. Microcyclospora quercina (CPC 10712). Line drawing showing conidiogenous 

cells and conidia formed in culture. Scale bar = 10 μm. 

0–1-septate; hila somewhat darkened. Secondary conidia present 

or absent; developing adjacent to primary conidia, pale olivaceous 

to subhyaline, aseptate, pyriform; conidium discharge active or 

passive (from Crous et al. 2009b). 

Type species: Dissoconium aciculare de Hoog, Oorschot & 

Hijwegen, Proc. K. Ned. Akad. Wet., Ser. C, Biol. Med. Sci. 86(2): 

198. 1983. 

Notes: Dissoconium has Mycosphaerella-like teleomorphs (Crous 

1998, Crous et al. 2004c) and was recently shown to represent a 

distinct family, Dissoconiaceae (Crous et al. 2009b). Species are 

different from other taxa in Capnodiales in that they form primary 

and secondary conidia that are actively discharged and anastomose 

on the agar surface shortly after germination (De Hoog et al. 1991). 

64


Fig. 8. Microcyclospora quercina (CPC 10712). A, B. Colony on oatmeal and potato-dextrose agar, respectively. C–E. Conidiogenous cells giving rise to conidia (arrows). F. 

Microcyclic conidiation. G, H. Conidia. Scale bars = 10 μm. 

Clade 8: Microcyclosporella and Zasmidium-like 

Microcyclosporella Jana Frank, Schroers & Crous, 

Persoonia 24: 101. 2010. 

Epiphytic on leaves and fruit. Mycelium consisting of pale brown, 

smooth to finely verruculose, branched, septate, 2–3.5 μm wide 

hyphae, at times covered in a mucoid layer, with integrated, lateral, 

truncate conidiogenous loci. Conidiophores mostly reduced to 

conidiogenous cells. Conidiogenous cells integrated, intercalary 

on hyphae, rarely terminal, cylindrical to doliiform, pale brown, 

but hyaline if occurring in yeast-like sectors of colonies, smooth, 

mono- or polyblastic, proliferating sympodially; loci inconspicuous, 

truncate, unthickened, not darkened, pale brown to hyaline. 

Conidia hyaline, smooth, subcylindrical to narrowly obclavate or 

narrowly fusoid with acutely rounded apex and obconically truncate 

base, guttulate, transversely 0–6-septate; microcyclic conidiation 

common. 

Type species: Microcyclosporella mali Jana Frank, Schroers & 

Crous, Persoonia 24: 101. 2010. 

Notes: Microcyclosporella was treated as part of the 

Pseudocercosporella generic complex (Batzer et al. 2005), but has 

since been shown to be polyphyletic within Mycosphaerellaceae 

(Crous 2009, Crous et al. 2003, 2009b, c, Frank et al. 2010). The 

clade accommodating Microcyclosporella contains many disjunct 

elements that vary in morphology from Microcyclosporella s. str. 

(hyaline structures) to pigmented structures, namely Zasmidiumlike 

(verrucuclose conidia) to Pseudocercospora-like (smooth 

conidia) (see Crous et al. 2009b). We suspect that these groups 

may eventually be recognised as distinct genera, but more taxa 

need to be examined to resolve this issue. 

Clade 9: Paracercospora and Pseudocercosporalike 

Paracercospora Deighton, Mycol. Pap. 144: 47. 1979. 

Foliicolous, phytopathogenic, causing leaf spots. Mycelium 

internal, hyaline to pale olivaceous. Stromata absent to poorly 

developed. Conidiophores fasciculate, smooth, subhyaline to pale 

olivaceous. Conidiogenous cells integrated, terminal, mono- to 

usually polyblastic, proliferating sympodially; conidiogenous loci 

moderately conspicuous, with narrow thickening along the rim. 

Conidia solitary, subcylindrical to obclavate-cylindrical, smooth, 

subhyaline to pale olivaceous, with a narrow thickening along the 

rim of the hilum. 

Type species: Paracercospora egenula (Syd.) Deighton, Mycol. 

Pap. 144: 48. 1979. 

Specimens examined: Japan, Shimane, on leaves of Solanum melongena, 5 Aug. 

1998, T. Mikami, CNS-415, cultures MUCC 883, MAFF 237766. South Korea, 

Hongcheon, on leaves of S. melongena, 26 Oct. 2005, H.D. Shin, CBS H-20836, 

culture CPC 12537. 

Notes: Stewart et al. (1999) conducted the first phylogenetic 

analysis of the Mycosphaerellaceae and concluded that the 

marginal thickening that occurs along the rims of conidial 

scars and hila, originally thought to be the main character to 

distinguish Paracercospora from Pseudocercospora, was not 

taxonomically significant and suggested that Paracercospora 

be reduced to synonymy with Pseudocercospora. The current 

study provides new evidence that Paracercospora is not a 

synonym of Pseudocercospora, but no consistent morphological 

characters that distinguish it from Pseudocercospora s. str. 

have been identified. Conidia of Paracercospora egenula are 

subhyaline to pale olivaceous with minimal marginal thickening 

of the conidiogenous loci (Fig. 9). Conidial scars and hila of Ps. 

fijiensis (Arzanlou et al. 2008) and Ps. basiramifera (Crous 1998) 

are marginally thickened. Both of the latter species, which belong 

to Pseudocercospora s. str., have pale to medium brown conidia. 

At present Paracercospora may be defined by a combination of 

the minimal marginal thickening of the conidiogenous loci and its 

subhyaline conidia. 

The taxonomic placement of Paracercospora is complicated 

by two other taxa that resolve in the clade together with it. These 

are Passalora brachycarpa (pale olivaceous, catenate conidia, and 


65

Crous et al. 

Fig. 9. Paracercospora egenula (CPC 12537). A. Leaf spots on upper and lower leaf surface. B. Close-up of lesion. C–F. Fascicles with conidiogenous cells. G. Conidia. Scale 

bars = 10 μm. 

Fig. 10. Pseudocercospora tibouchinigena (CBS 116462). A. Leaf spots on upper and lower leaf surface. B, C. Close-up of lesions. D–G. Fascicles with conidiogenous cells. 

H. Conidia. Scale bar = 10 μm. 

66


prominent, thickened, darkened scars; also visible when sporulating 

in culture), and Pseudocercospora tibouchinigena described below, 

which has subhyaline conidia, and unthickened hila and scars. This 

indicates that it is neither a species of Pseudocercospora s. str. 

(subhyaline conidia), nor Paracercospora (lacking any form of scar 

thickening). As a temporary solution, the species on Tibouchina 

is described in Pseudocercospora, although taxa in this subclade 

may eventually be shown to represent a distinct genus. 

Pseudocercospora tibouchinigena Crous & U. Braun, sp. 

nov. MycoBank MB564816. Fig. 10. 

Etymology: Name is derived from Tibouchina, the host on which it 

was collected. 

Leaf spots amphigenous, angular to irregular, 1–3 mm diam, up to 10 

mm long, medium brown, with raised, dark brown border. Mycelium 

internal, hyaline, smooth, consisting of septate, branched, smooth, 

1.5–2 μm diam hyphae. Caespituli fasciculate, predominantly 

hypophyllous, hyaline to pale olivaceous on leaves, up to 60 μm 

wide and 40 μm high. Conidiophores aggregated in dense fascicles, 

arising from the upper cells of a hyaline to subhyaline stroma, 

up to 50 μm wide and 20 μm high; conidiophores subcylindrical 

to ampulliform, 0–3-septate, straight to variously curved or 

geniculate-sinuous, unbranched, 15–25 × 3–5 μm. Conidiogenous 

cells terminal, unbranched, hyaline, smooth, tapering to flat-tipped 

apical loci, proliferating sympodially, 5–10 × 2.5–3.5 μm. Conidia 

solitary, subhyaline, smooth, guttulate or not, subcylindrical or 

narrowly obclavate, apex subobtuse, base obconically truncate, 

straight to variously curved, 3–10-septate, (15–)30–40(–60) × 

(1.5–)2–2.5(–3) μm; hila unthickened, not darkened nor refractive, 

1–1.5 μm diam; prominent microcyclic conidiation observed in vivo. 

Culture characteristics: Colonies after 1 mo at 24 ºC in the dark 

on MEA; erumpent, spreading, with moderate aerial mycelium, and 

smooth, lobate margins. Surface pale olivaceous-grey; reverse 

olivaceous-grey. Colonies reaching 30 mm diam. 

Specimen examined: New Zealand, Auckland, Princes Street, Auckland University 

Campus, on leaves of Tibouchina sp. (Melastomataceae), 9 Aug. 2004, C.F. Hill 

1061, holotype HAL 2359F, culture ex-type CBS 116462. 

Notes: Pseudocercospora tibouchinigena was initially reported 

from New Zealand as P. tibouchina (Braun et al. 2006), which is 

hitherto known only from Brazil. It differs from P. tibouchinae in that 

the latter species has narrowly subcylindrical conidia that are larger, 

40–120 × 2–3 μm (Viégas 1945), than those of P. tibouchinigena. 

The subhyaline conidia of P. tibouchinigena are not typical of 

Pseudocercospora s. str., but for the present, we choose to name 

it in Pseudocercospora until the clade in which it resides has been 

more fully resolved (Fig. 5). 

Clade 10: Cercospora, Miuraea, Phloeospora, 

Pseudocercosporella, Septoria, Xenocercospora 

Cercospora Fresen., in Fuckel, Hedwigia 1(15): 133. 1863 

and in Fuckel, Fungi Rhen. Exs., Fasc. II, No. 117. 1863. 

Mostly phytopathogenic producing conspicuous lesions, but also 

including saprobes. Mycelium internal, rarely also external; hyphae 

colourless or almost so to pigmented, branched, septate, smooth to 

faintly rough-walled. Stromata lacking to well-developed, subhyaline 

to usually pigmented, substomatal to intraepidermal. Conidiophores 

mononematous, macronematous, solitary to fasciculate, arising from 

internal hyphae or stromata, emerging through stomata or erumpent, 

very rarely arising from superficial hyphae, erect, continuous to 

pluriseptate, subhyaline to pigmented, smooth to faintly rough-walled, 

thin- to moderately thick-walled. Conidiogenous cells integrated, 

terminal or intercalary or conidiophores reduced to conidiogenous 

cells, monoblastic, determinate to usually polyblastic, sympodial, 

rarely with a few enteroblastically percurrent rejuvenations which 

are not connected with conidiogenesis; conidiogenous loci (scars) 

conspicuous, thickened and darkened, planate. Conidia solitary, 

very rarely catenate, scolecosporous, obclavate, cylindrical-filiform, 

acicular, hyaline or subhyaline (with a pale greenish tinge), mostly 

pluriseptate, euseptate, rarely with 0–1 or few septa, smooth or 

almost so, hila thickened and darkened, planate (from Crous & Braun 

2003). 

Type species: Cercospora penicillata (Ces.) Fresen., Beiträge zur 

Mykologie 3: 93. 1863. [= C. depazeoides (Desm.) Sacc.]. 

Cercospora sojina Hara, Nogyo Sekai, Tokyo 9: 28. 1915. 

Fig. 11. 

≡ Passalora sojina (Hara) H.D. Shin & U. Braun, Mycotaxon 58: 163. 

1996. 

Specimen examined: South Korea, Hongcheon, on Glycine soja (= G. max subsp. 

soja), 20 Jul. 2004, H.D. Shin, CBS H-20837, culture CPC 12322. 

Notes: Despite sparingly septate and broadly obclavatecylindrical 

conidia that tend to be subhyaline, this species is better 

accommodated in Cercospora than Passalora (Shin & Braun 1996) 

based on phylogenetic analysis. 

Cercospora eucommiae Crous, U. Braun & H.D. Shin, sp. 


Etymology: Name derived from Eucommia, the host on which it 

occurs. 

Leaf spots amphigenous, irregular to subcircular, 2–5 mm 

diam; surface grey-brown to brown with diffuse border; reverse 

olivaceous-brown with diffuse border. Mycelium internal, hyaline, 

consisting of septate, branched, smooth, 2–3 μm diam hyphae. 

Caespituli fasciculate, pale brown, amphigenous, up to 40 μm 

diam and 50 μm high (conidial mass white on leaf surface). 

Conidiophores aggregated in loose fascicles arising from the 

upper cells of a weakly developed brown stroma, up to 30 μm diam 

and 20 μm high, conidiophores pale brown, smooth, 1–3-septate, 

subcylindrical, straight to variously curved, unbranched, 20–50 × 

4–5 μm. Conidiogenous cells terminal, unbranched, pale brown, 

smooth, tapering to flat-tipped apical loci that are thickened, 

somewhat darkened, slightly refractive, 2 μm diam, 15–25 × 

4–5 μm, proliferating sympodially at apex. Conidia solitary, or 

in unbranched short chains, hyaline to pale olivaceous (with 

age), smooth, guttulate, obclavate, apex obtuse to subobtuse or 

clavate, base obconically subtruncate, straight to mildly curved, 

3–8-septate, (35–)60–75(–80) × (4–)5–6(–8) μm; hila thickened 

along the rim, but not darkened or planate, 1.5–2 μm diam. 

Culture characteristics: Colonies after 2 wk at 24 ºC in the dark 

on MEA; erumpent, spreading, with sparse aerial mycelium, and 


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Crous et al. 

Fig. 11. Cercospora sojina (CPC 12322). A. Leaf spots on upper and lower leaf surface. B. Close-up of lesion. C–G. Fascicles with conidiophores and conidiogenous cells. H. 

Conidia. Scale bars = 10 μm. 

smooth, lobate margins. Surface folded, dark mouse-grey with 

patches of dirty white; reverse fuscous black becoming greyish 

sepia at margin. Colonies reaching 12 mm diam. 

Specimens examined: South Korea, Chuncheon, on Eucommia ulmoides, 7 Oct. 

2003, H.D. Shin, holotype CBS H-20839, cultures ex-type CPC 10802 = CBS 

131932, CPC 10803, 10804; Chuncheon, on E. ulmoides, 11 Oct. 2002, H.D. Shin, 

CBS H-20838, culture CPC 10047. 

Notes: In the Korean material C. eucommiae occurred in mixed 

infections with a Pseudocercospora species (conidia 22–160 × 

4–7 μm) that resembles P. eucommiae (conidia 15–75 × 2–4 um), 

which is known from this host in China (Guo & Hsieh 1995). The 

description of C. eucommiae reveals the genus Cercospora to be 

paraphyletic. Morphologically C. eucommiae is distinct from other 

species in Cercospora in that the conidial hila and conidiogenous 

scars are different (thickened along the rim, not darkened and 

planate), and conidia also tend to occur in unbranched chains, 

which is not typical of Cercospora. Interestingly, it does not cluster 

with C. eremochloae, which also forms conidia in chains (Crous et 

al. 2011a). Although this species is not part of Cercospora s. str., 

we name it in this genus until further taxa are collected and studied 

to resolve the status of this subclade in relation to Cercospora s. str. 

Miuraea Hara, Byochugai-Hoten (Manual of Pests and 

Diseases): 779. 1948. 

Synonyms: See Braun (1995). 

Foliicolous, phytopathogenic, causing leaf spots. Mycelium internal 

and external, consisting of septate, branched, hyaline to subhyaline 

hyphae. Conidiophores semi-macronematous, mononematous, 

reduced to a single conidiogenous cell, integrated on hyphae, with 

small lateral peg-like protuberances; conidiogenesis holoblastic, 

monoblastic, determinate, occasionally polyblastic, proliferation 

sympodial or percurrent; conidiogenous loci more or less truncate, 

inconspicuous, unthickened, not darkened. Conidia solitary, 

ellipsoid-ovoid, subcylindrical-vermiform, obclavate, subclavate, 

somewhat asymmetrical, euseptate, transversely pluriseptate to 

muriformly septate, hyaline to faintly pigmented, thin-walled; hila 

truncate to somewhat convex, unthickened, not darkened (adapted 

from Braun 1995). 

Type species: Miuraea degenerans (Syd. & P. Syd.) Hara, 

Byochugai-Hoten (Manual of Pests and Diseases): 260. 1948. 

Notes: Morphologically Miuraea is intermediate between 

Pseudocercospora and Pseudocercosporella, which explains 

its phylogenetic position in this clade (Fig. 4). It differs from 

Pseudocercosporella in having superficial mycelium, and very 

broad, muriformly septate conidia. 

Miuraea persicae (Sacc.) Hara, Byochugai-Hoten (Manual 

of pests and diseases): 224. 1948. Fig. 13. 

Basionym: Cercospora persicae Sacc., Hedwigia 15: 119. 1876. 

Teleomorph: “Mycosphaerella” pruni-persicae Deighton, Trans. 

Brit. Mycol. Soc. 50: 328. 1967. 

Specimens examined: South Korea, Chuncheon, Prunus persica, 11 Oct. 2002, 

H.D. Shin, CBS H-20841, culture CPC 10069; Chuncheon, 7 Oct. 2003, P. 

armeniaca, H.D. Shin, CBS H-20840, CPC 10828–10830. 

68


Fig. 12. Cercospora eucommiae (CPC 10047). A. Leaf spots on upper and lower leaf surface. B. Close-up of lesion. C–G. Fascicles with conidiophores and conidiogenous 

cells. H, I. Conidia. Scale bars = 10 μm. 

Fig. 13. Miuraea persicae (CPC 10069). A. Leaf spots on upper and lower leaf surface. B, C. Close-up of fruiting (rather inconspicuous). D, E. Fascicles with conidiophores and 

conidiogenous cells. F. Conidia (note septation). Scale bars = 10 μm. 


69

Crous et al. 

Fig. 14. Pseudocercosporella arcuata (CPC 10050). A. Leaf spots on upper and lower leaf surface. B–D. Close-up of lesions. E, F. Fascicles with conidiophores and 

conidiogenous cells. G. Conidia. Scale bars = 10 μm. 

Phloeospora Wallr., Flora Cryptogamica Germaniae 2: 176. 

1833. 

Phytopathogenic, commonly associated with leaf spots, occurring 

on leaves and fruit. Mycelium immersed, consisting of hyaline, 

septate, branched hyphae. Conidiomata acervular, subepidermal, 

erumpent; wall of thin-walled textura angularis, opening by means 

of an irregular split. Conidiophores reduced to conidiogenous 

cells. Conidiogenous cells hyaline, smooth, cylindrical, discrete, 

indeterminate, proliferating via percurrent proliferations, or 

sympodially, formed from the upper cells of the acervulus. Conidia 

solitary, hyaline, smooth, septate, cylindrical, apex subobtuse to 

obtuse, base truncate, straight to curved. 

Type species: Phloeospora ulmi (Fr.) Wallr., Flora Cryptogamica 

Germaniae 2: 177. 1833. 

Specimens examined: Austria, Ulmus sp., H.A. van der Aa, CBS 613.81; Ulmus 

glabra, G. Verkley, CBS 344.97. Netherlands, Ulmus sp., H.A. van der Aa, CBS 

101564. 

Notes: Phloeospora is distinguished from Septoria by the production 

of conidia in acervuli, whereas conidiomata in the latter genus are 

pycnidial. Both genera are known to be polyphyletic (Verkley & 

Priest 2000, Quaedvlieg et al. 2011) and require further revision. 

Pseudocercosporella Deighton, Mycol. Pap. 133: 38. 1973. 

Foliicolous, phytopathogenic, causing discrete leaf spots. Mycelium 

mostly consistently internal, in some species with internal as well 

as external hyphae, hyaline to pale brown, septate, branched, 

smooth or almost so; stromata lacking or weakly to well-developed, 

substomatal to intraepidermal, usually colourless. Conidiophores 

solitary to fasciculate, emerging through stomata or erumpent 

through the cuticle, arising from inner hyphae or from stromata, 

sometimes formed as lateral branches of superficial hyphae, or 

aggregated in crustose to subglobose sporodochia; conidiophores 

simple, rarely branched, straight and subcylindrical to geniculatesinuous, 

hyaline, occasionally faintly pigmented at the base, rarely 

throughout, one-celled or septate. Conidiogenous cells integrated, 

terminal, or reduced to conidiogenous cells, mono- to polyblastic, 

sympodial; conidiogenous loci inconspicuous, unthickened, neither 

darkened nor conspicuously refractive. Conidia formed singly, rarely 

in simple or branched chains, subcylindrical, filiform, somewhat 

obclavate, 1–multi-euseptate, hyaline, thin-walled, mostly smooth, 

apex obtuse to subacute, base subtruncate, hilum unthickened, 

neither darkened, nor refractive (adapted from Braun 1995). 

Type species: Pseudocercosporella ipomoeae Deighton, Mycol. 

Pap. 133: 39. 1973. [= P. bakeri (Syd. & P. Syd.) Deighton, Mycol. 

Pap. 133: 41. 1973]. 

Note: Pseudocercosporella is polyphyletic (see Frank et al. 2010, 

Crous et al. 2011b) and new taxonomically useful morphological 

features will need to be determined to delineate all the genera 

presently accommodated in this clade. 

Pseudocercosporella arcuata S.K. Singh, P.N. Singh & 

Bhalla, Mycol. Res. 101: 542. 1997. Fig. 14. 

Specimen examined: South Korea, Chuncheon, on Rubus oldhamii (≡ R. pungens 

var. oldhamii), 11 Oct. 2002, H.D. Shin, CBS H-20842, culture CPC 10050. 

Pseudocercosporella capsellae (Ellis & Everh.) Deighton, 

Mycol. Pap. 133: 42. 1973. 

Basionym: Cylindrosporium capsellae Ellis & Everh., J. Mycol. 

3(11): 130. 1887. 

Additional synonyms in Braun (1995). 

Teleomorph: “Mycosphaerella” capsellae A.J. Ingman & Sivan., 

Mycol. Res. 95: 1339. 1991. 

Specimen examined: South Korea, Namyangju, Raphanus sativus, 22 Oct. 2007, 

H.D. Shin, CBS H-20843, cultures CPC 14773 = CBS 131896. 

Pseudocercosporella chaenomelis (Y. Suto) C. Nakash., 

Crous, U. Braun & H.D. Shin, comb. nov. MycoBank 

MB564818. Fig. 15. 

Basionym: Cercosporella chaenomelis Y. Suto, Mycoscience 40: 

513. 1999. 

= Mycosphaerella chaenomelis Y. Suto, Mycoscience 40: 513. 1999. 

Leaf spots amphigenous, irregular to angular, 5–20 mm diam, 

brown, delimited by leaf veins. Mycelium internal, hyaline, consisting 

of septate, branched, smooth, 1.5–2 μm diam hyphae. Caespituli 

70


Fig. 15. Pseudocercosporella chaenomelis (CPC 14795). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with white fruiting (rather inconspicuous). C–F. 

Fascicles with conidiophores and conidiogenous cells. G. Conidia. Scale bars = 10 μm. 

fasciculate to sporodochial, white, predominantly epiphyllous, up 

to 200 μm diam and 120 μm high. Conidiophores aggregated in 

dense fascicles, arising from the upper cells of a hyaline stroma, 

up to 180 μm diam and 100 μm high; conidiophores hyaline, 

smooth, subcylindrical to ampulliform, straight to variously curved, 

unbranched, reduced to conidiogenous cells, 5–12 × 3–4 μm, 

proliferating sympodially at apex. Conidia solitary, hyaline, smooth, 

guttulate to granular, subcylindrical to obclavate, apex subobtuse, 

base obconically truncate, straight to variously curved, 1–4-septate, 

(10–)30–38(–50) × (2–)2.5–3(–4) μm; hila unthickened, not 

darkened nor refractive, 1.5–2 μm diam; undergoing microcyclic 

conidiation on the host. Description based on CPC 14795. 

Culture characteristics: Colonies after 1 mo at 24 ºC in the dark on 

MEA; Colonies erumpent, spreading, with aerial mycelium sparse 

to absent, margins smooth, lobate. Surface irregularly folded, with 

a prominent network of ridges; folds appearing cinnamon, with 

surrounding areas and border brown-vinaceous; reverse sepia to 

chestnut, reaching up to 35 mm diam. 

Specimens examined: Japan, Shimane Pref., Matsue, on leaves of Chaenomeles 

sinensis, Y. Suto, 6 Nov. 1983, holotype SFH-917, in Herbarium of SPFRC; 

Mie Pref., Tsu, on leaves of C. sinensis, C. Nakashima, 29 Oct. 2011, epitype 

designated here TFM: FPH-8101, culture ex-epitype MUCC 1510 = CBS 132131. 

South Korea, Kimhae, C. speciosa (= C. lagenaria), 14 Nov. 2007, H.D. Shin, CBS 

H-20844, culture CPC 14795 = CBS 131897. 

Notes: Suto (1999) established the connection between 

Pseudocercosporella chaenomelis (as Cercosporella) and 

Mycosphaerella chaenomelis, which is the cause of a serious 

leaf spot disease referred to as frosty mildew on Chaenomeles 

sinensis in Japan. The fungus was found to overwinter by means 

of ascomata on fallen leaves, which provided the primary inoculum 

for new infections (April to June). Since the disease was previously 

known in Japan to be caused by a species of Cercosporella, Suto 

(1999) chose the latter genus to accommodate the anamorph. 

The hyaline conidia with unthickened conidial hila indicate that the 

fungus is better placed in Pseudocercosporella, and hence a new 

combination is proposed. Based on DNA sequence data from the 

ITS and ACT gene regions, strains from Japan and Korea appear 

identical (unpubl. data). 

Pseudocercosporella chaenomelis occurs in mixed infections 

with Pseudocercospora cydoniae. Pseudocercosporella 

chaenomelis is morphologically comparable only with Ps. gei, 

known on Geum spp. in North America and the Far East of Russia 

(Braun 1995). The latter species differs in having smaller stromata 

(20–45 μm diam) and much longer filiform-acicular conidia, 20–120 

× 1–3(–4) μm (Braun 1995). Pseudocercosporella crataegi on 

Crataegus spp. in North America is distinct, forming superficial 

hyphae with solitary conidiophores, and its much smaller stromata 

and much longer conidia, and P. potentillae on Potentilla sp. in 

Russia also differs by having very long conidia (Braun 1995). 

Pseudocercosporella koreana Crous, U. Braun & H.D. 

Shin, sp. nov. MycoBank MB564819. Fig. 16. 

Etymology: Name derived from the country where it was collected. 

Leaf spots amphigenous, indistinct, irregular, chlorotic, up to 6 mm 

diam. Mycelium internal, hyaline, consisting of septate, branched, 

smooth, 1.5–2.5 μm diam hyphae. Caespituli fasciculate, white, 

amphigenous, up to 60 μm diam and 90 μm high. Conidiophores 

aggregated in dense fascicles, on the upper cells of a pale brown 

to hyaline, usually substomatal stroma, up to 45 μm diam and 20 


71

Crous et al. 

Fig. 16. Pseudocercosporella koreana (CPC 11414). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with white fruiting. C. Substomatal stroma. D–H. 

Fascicles with conidiophores and conidiogenous cells. I. Conidia. Scale bars = 10 μm. 

μm high; conidiophores hyaline or pale brown at base, smooth, 

0–2-septate, but frequently reduced to conidiogenous cells, 

subcylindrical, straight to variously curved or geniculate-sinuous, 

unbranched or branched below, 15–25 × 4–5 μm, proliferating 

sympodially at apex. Conidia solitary, hyaline, smooth, prominently 

guttulate, narrowly obclavate, apex obtuse to subobtuse, base 

obconically subtruncate, straight to variously curved, 3–13-septate, 

(40–)60–80(–130) × (2.5–)3(–4) μm; hila unthickened, neither 

darkened nor refractive, 2 μm diam. 

Culture characteristics: Colonies after 2 wk at 24 ºC in the dark on 

MEA; surface folded with a prominent network of ridges, erumpent, 

spreading, with sparse aerial mycelium, and smooth, lobate 

margins. Surface olivaceous-grey to iron-grey; reverse iron-grey to 

greenish black. Colonies reaching 6 mm diam. 

Specimen examined: South Korea, Hoengseong, on Vicia amurensis, 4 Aug. 2004, 

H.D. Shin, holotype CBS H-20845, isotype HAL 1850 F, culture ex-holotype CPC 

11414. 

Notes: Braun (1995) listed several species of Pseudocercosporella 

on Fabaceae. None of these occur on Vicia, and only one, P. 

tephrosiae (on Tephrosia, Africa), has conidia of similar length (40– 

110 × 3–4.5 μm), although they are wider, subcylindrical-acicular, 

and have 3–6 septa. 

Pseudocercosporella oxalidis (Goh & W.H. Hsieh) U. 

Braun, Nova Hedwigia 55: 218. 1992. 

Basionym: Pseudocercospora oxalidis Goh & W.H. Hsieh, Bot. 

Bull. Acad. Sinica 30: 127. 1989. 

Specimen examined: Taiwan, Taipei, Wulai, on living leaves of Oxalis debilis (= O. 

corymbosa), R. Kirschner, 2258, 22 Feb. 2005, culture CBS 118758. 

Septoria Sacc., Syll. Fung. 3: 474. 1884. 

Synonyms: See Sutton (1980). 

Phytopathogenic and endophytic, occurring on leaves, fruit 

and stems, causing discrete lesions. Conidiomata pycnidial, 

immersed, separate or aggregated, globose, papillate or not, 

brown, with a thin wall of brown textura angularis. Ostiole single, 

circular, central, sometimes papillate. Conidiophores reduced 

to conidiogenous cells. Conidiogenous cells hyaline, smooth, 

ampulliform, doliiform or lageniform to short cylindrical, holoblastic, 

determinate or indeterminate, proliferating sympodially and/or 

percurrently; conidiogenous loci unthickened. Conidia solitary, 

hyaline, multiseptate, guttulate or not, thin-walled, filiform, smooth, 

continuous or constricted at the septa; hila unthickened. 

Type species: Septoria cytisi Desm. Ann. Sci. Nat., Bot., Sér. 3, 8: 

24. 1847. 

Note: Septoria is polyphyletic (Quaedvlieg et al. 2011). 

Clade 11: Sonderhenia 

Sonderhenia H.J. Swart & J. Walker, Trans. Brit. Mycol. Soc. 

90: 640. 1988. 

Foliicolous, phytopathogenic, causing discrete leaf spots. Leaf 

spots amphigenous, round to confluent and irregular, surrounded 

by a purple border when young, which becomes dark red to brown 

and raised with age. Ascomata pseudothecial, amphigenous, on 

one side of each lesion, often 1–3, intermingled with conidiomata, 

immersed, black, punctiform, globose to subglobose; apical 

ostiole substomatal; wall olive-brown, of 3–4 layers of textura 

angularis, subhymenium of 1–2 layers of colorless cells. Asci 

72


Fig. 17. Pallidocercospora heimii (CPC 1395). Asci, ascospores, germinating ascospores (after 24 h on malt extract agar), hyphae with conidiogenous loci, and conidia. Scale 

bar = 10 μm. 

fasciculate, bitunicate, subsessile, 8-spored, ovoid to obclavate, 

straight to incurved. Ascospores 2–3-seriate, hyaline, guttulate, 

straight or slightly curved, fusiform, 1-septate, widest just above 

median septum, slightly constricted at septum. Conidiomata 

pycnidial, amphigenous, subepidermal with central non-projecting 

ostiole, scattered, black, globose; wall of 2–3 layers of brown 

cells. Conidiogenous cells minute, olivaceous, proliferating 

enteroblastically and percurrently, lining the inner pycnidial wall 

layer. Conidia ellipsoid to cylindrical or ovoid, straight or bent, 

brown, 3-distoseptate, not constricted, verruculose, apex obtuse, 

base truncate with marginal frill (adapted from Crous 1998). 

Type species: Sonderhenia eucalyptorum H.J. Swart & J. Walker, 

Trans. Brit. Mycol. Soc. 90: 640. 1988. 

Notes: Sonderhenia includes taxa with Mycosphaerella-like 

teleomorphs and pycnidial anamorphs that form brown, transversely 

distoseptate conidia on brown, percurrently proliferating 

conidiogenous cells. Only two species, S. eucalypticola and S. 

eucalyptorum are known. 

Clade 12: Pallidocercospora, Scolecostigmina, 

Trochophora and Pseudocercospora-like 

Pallidocercospora Crous, gen. nov. MycoBank MB564820. 

Fig. 17. 

Etymology: The name reflects the pale brown Cercospora-like 

conidia in this genus. 


Ascomata single, black, immersed, globose, glabrous; wall of 

3–4 layers of medium brown textura angularis. Asci fasciculate, 

bitunicate, aparaphysate, subsessile, 8-spored, ellipsoid to 

obclavate or cylindrical, straight or curved, numerous. Ascospores 

2–multi-seriate, oblique, overlapping, straight ellipsoidal to obovoid, 

colourless, smooth, 1-septate. Mycelium predominantly immersed, 

consisting of olivaceous-brown hyphae, smooth, branched, septate, 

2–4 μm diam. Conidiophores in vivo fasciculate, or occurring 

singly on superficial mycelium as lateral projections, unbranched 

or branched, septate, cylindrical, straight to geniculate–sinuous, 

olivaceous-brown. Conidiogenous cells integrated, terminal, 


73

Crous et al. 

cylindrical, straight to geniculate-sinuous, olivaceous-brown, 

proliferating sympodially or percurrently; conidiogenous loci 

unthickened, not darker than the surrounding conidiogenous 

cell. Conidia solitary, straight to irregularly curved, guttulate, 

pale olivaceous to olivaceous-brown, subcylindrical to narrowly 

obclavate, multiseptate; hila neither thickened nor darkened. 

Type species: Pallidocercospora heimii (Crous) Crous, comb. nov. 

Notes: Species of Pallidocercospora have pale olivaceous, 

smooth conidia (generally referred to as the Mycosphaerella heimii 

complex; Crous et al. 2004c), and form red crystals when cultivated 

in agar (on WA, SNA, PDA, MEA), which distinguishes them from 

Pseudocercospora. Pseudocercospora has several synonyms 

(see Seifert et al. 2011). Cercoseptoria with its mostly acicular 

conidia, was correctly treated as synonym of Pseudocercospora 

by Deighton (1976). Other synonyms include Ancylospora Sawada 

(based on A. costi), now treated as P. costina; Cercocladospora G.P. 

Agarwal & S.M. Singh (based on C. adinae, nom. non rite publ.), 

now treated as P. adinicola; and Helicominia L.S. Olive (based on 

H. caperonia), now P. caperoniae, and Pantospora Cif. (based on P. 

guazumae) (see Ellis 1971, Deighton 1976), the muriformly septate 

conidia of the latter are similar to those of Pseudocercospora 

pseudostigmina-platani, though Pantospora has been shown to be 

a genus in its own right (Minnis et al. 2011). 

Pallidocercospora acaciigena (Crous & M.J. Wingf.) Crous 

& M.J. Wingf., comb. nov. MycoBank MB564821. 

Basionym: Pseudocercospora acaciigena Crous & M.J. Wingf., 

Stud. Mycol. 50: 464. 2004. 

Teleomorph: “Mycosphaerella” acaciigena Crous & M.J. Wingf., 

Stud. Mycol. 50: 463. 2004. 

Specimen examined: Venezuela, Acarigua, on leaves of Acacia mangium, May 

2000, M.J. Wingfield, CBS H-9873, holotype of M. acaciigena and P. acaciigena; 

cultures ex-type CBS 115432, 112515, 112516 = CPC 3836–3838. 

Pallidocercospora crystallina (Crous & M.J. Wingf.) Crous 


Basionym: Pseudocercospora crystallina Crous & M.J. Wingf., 

Mycologia 88: 451. 1996. 

Teleomorph: “Mycosphaerella” crystallina Crous & M.J. Wingf., 

Mycologia 88: 451. 1996. 

Specimens examined: South Africa, Kwazula-Natal Province,Umvoti, on leaves 

of Eucalyptus bicostata, Oct. 1994, M.J. Wingfield (holotypes PREM 51922, 

teleomorph; PREM 51923, anamorph, cultures ex-type CPC 800–802); Kwazula- 

Natal Province, leaf litter of E. grandis × camaldulensis, Jun. 1995, M.J. Wingfield 

(PREM 51937, cultures CPC 1178–1180). 

Pallidocercospora heimii (Crous) Crous, comb. nov. 


Basionym: Pseudocercospora heimii Crous, S. African For. J. 172: 

4. 1995. 

Teleomorph: “Mycosphaerella” heimii Crous, S. African For. J. 172: 

2. 1995. 

≡ “Mycosphaerella” heimii Bouriquet, Encycl. Mycol. 12: 418. 1946. nom. 

nud. 

Specimens examined: Madagascar, Moramanga, on leaves of Eucalyptus sp., Apr. 

1994, P.W. Crous, PREM 51749, holotype of teleomorph; PREM 51748, holotype 

of anamorph, cultures ex-type CPC 760–761 = CBS 110682. 

Pallidocercospora heimioides (Crous & M.J. Wingf.) Crous 


Basionym: Pseudocercospora heimioides Crous & M.J. Wingf., 

Can. J. Bot. 75: 787. 1997. 

Teleomorph: “Mycosphaerella” heimioides Crous & M.J. Wingf., 

Can. J. Bot. 75: 787. 1997. 

Specimens examined: Indonesia, N. Sumatra, Lake Toba area, leaves of Eucalyptus 

sp., Mar. 1996, M.J. Wingfield, holotype of teleomorph PREM 54966; holotype of 

anamorph PREM 54967; cultures ex-type CPC 1311, 1312 = CBS 111190). 

Pallidocercospora holualoana (Crous, Joanne E. Taylor & 

M.E. Palm) Crous, comb. nov. MycoBank MB564825. 

Basionym: “Mycosphaerella” holualoana Crous, Joanne E. Taylor & 

M.E. Palm, Mycotaxon 78: 458. 2001. 

Specimen examined: USA, Hawaii, Kona district, Holualoa, on a living leaf of 

Leucospermum sp., P.W. Crous & M.E. Palm, 17 Nov. 1998, holotype PREM 

56926, cultures ex-type CPC 2126–2128). 

Pallidocercospora irregulariramosa (Crous & M.J. Wingf.) 

Crous & M.J. Wingf., comb. nov. MycoBank MB564826. 

Basionym: Pseudocercospora irregulariramosa Crous & M.J. 

Wingf., Can. J. Bot. 75: 785. 1997. 

Teleomorph: “Mycosphaerella” irregulariramosa Crous & M.J. 

Wingf., Can. J. Bot. 75: 785. 1997. 

Specimens examined: South Africa, Northern Province, Tzaneen, on leaves of 

Eucalyptus saligna, Mar. 1996, M.J. Wingfield, holotype of teleomorph PREM 

54964; holotype of anamorph PREM 54965; cultures ex-type CPC 1360 = CBS 

114777). 

Pallidocercospora konae (Crous, Joanne E. Taylor & M.E. 

Palm) Crous, comb. nov. MycoBank MB564827. 

Basionym: “Mycosphaerella” konae Crous, Joanne E. Taylor & 

M.E. Palm, Mycotaxon 78: 459. 2001. 

Specimen examined: USA, Hawaii, Kona district, Holualoa, on a living leaf on 

Leucadendron cv. Safari Sunset, 17 Nov. 1998, P.W. Crous & M.E. Palm, holotype 

PREM 56921; ex-type cultures CPC 2123–2125. 

Scolecostigmina U. Braun, N. Z. J. Bot. 37: 323. 1999. Fig. 

18. 

Foliicolous, phytopathogenic, associated with leaf spots. Mycelium 

immersed, consisting of septate, branched, pigmented hyphae. 

Sporodochia immersed to erumpent; stromata subglobose to 

applanate, composed of brown, angular to subglobose cells. 

Conidiophores numerous, densely aggregated, arising from stroma, 

subcylindrical or somewhat tapered towards the apex, occasionally 

ampulliform, continuous or septate, pigmented, wall somewhat 

thickened, usually verruculose; conidiogenous cells integrated, 

terminal or at times conidiophores reduced to conidiogenous cells, 

holoblastic, proliferating percurrently via conspicuous annellations. 

Conidia solitary, scolecosporous, usually subcylindrical-obclavate, 

transversely pluriseptate, occasionally with few longitudinal or 

oblique septa, euseptate, rarely with few intermixed distosepta, 

thick-walled, pigmented, dark, smooth to verrucose, apex obtuse 

to subacute, base truncate or obconically truncate; secession 

schizolytic (adapted from Braun et al. 1999). 

Type species: Scolecostigmina mangiferae (Koord.) U. Braun & 

Mouch., N. Z. J. Bot. 37: 323. 1999. 

74


Fig. 18. Scolecostigmina mangiferae (CBS 125467). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C–E. Fascicles with conidiophores and 

conidiogenous cells (note rough percurrent proliferations). F. Conidia. Scale bars = 10 μm. 

Fig. 19. Trochophora simplex (CBS 124744). A. Leaf spots on upper and lower leaf surface. B, C. Close-up of leaf spot with fruiting. D–G. Fascicles with conidiophores and 

conidiogenous cells. H, I. Conidia. Scale bars = 10 μm. 

Specimen examined: Australia, Queensland, Mareeba, S16º58’75.5” 

E145º20’60.8”, leaves of Mangifera indica, 10 Aug. 2009, P.W. Crous & R.G. Shivas, 

CBS H-20846, culture CPC 17352, 17351 = CBS 125467. 

Trochophora R.T. Moore, Mycologia 47: 90. 1955. Fig. 19. 

Foliicolous, but pathogenicity unproven. Colonies hypophyllous, 

medium to dark brown, consisting of numerous synnemata. Stroma 

absent, but with a superficial network of hyphae linking the various 


75

Crous et al. 

synnemata. Conidiophores synnematous, mostly unbranched and 

straight, or with 1–2 short branches, straight or curved, cylindrical, 

individual conidiophores tightly aggregated, but separating near 

the apex, pale to medium brown, smooth. Conidiogenous cells 

polyblastic, integrated, terminal, determinate to sympodial, with 

visible unthickened scar, clavate. Conidia solitary, terminal or 

lateral on conidiogenous cells, prominently curved to helicoid, pale 

to medium brown, smooth, transversely euseptate with a darkened, 

thickened band at the septa (adapted from Crous et al. 2009a). 

Type species: Trochophora simplex (Petch) R.T. Moore, Mycologia, 

47: 90. 1955. 

Specimens examined: Japan, Shimane, on Daphniphyllum teijsmannii, 26 April 

2008, C. Nakashima & I. Araki, MUMH 11134, culture MUCC 952. South Korea, 

Jeju, Halla arboretum, on D. macropodum, 29 Oct. 2005, H.D. Shin, CBS H-20847, 

culture CBS 124744. 

Notes: Other Pseudocercospora-like species found in this clade are 

P. colombiensis (foliar pathogen of Eucalyptus; Crous 1998), and P. 

thailandica (foliar pathogen of Acacia; Crous et al. 2004d), both 

also having Mycosphaerella-like teleomorphs. Morphologically, 

these taxa appear typical members of Pseudocercospora s. 

str. so it would be difficult to identify these as different from 

Pseudocercospora without the aid of DNA sequence comparisons. 

Clade 13: Passalora-like 

Notes: This clade is represented by Passalora eucalypti, which was 

originally described as a leaf spot pathogen of Eucalyptus saligna 

in Brazil (Crous 1998, Crous & Braun 2003). Recently, a second 

species was found to belong to this clade, namely Passalora 

leptophlebiae, which was described from Eucalyptus leptophlebia 

leaves collected in Brazil (Crous et al. 2011a). Both species are 

charaterised by fasciculate conidiophores and catenate, pale brown 

conidia, with thickened, darkened and refractive scars and hila. 

Clade 14: Pseudocercospora s. str. 

Pseudocercospora Speg., Anales Mus. Nac. Hist. Nat. 

Buenos Aires, Ser. 3, 20: 437. 1910. 

Foliicolous, chiefly phytopathogenic, but also endophytic; 

commonly associated with leaf spots, but also occurring on fruit. 

Mycelium internal and external, consisting of smooth, septate, 

subhyaline to brown, branched hyphae. Stroma absent to welldeveloped. 

Conidiophores in vivo arranged in loose to dense 

fascicles, sometimes forming distinct synnemata or sporodochia, 

emerging through stomata or erumpent through the cuticle, often 

arising from substomatal or subcuticular to intraepidermal stromata, 

or occurring singly on superficial hyphae, short to long, septate or 

continuous, i.e. conidiophores may be reduced to conidiogenous 

cells, simple to branched and straight to geniculate-sinuous, pale 

to dark brown, smooth to finely verruculose. Conidiogenous cells 

integrated, terminal, occasionally intercalary, polyblastic, sympodial, 

or monoblastic, proliferating percurrently via inconspicuous or 

darkened, irregular annellations, at times denticulate, pale to dark 

brown; scars inconspicous, or only thickened along the rim, or flat, 

and slightly thickened and darkened, but never pronounced. Conidia 

solitary, rarely in simple chains, subhyaline, olivaceous, pale to 

dark brown, usually scolecosporous, i.e. obclavate–cylindrical, 

filiform, acicular, and transversely plurieuseptate, occasionally 

also with oblique to longitudinal septa, conidia rarely amero- to 

phragmosporous, short subcylindrical or ellipsoidal-ovoid, aseptate 

or only with few septa, apex subacute to obtuse, base obconically 

truncate to truncate, or bluntly rounded, with or without a minute 

marginal frill, straight to curved, rarely sigmoid, smooth to finely 

verruculose; hila usually unthickened, not darkened, at most 

somewhat refractive, occasionally slightly thickened along the 

rim, or rarely flat, and slightly thickened and darkened, but never 

pronounced. 

Type species: P. vitis (Lév.) Speg., Anales Mus. Nac. Hist. Nat. 

Buenos Aires, Ser. 3, 20: 438. 1910. 

Specimens examined: South Korea, Namyangju, on Vitis vinifera, 30 Sep. 2004, 

H.D. Shin, CBS H-20848, CPC 11595 = CBS 132012; V. vinifera, 1 Oct. 2007, H.D. 

Shin, CPC 14661 = CBS 132112. 

Pseudocercospora abelmoschi (Ellis & Everh.) Deighton, 

Mycol. Pap. 140: 138. 1976. Fig. 20. 

Basionym: Cercospora abelmoschi Ellis & Everh., J. Inst. Jamaica 

1: 347. 1893. 

= Cercospora hibisci Tracy & Earle, Bull. Torrey Bot. Club 22: 179. 1895. 

= Cercospora hibisci-manihotis Henn., Hedwigia 43: 146. 1904. 

Specimen examined: South Korea, Suwon, on Hibiscus syriacus, 2 Oct. 2007, H.D. 

Shin, CBS H-20849, CPC 14478 = CBS 132103. 

Pseudocercospora ampelopsis Crous, U. Braun & H.D. 


Etymology: Name derived from the host Ampelopsis, from which it 

was collected. 

Leaf spots amphigenous, irregular to subcircular, 2–8 mm diam, 

dark brown on upper surface, dull brownish green on lower surface. 

Mycelium internal and external, pale brown to brown, consisting of 

septate, branched, smooth, 1.5–4 μm diam hyphae, anastomosing 

on surface. Caespituli fasciculate, brown, amphigenous, emerging 

through stomata (but stromata lacking). Conidiophores aggregated 

in loose fascicles, or solitary, arising from superficial mycelium, 

medium to dark brown, smooth to finely verruculose, 3–6-septate, 

subcylindrical, straight to variously curved, unbranched, 20–80 

× (2.5–)3–5(–6) μm. Conidiogenous cells terminal, unbranched, 

brown, finely verruculose, tapering to flat-tipped apical loci, 

proliferating sympodially, 10–15 × 4–5 μm. Conidia solitary, dark 

brown, finely verruculose, guttulate, obclavate-cylindrical, apex 

obtuse, base obconically subtruncate, straight to gently curved, 

3–12-septate, (35–)40–90(–110) × 3–5(–6) μm; hila unthickened, 

neither darkened nor refractive, 2 μm diam. 


on MEA; surface folded, erumpent, spreading, with sparse aerial 

mycelium, and smooth, lobate margins. Surface olivaceous-grey; 

reverse iron-grey. Colonies reaching 7 mm diam. 

Specimen examined: South Korea, Hongcheon, on Ampelopsis glandulosa var. 

heterophylla, 24 Oct. 2004, H.D. Shin, holotype CBS H-20850, isotype HAL 1866 

F, culture ex-type CPC 11680 = CBS 131583. 

Notes: Pseudocercospora brachypus, which also occurs on 

Ampelopsis, has much shorter and narrower conidia, 25–60 × 

2–3.5 μm (Guo & Hsieh 1995). Pseudocercospora ampelopsis is 

morphologically close to P. riachuelii var. horiana on Ampelocissus, 

76


Fig. 20. Pseudocercospora abelmoschi (CPC 14478). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C–G. Hyphae giving rise to 

conidiogenous cells and conidia. H. Conidia. Scale bars = 10 μm. 

Fig. 21. Pseudocercospora ampelopsis (CPC 11680). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C. Conidiophores and conidiogenous 

cells. D. Conidia. Scale bar = 10 μm. 

Cissus and Parthenocissus species (Crous & Braun 2003). The 

two are similar in that conidiophores are solitary and form in 

fascicles and arise from superficial hyphae, and conidia of the two 

taxa are similar in size. Pseudocercospora ampelopsis differs in 

having much longer pluriseptate conidiophores whereas those of P. 

riachuelii var. horiana are much shorter and 0–1-septate. 

Pseudocercospora angolensis (T. Carvalho & O. Mendes) 

Crous & U. Braun, Sydowia 55: 301. 2003. 

Basionym: Cercospora angolensis T. Carvalho & O. Mendes, Bol. 

Soc. Brot. 27: 201. 1953. 

≡ Phaeoramularia angolensis (T. Carvalho & O. Mendes) P.M. Kirk, 

Mycopathologia 94: 177. 1986. 

≡ Pseudophaeoramularia angolensis (T. Carvalho & O. Mendes) U. 

Braun, Cryptog. Mycol. 20: 171. 1999. 

Specimens examined: Angola, Mozambique Province, on leaves of Citrus × 

aurantium (= ×sinensis), Dec. 1951, Carvalho & O. Mendes, BPI 432660, BPI 

442839 (paratypes), BPI 442837 (holotype), IMI 56597 (isotype). Camaroon, 

Yaoundé, on leaves of C. × aurantium, 17 Mar. 1978, E. Milla, IMI 252792. Ethiopia, 

on leaves of Citrus sp., IMI 361170. Kenya, on leaves of C. × aurantium, 15 Nov. 

1991, A. Seif W3753, IMI 351626. Uganda, on leaves of C. × aurantium, 14 Jun. 

1991, W.T.H. Peregrine, IMI 384297. West Africa, intercepted at San Pedro, 

California, USA, on leaves of Citrus sp., 2 Oct. 1953, L.A. Hart, BPI 432661, BPI 

432659. Zambia, on leaves of Citrus sp., 18 Jun. 1973, R.H. Raemakers 7837, 

IMI 176562; Chilanga, on leaves of C. × aurantium, 28 Sep. 1983, D.M. Naik, IMI 

280618; Chilanga, on leaves of Citrus sp., 18 Jul. 1975, B.K. Patel, IMI 196889; 

Lusaka, on leaves of Citrus sp., 17 June 1977, I. Javaid, IMI 214501. Zimbabwe, 


77

Crous et al. 

Fig. 22. Pseudocercospora araliae (CPC 10154). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C–E. Fascicles with conidiophores and 

conidiogenous cells. F. Conidia. Scale bars = 10 μm. 

Fig. 23. Pseudocercospora atromarginalis (CPC 11372–11374). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with 

conidiophores and conidiogenous cells. E. Conidia. Scale bar = 10 μm. 

Bindura, on leaves of Citrus sp., 13 Aug. 1979, A. Rothwell, IMI 240682; on leaves 

of Citrus sp., Sep. 2000, M.C. Pretorius, epitype designated here CBS H-20851, 

culture ex-epitype CPC 4112–4118, 4111 = CBS 112933. 

Pseudocercospora araliae (Henn.) Deighton, Mycol. Pap. 

140: 19. 1976. Fig. 22. 

Basionym: Cercospora araliae Henn., Bot. Jahrb. Syst. 31: 742. 

1902; also 37: 165. 1906. 

≡ Cercosporiopsis araliae (Henn.) Miura, Fl. Manchuria & E. Mongolia, 

27, 3: 533. 1928. 

= Cercospora atromaculans auct., non Ellis & Everh. 

Specimens examined: Japan, Tosa, Ushioe-yama, on Aralia elata var. glabrescens, 

Aug. 1901, T. Yoshinaga, holotype B 700015014; A. elata, T. Kobayashi & C. 

Nakashima, epitype designated here TFM: FPH-8094, ex-epitype cultures MUCC 

873, MAFF 238192. South Korea, Jeju, Halla Arboretum, on A. elata, 14 Sep. 2002, 

H.D. Shin, CBS H-20852, culture CPC 10154; Wonju, on A. elata, 21 Sep. 2003, 

H.D. Shin, CBS H-20853, cultures CPC 10782–10784. 

Pseudocercospora atromarginalis (G.F. Atk.) Deighton, 

Mycol. Pap. 140: 139. 1976. Fig. 23. 

Basionym: Cercospora atromarginalis G.F. Atk. (atramarginalis), J. 

Elisha Mitchell Sci. Soc. 8: 59. 1892. 

= Cercospora rigospora G.F. Atk., J. Elisha Michell Sci. Soc. 8: 65. 1892. 

= Cercospora tosensis Henn., Bot. Jahrb. Syst. 34: 605. 1905. 

= Cercospora nigri Tharp, Mycologia 9: 112. 1917. 

= Cercospora solani-biflori Sawada, Formosan Agric. Rev. 39: 701. 1942 

(nom. inval.). 

Specimens examined: Japan, Prov. Tosa, Aki-machi, on Solanum nigrum, Oct. 

1903, Yoshinaga No. 43, (holotype of C. tosensis, B 700015016). South Korea, 

Namyangju, on S. nigrum, 27 Jul. 2004, H.D. Shin, CBS H-20854, CPC 11372– 

11374. New Zealand, Auckland, Jan. 2004, C.F. Hill 970, CBS 114640. 

Notes: Pseudocercospora atromarginalis was described from 

Solanum collected in Auburn Alabama, USA. Material studied here 

from New Zealand and Korea represents the same species, which 

might be authentic for the name. Fresh material from Solanum in the 

USA, and a detailed study of the synonyms listed by Chupp (1954) 

would resolve this issue. An isolate identified as P. chengtuensis 

(on Lycium, Solanaceae) appears identical to Pseudocercospora 

atromarginalis. 

78


Fig. 24. Pseudocercospora balsaminae (CPC 10044). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C–F. Fascicles and solitary 

conidiophores with conidiogenous cells. G. Conidia. Scale bars = 10 μm. 

Pseudocercospora balsaminae (Syd.) Deighton, Mycol. 

Pap. 140: 139. 1976. Fig. 24. 

Basionym: Cercoseptoria balsaminae Syd., Ann. Mycol. 33: 69. 

1935. 

Specimens examined: South Korea, Chuncheon, on Impatiens textorii, 11 Oct. 

2002, H.D. Shin, CBS H-20856, CPC 10044 = CBS 131882; Dongducheon, on I. 

textorii, 11 Oct. 2004, H.D. Shin, CBS H-20855, CPC 10699–10701. 

Pseudocercospora callicarpae (Cooke) Y.L. Guo & W.X. 

Zhao, Acta Mycol. Sin. 8: 118. 1989. 

Basionym: Cercospora callicarpae Cooke, Grevillea 6: 140. 1878. 

= ? Cercospora callicarpicola Naito, Mem. Coll. Agric. Kyoto Imp. Univ. 47: 49. 

1940. 

Specimen examined: Japan, Ibaraki, on Callicarpa japonica, 11 Sep. 1998, T. 

Kobayashi, MUCC 888, MAFF 237784, CNS-442. 

Pseudocercospora catalpigena U. Braun & Crous, Mycol. 

Progr. 2: 198. 2003. 

Specimen examined: Japan, Wakayama, on Catalpa ovata, 30 Oct. 2007, C. 

Nakashima & I. Araki, MUMH 10868, culture MUCC 743. 

Pseudocercospora catappae (Henn.) X.J. Liu & Y.L. Guo, 

Mycosystema 2: 230. 1989. 

Basionym: Cercospora catappae Henn., Bot. Jahrb. Syst. 34: 56. 

1905. 

= Pseudocercospora catappae Goh & W.H. Hsieh, in Hsieh & Goh, Cercospora 

and similar fungi from Taiwan: 57. 1990, homonym of P. catappae (Henn.) X.J. 

Liu & Y.L. Guo, 1989. 

= Ramularia catappae Racib., Paras. Algen u. Pilze Javas II, Batavia: 41. 

1900. 

= Cercospora terminaliae Sawada (terminariae), Taiwan Agric. Rev. 38: 701. 

1942 (nom. illeg.), homonym of C. terminaliae Syd. 1929. 

Specimens examined: Tanzania, Zanzibar, Dar-es-Salam, on Terminalia catappa, 

26 Oct. 1901, Stuhlmann holotype B 700015015. Japan, Okinawa, on T. catappa, 

17 Nov. 2007, C. Nakashima & T. Akashi, MUMH 10913, culture MUCC 809. 

Pseudocercospora cercidicola Crous, U. Braun & C. 

Nakash., sp. nov. MycoBank MB564829. Fig. 25. 

Etymology: Name reflects the host Cercis, from which it was 

collected. 

Leaf spots amphigenous, irregular to angular, 1–5 mm diam, 

confined by leaf veins, brown on upper surface, with raised, dark 

brown border, on lower surface medium brown, with indistinct 

borders. Mycelium internal, consisting of pale brown, smooth, 

septate, branched, 2–3 μm diam hyphae. Caespituli fasciculate to 

sporodochial, amphigenous, but predominantly epiphyllous, greybrown 

on leaves, up to 130 μm wide and 150 μm high. Conidiophores 

aggregated in dense fascicles arising from the upper cells of a 

brown stroma up to 80 μm wide and 60 μm high; conidiophores 

brown, finely verruculose, 2–6-septate, subcylindrical, straight 

to variously curved, unbranched or branched above, 20–50 × 

3–5 μm. Conidiogenous cells terminal or lateral, unbranched, 

medium brown, finely verruculose, tapering to flat-tipped apical 

loci, proliferating sympodially, 10–20 × 2–3 μm. Conidia solitary, 

medium brown, smooth, guttulate, subcylindrical to narrowly 

obclavate, apex subobtuse, base long obconically subtruncate, 

straight to variously curved, (0–)3–6-septate, (27–)30–50(–60) × 

(2.5–)3(–3.5) μm; hila neither thickened, nor darkened-refractive, 

1.5–2 μm diam. 

Culture characteristics: Colonies on MEA 10–15 mm after 2 wk at 

20 °C in the dark, restricted, with margin mildly lobed, felty, pale 

olivaceous or greyish olivaceous, surrounded by greyish margin; 

reverse olivaceous. 

Specimens examined: Japan, Ibaraki, on Cercis chinensis, 10 Sep. 1998, T. & Y. 

Kobayashi, holotype CBS H-20895, culture ex-type MUCC 896, MAFF 237791 

= CBS 132041; Tokyo, Koishikawa Botanical Garden, on Cercis chinensis, 10 Nov. 

2007, I. Araki & M. Harada, MUMH 11108, culture MUCC 937; Japan, Kanagawa, 

on Cercis chinensis, May 1992, K.Kishi, culture MAFF 237128. 

Notes: Asian collections of cercosporoid fungi on Cercis chinensis 

were considered as representative of Cercospora chionea by 

Chupp (1954). The latter species was shown to be a member of 

Passalora by Braun (1993). Shin & Braun (2000) introduced a new 

species of Pseudocercospora for the taxon occurring on Cercis 

in Asia, namely P. cercidis-chinensis, based on material collected 

in Korea. Phylogenetic data obtained in the present study (Fig. 

5) show that the Japanese collections are distinct. As the name 


79

Crous et al. 

Fig. 25. Pseudocercospora cercidicola (CBS H-20895). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with conidiophores 

and conidiogenous cells. E, F. Conidiophores on superficial hyphae. G. Conidia. Scale bars = 10 μm. 

Fig. 26. Pseudocercospora cercidis-chinensis (CPC 14481). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with 

conidiophores and conidiogenous cells. E. Conidia. Scale bars = 10 μm. 

Cercospora cercidis Nishikado is illegitimate, a new name, P. 

cercidicola is introduced for the species occurring on Cercis in 

Japan. Pseudocercospora cercidicola is morphologically very 

close to P. cercidis-chinensis but superficial hyphae with solitary 

conidiophores are not formed and the conidia are shorter. 

Pseudocercospora cercidis-chinensis H.D. Shin & U. 

Braun, Mycotaxon 74: 109. 2000. Fig. 26. 

Specimens examined: South Korea, Kyeongju, on Cercis chinensis, 26 Aug. 1998, 

H.D. Shin, holotype KUS-F 14914, isotype HAL; Suwon, C. chinensis, 2 Oct. 2007, 

H.D. Shin, epitype designated here CBS H-20857, culture ex-epitype CPC 14481 

= CBS 132109. 

Note: See P. cercidicola. 

Pseudocercospora chengtuensis (F.L. Tai) Deighton, 

Mycol. Pap. 140: 141. 1976. Fig. 27. 

Basionym: Cercospora chengtuensis F.L. Tai, Lloydia 11: 40. 1948. 

Specimens examined: China, Szechuan, Chengtu, Lycium chinense, Lee Ling No. 

126, 1943, holotype (not seen). South Korea, Dongducheon, Lycium chinense, 28 

Sep. 2003, H.D. Shin, CBS H-20858, culture CPC 10696–10698. 

Notes: The isolate identified here as P. chengtuensis appears to 

be identical to P. atromarginalis (also on Solanaceae) based on 

phylogenetic analysis and the two are morphologically similar. 

Study of of additional collections of both are needed to determine 

whether they are synonymous or distinct species. 

Pseudocercospora chionanthi-retusi Goh & W.H. Hsieh, 

in Hsieh & Goh, Cercospora and similar fungi from Taiwan: 

249. 1990. Fig. 28. 

80


Fig. 27. Pseudocercospora chengtuensis (CPC 10696–10698). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C–G. Fascicles with 

conidiophores and conidiogenous cells. H. Conidia. Scale bars = 10 μm. 

Fig. 28. Pseudocercospora chionanthi-retusi (CPC 14683). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C–H. Fascicles and solitary 

conidiophores with conidiogenous cells. I. Conidia. Scale bars = 10 μm. 

= Cercospora chionanthi-retusi Togashi & Katsuki, Sci. Rep. Yokohama Nat. 

Univ. Sect. II, 1: 1. 1952. 

≡ Pseudocercospora chionanthi-retusi (Togashi & Katsuki) Nishijima, 

C. Nakash. & Tak. Kobay., Mycoscience 40: 270. 1999 (nom. illeg.), 

homonym of P. chionanthi-retusi Goh & Hsieh, 1990. 

= Pseudocercospora chionanthicola C. Nakash. & Tak. Kobay., 

Mycoscience 43: 98. 2002. 

Specimen examined: South Korea, Osan, on Chionanthus retusus, 30 Oct. 2007, 

H.D. Shin, CBS H-20859, culture CPC 14683 = CBS 132110. 

Pseudocercospora chrysanthemicola (J.M. Yen) Deighton, 

Mycol. Pap. 140: 141. 1976. 

Basionym: Cercospora chrysanthemicola J.M. Yen, Rev. Mycol. 29: 

216. 1964. 

Specimen examined: South Korea, Seoul, on Chrysanthemum sp., 6 Sep. 2003, 

H.D. Shin, CPC 10633. 

Pseudocercospora contraria (Syd. & P. Syd.) Deighton, 

Mycol. Pap. 140: 30. 1976. Fig. 29. 

Basionym: Cercospora contraria Syd. & P. Syd., Ann. Mus. Congo, 

Bot., Ser. V, 3: 21. 1909. 

= Cercospora wildemanii Syd. & P. Syd., Ann. Mus. Congo, Bot., Ser. V, 3: 

21. 1909. 

= Mycosphaerella contraria Hansf., Proc. Linn. Soc. London 153: 22. 1941. 

Specimen examined: South Korea, Bukjeju, Jeolmul recreation forest, on 

Dioscorea quinqueloba, 2. Nov. 2007, H.D. Shin, CBS H-20861, CPC 14714 



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Crous et al. 

Fig. 29. Pseudocercospora contraria (CPC 14714). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C–E. Fascicles with conidiophores, and 

solitary loci on hyphae. F. Conidia. Scale bars = 10 μm. 

Notes: This fungus was first reported from Korea by Shin & Kim 

(2001). Conidial measurements (16–75 × 2.5–4.5 μm) are smaller 

than those of the type collected in the Democratic Republic of the 

Congo (20–120 × 5–8 μm, Chupp 1954), and the Korean material 

may eventually be shown to represent a distinct species. 

Pseudocercospora coriariae (Chupp) X.J. Liu & Y.L. Guo, 

Mycosystema 2: 232. 1989. 

Basionym: Cercospora coriariae Chupp, J. Dept. Agric. Puerto Rico 

14: 285. 1930. 

= Cercospora coriariae F.L. Tai, Lloydia 11: 43. 1948 (nom. illeg.), 

homonym of C. coriariae Chupp, 1930. 

Specimen examined: Japan, Tokyo, on Coriaria japonica, 10 Nov. 2007, I. Araki & 

M. Harada, MUMH 10942, culture MUCC 840. 

Pseudocercospora cornicola (Tracy & Earle) Y.L. Guo & 

X.J. Liu, Mycosystema 2: 232. 1989. 

Basionym: Cercospora cornicola Tracy & Earle, Bull. Torrey Bot. 

Club 23: 205. 1896. 

Specimen examined: Japan, Tokyo, Cornus alba var. sibirica, 7 Nov. 1998, C. 

Nakashima & E. Imaizumi, CNS-494, culture MUCC 909, MAFF 237773. 

Pseudocercospora corylopsidis (Togashi & Katsuki) C. 

Nakash. & Tak. Kobay., Mycoscience 40: 270. 1999. 

≡ Cercospora corylopsidis Togashi & Katsuki, Bot. Mag. (Tokyo) 65: 20. 

1952. 

= Cercospora hamamelidis auct.; sensu Togashi & Katsuki, Bot Mag. (Tokyo) 

65: 21. 1952, non (Peck) Ellis & Everh. 

Specimens examined: Japan, Kagoshima, on Corylopsis pauciflora, 26 Oct. 1949, 

S. Katsuki, holotype YNU, Isotype TNS-F-243824; Ibaraki, Tsukuba Botanical 

Garden, on C. pauciflora, Oct. 1996, T. Kobayashi; Ibaraki, on C. pauciflora, 9 Nov. 

1998, T. Kobayashi; Tokyo, Todori, on C. pauciflora, 12 Oct. 1979, M. Kusunoki, 

TFM:FPH-6152; Tokyo, Jindai Bot. Park, on C. spicata, 7 Nov. 1998, C. Nakashima 

& E. Imaizumi, epitype designated here TFM: FPH-8095, ex-epitype cultures 

MUCC 908, MAFF 237795; Saitama, isolated from C. pauciflora, Nov. 1995, 

MUCC1249, MAFF 237302; Kagoshima, 26 Oct. 1949, on Hamamelis japonica, 

S. Katsuki, SK2077; Shizuoka, 2 Nov. 1996, on H. japonica, T. Koboyashi & C. 

Nakashima, CNS-114, cultures MAFF 237632, MUCC 874. 

Notes: Isolate MUCC 874, which was isolated from Hamamelis 

japonica (Hamamelidaceae), appears to be phylogenetically 

identical to P. corylopsidis. Based on morphology, there is little 

difference between these specimens other than the presence or 

absence of external mycelium. 

Togashi & Katsuki (1952) reported a fungus on Hamamelis 

japonica as Cercospora hamamelidis (Peck) Ellis & Everh. based 

on a specimen collected in Kagoshima (SK2077). Recently, C. 

hamamelidis was transferred to the genus Passalora (Crous & 

Braun 2003). The Japanese specimens of C. hamamelidis are 

morphologically and phylogenetically identical to Pseudocercospora 

corylopsidis. We conclude that the fungus on Corylopsis and 

Hamamelis in Japan represents P. corylopsidis. In addition, a species 

of Pseudocercospora collected in Tokyo (TFM:FPH-4348, isolate 

MAFF 410032) was recognised as a distinct taxon on Corylopsis 

plants, based on its longer and narrower conidia, and DNA phylogeny. 

Pseudocercospora cotoneastri (Katsuki & Tak. Kobay.) 

Deighton, Trans. Brit. Mycol. Soc. 88: 389. 1987. 

Basionym: Cercospora cotoneastri Katsuki & Tak. Kobay. 

(cotoneasteris), Trans. Mycol. Soc. Japan 17: 276. 1976. 

Specimens examined: Japan, Tokyo, Asakawa Experimental Forest Station, on 

Cotoneaster dammeri, 13 Aug. 1974, T. Kobayashi, holotype TFM:FPH-4185, exholotype 

culture MAFF 410089; Tokyo, Tokyo Agric. Exp. Stn., on C. franchetii, 27 

Sep. 1978, T. Kobayashi, TFM:FPH-4924; Tokyo, Jindai Bot. Park, on C. horizontalis, 

4 Sep. 1975, H. Horie, TFM: FPH-4417; Tokyo, on C. horizontalis, 23 Oct. 1975, 

K. Sasaki, TFM:FPH-4798; Tokyo, culture isolated from Cotoneater sp., 1977, H. 

Horie, culture MAFF 305633; Fukuoka, Kitakyushu, on C. horizontalis,4 Oct. 1975, 

S. Ogawa (TFM:FPH-4401); Shizuoka, Hamamatsu, on C. salicifolius, 1 Nov. 1996, 

T. Kobayashi & C. Nakashima, CNS-126, culture MUCC 876, MAFF 237629. 

Note: Three isolates including the ex-holotype, MAFF 410089, 

305633 and 237629, were identical based on ACT gene sequence 

data (data not shown). 

Pseudocercospora crispans G.C. Hunter & Crous, sp. 


Etymology: Name reflects the characteristic curling or undulate 

nature of the conidia produced by this fungus. 

Leaf spots amphigenous, angular to irregular, predominantly 

occurring next to or close to the mid-rib, 2–15 mm diam, pale 

82


Fig. 30. Pseudocercospora crispans (CPC 14883). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C–F. Fascicles with conidiophores and 

conidiogenous cells. G, H. Conidia. Scale bars = 10 μm. 

brown on the upper side of the leaf, and pale to darker brown 

on the bottom side of the lesion, surrounded by a raised, dark 

brown border with a diffuse red pigment emanating away from 

the border; single, discrete lesions may coalesce to form larger 

lesions. Mycelium smooth, septate, guttulate, thick-walled, 

branched, internal and external, pale brown, 2–4 mm wide. 

Caespituli amphigenous, sparsely scattered over lesion, floccose, 

whitish. Stromata hypophyllous, brown, well-developed, immersed, 

globular to irregular, 40–120 mm diam. Conidiophores brown at 

base, becoming paler toward apex, arising from cells of brown 

stroma; arranged in loose fascicles, smooth, thick-walled, guttulate, 

unbranched, straight to curved, 0–4-septate, straight to geniculate– 

sinuous, (14–)17–31(–42) × (2–)3–4(–5) μm. Conidiogenous cells 

terminal, unbranched, smooth, guttulate, pale brown, straight to 

geniculate to geniculate–sinuous, proliferating sympodially and 

percurrently, tapering toward apex; apex obtuse to truncate, (8–)9– 

15(–19) × (2–)3(–4) μm. Conidia solitary, smooth, guttulate, curved 

to undulate, pale brown, 3–9-septate, apex acute to subacute, base 

truncate, (40–)65–96(–102) × (2–)3(–4) μm; hila unthickened, not 

darkened. 

Culture characteristics: Colonies on MEA reaching 54 mm diam 

after 30 d at 24 °C. Colonies circular, flat to slightly convex, with 

a feathery margin and profuse aerial mycelium; lavender-grey to 

glaucous-grey (surface) and olivaceous-grey (reverse). 

Specimen examined: South Africa, Western Cape Province, Knysna, on leaves 

of Eucalyptus sp., Jan. 2008, P.W. Crous, holotype CBS H-20392, culture ex-type 

CPC 14883 = CBS 125999. 

Notes: Pseudocercospora crispans is phylogenetically distinct 

from other taxa described from Eucalyptus (Crous et al. 1989, 

Crous & Alfenas 1995, Crous & Wingfield 1997, Crous 1998, 

Braun & Dick 2002, Hunter et al. 2006a), and can be distinguished 

morphologically by its prominently curled conidia. 

Pseudocercospora crocea Crous, U. Braun, G.C. Hunter & 

H.D. Shin, sp. nov. MycoBank MB564831. Fig. 31. 

Etymology: Name reflects the typical diffuse yellow border 

surrounding leaf lesions caused by this fungus. 

Leaf spots distinct, scattered and at the leaf margin, pale brown 

to brown, circular to irregular, 2–5 mm diam, indefinite border, 

with a pale yellow diffuse halo. Mycelium, internal and external, 

subhyaline, septate, branched, smooth, 2–5 mm wide. Caespituli 

amphigenous, grey, scattered over the lesion surface, arachnoid. 

Stromata well-developed, 40–100 mm diam, subimmersed, 

globular, dark brown. Conidiophores fasciculate, brown, becoming 

paler toward the apex, 0–1-septate, smooth, unbranched, straight 

to curved, apex truncate to subtruncate, 0–1-septate, (14–)17–24(– 

32) × (3–)4(–5) μm. Conidiogenous cells terminal, unbranched, pale 


83

Crous et al. 

Fig. 31. Pseudocercospora crocea (CPC 11668). A, B. Leaf spots on upper and lower leaf surface. C. Close-up of leaf spot with fruiting. D, E. Fascicles with conidiophores and 

conidiogenous cells. F–I. Conidia. Scale bars = 10 μm. 

brown, smooth to slightly verruculose, proliferating percurrently, (9–) 

13–18(–21) × (3–)4(–5) μm. Conidia solitary, 4–10-septate, straight 

to curved, sparsely guttulate, narrowly obclavate, apex subobtuse, 

base obconically truncate to long obconically truncate, smooth, 

subhyaline, (67–)79–94(–104) × (3–)4(–5)μm, hila unthickened not 

darkened. 


after 30 d at 24 °C. Colonies circular with feathery margin, flat 

to slightly convex, some folding occurs, with a darker radial ring 

toward the colony margin, aerial mycelium medium; iron-grey to 

olivaceous-grey (surface) and iron-grey (reverse). 

Specimen examined: South Korea, Suwon, on leaves of Pilea hamaoi (≡ P. pumila 

var. hamaoi), 5 Nov. 2004, H.D. Shin, holotype CBS H-20387, isotype HAL 1860 F, 

cultures ex-type CPC 11668 = CBS 126004. 

Notes: Singh et al. (1996) provide an account of the 

Pseudocercospora spp. present on members of Urticaceae. 

Of these, P. crocea is most similar to P. pileae as it also has a 

well-developed stroma. Pseudocercospora pileae is distinct from 

P. crocea, which lacks stromata and has conidiophores that are 

consistently solitary, arising from superficial hyphae. 

Pseudocercospora cydoniae (Ellis & Everh.) Y.L. Guo & 

X.J. Liu, Mycosystema 5: 103. 1992. Fig. 32. 

Basionym: Cercospora cydoniae Ellis & Everh., J. Mycol. 8: 72. 

1902. 

≡ Cercosporina cydoniae (Ellis & Everh.) Sacc., Syll. Fung. 25: 915. 1931. 

≡ Pseudocercospora cydoniae (Ellis & Everh.) U. Braun & H.D. Shin, 

Mycotaxon 49: 356. 1993. 

Specimens examined: South Korea, Seoul, on Chaenomeles speciosa (= C. 

lagenaria), 17 Sep. 2003, H.D. Shin, cultures CPC 10678 = CBS 131923; Jeonju, 

C. sinensis, 15 Oct. 2003, H.D. Shin, CBS H-20863. 

Pseudocercospora dovyalidis (Chupp & Doidge) Deighton, 

Mycol. Pap. 140: 143. 1976. Fig. 33. 

Basionym: Cercospora dovyalidis Chupp & Doidge, Bothalia 4: 

885. 1948. 

≡ Pseudocercosporella dovyalidis (Chupp & Doidge) B. Sutton, Mycol. 

Pap. 138: 99. 1975. 

Leaf spots amphigenous, distinct, 1–3 lesions per leaf, scattered 

over the leaf, 3–10 mm diam, pale brown surrounded by a 

dark brown to black border. Mycelium internal, consisting of 

pale brown, septate, smooth, 2–6 μm diam hyphae. Caespituli 

hypophyllous, evenly distributed over the leaf spot, floccose 

to punctiform, olivaceous to black. Stromata well-developed, 

subimmersed to erumpent, globular, dark brown, 40–100 mm 

diam. Conidiophores fasciculate, emerging from the upper cells 

of stromata, brown, becoming paler toward the apex, smooth, 

0–2-septate, straight to variously curved, guttulate, apex 

84


Fig. 32. Pseudocercospora cydoniae (CPC 10678). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C. Fascicle with conidiophores and 

conidiogenous cells. D, E. Conidiogenous cells. F. Conidia. Scale bars = 10 μm. 

Fig. 33. Pseudocercospora dovyalidis (CPC 13771–13773). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with conidiophores 

and conidiogenous cells. E, F. Conidiogenous cells. G–K. Conidia. Scale bars = 10 μm. 


85

Crous et al. 

Fig. 34. Pseudocercospora flavomarginata (CPC 14142). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C. Fascicle with conidiophores 

and conidiogenous cells. D–F. Conidiogenous cells. G–J. Conidia. Scale bars = 10 μm. 

rounded, conidiophores rarely branched below, (12–)13–22(–34) 

× (3–)3–5(–6) μm. Conidiogenous cells terminal, pale brown, 

smooth, guttulate, proliferating percurrently, (4–)6–12(–15) × (2–) 

3–4(–5) μm. Conidia solitary, pale brown or subhyaline, smooth, 

distinctively guttulate, 1–10-septate, thick-walled, straight to 

curved, broadly filliform to cylindrical, apex rounded to subacute, 

base long obconically truncate, (20–)30–70(–84) × (3–)3–5(–6) 

μm; hila neither thickened nor darkened. 


after 30 d at 24 °C. Colonies circular with a smooth margin, 

either flat with excessive folding into the media or convex, aerial 

mycelium moderate, margin of colony darker than colony interior; 

greenish glaucous to olivaceous-grey (surface) and olivaceousgrey 

(reverse). 

Specimens examined: South Africa, Gauteng, Pretoria, Groenkloof, on Dovyalis 

zeyheri, 18 Feb. 1914, E.M. Doidge, holotype PREM 7398; Gauteng, Walter 

Susulu Botanical Garden, on leaves of D. zeyheri, 2 Mar. 2007, P.W. Crous, epitype 

designated here CBS H-20389, culture ex-type CPC 13771 = CBS 126002. 

Pseudocercospora eucalyptorum Crous, M.J. Wingf., 

Marasas & B. Sutton, Mycol. Res. 93: 394. 1989. 

= Pseudocercospora pseudoeucalyptorum Crous, Stud. Mycol. 50: 210. 2004. 

Specimens examined: South Africa, Western Cape Province, Stellenbosch, 

Stellenbosch Mountain, on leaves of E. nitens, 21 Dec. 1987, P.W. Crous, holotype 

of P. eucalyptorum PREM 49112, cultures ex type CPC16 = CBS 110777. Spain, 

Pontevedra, Lourizán, Areeiro, on leaves of E. globulus, 2003, J.P. Mansilla, 

holotype of P. pseudoeucalyptorum CBS H-9893, culture ex-type CPC 10390 


Note: Pseudocercospora pseudoeucalyptorum is reduced to 

synonymy with P. eucalyptorum on the basis of the phylogeny 

obtained here and similarity in pigmentation (Crous et al. 2004c). 

Pseudocercospora exosporioides (Bubák) B. Sutton & 

Hodges, Mycologia 82: 320. 1990. 

Basionym: Cercospora exosporioides Bubák, Ann. Mycol. 13: 33. 

1915. 

Specimen examined: Japan, Ibaraki, on Sequoia sempervirens, 11 Sep. 1998, T. 

Kobayashi, CNS-448, cultures MUCC 893, MAFF 237788. 

Pseudocercospora flavomarginata G.C. Hunter, Crous & 

M.J. Wingf., Fungal Diversity 22: 80. 2006. Fig. 34. 

Specimens examined: Thailand, Chang Gao Province near Pratchinburi, on 

leaves of Eucalyptus camaldulensis, 2004, M.J. Wingfield, holotype PREM 58952, 

cultures ex-type CBS 118841, 118823, 118824; Chachoengsao Province, on leaves 

of E. camaldulensis, 2001, W. Himaman, CBS H-20388, culture CPC 13492–13494. 

86


Fig. 35. Pseudocercospora fukuokaensis (CPC 14689). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with conidiophores 

and conidiogenous cells. E. Conidia. Scale bars = 10 μm. 

China, on leaves of Eucalyptus sp., 2003, X. Zhou, CBS H-20390, culture ex-type 

CPC 14142 = CBS 126001. 

Notes: Pseudocercospora flavomarginata was described as the 

causal agent of a prominent leaf spot disease of E. camaldulensis 

in Thailand (Hunter et al. 2006a). Based on this study it appears 

that it is present also on this host in China. 

Pseudocercospora fukuokaensis (Chupp) X.J. Liu & Y.L. 

Guo, Mycosystema 5: 103. 1992. Fig. 35. 

Basionym: Cercospora fukuokaensis Chupp, Sci. Rep. Yokahama 

Natl. Univ., Sect. II, Biol. Sci. 1: 2. 1952. 

Specimens examined: Japan, Fukuoka, Futsukaichi-machi, on Styrax japonicus, 

5 Sep. 1951, S. Katsuki, holotype TNS-F243813; Ibaraki, on S. japonicus, 11 

Sep. 1998, T. Kobayashi & C. Nakashima, epitype designated here TFM: FPH- 

8096, ex-epitype cultures MUCC 887, MAFF 237768; Ibaraki, Ibaraki Nat. Mus., 

on S. japonicus, 10 Sep. 1998, T. & Y. Kobayashi; Fukuoka, Fukuoka For. Exp. 

Stn., on S. japonicus, 30 Jul. 1975, S. Ogawa (TFM: FPH-4356); Kaogshima, 

Tanegashima Is., on S. japonicus, 18 Oct. 1997, T. Kobayashi & C. Nakashima 

(culture: MAFF238203); Kagoshima, Tokunoshima Is., on S. japonicus, 8 Nov. 

1993, T. Kobayashi & T. Hosoya (Culture: MAFF236995); Okinawa, Kunigami, on 

S. japonicus, 18 Nov. 1999, T. Kobayashi & C. Nakashima; Fukuoka, Fukuoka For. 

Exp. Stn., on S. obassia, 14 Sep. 1978, S. Ogawa (TFM: FPH -4941); Fukuoka, 

on S. grandiflora (= S. japonicus var. kotoensis), Oct. 2001, T. Kobayashi (MAFF 

238480); Yamaguchi, on S. japonicus, Dec. 1996, T. Kobayashi (MAFF 237634); 

Saitama, on S. japonicus, Sep. 2002, T. Kobayashi & Y.Ono (MAFF 239411). South 

Korea, Osan, S. japonicus, 30 Oct. 2007, H.D. Shin, culture CPC 14689 = CBS 

132111. 

Notes: DNA sequence data for different isolates from Styrax 

japonica collected in Japan are identical, and distinct from the strain 

collected in Korea, suggesting that the Korean material represents 

a different taxon. 

Pseudocercospora fuligena (Roldan) Deighton, Mycol. 

Pap. 140: 144. 1976. 

Basionym: Cercospora fuligena Roldan, Philipp. J. Sci. 66: 8. 1938. 

Holotype: Philippines, Luzon, Laguna, College of Agriculture 

Campus, on Solanum lycopersicum (≡ Lycopersicon esculentum), 

E.F. Roldan No 32, holotype (not seen). 

Specimens examined: Thailand, on Solanum lycopersicum (variety FMMT260), 28 

Aug. 2005, Z. Mersha, CBS H-20864, culture CPC 12296 = CBS 132017. Japan, 

Mie, on Lycopersicon esculentum, 6 Feb. 2007, C. Nakashima, MUCC 533. 

Notes: DNA sequence data (ITS and EF-1α) for 40 Japanese 

isolates revealed variation in only one position (data not shown) and 

the culture from Thailand is very similar genetically. The collections 

of P. fuligena treated in this study are also morphologically similar 

to the description of the holotype specimen, which was collected 

in the Philippines. Chupp (1954) did not see the holotype, nor did 

Deighton (1976) refer to it. Fresh collections from the type location 

are needed to resolve this apparent species complex. 

Pseudocercospora glauca (Syd.) Y.L. Guo & X.J. Liu, Acta 

Mycol. Sin. 11: 132. 1992. Fig. 36. 

Basionym: Cercospora glauca Syd., Ann. Mycol. 27: 432. 1929. 

Specimen examined: South Korea, Wando, Wando arboretum, on Albizzia 

julibrissin, 9 Nov. 2002, H.D. Shin, CBS H-20865, culture CPC 10062 = CBS 

131884. 

Pseudocercospora guianensis (F. Stevens & Solheim) 

Deighton, Mycol. Pap. 140: 145. 1976. 

Basionym: Cercospora guianensis F. Stevens & Solheim, Mycologia 

23: 375. 1931. 

Specimen examined: Japan, Tateyama, Chiba, on Lantana camara, 4 June 1997, 

C. Nakashima CNS-162, cultures MUCC 879, MAFF 238239. 

Pseudocercospora haiweiensis Crous & X. Zhou, sp. nov. 


Etymology: Name is derived from Hai Wei, China, where this 

fungus was collected. 

Leaf spots amphigenous, irregular to subcircular or angular, 2–4 

mm diam, brown, with raised border, and at times with a red-purple 

margin. Mycelium internal, subhyaline, consisting of septate, 

branched, smooth, 2–3 μm diam hyphae. Caespituli fasciculate to 

sporodochial, amphigenous, breaking through epidermis, appearing 


87

Crous et al. 

Fig. 36. Pseudocercospora glauca (CPC 10062). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C. Fascicle with conidiophores and 

conidiogenous cells. D. Conidiophores. E, F. Conidia. Scale bars = 10 μm. 

Fig. 37. Pseudocercospora haiweiensis (CPC 14084). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C. Fascicle with conidiophores and 

conidiogenous cells. D–F. Conidiophores. G. Conidia. Scale bars = 10 μm. 

almost acervular, grey-brown on leaves, up to 90 μm wide and 50 

μm high. Conidiophores aggregated in dense fascicles arising from 

the upper cells of a brown stroma up to 60 μm wide and 30 μm high; 

conidiophores brown, smooth to finely verruculose, 0–2-septate, 

subcylindrical, straight to variously curved or geniculate-sinuous, 

unbranched, 10–25 × 3–4 μm. Conidiogenous cells terminal, 

unbranched, brown, subcylindrical, smooth to finely verruculose, 

tapering to flat-tipped apical loci, proliferating sympodially, rarely 

percurrently near apex, 10–15 × 2.5–3.5 μm. Conidia solitary, 

brown, finely verruculose, guttulate, subcylindrical, apex obtuse, 

base obconically subtruncate to truncate, straight to gently curved, 

3(–5)-septate, (25–)30–40(–45) × 3(–4) μm; hila unthickened, 

neither darkened nor refractive, 1.5 μm wide. 


MEA; surface folded, erumpent, spreading, with moderate aerial 

mycelium, and smooth, lobate margins. Surface olivaceous-grey 

with patches of pale olivaceous-grey; reverse olivaceous-grey. 

Colonies reaching 12 mm diam. 

Specimen examined: China, Hai Wei, on leaves of Eucalyptus sp. (APP 21), 3 June 

2007, X. Zhou, holotype CBS H-20866, culture ex-type CPC 14084 = CBS 131584. 

Notes: A combination of relatively short conidia (1–3-septate, 

25–45 × 3–4 μm) that are subcylindrical in shape, the absence 

of superficial mycelium, and dense fascicles with well-developed 

stromata, distinguish this new species on Eucalyptus from other 

taxa known from this host (Crous 1998, Braun & Dick 2002). 

Pseudocercospora hakeae (U. Braun & Crous) U. Braun & 

Crous, comb. et stat. nov. MycoBank MB564833. 

Basionym: Cercostigmina protearum var. hakeae U. Braun & 

Crous, Sydowia 46: 206. 1994. 

≡ Pseudocercospora protearum var. hakeae (U. Braun & Crous) U. Braun 

& Crous, Mycol. Progr. 1: 22. 2002. 

88


Fig. 38. Pseudocercospora humulicola (CPC 11358). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C–E. Fascicles with conidiophores and 

conidiogenous cells. F. Conidia. Scale bars = 10 μm. 

Specimens examined: South Africa, Northern Province, Louis Trichardt, Hangklip 

Forest Station, on leaves of Hakea salicifolia (= H. saligna), Apr. 1988, C. Roux, 

holotype PREM 51117. Australia, New South Wales, Mount Annan Botanic 

Gardens, on leaves of Grevillea sp., Aug. 1999, P.W. Crous & B. Summerell, JT 

926, DAR 74861, CPC 2968; Mount Tomah Botanic Gardens, on leaves of Grevillea 

sp., Aug. 1999, P.W. Crous & B. Summerell, JT 873, DAR 74862, CPC 3145 = CBS 

112226. 

Note: No culture from Hakea is presently available, and thus 

the position of this taxon on Hakea and Grevillea has yet to be 

confirmed based on DNA sequence comparisons. 

Pseudocercospora humuli (Hori) Y.L. Guo & X.J. Liu, Acta 

Mycol. Sin., Suppl. 1: 345. (1986) 1987. 

Basionym: Cercospora humuli Hori, in S. Takimoto, Trans. Agric. 

Assoc. Chosen 13(12): 34. 1918. 

≡ Cercospora humuli Hori, in Salmon & Wormald. J. Bot. (London) 61: 

135. 1923. 

= Cercospora humuli-japonici Sawada, Taiwan Agric. Rev. 38: 697. 1942 

(nom. inval.). 

≡ Pseudocercospora humuli-japonici Sawada ex Goh & W.H. Hsieh, in 

Hsieh & Goh, Cercospora and similar fungi from Taiwan: 239. 1990. 

Specimens examined: Japan, Tokyo, Nishigahara, on Humulus scandens, 28 Sep. 

1915, S. Hori, holotype NIAES herbarium C-487; Wakayama, on H. lupulus var. 

lupulus, 30 Oct. 2007, C. Nakashima & I. Araki, epitype designated here TFM: 

FPH-8097, ex-epitype culture MUCC 742. 

Pseudocercospora humulicola Crous, U. Braun & H.D. 


Etymology: Name derived from Humulus, the plant on which it was 

collected. 

Leaf spots amphigenous, irregular to angular, 0.5–1.5 mm diam, 

brown, with raised border and wide chlorotic halo. Mycelium 

internal, subhyaline, consisting of septate, branched, smooth, 

2–3 μm diam hyphae. Caespituli fasciculate to sporodochial, 

amphigenous, predominantly epiphyllous, pale brown on leaves, 

up to 90 μm wide and 200 μm high. Conidiophores aggregated 

in dense fascicles arising from the upper cells of a brown stroma 

up to 80 μm wide and 30 μm high; conidiophores pale brown, 

smooth, 2–5-septate, subcylindrical, straight to variously curved or 

geniculate-sinuous, unbranched, 40–90 × 3–4 μm. Conidiogenous 

cells terminal, unbranched, subhyaline to pale brown, subcylindrical, 

smooth, tapering to flat-tipped apical conidiogenous loci, 2 μm 

diam, proliferating sympodially, 10–30 × 3–4 μm. Conidia solitary, 

subhyaline, smooth, finely granular, subcylindrical, apex obtuse, 

base truncate, straight to gently curved, 3–12-septate, (70–)80– 

95(–120) × 2.5(–3) μm; hila unthickened, neither darkened nor 

refractive, 2–3 μm wide. 



mycelium, and smooth, lobate margins. Surface pale olivaceousgrey; 

reverse olivaceous-grey. Colonies reaching 10 mm diam. 

Specimens examined: South Korea, Hongchon, on leaves of Humulus scandens, 

9 Jul. 2004, H.D. Shin, holotype CBS H-20867, culture ex-type CPC 11358 = CBS 

131585; Chuncheon, on H. scandens, 11 Oct. 2002, H.D. Shin, CBS H-20868, 

culture CPC 10049 = CBS 131883; Cheongju, on H. scandens, 4 June 2004, H.D. 

Shin, CBS H-20869, culture CPC 10002. 

Notes: Pseudocercospora humulicola is very similar to P. humuli, 

originally described from Japan, but it is distinct based on DNA 

sequence comparisons. In P. humuli conidia are obclavatecylindrical, 

35–120 × 2.5–4 μm (Chupp 1954), while conidia of P. 

humulicola are subcylindrical, and on average longer than 80 μm. 

Furthermore, P. humuli has shorter conidiophores (10–55 μm long, 

0–2-septate) than those of P. humulicola, which are 2–5-septate, 

and 40–90 μm long. 

Pseudocercospora jussiaeae (G.F. Atk.) Deighton, Mycol. 

Pap. 140: 146. 1976. Fig.39. 

Basionym: Cercospora jussiaeae G.F. Atk., J. Elisha Mitchell Sci. 

Soc. 8: 50. 1892. 

= Cercospora ludwigiae G.F Atk., J. Elisha Mitchell Sci. Soc. 8: 58. 1892. 

Specimen examined: South Korea, Hongcheon, on Ludwigia prostrata, 9 Oct. 

2007, H.D. Shin, KUS-F22981, CBS H-20870, culture CPC 14625 = CBS 132117. 


89

Crous et al. 

Fig. 39. Pseudocercospora jussiaeae (CPC 14625). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with conidiophores and 

conidiogenous cells. E. Conidia. Scale bars = 10 μm. 

Fig. 40. Pseudocercospora kaki (CPC 10837–10839). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with conidiophores 

and conidiogenous cells. E, F. Conidiogenous cells. G. Conidia. Scale bars = 10 μm. 

Pseudocercospora kaki Goh & W.H. Hsieh, in Hsieh & Goh, 

Cercospora and similar fungi from Taiwan: 109. 1990. Fig. 

40. 

Specimens examined: Japan, Toyama, Kureha, on Diospyros kaki, 25 Sep. 1998, 

T. Kobayashi & E. Imaizumi, CNS-472, culture MAFF 238214; Chiba, on D. kaki, 

18 Sep. 1998, S. Uematsu & C. Nakashima, CNS-464, cultures MUCC 900, MAFF 

238238; Chiba, on D. kaki, Nov. 1993, T. Kobayashi, cultures MAFF 237013. South 

Korea, Gongju, on D. lotus, 28 Oct. 2003, H.D. Shin, CBS H-20871, cultures CPC 

10837–10839. 

Additional isolates examined (representing a different lineage): Japan, Kagoshima, 

Oshima Is., on D. kaki, 11 Nov. 1993, T. Kobayashi, CNS-993, culture MAFF 236999; 

Chiba, on D. kaki, Oct. 1991, T. Kobayashi, culture MAFF 235880. 

Notes: The type specimen of this species is from Taiwan but 

the type was not cultured or sequenced. It may be synonymous 

with Cercospora kaki, which is based on material from the USA. 

The Japanese material studied here is different from the Korean 

material based on DNA sequence data. Actin sequences generated 

for additional Japanese isolates resolved two different lineages, 

one of which may be attributed to Cercospora kakivora, but this 

can only be resolved once fresh collections from Taiwan and the 

USA have been obtained. 

Pseudocercospora kiggelariae (Syd.) Crous & U. Braun, 

Sydowia 46: 215. 1994. 

Basionym: Cercospora kiggelariae Syd., Ann. Mycol. 22: 434. 1924. 

Holotype: South Africa, Western Cape Province, Stellenbosch, on 

leaves of Kiggelaria africana, May 1924, C.K. Brain No 1449 (not 

preserved). 

Specimens examined: South Africa, Gauteng, Walter Susulu Botanical Garden, 

on leaves of K. africana, Jan. 2005, W. Gams, neotype designated here CBS 

H-20872, cultures ex-neotype CPC 11853 = CBS 132016; Western Cape Province, 

Hermanus, Fernkloof Botanical Garden, S34º23’52.1” E19º15’58.5”, K. africana, 2 

May 2010, P.W. Crous, CBS H-20873, CPC 18286, 18287. 

90


Fig. 41. Pseudocercospora lythracearum (CPC 10707). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with conidiophores 

and conidiogenous cells. E. Conidiophore with conidiogenous cells. F. Conidia. Scale bars = 10 μm. 

Pseudocercospora latens (Ellis & Everh.) Y.L. Guo & X.J. 

Liu, Mycosystema 2: 236. 1989. 

Basionym: Cercospora latens Ellis & Everh., J. Mycol. 4: 3. 1888. 

≡ Pseudocercospora latens (Ellis & Everh.) U. Braun, Trudy Bot. Inst. im. 

V.L. Komarova 20: 67. 1997 (comb. superfl.). 

Specimen examined: Japan, Okinawa, on Lespedeza wilfordii (= L. thunbergii 

subsp. formosa), 18 Nov. 2007, C. Nakashima & T. Akashi, MUMH 10815, culture 

MUCC 763. 

Pseudocercospora leucadendri (Cooke) U. Braun & Crous, 

comb. et stat. nov. MycoBank MB564835. 

Basionym: Cercospora protearum var. leucadendri Cooke, 

Grevillea 12: 39. 1883. 

≡ Stigmina protearum var. leucadendri (Cooke) M.B. Ellis, Mycol. Pap. 

131: 7. 1972. 

≡ Cercostigmina protearum var. leucadendri (Cooke) U. Braun & Crous, 

in Crous & Braun, Sydowia 46: 206. 1994. 

≡ Pseudocercospora protearum var. leucadendri (Cooke) U. Braun & 

Crous, Mycol. Progr. 1: 22. 2002. 

= Passalora protearum Kalchbr. & Cooke, Grevillea 19: 6. 1890. 

Specimen examined: South Africa, Western Cape Province, Stellenbosch, Devon 

Valley, Protea Heights, on Leucadendron sp., 3 Apr. 1998, S. Denman & P.W. Crous, 

specimen JT-178, culture CPC 1869 (no longer viable). 

Note: Pseudocercospora protearum has three varieties on 

Proteaceae, viz. protearum, leucadendri and hakeae (Braun & Hill 

2002), that should be recognised as distinct species (Crous et al. 

2004a) as shown here (Fig. 5). 

Pseudocercospora lonicericola (W. Yamam.) Deighton, 

Mycol. Pap. 140: 146. 1976. 

Basionym: Cercospora lonicericola W. Yamam. J. Soc. Trop. Agric. 

6: 604. 1934. 

Holotype: Taiwan, Taihoku, on Lonicera japonica var. sempervillosa, 

3 Nov. 1933, W. Yamamoto (holotype could not be located, and is 

probably lost). 

Specimens examined: Japan, Tokyo, Jindai Bot. Park, on L. japonica, 21 Oct. 1976, 

T. Kobayashi, TFM: FPH-4479; Chiba, Matsudo, on L. japonica, 14 Sep. 1951, E. 

Kurosawa, SK -2207; Fukuoka, Yame, on L. japonica, 29 Nov. 1949, S. Katsuki, 

SK -2206; Kagoshima, Yaku Is., on L. japonica, 29 Dec. 1952, S. Katsuki, SK -392; 

Ibaraki, L. gracilipes var. glabra, 11 Sep. 1998, T. Kobayashi, neotype designated 

here TFM: FPH-8098, ex-neotype cultures MUCC 889, MAFF 237785. 

Pseudocercospora lyoniae (Katsuki & Tak. Kobay.) 


Basionym: Cercospora lyoniae Katsuki & Tak. Kobay., Trans. 

Mycol. Soc. Japan 16: 3. 1975. 

Specimens examined: Japan, Tokyo, Asakawa Experimental Forest, Government 

Forest Experimental Station, on Lyonia ovalifolia var. elliptica, 21 Sep. 1973, H. 

Horie, holotype TFM: FPH-3999; Tokyo, Jindai Bot. Garden, on L. ovalifolia var. 

elliptical, 25 Sep. 1974, T. Kobayashi, TFM: FPH -4202; Tokyo, Jindai Bot. Garden, 

on L. ovalifolia var. elliptica, 7 Nov. 1998, C. Nakashima & E. Imaizumi, epitype 

designated here TFM: FPH-8100, ex-epitype cultures MUCC 910, MAFF 237775. 

Pseudocercospora lythracearum (Heald & F.A. Wolf) X.J. 

Liu & Y.L. Guo, Acta Mycol. Sin. 11: 294. 1992. Fig. 41. 

Basionym: Cercospora lythracearum Heald & F.A. Wolf, Mycologia 

3: 18. 1911. 

≡ Cercosporina lythracearum (Heald & F.A. Wolf) Sacc., Syll. Fung. 25: 

909. 1931. 

= Cercospora lagerstroemiae Syd. & P. Syd., Ann. Mycol. 12: 203. 1914. 

= Cercospora lagerstroemiae-subcostatae Sawada, Taiwan Agric. Res. Inst. 

Rept. 51: 129. 1931. 

≡ Pseudocercospora lagerstroemiae-subcostatae (Sawada) Goh & W.H. 

Hsieh, in Hsieh & Goh, Cercospora and similar fungi from Taiwan: 212. 

1990. 

= Cercospora lagerstroemiicola Sawada, Taiwan Agric. Res. Inst. Rept. 85: 

112. 1943 (nom. inval.). 

Specimens examined: Japan, Ibaraki, on Lagerstroemia indica, 11 Sep. 1998, T. 

Kobayashi, CNS-444, cultures MUCC 890, MAFF 237786; Kanagawa, isolated 

from L. subcostata, collection date unknown, T. Kobayashi, MAFF 410017; Ibaraki, 

isolated from L. subcostata, Oct. 1994, T. Nishijima, MAFF 237185; Chiba, isolated 

from L. subcostata, Oct. 1993, T. Kobayashi, MAFF 236964. South Korea, Jinju, L. 

indica, 15 Oct. 2003, H.D. Shin, CBS H-20874, KUS-F 19899, culture CPC 10707 


Notes: The material collected from Korea is genetically similar to 

that from Japan (Fig. 5). However, fresh collections from the USA 

are required to determine if the Asian material is the same as 

that from the USA. The synonyms cited by Chupp (1954) could 

represent different species. 


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Crous et al. 

Fig. 42. Pseudocercospora lythri (CPC 14588). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with conidiophores and 

conidiogenous cells. E. Conidia. Scale bars = 10 μm. 

Pseudocercospora lythri H.D. Shin & U. Braun, Mycotaxon 

74: 111. 2000. Fig. 42. 

Specimens examined: Japan, Tokyo, on Lythrum salicaria (incl. L. anceps) 10 

Nov. 2007, I. Araki & M. Harada, MUMH 11104, culture MUCC865. South Korea, 

Chuncheon, on L. salicaria, 21 Sep. 1991, H.D. Shin, holotype KUS-F 11109; 

Yangku, on L. salicaria, 28 Sep. 2007, H.D. Shin, epitype designated here CBS 

H-20875, culture ex-epitype CPC 14588 = CBS 132115. 

Pseudocercospora marginalis G.C. Hunter, Crous, U. 

Braun & H.D. Shin, sp. nov. MycoBank MB564836. Fig. 43. 

Etymology: Margo, marginalis, referring to border or margin; 

indicating leaf spots that extend along the leaf margin. 

Leaf spots distinct, 2–5 mm diam, also predominantly forming 

larger blotches extending along the length of the leaf margin, 

brown, irregular; border indefinite. Mycelium internal and external, 

septate, smooth, subhyaline, branched, 2–4 μm wide. Caespituli 

epiphyllous, aggregated along leaf veins, floccose, olivaceous, 

emerging from stomata. Stromata well-developed, subimmersed 

to erumpent, globular to elongated, brown, 20–75 μm diam. 

Conidiophores fasciculate, pale brown to brown, straight to curved 

to undulate, cylindrical, unbranched, apex rounded to subtruncate, 

smooth, finely guttulate, 0–4-septate, (15–)18–31(–41) × (3–)4(–5) 

μm. Conidiogenous cells terminal, unbranched, smooth, finely 

guttulate, pale brown, straight to curved, cylindrical, apex rounded 

to subtruncate, proliferating sympodially or percurrently, (5–)8–11(– 

14) × 3(–4) μm. Conidia solitary, smooth, cylindrical to narrowly 

obclavate, guttulate, thick-walled, straight to curved, pale brown 

to pale olivaceous, apex rounded to obtuse, base obconic to long 

obconically truncate, 1–7-septate, (19–)30–48(–58) × (3–)4(–5) 

μm; hila neither thickened nor darkened. 


on MEA; erumpent, spreading, with moderate aerial mycelium, 

and smooth, even margins. Surface pale olivaceous-grey; reverse 


Specimen examined: South Korea, Jeju, Halla arboretum, on leaves of Fraxinus 

rhynchophylla (≡ F. chinensis subsp. rhynchophylla), 29 Oct. 2005, H.D. Shin, holotype 

CBS H-20397, culture ex-type CPC 12497 = CBS 131582, CPC 12498, 12499. 

Specimens examined of P. fraxinites: South Korea, Jinju, on Fontanesia 

phillyreoides, 15 Oct. 2003, H.D. Shin, CBS H-20876, cultures CPC 10743–10745. 

Japan, Ibaraki, on Fraxinus excelsior, 11 Sep. 1998, T. Kobayashi, CNS-445, 

cultures MUCC 891, MAFF 237787. 

Notes: Although similar to P. fraxinites (conidia 20–60 × 1.5–3 

μm; Chupp 1954) (Fig. 44), conidia of P. marginalis are wider 

and cluster apart from isolates of P. fraxinites on Fontanesia from 

Korea (CPC 10743–10745) and Fraxinus from Japan (MUCC 891). 

Pseudocercospora fraxinites was originally described from Fraxinus 

in the USA. Morphological and molecular characterisation of new 

collections and cultures from this host in the USA are needed to 

clarify the limits of P. fraxinites and P. marginalis. 

Pseudocercospora melicyti U. Braun & C.F. Hill, Australas. 

Pl. Pathol. 33: 489. 2004. 

Specimen examined: New Zealand, Auckland, Waiatarua, on Melicytus 

macrophyllus, 13 Mar. 2003, C.F. Hill, holotype HAL 1787 F (isotype PDD 77567), 

culture ex-type ICMP 14984 = CBS 115023. 

Pseudocercospora myrticola (Speg.) Deighton, Mycol. 

Pap. 140: 148. 1976. 

Basionym: Cercospora myrticola Speg., Anales Soc. Ci. Argent. 16: 

167. 1883. 

= Cercospora myrti Erikss., Bidrag Känn. om vara odlade Vaxters s jukdomar, 

Stockholm 8: 79. 1885 and Rev. Mycol. 8: 60. 1886. 

= Cercospora saccardoana Scalia, Atti Accad. Gioenia Sci. Nat. Catania, Ser. 

4, 14: 35. 1901. 

= Cercospora amadelpha Syd., Ann. Mycol. 30: 89. 1932. 

= Fusariella cladosporioides P. Karst., Hedwigia 30: 248. 1891. 

Specimen examined: Japan, Kagoshima, on Myrtus communis, 29 May 2007, C. 

Nakashima & K. Motohashi, MUMH 10572, culture MUCC 632. 

Pseudocercospora ocimi-basilici Crous, M.E. Palm & U. 

Braun, sp. nov. MycoBank MB564837. Fig. 45. 

Etymology: Name derived from Ocimum basilicum, the host from 

which it was collected. 

92


Fig. 43. Pseudocercospora marginalis (CPC 12497). A, B. Leaf spots on upper and lower leaf surface. C, D. Fascicles with conidiophores and conidiogenous cells. E. Close-up 

of leaf spot with fruiting. F–J. Conidia. Scale bars = 10 μm. 

Fig. 44. Pseudocercospora fraxinites (CPC 10743–10745). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C–E. Fascicles with conidiophores 

and conidiogenous cells. F. Conidiogenous cells. G. Conidia. Scale bars = 10 μm. 

Leaf spots amphigenous, subcircular, circular or somewhat irregular, 

2–10 mm diam, greyish green, dull grey to dark brown, border 

indistinct, at times raised. Mycelium internal, pale brown, consisting 

of septate, branched, smooth, 2–3 μm diam hyphae. Caespituli 

fasciculate to sporodochial, brown, predominantly hypophyllous, up 

to 90 μm diam and 70 μm high. Conidiophores aggregated in mostly 


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Crous et al. 

Fig. 45. Pseudocercospora ocimi-basilici (CPC 10283–10285). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C–E. Fascicles with 

conidiophores and conidiogenous cells. F. Conidia. Scale bars = 10 μm. 

dense, small to large, sometimes almost sporodochial fascicles, 

emerging through stomata or erumpent through the cuticle, 

arising from the upper cells of a brown, substomatal to mostly 

intraepidermal stroma, 10–80 μm; conidiophores pale to medium 

brown or olivaceous-brown, smooth, thin-walled, 0–2-septate, 

subcylindrical or attenuated towards the tip, straight to moderately 

geniculate-sinuous, unbranched or branched above, 5–35 × 2–5 

μm. Conidiogenous cells integrated, terminal or conidiophores 

reduced to conidiogenous cells, pale olivaceous-brown, smooth, 

tapering to flat-tipped apical loci, 1–2 μm wide, proliferating 

sympodially, 5–20 × 2–4 μm. Conidia solitary, subhyaline to pale 

olivaceous-brown, smooth, guttulate, shape and size variable, 

small conidia short obclavate-cylindrical to fusiform, longer conidia 

narrowly obclavate-filiform, sometimes acicular, apex subacute to 

subobtuse, base short to long obconically truncate to truncate in 

acicular conidia, straight to curved, 3–12-septate, (25–)30–120(– 

130) × (2–)2.5–5(–5.5) μm; hila unthickened, neither darkened nor 

refractive, 1.5–2.5 μm diam. 

Specimens examined: Fiji (intercepted at the Auckland International Airport, on basil 

foliage imported from Fiji), on Ocimum basilicum, 24 Feb. 2002, C.F. Hill 529, HAL. 

Mexico, on O. basilicum, Dec. 2001, without collector (cultured as MEP 1515), BPI 

841445; (intercepted at Los Angeles), 2 Nov. 2002, L.C. Lastra 1395 A, BPI 747831; 

6 Dec. 2002, M.E. Palm, holotype CBS H-20877, culture ex-type CPC 10283– 

10285 (unfortunately no longer viable). New Zealand, Auckland, Botanical Garden, 

on O. basilicum, 9 Mar. 2002, C.F. Hill 546, HAL. Vanuatu, Efate, Vanuatu Tropical 

Products, on O. basilicum, 25 Oct. 1996, E. McKenzie, PDD 66438; Rainbow 

Garden, on O. basilicum, 22 Oct. 1996, E. McKenzie, PDD 66537. 

Notes: Braun et al. (2003b) examined Pseudocercospora 

collections on Ocimum basilicum from Fiji, New Zealand, and 

Vanuatu and identified those collections as P. ocimicola, in spite 

of some morphological differences observed. Pseudocercospora 

ocimicola differs from collections on Ocimum basilicum, herein 

described as P. ocimi-basilici, in having shorter conidia (about 

25–80 μm long), conidiophores in small, loose fascicles as well as 

solitary conidiophores arising from superficial hyphae, and lacking 

or almost lacking stromata. 

The description of Cercospora ocimicola provided by Chupp 

(1954) covers type material of this species as well as material on 

O. basilicum. Based on type material and additional collections, C. 

ocimicola is redescribed as P. ocimicola in the current study (see 

below). 

Pseudocercospora ocimicola (Petr. & Cif.) Deighton, 

Mycol. Pap. 140: 149. 1976. 

Basionym: Cercospora ocimicola Petr. & Cif., Ann. Mycol. 30: 324. 

1932. 

= C. hyptidicola (“hypticola”) Chupp & A.S. Mull., Bol. Soc. Venez. Ci. Nat. 8: 

47. 1942. nom. inval. 

Leaf spots lacking or almost so to indistinct or angular-irregular, 

yellowish ochraceous, olivaceous to brownish, centre finally 

sometimes paler, dingy greyish brown to grey, 1–10 mm diam., 

margin indefinite. Mycelium internal and external, superficial, 

hyphae emerging through stomata, sparingly branched, septate, 

subhyaline to olivaceous-brown, 1–3 μm wide, thin-walled, smooth. 

Stromata lacking or small, mostly substomatal, occasionally 

intraepidermal, 10–30 μm diam. Caespituli amphigenous, usually 

not very conspicuous, olivaceous-brown, finely punctiform to 

subeffuse. Conidiophores in small, loose to moderately large and 

denser fascicles, arising from stromata or internal hyphae, through 

stomata or erumpent through the cuticle, or conidiophores solitary, 

arising from superficial hyphae, lateral or occasionally terminal, 

straight and subcylindrical to conical or usually geniculatesinuous, 

unbranched or occasionally branched, pale olivaceous to 

olivaceous-brown, 0–3-septate, thin-walled, smooth, 5–50 × (2–)3– 

5 μm. Conidiogenous cells integrated, terminal or conidiophores 

reduced to conidiogenous cells, 5–20 × 2–4 μm, proliferating 

sympodially, with a single or several inconspicus to flat-tipped 

conidiogenous loci, 1–2 μm wide. Conidia solitary, subhyaline 

to pale olivaceous or olivaceous-brown, thin-walled, smooth, 

obclavate-subcylindrical, apex obtuse to subacute, base truncate 

to obconically truncate, 1–8-septate, (15–)25–75(–85) × 2–4 μm, 

hila unthickened, neither darkened nor refractive, 1–2 μm diam. 

Specimens examined: Brazil, State of Ceará, Pentecoste County, on Ocimum 

sp., 2 Mar. 2001, F. Freire, HAL; State of Ceará, Cascavel County, Preaoca, on 

Marsypianthes chamaedrys; 12 June 1999, F. Freire, HAL. Cuba, Habana, Santiago 

de las Vegas, on Ocimum gratissimum, 6 Sep. 1988, R.F. Castañeda [C88/316], 

HAL; Habana, Santiago de las Vegas, on O. sanctum, 28 Dec. 1987, R.F. Castañeda 

[C87/382], HAL. Dominican Republic, Santiago, Valle del Cibao, Prov. Santiago, Hato 

94


Fig. 46. Pseudocercospora oenotherae (CPC 10290, 10041). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with 


Fig. 47. Pseudocercospora paederiae (CPC 10007). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C. Broken base of detached fascicle. 

D–G. Synnematal fascicles with conidiophores and conidiogenous cells. H. Conidia. Scale bars = 10 μm. 

del Yonque, on O. campechianum (= O. micranthum), 26 Nov. 1930, E.L. Ekman, Cif., 

Mycofl. Doming. Exs. 359, lectotype designated here BPI 845245 and isolectotype 

BPI 438987. India, Midnapur, Daspur, on O. sanctum, 3 Dec. 1967, M. Mandal, BPI 

438988. Venezuela, Les Tincheras, Edo Carabobo, on Hyptis sp., 24 Feb. 1940, M.F. 

Barrus & A.S. Muller, type of Cercospora hyptidicola, CUP-VZ 3863; La Cuchilla, Río 

Claro, Lara, on Hyptis suaveolens, June 2007, R. Urtiaga, HAL. 

Notes: Chupp (1954) reduced C. hyptidicola, described from 

Venezuela on Hyptis sp., to synonymy with C. lycopodis, and 

Crous & Braun (2003) followed this treatment. Braun & Urtiaga 

(2008) examined type material of this species and an additional 

new collection from Venezuela and considered C. hyptidicola a 

synonym of C. ocimicola since the two species are morphologically 

indistinguishable. Both also occur on two closely related plants, 

Hyptis and Ocimum, in the Lamiaceae subfam. Ocimoideae. 

Pseudocercospora collections on Marsypianthes (subfam. 

Ocimoideae) in Brazil, is morphologically also indistinguishable 

from collections on Ocimum spp. and was assigned to P. ocimicola 

by Braun & Freire (2002). 

Pseudocercospora oenotherae (Ellis & Everh.) Y.L. Guo & 

X.J. Liu, Acta Mycol. Sin. 11: 297. 1992. Fig. 46. 

Basionym: Cercospora oenotherae Ellis & Everh., Proc. Acad. Nat. 

Sci. Philadelphia 46: 380. 1894. 

Specimens examined: South Korea, Seoul, Oenothera odorata, 6 Sep. 2003, H.D. 

Shin, KUS-F 19606, CPC 10630 = CBS 131920; O. odorata, 2 Oct. 2002, H.D. 

Shin, CBS H-20878, cultures CPC 10290 = CBS 131885, CPC 10041. 

Pseudocercospora paederiae Goh & W.H. Hsieh, 

Cercospora and similar fungi from Taiwan: 291. 1990. Fig. 

47. 


pale brown in centre, with raised, dark brown border, at times 

with concentric zones delimited by dark borders. Mycelium 

internal, occasionally in addition with a few external hyphae 


95

Crous et al. 

Fig. 48. Pseudocercospora pallida (CPC 10776–10778). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with conidiophores 


emerging through stomata, pale to medium brown, consisting of 

septate, branched, smooth to finely verruculose, 3–4 μm diam 

hyphae. Caespituli predominantly hypophyllous, synnematous, 

dark brown on leaves, 25–50 μm wide and 100–200 μm high. 

Conidiophores aggregated in dense synnemata arising from 

the upper cells of a brown substomatal stroma 20–40 μm diam; 

individual conidiophores subhyaline to olivaceous-brown, smooth, 

multiseptate, subcylindrical-filiform, straight to gently curved, 

unbranched, 80–200 × 3–5 μm. Conidiogenous cells terminal, 

unbranched, brown, subcylindrical to clavate, smooth, tapering to 

flat-tipped apical loci, neither thickened nor darkened, proliferating 

sympodially, or rarely percurrently near apex, 20–35 × 2–5 μm. 

Conidia solitary, subhyaline, greenish yellow to pale brown, smooth 

to finely verruculose, guttulate, obclavate, short conidia sometimes 

cylindrical or fusiform, apex obtuse to subobtuse, base obconically 

truncate, straight to curved, 1–10-septate, (20–)40–60(–70) × 3–7 

μm; hila not thickened nor darkened or refractive, 1–2 μm diam. 

Specimen examined: South Korea, Pocheon, National Arboretum, Paederia foetida 

(= P. scandens), 23 Oct. 2002, H.D. Shin, CBS H-20879, culture CPC 10007 

(unfortunately no longer viable). 

Notes: A brown leaf spot on P. scandens was reported from the 

Keryong Mountain in Chungnam district, South Korea, including 

the southern districts, Chonnam, Kyeongnam, and Jeju Island 

by Lee et al. (2001). The associated fungus was identified as 

Pseudocercospora paederiae. Characteristics of the Korean 

material are consistent with the original description of P. paederiae 

(from Taiwan), except for longer conidiophores and shorter conidia 

that are up to 10-septate. All characterstics overlap, and the 

Korean collections are tentatively assigned to P. paederiae. New 

collections from Taiwan, together with cultures and sequence 

data are necessary to reassess Pseudocercospora on Paederia 

scandens in Asia. 

Pseudocercospora pallida (Ellis & Everh.) H.D. Shin & U. 

Braun, Mycotaxon 74: 114. 2000. Fig. 48. 

Basionym: Cercospora pallida Ellis & Everh., J. Mycol. 3: 21. 

1887. 

≡ Cercospora langloisii Sacc., Syll. Fung. 10: 647. 1892 (nom. superfl.). 

= Cercospora duplicata Ellis & Everh., J. Mycol. 5: 70. 1889. 

= Cercospora capreolata Ellis & Everh., J. Mycol. 8: 70. 1902. 

Specimen examined: South Korea, Suwon, on Campsis grandiflora, 14 Oct. 2003, 

H.D. Shin, KUS-F 19888, CBS H-20880, CPC 10776 = CBS 131889. 

Pseudocercospora paraguayensis (Kobayashi) Crous, 

Mycotaxon 57: 270. 1996. 

Basionym: Cercospora paraguayensis Kobayashi, Trans. Mycol. 

Soc. Japan 25: 263. 1984. 

Specimen examined: Brazil, São Paulo, Susano clonal orchard, leaves of 

Eucalyptus nitens, Jun. 1996, P.W. Crous, CPC 1458 = CBS 111317. 

Pseudocercospora pini-densiflorae (Hori & Nambu) 


Basionym: Cercospora pini-densiflorae Hori & Nambu, J. Pl. 

Protect. (Tokyo) 4: 353. 1917. 

≡ Cercoseptoria pini-densiflorae (Hori & Nambu) Deighton, Mycol. Pap. 

140: 167. 1976. 

Teleomorph: “Mycosphaerella” gibsonii H.C. Evans, Mycol. Pap. 

153: 61. 1984. 

Specimens examined: Japan, C-511, NIAES herbarium; Shizuoka, Kanaya, on P. 

densiflora, 6 Mar. 1976, K. Kasai, TFM: FPH-4544; Kumamoto, isolated from P. 

thunbergii, 24 April 1964, Y. Tokushige, MUCC 534. 

Pseudocercospora plectranthi G.C. Hunter, Crous, U. 


Etymology: Name derived from the host genus Plectranthus, from 


Leaf spots distinct, scattered over leaf surface and along leaf 

border, amphigenous, subcircular to irregular, 2–12 mm diam, 

brown to pale brown. Mycelium internal and external, pale 

brown to hyaline, branched, smooth, 1.5–4 mm diam. Caespituli 

amphigenous, predominantly epiphyllous, black, distributed 

evenly over the leaf spot, punctiform. Stromata almost absent, 

weakly developed, subimmersed, globular, olivaceous-brown, 

20–70 μm diam. Conidiophores fasciculate, brown to pale brown, 

straight to curved, smooth, unbranched, apex rounded to truncate, 

0–2-septate, (18–)22–35(–45) × (3–)4(–5) μm. Conidiogenous cells 

integrated, terminal, unbranched, brown to pale brown, smooth, 

proliferating sympodially, (9–)14–21(–25) × (2–)3–4(–5) μm. 

Conidia solitary, pale brown to subhyaline, guttulate, 2–10-septate, 

slightly constricted at septa, filiform, apex obtuse to subobtuse, 

base obconic to long obconic, (41–)62–98(–112) × (3–)4(–5) μm, 

hila unthickened, not darkened. 

96


Fig. 49. Pseudocercospora plectranthi (CPC 11462). A. Leaf spots on lower leaf surface. B. Close-up of leaf spot with fruiting. E. Fascicle with conidiophores and conidiogenous 

cells. C, D, F–H. Conidia. Scale bars = 10 μm. 

Fig. 50. Pseudocercospora profusa (CPC 10055). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Conidiophores and conidiogenous 

cells. E. Conidia. Scale bars = 10 μm. 


on MEA; erumpent, spreading, with moderate aerial mycelium, and 


iron-grey. Colonies reaching 8 mm diam. 

Specimen examined: South Korea, Jeonju, on leaves of Plectranthus sp., 1 July 


131586, CPC 11463. 

Notes: No species of Pseudocercospora are presently known from 

Plectranthus and allied genera, and as P. plectranthi does not 

correspond to any sequences available in GenBank at present, 

it is described as a new species. Numerous Pseudocercospora 

species have been described from hosts in the Lamiaceae, e.g. P. 

anisomelicola, P. colebrookiae, P. colebrookiicola, P. lamiacearum, 

P. leucadis, P. lycopodis, P. ocimicola, P. perillulae, P. pogostemonis, 

P. salvia, and P. scutellariae, but all of them are morphologically 

easily distinguishable from P. plectranthi by having different conidial 

shapes (mostly obclavate-cylindrical), smaller or no stromata 

or abundant superficial mycelium with solitary conidiophores. 

Pseudocercospora salvia has filiform conidia similar to those of P. 

plectranthi but in the former they are narrower (Hsieh & Goh 1990) 

and conidiophores are not fasciculate. 

Pseudocercospora profusa (Syd. & P. Syd.) Deighton, 

Trans. Brit. Mycol. Soc. 88: 388. 1987. Fig. 50. 

Basionym: Cercospora profusa Syd. & P. Syd., Ann. Mycol. 7(2): 

175. 1909. 

≡ Cercosporiopsis profusa (Syd. & P. Syd.) Miura, in: M. Miura, Flora 

of Manchuria and East Mongolia. Part III. Cryptogams, fungi 3: 530. 

1928. 

Specimens examined: South Korea, Seoul, Acalypha australis, 17 Sep. 2003, H.D. 

Shin, CBS H-20882, culture CPC 10713–10715; Wonju, A. australis, 18 Oct. 2002, 

H.D. Shin, CBS H-20881, culture CPC 10055. 


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Crous et al. 

Fig. 51. Pseudocercospora proteae (CPC 15217). A. Fascicle with conidiophores and conidiogenous cells. B. Conidiogenous cell giving rise to a conidium. C–F. Conidia. Scale 

bars = 10 μm. 

Fig. 52. Pseudocercospora prunicola (CPC 14511). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with conidiophores with 

conidiogenous cells. E. Hypha with conidiogenous loci. F. Conidia. Scale bars = 10 μm. 

Pseudocercospora proteae Crous, sp. nov. MycoBank 

MB564840. Fig. 51. 

Etymology: Name derived from Protea, the host genus from which 

it was collected. 

Leaf spots absent, with sporulation on adaxial leaf surface, 

prominent among leaf hairs. Mycelium internal and external, pale 

brown, consisting of septate, branched, smooth, 1.5–2 μm diam 

hyphae. Caespituli fasciculate, brown, hypophyllous, up to 120 

μm diam and 40 μm high. Conidiophores aggregated in dense 

fascicles, arising from the upper cells of a brown stroma, up to 

100 μm diam and 20 μm high; conidiophores pale brown to brown, 

smooth, 0–2-septate, subcylindrical to somewhat doliiform at base, 

straight to geniculate-sinuous, unbranched or branched above, 

15–40 × 3–6 μm. Conidiogenous cells terminal, unbranched, pale 

brown to brown, smooth, proliferating sympodially near apex, with 

flat-tipped loci, 10–15 × 2.5–5 μm. Conidia solitary, pale brown, 

smooth, guttulate, subcylindrical, straight to curved, apex obtuse, 

base truncate, (3–)8–12-septate, (35–)70–85(–100) × 3(–3.5) 

μm; hila unthickened, neither darkened nor refractive, 2.5–3 μm 

diam. 



smooth, even margins. Surface olivaceous-grey; reverse iron-grey. 


Specimen examined: South Africa, Western Cape Province, Stellenbosch, 

Assegaaibos, on leaves of Protea mundii, 16 Apr. 2008, F. Roets, holotype CBS 

H-20883, culture ex-type CPC 15216 = CBS 131587, CPC 15218, 15217. 

Notes: The long, multi-septate, subcylindrical conidia of P. proteae 

are distinct from those of P. stromatosa (25–40 × 2.5–3 μm), and 

from the shorter, verruculose conidia of P. protearum (Taylor & 

Crous 2000, Crous et al. 2004a). 

Pseudocercospora prunicola (Ellis & Everh.) U. Braun, in: 

Braun & Mel’nik, Trudy Bot. Inst. Im. V.L. Komarova 20: 82. 

1997. Fig. 52. 

Basionym: Cercospora prunicola Ellis & Everh., J. Mycol. 3: 17. 

1887. 

≡ Cercoseptoria prunicola (Ellis & Everh.) J.M. Yen, Bull. Trimest. Soc. 

Mycol. France 97: 92. 1981. 

= Cercospora pruni-yedoensis Sawada, Rep. Gov. Agric. Res. Inst. Taiwan 85: 

120. 1943, nom. inval. 

98


Fig. 53. Pseudocercospora pseudostigmina-platani (CPC 11726). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, E, F. Fascicles with 

conidiophores and conidiogenous cells, giving rise to dimorphic conidia. D. Pseudocercospora conidia. G. Conidia of Stigmina-like synanamorph. Scale bars = 10 μm. 

≡ Pseudocercospora pruni-yedoensis Goh & W.H. Hsieh, in Hsieh & Goh, 

Cercospora and similar genera from Taiwan: 282. 1990. 

= Cercospora pruni-persicae J.M. Yen, Bull. Trimest. Soc. Mycol. France 94: 

61. 1978 and Rev. Mycol. 42: 59. 1978. 

≡ Cercoseptoria pruni-persicae (J.M. Yen) J. M. Yen, Bull. Trimest. Soc. 

Mycol. France 97: 92. 1981. 

Misapplied name: Pseudocercospora circumscissa (Sacc.) Y.L. 

Guo & X.J. Liu, Mycosystema 2: 231. 1989. 

Descriptions: Hsieh & Goh (1990: 282–283, as Pseudocercospora 

pruni-yedoensis), Braun & Mel’nik (1997: 82–83). 

Illustrations: Hsieh & Goh (1990: 283, fig. 216, as Pseudocercospora 

pruni-yedoensis), Braun & Mel’nik (1997: 121, fig. 48). 

Specimens examined: South Korea, Suwon, on Prunus yedoensis (≡ Cerasus 

yedoensis), 2 Oct. 2007, H.D. Shin, CBS H-20860, CPC 14511 = CBS 132107. 

Taiwan, Taipei, on Prunus yedoensis, 30 Nov. 1930, K. Sawada, holotype of 

Pseudocercospora pruni-yedoensis, NTU-PPE. USA, Louisiana, Point a la Hache, 

Langlois 542, holotype of Cercospora prunicola, NY (also Ellis & Everh., North 

American Fungi 1771, NY, isotype). 

Notes: Braun & Mel’nik (1997) discussed the intricate taxonomy 

of Passalora and Pseudocercospora on species of Prunus s. lat. 

in detail and demonstrated, based on type material and other 

collections, that two distinct species are involved. Cercospora 

circumscissa is a true Passalora with somewhat thickened and 

darkened conidiogenous loci and hila. Its placement in Passalora 

s. str. has recently been confirmed based on molecular data 

(unpubl.). Superficial mycelium with solitary conidiophores is 

lacking, and the conidia are mostly somewhat rough-walled. 

Passalora circumscissa is also known from Asia, e.g. China, Iran 

and Japan. Some Chinese collections deposited at HMAS have 

been examined and proved to be true Passalora circumscissa 

(e.g. on Prunus mandshurica º Armeniaca mandshurica, Yanji, 

Jilin, HMAS 55845). Other collections belong to Pseudocercospora 

prunicola (e.g. on Prunus yedoensis, Nanjing, Jiangsu, HMAS 

06632, and Changshan, Hunan, HMAS 55847). The Chinese 

authors misapplied the name Pseudocercospora circumscissa. 

The published descriptions of “Pseudocercospora circumscissa” 

in Guo & Hsieh (1995) and Guo & Liu (1998) cover both species, 

namely Passalora circumscissa as well as Pseudocercospora 

prunicola, but the illustrations seem to be based on material of the 

true Pseudocercospora on Prunus. Pseudocercospora prunicola is 

morphologically easily distinguishable from Passalora circumscissa 

by its inconspicuous, unthickened, not darkened conidiogenous 

loci and hila, well-developed superficial hyphae with solitary 

conidiophores and smooth conidia. The position of P. prunicola 

within the Pseudocercospora clade has been confirmed on the 

basis of sequence data retrieved from the present Korean culture. 

Pseudocercospora pseudostigmina-platani Crous, U. 


Etymology: Name reflects its morphological similarity to the 

Pseudocercospora anamorph of Mycosphaerella stigmina-platani. 


medium brown with a wide chlorotic margin. Mycelium predominantly 

internal, pale brown, consisting of septate, branched, smooth, 2–3 

μm diam hyphae. Caespituli fasciculate to sporodochial, brown, 

predominantly hypophyllous, up to 60 μm diam and 30 μm high. 

Conidiophores aggregated in loose to dense fascicles, arising from 

the upper cells of a brown stroma, up to 50 μm diam and 20 μm high; 

conidiophores brown, verruculose, 0–1-septate, subcylindrical to 

somewhat doliiform, straight to slightly curved, unbranched, 10–20 

× 7–10 μm. Conidiogenous cells terminal, unbranched, brown, 

verruculose, proliferating percurrently near apex, with 1–4 irregular 


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Crous et al. 

Fig. 54. Pseudocercospora pyracanthigena (CPC 10808). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with conidiophores 

and conidiogenous cells. E. Conidiogenous cell giving rise to a conidium. F. Conidia. Scale bars = 10 μm. 

proliferations, 8–20 × 5–8 μm. Conidia dimorphic: Cercostigminalike 

conidia fusoid-ellipsoidal to obclavate, straight to curved, 

apex obtuse, base obconically subtruncate, brown, verruculose, 

3–5-septate, at times constricted at septa, (28–)30–35(–38) × 

(5–)7–8(–9) μm; Stigmina-like conidia broadly ellipsoid, straight 

to curved, apex obtuse, base obconically subtruncate, brown, 

verruculose, 3-septate, at times constricted at septa, which can also 

be darkened, and wall can appear thick though not distoseptate 

sensu stricto, (17–)21–25(–28) × (9–)10–12 μm; hila unthickened, 

neither darkened nor refractive, 3–3.5 μm diam. 



mycelium, and smooth, lobate margins. Surface pale olivaceousgrey, 

with thin, olivaceous-grey margin; reverse iron-grey. Colonies 

reaching 7 mm diam. 

Specimen examined: South Korea, Suwon, on leaves of Platanus occidentalis, 7 

Nov. 2007, H.D. Shin, holotype CBS H-20884, culture ex-type CPC 11726 = CBS 

131588. 

Notes: Pseudocercospora pseudostigmina-platani resembles the 

Pseudocercospora/Stigmina synanamorphs of Mycosphaerella 

stigmina-platani on Platanus in the USA, although its conidia 

are larger in size. The Stigmina-like anamorph has conidia that 

are 3–6-septate, (15–)23–30(–45) × (6–)8–9(–10) μm, and the 

Pseudocercospora conidia are 3–7-septate, (35–)45–60(–100) × 

(4–)4.5–6(–6.5) μm (Crous & Corlett 1998). Based on DNA sequence 

comparisons, the genus Stigmina was treated as synonym of 

Pseudocercospora (Crous et al. 2006). The two species occurring 

on Platanus both with Pseudocercospora/Stigmina synanamorphs 

treated here, further support this synonymy. 

Pseudocercospora pyracanthae (Katsuki) C. Nakash. & 

Tak. Kobay., Ann. Phytopathol. Soc. Japan 63: 313. 1997. 

Basionym: Cercospora pyracanthae Katsuki, Bull. Agric. Improv. 

Sect. Econ. Dept. Fukuoka Pref. 1: 19. 1949. 

Specimens examined: Japan, Fukuoka, Kurume, on Pyracantha angustifolia, 6 

Nov. 1947, S Katsuki, holotype TNS-F-243829; Chiba, Sanbu, October 1976, E. 

Ishizawa, TFM: FPH-4432; Okayama, Okayama, on P. angustifolia, 20 Nov. 1960, H. 

Tanaka, TFM: FPH-3247; P. angustifolia, T. Koboyashi & C. Nakashima, CNS-446, 

culture MUCC892; Ibaraki, on P. angusti, Nov. 1994, T. Nishijima, culture MAFF 

237140; Kumamoto, on P. crenulata, 1973, T. Kobayashi, culture MAFF 410022. 

Notes: DNA sequence data obtained for Japanese isolates of this 

species indicate at least two different taxa. Further research is 

required to select a specimen and isolate that is authentic for the 

name, while other collections probably represent a novel species. 

Pseudocercospora pyracanthigena Crous, U. Braun & 


Etymology: Name derived from the host plant Pyracantha, from 


Leaf spots amphigenous, irregular to angular, up to 7 mm diam, 

brown, with inconspicuous border. Mycelium internal, hyaline 

to pale brown, consisting of septate, branched, smooth, 2–3 μm 

diam hyphae. Caespituli fasciculate to sporodochial, amphigenous, 

but predominantly epiphyllous, olivaceous on leaves, up to 150 

μm wide and 60 μm high. Conidiophores aggregated in dense 

fascicles arising from the upper cells of a brown stroma up to 120 

μm wide and 35 μm high; conidiophores medium brown, smooth, 

0–1-septate, subcylindrical to ampulliform, straight, unbranched, 

mostly reduced to conidiogenous cells, tapering to flat-tipped apical 

loci, proliferating sympodially or percurrently near apex, 7–15 × 2–3 

μm. Conidia solitary, brown, smooth, guttulate, subcylindrical to 

narrowly obclavate, apex subobtuse, base obconically subtruncate 

to truncate, straight to gently curved, 1–4-septate, (30–)35–40(– 

45) × (2.5–)3(–3.5) μm; hila unthickened, neither darkened nor 

refractive, 1.5 μm wide. 



mycelium, and smooth, lobate margins. Surface smoke-grey; 

reverse olivaceous-grey. Colonies reaching 15 mm diam. 

Specimen examined: South Korea, Jeju, Halla arboretum, on leaves of Pyracantha 

angustifolia, 1 Nov. 2007, M.J. Park, holotype CBS H-20885, culture ex-type CPC 

10808 = CBS 131589. 

100


Fig. 55. Pseudocercospora ranjita (CPC 11141). A. Leaf spots on upper leaf surface. B, C. Close-up of leaf spots with fruiting. D–F. Fascicles with conidiophores and 

conidiogenous cells. H. Branched conidiophore. G, I–K. Conidia. Scale bars = 10 μm. 

Notes: Pseudocercospora pyracanthigena is distinct from P. 

pyracanthae (conidia 25–65 × 2.4–4 μm, conidiophores 15–40 × 

2.5–3 μm; Chupp 1954) in having shorter conidia and conidiophores. 

A second species has been recorded on Pyracantha angustifolia in 

Korea (CPC 14711–14713), for which a new name is required. 

Pseudocercospora ranjita (S. Chowdhury) Deighton, 

Mycol. Pap. 140: 151. 1976. Fig. 55. 

Basionym: Cercospora ranjita S. Chowdhury, Lloydia 21: 155. 

1958. 

Leaf spots epiphyllous, distinct, scattered, white to pale brown, 

irregular, 1–4 mm diam, definite raised brown border, surrounded 


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Crous et al. 

Fig. 56. Pseudocercospora ravenalicola (CBS 122468). A. Leaf spots on upper leaf surface. B, C. Close-up of leaf spots. D–G. Fascicles with conidiophores and conidiogenous 

cells. H–L. Conidia. Scale bars = 10 μm. 

entirely or partly by brown to dark brown irregular halo. Mycelium 

internal and external, 2–5 mm wide, branched, smooth, septate, 

subhyaline to pale brown. Caespituli epiphyllous, few in number, 

distributed over the leaf spot, dark brown to black. Stromata 

well-developed, intraepidermal to subimmersed, brown, globular 

to irregular, 40–90 μm diam. Conidiophores fasciculate, arising 

from the upper cells of stromata, pale brown, straight to curved, 

unbranched and branched, 1–4-septate, irregular in width, apex 

truncate, (20–)27–38(–42) × (3–)3.5–4.5(–5) μm. Conidiogenous 

cells terminal, unbranched, pale brown, smooth to finely verrucose, 

proliferating percurrently, (8–)9–15(–19) × 3(–4) μm. Conidia 

solitary, cylindrical to obclavate, 2–9-septate, subhyaline to pale 

brown, smooth, apex rounded to subobtuse, base obconically to 

long obconically truncate, (26–)44–67(–84) × (3–)4–5(–6) μm; hila 

unthickened nor darkened. 

Culture characteristics: Colonies on MEA reaching 27 mm diam after 

30 d at 24 °C on MEA. Colonies circular with a smooth margin, that is 

darker than the colony centre, slight folding; aerial mycelium moderate; 

greyish blue to olivaceous-grey (surface) and iron-grey (reverse). 

Specimen examined: Indonesia, Northern Sumatra, on leaves of Gmelina sp., Mar. 

2004, M.J. Wingfield, CBS H-20386, culture CPC 11141 = CBS 126005. 

Note: The present collection closely matches the morphological 

description of the type specimen, which was collected from India 

(Chowdhury 1958). 

Pseudocercospora ravenalicola G.C. Hunter & Crous, sp. 


Etymology: Name derived from the plant host Ravenala, from which 

this fungus was isolated. 

102


Fig. 57. Pseudocercospora rhamnellae (CPC 12500–12502). A. Leaf spots on upper leaf surface. B, C. Close-up of leaf spots with fruiting. D, E. Fascicles with conidiophores 

and conidiogenous cells. F–J. Conidia. Scale bars = 10 μm. 

Leaf spots amphigenous, distinct, brown to pale, predominantly 

at leaf margin, but smaller spots are scattered over the whole 

leaf, elongated to irregular; border definite, raised, with dark 

brown to black border. Caespituli amphigenous, sparsely 

scattered over the leaf spot and aggregated toward the lesion 

margin, flocculose, pale to pale olivaceous. Stromata erumpent 

to superficial, globular, pale to dark brown, 30–80 μm diam. 

Conidiophores fasciculate, arising from the stromata, brown, 

becoming paler toward the apex, smooth, 0–3-septate, straight to 

curved, apex subtruncate to rounded, predominantly unbranched, 

sometimes branched below, (14–)17–25(–32) × (3–)4–5(–6) 

μm. Conidiogenous cells terminal, pale brown, smooth, straight 

to geniculate, tapering to a truncate to blunt apex, proliferating 

sympodially and percurrently, (7–)13(–15) × (3–)3.5(–4) μm. 

Conidia solitary, cylindrical, straight to curved, smooth, subhyaline 

to pale brown, 1–6-septate, infrequently constricted at the septa, 

apex obtuse to narrowly rounded, base obconically truncate to 

long obconically truncate, (16–)25–47(–60) × (3–)4(–5) μm; hila 

unthickened, nor darkened. 

Culture characteristics: Colonies after 1 mo at 24 ºC in the dark 


and smooth, lobate margins. Surface smoke-grey in centre, pale 

olivaceous-grey in outer region; reverse olivaceous-grey. Colonies 


Specimen examined: India, Chandigarh, on leaves of Ravenala madagascariensis, 

2 Mar. 2004, W. Gams, holotype CBS H-20394, culture ex-type CBS 122468. 

Note: Pseudocercospora ravenalicola represents the first species 

of Pseudocercospora known from this host and the Strelitziaceae. 

Pseudocercospora rhabdothamni U. Braun & C.F. Hill, 

Australas. Plant Pathol. 33: 489. 2004. 

Specimen examined: New Zealand, Auckland, University Campus, Princes Street, 

on Rhabdothamnus solanderi, 9 Nov. 2003, C.F. Hill, holotype HAL 1790 F, isotype 

PDD 80279, culture ex-isotype CBS 114872, ICMP 15289. 

Note: Two strains have been deposited in CBS under the name P. 

rabdothamni. 

Pseudocercospora rhamnellae G.C. Hunter, H.D. Shin, U. 

Braun & Crous, sp. nov. MycoBank MB564844. Fig. 57. 

Etymology: Name derived from the plant host Rhamnella, from 

which this fungus was isolated. 

Leaf spots distinct, amphigenous, subcircular to irregular, pale to 

dark brown, dark brown to black raised border with effuse spreading 

pale to dark brown halo, solitary or sometimes coalescing, 2–11 


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Crous et al. 

Fig. 58. Pseudocercospora rhododendri-indici (CPC 10822–10824). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with 


mm diam. Mycelium smooth, branched, internal and external, pale 

brown, septate 2–4 μm diam. Caespituli amphigenous, on adaxial 

surface single, scattered to slightly aggregated, pale to light brown, 

on abaxial surface significantly more dense, mostly aggregated 

over the lesions surface, light brown to light olive-green. Stromata 

medium to large, well-developed, superficial to intraepidermal, pale 

to dark brown, 30–85 μm diam. Conidiophores fasciculate, straight 

to curved, brown, becoming paler to the apex, unbranched, smooth 

to finely verruculose, subcylindrical, 0–1-septate, (10–)13–19(–23) 

× (2–)3–4(–5) μm. Conidiogenous cells terminal, unbranched, pale 

brown, smooth to slightly verruculose, proliferating sympodially or 

percurrently near apex, (3–)5–10(–15) × (2–)3–4(–5) μm. Conidia 

solitary, guttulate, straight to curved, apex obtusely rounded, base 

truncate, solitary, pale brown, thin-walled, smooth, subcylindrical 

to narrowly obclavate, 1–12-septate, (17–)33–57(–80) × (2–)3(–4) 

μm, hila neither thickened, nor darkened or refractive, 2–3 μm diam. 



mycelium, and smooth, lobate margins. Surface olivaceous-grey 

with patches of pale olivaceous-grey; reverse iron-grey. Colonies 


Specimen examined: South Korea, Jeju, Halla arboretum, on leaves of Rhamnella 

franguloides, 29 Oct. 2005, H.D. Shin, holotype CBS H-20395, culture ex-type CPC 

12500 = CBS 131590, CPC 12501, 12502. 

Notes: No species of Pseudocercospora are presently known 

to occur on Rhamnella (Rhamnaceae). Pseudocercospora 

rhamnellae is distinct from P. rhamnaceicola (on Paliurus, Rhamnus 

and Zizyphus; conidia 18–85 × 1.5–2.5 μm, apex pointed, base 

obconically truncate, Hsieh & Goh 1990) by having wider conidia, 

which are subcylindrical-obclavate with an obtusely rounded apex 

and truncate base. The conidiophores are also shorter and wider. 

Further collections are needed to determine whether isolates from 

other hosts in the Rhamnaceae all represent P. rhamnaceicola. 

Pseudocercospora rhododendri-indici Crous, U. Braun & 


Etymology: Name derived from the plant host Rhododendron 

indicum, from which it was collected. 

Leaf spots amphigenous, subcircular to circular, 2–3 mm diam, 

medium brown with a raised, dark brown border. Mycelium internal, 

pale brown, consisting of septate, branched, smooth, 2–3 μm diam 

hyphae. Caespituli fasciculate to sporodochial, olivaceous-brown, 

predominantly epiphyllous, up to 100 μm diam and 80 μm high. 

Conidiophores aggregated in dense fascicles, arising from the 

upper cells of a brown stroma, up to 80 μm diam and 40 μm high; 

conidiophores pale brown, smooth, 0–2-septate, subcylindrical, 

straight to geniculate-sinuous, unbranched, 10–30 × 3–4 μm. 

Conidiogenous cells terminal, pale brown, smooth, tapering to 

flat-tipped apical loci, proliferating sympodially, 10–15 × 3–3.5 μm. 

Conidia solitary, pale brown, smooth, guttulate, subcylindrical, apex 

subobtuse, base truncate, straight to variously curved, 1–4-septate, 

(35–)40–55(–65) × (2–)3 μm; hila unthickened, neither darkened 

nor refractive, 2–3 μm diam. 



and smooth, lobate margins. Surface olivaceous-grey in centre, 

pale olivaceous-grey in outer region; reverse iron-grey. Colonies 


Specimen examined: South Korea, Seoul, on Rhododendron indicum, 27 Oct. 


131591, CPC 10823, 10824. 

Notes: Of the species occurring on Rhododendron, P. rhododendriindici 

differs from P. handelii (conidia narrowly linear to obclavate, 

indistinctly multiseptate, 12–140 × 1.5–3 μm; Chupp 1954) by its 

subcylindrical, 1–4-septate conidia with truncate base and obtuse 

apex, and phylogentic position (Fig. 5). The description and 

illustration of P. handelii based on Chinese material (Guo & Hsieh 

1995) agrees well with Chupp’s (1954) description. The identity of 

Korean collections on Rhododendron indicum described in Shin & 

Kim (2001), characterised by much longer acicular-filiform conidia 

with truncate base, is unclear. Pseudocercospora rhododendriindici 

differs from P. rhododendricola (conidia 54–96 × 2–2.5 μm; 

Yen 1966) by its shorter conidia. Beside epiphyllous colonies, 

P. rhododendricola forms hypophyllous colonies composed of 

small, loose fascicles of conidiophores that emerge through 

stomata, together with superficial hyphae that give rise to solitary 

conidiophores. The hypophyllous fruiting was neither mentioned in 

the original description nor in Yen & Lim (1980). It was observed 

during the re-examination of type material (Singapore, Botanic 

Gardens, on Rhododendron sp., 13 Apr. 1965, S.H. Yen No. 112, 

holotype PC). 

Pseudocercospora rhoina (Cooke & Ellis) Deighton, Mycol. 

Pap. 140: 152. 1976. Fig. 59. 

Basionym: Cercospora rhoina Cooke & Ellis, Grevillea 6: 89. 1878. 

= Cercospora copallina Cooke, Grevillea 12: 31. 1883. 

= Cercospora rhoina var. nigromaculans Peck, Rep. (Annual) New York State 

Mus. Nat. Hist. 42: 129. 1889. 

Specimen examined: South Korea, Namhae, on Rhus chinensis, 30 Jun. 2004, 

H.D. Shin, CBS H-20887, KUS-F 20367, CPC 11464 = CBS 131891. 

104


Fig. 59. Pseudocercospora rhoina (CPC 11464–11465). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C–E. Fascicles with conidiophores 

and conidiogenous cells. F. Conidia. Scale bars = 10 μm. 

Fig. 60. Pseudocercospora sambucigena (CPC 14397–14399). A, B. Leaf spots on upper and lower leaf surface. C, D. Close-up of leaf spots with fruiting. E, F. Fascicles with 

conidiophores and conidiogenous cells. G. Conidiogenous cells. H–L. Conidia. Scale bars = 10 μm. 

Pseudocercospora sambucigena U. Braun, Crous & K. 

Schub., Mycotaxon 92: 400. 2005. Fig. 60. 

Leaf spots distinct, scattered over leaf surface, amphigenous, 

upper surface pale brown to grey, with definite border that is raised 

and dark brown in colour; lower surface pale grey to pale brown, 


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Crous et al. 

Fig. 61. Pseudocercospora securinegae (CPC 10793). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with conidiophores 


with distinctly raised, brown border, 2–10 mm diam. Mycelium 

smooth, internal and external, consisting of branched, subhyaline, 

2–4 μm diam hyphae. Caespituli amphigenous, predominantly 

occurring on the abaxial lesion surface, evenly distributed over the 

lesion, punctiform, grey to dark brown. Stromata well-developed, 

subimmersed becoming erumpent, globular, dark brown, 45–100 

mm diam. Conidiophores fasciculate, emerging from stomata, 

brown, becoming paler toward the apex, unbranched, straight 

to curved, cylindrical, uniform or irregular in width, rounded 

apex, indistinctly 0–3-septate, (25–)35–51(–60) × (4–)5(–7) μm. 

Conidiogenous cells terminal, unbranched, smooth, pale brown, 

proliferating sympodially and percurrently, conidiogenous loci 

(scars) unthickened to slightly thickened, but not darkened, (10–) 

19–34(–46) × (3–)5 μm. Conidia solitary, pale olivaceous to pale 

brown, smooth, guttulate, apex obtuse, base long obconically 

truncate, shape variable from cylindrical to obclavate, 1–7-septate, 

(40–)68–117(–156) × (4–)5–6(–7) μm; hila unthickened to slightly 

thickened, but not darkened. 


after 30 d in the dark at 24 °C. Colonies circular to subcircular, 

smooth to slightly irregular margin, prominently convex, moderate 

aerial mycelium; pale greenish grey to pale olivaceous-grey 

(surface) and olivaceous-black (reverse). 

Specimens examined: Italy, Parma, on leaves of Sambucus nigra, G. Passerini, 

paratype B 70-6710. Netherlands, Milingerwaard on leaves of Sambucus nigra, 

2007, P.W. Crous, epitype designated here CBS H-20391, cultures ex-epitype 

CPC 14397 = CBS 126000. USA, Pennsylvania, Dauphin Co., on leaves of 

Sambucus pubens, 21 Aug. 1921, O.E. Jennings, Acc. 6736, holotype NY. 

Pseudocercospora securinegae (Togashi & Katsuki) 

Deighton, Mycol. Pap. 140: 152. 1976. Fig. 61. 

Basionym: Cercospora securinegae Togashi & Katsuki, Ann. 

Phytopathol. Soc. Japan 17: 7. 1952. 

Specimen examined: South Korea, Yangpyong, on Flueggea suffruticosa 

(≡ Securinega suffruticosa), 30 Sep. 2003, H.D. Shin, CBS H-20888, culture CPC 

10793 = CBS 131930. 

Pseudocercospora snelliana (Reichert) U. Braun, H.D. 

Shin, C. Nakash. & Crous, comb. nov. MycoBank MB564846. 

Figs 62, 63. 

Basionym: Cercospora snelliana Reichert, Bot. Jahrb. Syst. 56: 

724. 1921. 

= Clasterosporium mori Syd. & P. Syd., Mem. Herb. Boiss. 4: 6. 1900. 

≡ Sirosporium mori (Syd. & P. Syd.) M.B. Ellis, Mycol. Pap. 87: 7. 1963. 

≡ Cercospora kusanoi Sawada, Rep. Dept. Agric. Gov. Res. Inst. 

Formosa 35: 109. 1928 (nom. nov.), non Cercospora mori Hara, 1918. 

= Cercospora bremeri Petr., Sydowia 2: 312. 1948. 

= Cercospora flexuosa Tanaka, unknown, nom. nud., non Tracy & Earle, 1895. 

Leaf spots lacking or amphigenous, but inconspicuous on upper 

leaf surface, chlorotic, irregular, as small speckles, up to 8 

mm diam, or effuse and much larger, forming large blotches or 

covering large portions of the hypophyllous surface with blackish 

colonies. Mycelium internal and external; internal hyphae pale 

olivaceous to pale brown, smooth, 3–4 μm diam, arising through 

stomata, giving rise to external mycelium that is pale yellowish 

green, olivaceous to brown, smooth, thin-walled, 1.5–5 μm 

diam. Conidiophores arising singly from superficial mycelium 

and in small, divergent fascicles from a few substomatal swollen 

hyphal cells, 2–8 μm diam., emerging through stomata, brown, 

smooth, becoming roughened towards apex, wall up to 1 μm 

thick, 1–12-septate, subcylindrical to often subclavate, i.e. width 

somewhat increasing towards the apex, straight to variously 

curved or geniculate-sinuous, unbranched or branched above, 

15–100 × 3–6 μm. Conidiogenous cells terminal or lateral, 

unbranched, brown, becoming paler towards the tip, roughened, 

tapering towards flat-tipped loci, 2–3 μm diam, proliferating 

sympodially (lateral scars as illustrated by Ellis 1971 observed), 

or percurrently near apex, 10–30 × 4–7 μm. Conidia solitary, 

medium to dark olivaceous-brown or brown, small young conidia 

sometimes subhyaline to pale olivaceous, wall up to 1 μm thick, 

smooth or almost so to verruculose, guttulate, smaller conidia 

ellipsoid-ovoid, subcylindrical, larger conidia usually distinctly 

obclavate, apex obtuse, base obconically truncate, subtruncate 

or sometimes rounded, straight to gently curved, 1–10-septate 

(septa somewhat refractive, at times also 1(–2) oblique or 

vertical septa present), (15–)30–70(–80) × (3–)4–6(–7) μm; 

106


Fig. 62. Pseudocercospora snelliana (B 700014740, holotype). Sparse fascicles, and solitary conidiophores on superficial mycelium giving rise to muriformly septate, thickwalled 

conidia. Scale bar = 10 μm. 

Fig. 63. Pseudocercospora snelliana (CPC 11654–11656). A. Leaf spots on the lower leaf surface. B. Close-up of leaf spot with fruiting. C–E. Solitary conidiophores and 

conidiogenous cells. F–H. Conidia. Scale bars = 10 μm. 


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Crous et al. 

hila neither thickened, nor darkened or refractive, 1–1.5(–2) μm 

diam. 





Specimens examined: Egypt, Kahirahm, near Bahtim, on Morus alba, Nov. 1913, 

Snell, holotype B 700014740. South Korea, Hoengseong, on Morus bombycis, 

11 Oct. 2004, H.D. Shin, CBS H-20889, HAL 1867 F, culture CPC 11654 = CBS 

131592, CPC 11655, 11656. 

Notes: Cercospora kusanoi is based on the same type specimen 

used by Sydow to describe Clasterosporium mori. Sawada (1928) 

considered this fungus a species of Cercospora. He introduced the 

name Cercospora kusanoi because the species epithet mori was 

occupied in Cercospora. The Korean material we studied closely 

resembles the description of the type, which was originally described 

on Morus alba from Japan (Sawada 1928). Pseudocercospora 

mori is also already occupied so type material of P. snelliana, the 

next available epithet, was re-examined. We determined it to be 

conspecific with C. kusanoi, so P. snelliana is introduced as a new 

combination. 

Pseudocercospora stephanandrae (Tak. Kobay. & H. 

Horie) C. Nakash. & Tak. Kobay., Mycoscience 41: 27. 2000. 

Basionym: Cercospora stephanandrae Tak. Kobay. & H. Horie, 

Trans. Mycol. Soc. Japan 20: 331. 1979. 

Specimens examined: Japan, Tokyo, Jindai Bot. Park, on Stephanandra incisa, 21 

Oct. 1976, T. Kobayashi & H. Horie TFM: FPH-4712; Tokyo, Jindai Botanical Park, 

Chofu-City, on S. incisa, 26 Oct. 1974, H. Horie, holotype TFM: FPH 4411; Tokyo, 

Jindai Bot. Park, on S. incisa, 7 Nov. 1998, C. Nakashima & E. Imaizumi, epitype 

designated here TFM: FPH-8099, ex-epitype cultures MUCC 914, MAFF 237799. 

Pseudocercospora timorensis (Cooke) Deighton, Mycol. 

Pap. 140: 154. 1976. 

Basionym: Cercospora timorensis Cooke, Grevillea 12: 38. 1883. 

= Ramularia batatae Racib., Paras. Algen Pilze Javas, Batavia 1: 35. 1900. 

= Cercospora batatae A. Zimmerm., Ber. Land.-Forstw. Deutsch Ostafrikas 2: 

28. 1904. 

= Cercospora batatae Henn., Bot. Jahrb. Syst. 38: 118. 1907 (nom. illeg.), 

homonym of C. batatae A. Zimmerm., 1904. 

= Cercospora ipomoeae-purpureae J.M. Yen, Rev. Mycol. 30: 173. 1965. 

≡ Pseudocercospora ipomoea-purpureae (J.M. Yen) J.M. Yen, in Yen & 

Lim, Gard. Bull., Singapore 33: 177. 1980. 

Specimen examined: Japan, Okinawa, Ipomoea indica, 19 Nov. 2007, C. Nakashima 

& T. Akashi, MUMH 10923, culture MUCC 819. 

Pseudocercospora udagawana (Katsuki) X.J. Liu & Y.L. 

Guo, Mycosystema 2: 238. 1989. Fig. 64. 

Basionym: Cercospora udagawana Katsuki, Ann. Phytopathol. 

Soc. Japan 20(2–3): 72. 1955. 

Specimen examined: South Korea, Dongducheon, on Hovenia dulcis, 28 Sep. 

2003, H.D. Shin, CBS H-20890, CPC 10799 = CBS 131931. 

Pseudocercospora viburnigena U. Braun & Crous, Mycol. 

Progr. 1: 23. 2002. Fig. 65. 

Basionym: Cercospora tinea Sacc., Michelia 1(2): 268. 1878 (non 

P. tinea Y.L. Guo & W.H. Hsieh, 1994). 

≡ Cercoseptoria tinea (Sacc.) Deighton, Mycol. Pap. 140: 167. 1976. 

≡ Cercostigmina tinea (Sacc.) U. Braun, Cryptog. Bot. 4: 108. 1993. 

Leaf spots distinct, scattered, amphigenous, 4–15 mm diam, 

lesions on abaxial surface dark to pale brown, subcircular to 

irregular, surrounded by a slightly raised dark brown border, 

lesions on adaxial surface dark to pale brown, surrounded by 

a dark brown border with a light red diffuse pigment extending 

outward from the border in older lesions. Mycelium internal 

and external, smooth, subhyaline, branched, 1.5–4 μm wide. 

Caespituli amphigenous, but predominantly hypophyllous, 

evenly distributed over the leaf spot, velvety, olivaceous. 

Stromata well-developed, subimmersed, globular, dark 

brown, 30–80 μm diam. Conidiophores fasciculate, smooth, 

0–2-septate, emerging from the upper cells of the stroma, pale 

brown, straight to curved, irregular in width, apex subtruncate 

to rounded, (14–)17–24(–30) × (3–)4–5(–6) μm. Conidiogenous 

cells integrated, terminal, inconspicuously proliferating 

percurrently, cylindrical, straight, pale brown, at times slightly 

verruculose, (5–)9–15(–19) × (2–) 3(–4) μm. Conidia solitary, 

pale brown, smooth, guttulate, apex obtusely rounded, base 

narrowly truncate, narrowly ellipsoidal to acicular, curved or 

sigmoid, 5–11-septate, (68–)87–110(–120) × (2–)3–4(–5) μm, 

hila unthickened. 


after 30 d at 24 °C in the dark. Colonies circular, convex, smooth 

margin that is distinctly darker than the rest of the colony, slight 

folding occurs toward the edge of the colony, moderate to profuse 

aerial mycelium; olivaceous-grey (surface) and greenish black 

(reverse). 

Specimens examined: Italy, Padova, Viburnum tinus, Oct. 1877, Bizzozera, Sacc., 

Mycoth. Venet. 1252, syntype HAL. Netherlands, Bilthoven, Sweelincklaan 87, on 

leaves of Viburnum davidii, 26 May 2008, M.K. Crous, epitype designated here 

CBS H-20393, culture ex-epitype CPC 15249 = CBS 125998. 

Note: The epitype closely matches the morphology of the holotype 

(Braun & Hill 2002), representing a species that is common on 

Viburnum in Europe. 

Pseudocercospora viticicola (J.M. Yen & Lim) J.M. Yen, 

Gardens Bulletin, Singapore 33: 190. 1980. 

Basionym: Cercospora viticicola J.M. Yen & Lim, Cah. Pacifique 

17: 104. 1973. 

= Cercospora viticis Ellis & Everh. (as viteae), J. Mycol. 3: 18. 1887, non 

Pseudocercospora viticis Goh & W.H. Hsieh, 1989. 

≡ Pseudocercosporella viticis (Ellis & Everh.) B.K. Gupta & Kamal, Indian 

Phytopathol. 42: 388. 1989 (nom. inval.). 

≡ Pseudocercospora viticicola U. Braun, Mycotaxon 48: 296. 1993 (nom. 

illeg.), homonym of P. viticicola (J.M. Yen & Lim) J.M. Yen, 1980. 

= Cercospora viticis Sawada, Rep. Gov. Agric. Res. Inst. Taiwan 87: 90. 1944 

(nom. illeg.), homonym of C. viticis Ellis & Everh., 1887. 

≡ Pseudocercospora viticis Goh & W.H. Hsieh, Trans. Mycol. Soc. 

Republ. China 4: 11. 1989. 

= Cercospora viticis-quinatae J.M. Yen, Bull. Trimestriel Soc. Mycol. France 

93: 158. 1977. 

≡ Pseudocercospora viticis-quinatae (J.M. Yen) J.M. Yen, Bull. Trimestriel 

Soc. Mycol. France 94: 388. (1978) 1979. 

= Pseudocercospora viticigena J.M. Yen, A.K. Kar & B.K. Das, Mycotaxon 16: 

68. 1982. 

Specimens examined: Japan, Okinawa, Okinawa Is, on Vitex trifolia, 19 Nov. 2007, 

C. Nakashima, MUMH 10828, culture MUCC 777; Chiba, Matsudo, on V. agnuscastus, 

7 Nov. 1987, M. Nagashima & T. Kobayashi, TFM: FPH-6912; Shizuoka, 

Kanzanji, on V. agnus-castus, 1 Nov. 1996, T. Kobayashi & C. Nakashima, CNS- 

101, culture MUCC 1069, MAFF 237866; Kuroki, Fukuoka, on V. cannabifolia (≡ V. 

negundo var. cannabifolia), 25 Sep. 1974, S. Ogawa, TFM: FPH-4193. 

108


Fig. 64. Pseudocercospora udagawana (CPC 10799–10801). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C, D. Fascicles with 

conidiophores and conidiogenous cells. E. Solitary conidiogenous cell on superficial hypha. F. Conidia. Scale bar = 10 μm. 

Fig. 65. Pseudocercospora viburnigena (CPC 15249). A. Leaf spots on upper leaf surface. B, C. Close-up of leaf spots with fruiting. D, E. Fascicles with conidiophores and 

conidiogenous cells. F–H. Conidia. Scale bars = 10 μm. 


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Crous et al. 

Fig. 66. Pseudocercospora xanthocercidis (CPC 11665–11667). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C–E. Fascicles with 

conidiophores and conidiogenous cells. F. Conidia. G. Colony on malt extract agar. H, I. Conidia formed in culture. Scale bars = 10 μm. 

Pseudocercospora weigelae (Ellis & Everh.) Deighton, 

Trans. Brit. Mycol. Soc. 88: 389. 1987. 

Basionym: Cercospora weigelae Ellis & Everh., Proc. Acad. Nat. 

Sci. Philadelphia 45: 170. 1893. 

Specimen examined: Japan, Ibaraki, on Weigela coraeensis, 10 Sep. 1998, T. & Y. 

Kobayashi, CNS-455, culture MUCC 899, MAFF 237794. 

Pseudocercospora xanthocercidis Crous, U. Braun & A. 

Wood, sp. nov. MycoBank MB564847. Fig. 66. 

Etymology: Name derived from the plant host Xanthocercis, from 



pale to medium brown, with indistinct border. Mycelium internal, 

pale brown, consisting of septate, branched, smooth, 2–3 μm diam 

hyphae. Caespituli sporodochial, hypophyllous, also occurring 

on green leaf tissue, prominent, appearing like insect galls, 

olivaceous-brown on leaves, up to 400 μm wide and 300 μm high. 

Conidiophores aggregated in dense sporodochial fascicles arising 

from the upper cells of a brown stroma up to 300 μm wide and 

250 μm high; conidiophores brown, finely verruculose, 1–2-septate, 

subcylindrical, straight to slightly curved, 20–30 × 5–7 μm. 

Conidiogenous cells terminal, unbranched, brown, subcylindrical, 

finely verruculose, proliferating percurrently near apex, with 

several irregular, rough proliferations, 7–12 × 5–6 μm. Conidia 

solitary, brown, finely verruculose, guttulate, narrowly obclavate, 

apex obtuse, base obconically subtruncate to truncate, straight to 

gently curved, 5–8-septate, (25–)28–36(–40) × (5–)6–7 μm; hila 

unthickened, neither darkened nor refractive, 3–4 μm diam, with 

minute marginal frill visible. 


MEA; surface irregular, folded, erumpent, spreading, with sparse 

aerial mycelium, and smooth, irregularly lobate margins. Surface 

olivaceous-grey, with patches of iron-grey; reverse iron-grey. 


Specimen examined: South Africa, Mpumalanga, Nelspruit, Lowveld National 

Botanical Garden, on Xanthocercis zambesiaca, 14 Sep. 2004, A. Wood, holotype 

HAL 1859 F, isotype CBS H-20891, culture ex-type CPC 11665 = CBS 131593, 

CPC 11666, 11667. 

Notes: No other species of Pseudocercospora are known 

from this host. Pseudocercospora xanthocercidis differs from 

other Pseudocercospora species on legumes by its very 

large sporodochial conidiomata with percurrently proliferating 

conidiogenous cells and verruculose conidia with visible marginal 

frill at the base. There is no comparable species on legumes. 

Pseudocercospora xanthoxyli (Cooke) Y.L. Guo & X.J. Liu, 

Mycosystema 4: 115. 1991. Fig. 67. 

Basionym: Cercospora xanthoxyli Cooke, Grevillea 12: 30. 1883. 

= Cercospora fagaricola Sawada (fagariae), Rep. Gov. Agric. Res. Inst. Taiwan 

85: 105. 1943 (nom. inval.). 

≡ Pseudocercospora fagaricola Goh & W.H. Hsieh, in Hsieh & Goh, 

Cercospora and similar species from Taiwan: 294. 1990. 

Specimen examined: South Korea, Wando, Wando Arboretum, on Xanthoxylum 

ailanthoides, 9 Nov. 2002, H.D. Shin, CBS H-20892, CPC 10009, 10064–10065. 

Pseudocercospora zelkovae (Hori) X.J. Liu & Y.L. Guo, 

Acta Mycol. Sin. 12: 33. 1993. Fig. 68. 

Basionym: Cercospora zelkowae Hori, Nambu N. Jour. Plant 

Protection 8: 492. 1921. 

110


Fig. 67. Pseudocercospora xanthoxyli (CPC 10009, 10064–10065). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C–E. Fascicles with 

conidiophores and conidiogenous cells. F. Close-up of conidiogenous cells. G. Conidia. Scale bars = 10 μm. 

Fig. 68. Pseudocercospora zelkovae (CPC 14484). A. Leaf spots on upper and lower leaf surface. B. Close-up of leaf spot with fruiting. C–E. Fascicles with conidiophores and 

conidiogenous cells. F. Conidia. Scale bar = 10 μm. 

Holotype: Japan, Tokyo, Forest Experimental Station, on Zelkova 

serrata, Jun. 1920 (not preserved). 

Specimens examined: Japan, Yamagata, Kamabuchi, on Z. serrata, 5 July 1956, K. 

Ito, neotype designated here TFM:FPH169, cultures ex-neotype MAFF 410008, 

MUCC 1398. South Korea, Suwon, on Z. serrata, 2 Oct. 2007, H.D. Shin, CBS 

H-20893, culture CPC 14484 = CBS 132106; Osan, on Z. serrata, 30 Oct. 2007, 

H.D. Shin, CBS H-20894, CPC 14717 = CBS 132118. 

DISCUSSION 

This study provides a broad framework and phylogeny for the 

genus Pseudocercospora. These fungi are very common and 

the foundation that has been set will form the basis for additional 

species to be described and for specific groups to be more 

thoroughly investigated. Although the results clarify several issues 

relating to the taxonomy of Pseudocercospora s. str., the study 

also highlights many remaining taxonomic questions relating to 

this complex. To resolve these issues many species will need to 

be recollected, cultured, and sequenced so that they can be placed 

into this phylogenetic backbone. This is especially true for species 

described in some of the obscure genera treated by Braun (1995) 

and Crous & Braun (2003), many of which (or their type species) 

are not currently known from culture, and thus DNA sequence 

comparisons and phylogenetic inference has not been possible. 

Amongst the cercosporoid fungi, it appears possible and even 

probable that the approximately 1 500 names in Pseudocercospora 

represent the tip of the iceberg in terms of biodiversity. Indeed it 

seems likely that this could emerge as the largest genus of 

cercosporoid fungi known. A significant result of this study was 

the determination that names based on American or European 

type specimens could in most cases not be used when identifying 


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Crous et al. 

identical diseases on the same hosts in Asia, Africa or South 

America. In this regard, it was surprising to find diversity even 

within a region such as Asia, where isolates from the same host 

and disease symptoms from Korea frequently differed from similar 

collections made in Japan. These important issues, which have 

significant ramifications pertaining to plant health and quarantine, 

will only be resolved when fresh collections from the American 

and European type locations have been made, thus allowing 

DNA sequence based comparisons. Furthermore, it emphasises 

the need to ensure that a DNA sequence has been provided for 

all novel taxa in this complex and that an authentic DNA barcode 

(Schoch et al. 2012) is available. The ITS gene region was found to 

be capable of differentiating only 25 of the 146 Pseudocercospora 

taxa (17 %) to species level in the present study. Where the ITS 

locus fails to provide acceptable resolution, it can be supplemented 

with sequences from the ACT or EF-1α gene regions (Fig. 5), 

though these loci still proved relatively conserved, and 57 taxa had 

less than 1% variation from their closest neighbours, suggesting 

that additional loci still have to be found to provide a more robust 

identification of Pseudocercospora species. 

Focused studies on specific crops such as those on Eucalyptus 

(Crous 1998, Hunter et al. 2006b), Musa (Arzanlou et al. 2007, 

2008, 2010), Chromolaena (Den Breeÿen et al. 2006) and Citrus 

(Pretorius et al. 2003) will undoubtedly confirm the already 

emerging view that many plant species are infected by a complex 

of Pseudocercospora spp. Some of these will clearly be specific 

to the host from which they were isolated, while others reflect 

chance occurrences or infections or broader host ranges (Crous 

& Groenewald 2005). In some instances, these chance infections 

may be caused by fungi that are major pathogens of other, 

completely unrelated hosts (Crous & Groenewald 2005, Arzanlou et 

al. 2008). Although the present study has succeeded in delineating 

Pseudocercospora within the Mycosphaerellaceae, and in the 

process has also delineated several other Pseudocercospora-like 

genera, the question relating to host specificity still remains largely 

unanswered. 

The taxa investigated during this study represent the largest 

collection of Pseudocercospora and Pseudocercospora-like taxa 

ever subjected to DNA sequence analysis. Of these, the vast 

majority appear to be host-specific. Of the 146 taxa subjected to 

multi-gene analysis, only four were found to occur on more than 

one host. These include P. norchiensis (Myrtaceae and Rosaceae), 

P. fraxinites (Oleaceae), P. atromarginalis (Solanaceae) and P. 

corylopsidis (Hamamelidaceae). In the latter three examples, 

the same species was found on different host genera within the 

same plant family, but never on unrelated hosts. This result was 

somewhat surprising as we initially expected to find at least some 

examples where species are generalists and occur on many hosts 

which are unrelated such as those in the Cercospora apii complex 

(Groenewald et al. 2006, 2007). The occurrence of P. norchiensis 

(a foliar pathogen of Eucalyptus in Italy; Crous et al. 2007c) on 

Rubus in New Zealand (CBS 114641), was highly unexpected, 

and further collections on Rubus from New Zealand will have to 

be made to resolve if this was a mere chance occurrence (Crous & 

Groenewald 2005), or true indication of its host range. 

In future studies of Pseudocercospora, additional taxa should 

be included in the analyses, and further loci screened to obtain a 

better separation of species. There is an urgent need to conduct 

inoculation tests to confirm inferences from taxonomic studies 

about host specificity in this important group of predominantly plant 

pathogenic fungi. For example, it remains to be shown whether 

isolates from different hosts with identical DNA barcodes and 

similar morphology have the ability to cross-infect hosts under 

natural conditions in the field. It appears that for the most part, F.C. 

Deighton was correct in his statement “If a sparrow flies to a cherry 

tree, it’s a cherry tree sparrow. If the same sparrow sits in an apple 

tree, it is an apple tree sparrow”. 

ACKNOWLEDGEMENTS 

We thank the technical staff, Arien van Iperen (cultures), Marjan Vermaas 

(photographic plates), and Mieke Starink-Willemse (DNA isolation, amplification and 

sequencing) for their invaluable assistance. 

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