Phylogenetic lineages in Pseudocercospora

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.

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 pseudostigmina-platani 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.

INTRODUCTION

Until recently, Pseudocercospora was treated as an anamorphic genus linked to Mycosphaerella (Mycosphaerellaceae, Capnodiales), along with approximately 30 other anamorphic 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 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.

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.

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).

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).

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).

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).

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.

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).

Table 1.

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

Species	Culture accession numbers1	Collector	Host	Family	Country	GenBank accession numbers2
						LSU	ITSy	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
	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
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
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	Genome3	Genome3
	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
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
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
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 × yedoensis	Rosaceae	South Korea	GU253723	GU269676	GU384393	GU320382
Pseudocercospora pseudostigmina-platani	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
	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
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
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

¹CBS: 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.

²LSU: 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.

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

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™ 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™ 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™ 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 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 branch-swapping 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 well-supported 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.

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).

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).

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).

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 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.

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 pseudocercospora-like 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.

Treatment of species

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, 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.

Foliicolous, phytopathogenic, causing discrete leaf spots. 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 the 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 the 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, hyphopodia-like 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.

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.

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.

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 the 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 the 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 the 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. 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 two-layered. 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 rough-walled, 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 heterosporium-like 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.

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.

Fig. 7.

Microcyclospora quercina (CPC 10712). Line drawing showing conidiogenous cells and conidia formed in culture. Scale bar = 10 μm.

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.

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 olivaceous-black 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 ellipsoid-fusoid, 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, 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).

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 zasmidium-like (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 pseudocercospora-like

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.

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.

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 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.

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.

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 °C 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.

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.

• ≡ 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 obclavate-cylindrical 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. nov. MycoBank MB564817. Fig. 12.

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.

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 the 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 °C in the dark on MEA; erumpent, spreading, with sparse aerial mycelium, and 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.

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.

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.

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 geniculate-sinuous, 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.

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.

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.

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.

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 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 the 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 °C 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 Ps. 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.

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.

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 μm high; conidiophores hyaline or pale brown at the 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 the 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 °C 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, Ps. 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 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.

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.

Etymology: The name reflects the pale brown cercospora-like conidia in this genus.

Foliicolous, phytopathogenic, causing discrete leaf spots. 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, 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 & M.J. Wingf., comb. nov. MycoBank MB564822.

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. MycoBank MB564823. Fig. 17.

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 & M.J. Wingf., comb. nov. MycoBank MB564824.

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.

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.

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.

Specimen examined: Australia, Queensland, Mareeba, S16°58’75.5” E145°20’60.8”, 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.

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.

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 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 well-developed. 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.

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.

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. Shin, sp. nov. MycoBank MB564828. Fig. 21.

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.

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.

Culture characteristics: Colonies after 2 wk at 24 °C in the dark 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, 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, 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.

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.

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.

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.

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.

Pseudocercospora balsaminae (Syd.) Deighton, Mycol. Pap. 140: 139. 1976. Fig. 24.

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.

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.

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.

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, grey-brown 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 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.

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.

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.

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.

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.

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.

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.

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 = CBS 132108.

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. (as “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, ex-holotype 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. nov. MycoBank MB564830. Fig. 30.

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.

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 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 the 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.

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.

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 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.

Culture characteristics: Colonies on MEA reaching 53 mm diam 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.

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.

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.

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.

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 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.

Culture characteristics: Colonies on MEA reaching 32 mm diam 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 olivaceous-grey (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 = CBS 114242.

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.

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.

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. 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.

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.

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.

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.

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. MycoBank MB564832. Fig. 37.

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.

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 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.

Culture characteristics: Colonies after 2 wk at 24 °C in the dark on 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.

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. Shin, sp. nov. MycoBank MB564834. Fig. 38.

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.

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.

Culture characteristics: Colonies after 2 wk at 24 °C in the dark on MEA; surface folded, erumpent, spreading, with sparse aerial mycelium, and smooth, lobate margins. Surface pale olivaceous-grey; 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 obclavate-cylindrical, 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.

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.

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.

Pseudocercospora kaki Goh & W.H. Hsieh, in Hsieh & Goh, Cercospora and similar fungi from Taiwan: 109. 1990. Fig. 40.

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.

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, S34°23’52.1” E19°15’58.5”, K. africana, 2 May 2010, P.W. Crous, CBS H-20873, CPC 18286, 18287.

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.) Deighton, Trans. Brit. Mycol. Soc. 88: 389. 1987.

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.

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.

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 = CBS 131925.

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.

Pseudocercospora lythri H.D. Shin & U. Braun, Mycotaxon 74: 111. 2000. Fig. 42.

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.

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.

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.

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.

Culture characteristics: Colonies after 2 wk at 24 °C in the dark on MEA; erumpent, spreading, with moderate aerial mycelium, and smooth, even margins. Surface pale olivaceous-grey; reverse olivaceous-grey. Colonies reaching 10 mm diam.

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.

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.

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.

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.

Etymology: Name derived from Ocimum basilicum, the host from which it was collected.

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 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 (as “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 geniculate-sinuous, 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 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.

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 conidiophores and conidiogenous cells. E. Conidia. Scale bars = 10 μm.

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.

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.

Leaf spots amphigenous, irregular to subcircular, 3-7 mm diam, 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 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.

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 and conidiogenous cells. E. Conidia. Scale bars = 10 μm.

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) Deighton, Trans. Brit. Mycol. Soc. 88: 390. 1987.

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. Braun & H.D. Shin, sp. nov. MycoBank MB564839. Fig. 49.

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.

Etymology: Name derived from the host genus Plectranthus, from which it was collected.

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.

Culture characteristics: Colonies after 2 wk at 24 °C in the dark on MEA; erumpent, spreading, with moderate aerial mycelium, and smooth, lobate margins. Surface pale olivaceous-grey; reverse iron-grey. Colonies reaching 8 mm diam.

Specimen examined: South Korea, Jeonju, on leaves of Plectranthus sp., 1 July 2004, H.D. Shin, holotype CBS H-20396, cultures ex-type CPC 11462 = CBS 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.

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.

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.

Pseudocercospora proteae Crous, sp. nov. MycoBank MB564840. Fig. 51.

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.

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 the 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.

Culture characteristics: Colonies after 2 wk at 24 °C in the dark on MEA; erumpent, spreading, with sparse aerial mycelium, and smooth, even margins. Surface olivaceous-grey; reverse iron-grey. Colonies reaching 10 mm diam.

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.

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.

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.

  • ≡ 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 × 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. Braun & H.D. Shin, sp. nov. MycoBank MB564841. Fig. 53.

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.

Etymology: Name reflects its morphological similarity to the Pseudocercospora anamorph of Mycosphaerella stigmina-platani. Leaf spots amphigenous, irregular to subcircular, 5-10 mm diam, 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 proliferations, 8-20 × 5-8 μm. Conidia dimorphic: cercostigmina-like 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.

Culture characteristics: Colonies after 2 wk at 24 °C in the dark on MEA; surface folded, erumpent, spreading, with sparse aerial mycelium, and smooth, lobate margins. Surface pale olivaceous-grey, 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 & H.D. Shin, sp. nov. MycoBank MB564842. Fig. 54.

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.

Etymology: Name derived from the host plant Pyracantha, from which it was collected.

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.

Culture characteristics: Colonies after 2 wk at 24 °C in the dark on MEA; surface folded, erumpent, spreading, with sparse aerial 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.

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.

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.

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 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. nov. MycoBank MB564843. Fig. 56.

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.

Etymology: Name derived from the plant host Ravenala, from which this fungus was isolated.

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 °C in the dark on MEA; erumpent, spreading, with moderate aerial mycelium, and smooth, lobate margins. Surface smoke-grey in centre, pale olivaceous-grey in outer region; reverse olivaceous-grey. Colonies reaching 35 mm diam.

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 Ps. rhabdothamni.

Pseudocercospora rhamnellae G.C. Hunter, H.D. Shin, U. Braun & Crous, sp. nov. MycoBank MB564844. Fig. 57.

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.

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 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.

Culture characteristics: Colonies after 2 wk at 24 °C in the dark on MEA; surface folded, erumpent, spreading, with sparse aerial mycelium, and smooth, lobate margins. Surface olivaceous-grey with patches of pale olivaceous-grey; reverse iron-grey. Colonies reaching 10 mm diam.

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 & H.D. Shin, sp. nov. MycoBank MB564845. Fig. 58.

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 conidiophores and conidiogenous cells. E. Conidia. Scale bars = 10 μm.

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.

Culture characteristics: Colonies after 2 wk at 24 °C in the dark on MEA; erumpent, spreading, with moderate aerial mycelium, and smooth, lobate margins. Surface olivaceous-grey in centre, pale olivaceous-grey in outer region; reverse iron-grey. Colonies reaching 14 mm diam.

Specimen examined: South Korea, Seoul, on Rhododendron indicum, 27 Oct. 2003, H.D. Shin, holotype CBS H-20886, cultures ex-type CPC 10822 = CBS 131591, CPC 10823, 10824.

Notes: Of the species occurring on Rhododendron, P. rhododendri-indici 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 rhododendri-indici 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.

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.

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.

Pseudocercospora sambucigena U. Braun, Crous & K. Schub., Mycotaxon 92: 400. 2005. Fig. 60.

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.

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, 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.

Culture characteristics: Colonies on MEA reaching 16 mm diam 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.

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 and conidiogenous cells. E. Conidia. Scale bars = 10 μm.

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.

Fig. 62.

Pseudocercospora snelliana (B 700014740, holotype). Sparse fascicles, and solitary conidiophores on superficial mycelium giving rise to muriformly septate, thick-walled 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.

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; hila neither thickened, nor darkened or refractive, 1-1.5(-2) μm diam.

Culture characteristics: Colonies after 2 wk at 24 °C in the dark on MEA; erumpent, spreading, with sparse aerial mycelium, and smooth, lobate margins. Surface pale olivaceous-grey; reverse olivaceous-grey. Colonies reaching 7 mm 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.

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.

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.

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.

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.

Culture characteristics: Colonies on MEA reaching 23 mm diam 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. agnus-castus, 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.

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.

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.

Etymology: Name derived from the plant host Xanthocercis, from which it was collected.

Leaf spots amphigenous, irregular to subcircular, 3-8 mm diam, 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.

Culture characteristics: Colonies after 2 wk at 24 °C in the dark on 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. Colonies reaching 5 mm diam.

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.

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.

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.

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.

Basionym: Cercospora zelkowae Hori, Nambu N. Jour. Plant Protection 8: 492. 1921.

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 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”.