Phylogeny and taxonomy of obscure genera of microfungi - Persoonia

Persoonia 22, 2009: 139–161 

www.persoonia.org 

RESEARCH ARTICLE 

doi:10.3767/003158509X461701 

Phylogeny and taxonomy of obscure genera of microfungi 

P.W. Crous 1 , U. Braun 2 , M.J. Wingfield 3 , A.R. Wood 4 , H.D. Shin 5 , B.A. Summerell 6 , 

A.C. Alfenas 7 , C.J.R. Cumagun 8 , J.Z. Groenewald 1 

Key words 

Brycekendrickomyces 

Chalastospora 

Cyphellophora 

Dictyosporium 

Edenia 

phylogeny 

taxonomy 

Thedgonia 

Trochophora 

Verrucisporota 

Vonarxia 

Xenostigmina 

Abstract The recently generated molecular phylogeny for the kingdom Fungi, on which a new classification scheme 

is based, still suffers from an under representation of numerous apparently asexual genera of microfungi. In an 

attempt to populate the Fungal Tree of Life, fresh samples of 10 obscure genera of hyphomycetes were collected. 

These fungi were subsequently established in culture, and subjected to DNA sequence analysis of the ITS and LSU 

nrRNA genes to resolve species and generic questions related to these obscure genera. Brycekendrickomyces 

(Herpotrichiellaceae) is introduced as a new genus similar to, but distinct from Haplographium and Lauriomyces. 

Chalastospora is shown to be a genus in the Pleosporales, with two new species, C. ellipsoidea and C. obclavata, 

to which Alternaria malorum is added as an additional taxon under its oldest epithet, C. gossypii. Cyphellophora 

eugeniae is newly described in Cyphellophora (Herpotrichiellaceae), and distinguished from other taxa in the genus. 

Dictyosporium is placed in the Pleosporales, with one new species, D. streliziae. The genus Edenia, which was 

recently introduced for a sterile endophytic fungus isolated in Mexico, is shown to be a hyphomycete (Pleosporales) 

forming a pyronellea-like synanamorph in culture. Thedgonia is shown not to represent an anamorph of 

Mycosphaerella, but to belong to the Helotiales. Trochophora, however, clustered basal to the Pseudocercospora 

complex in the Mycosphaerellaceae, as did Verrucisporota. Vonarxia, a rather forgotten genus of hyphomycetes, 

is shown to belong to the Herpotrichiellaceae and Xenostigmina is confirmed as synanamorph of Mycopappus, 

and is shown to be allied to Seifertia in the Pleosporales. Dichotomous keys are provided for species in the various 

genera treated. Furthermore, several families are shown to be polyphyletic within some orders, especially in the 

Capnodiales, Chaetothyriales and Pleosporales. 

Article info Received: 2 April 2009; Accepted: 23 April 2009; Published: 9 June 2009. 

Introduction 

The recent ‘Deep Hypha’ issue of Mycologia (vol. 98, 2006) 

included 21 phylogenetic studies employing multi-gene phylogenies 

to resolve major groups of Fungi. These papers provided 

the foundation for the study of James et al. (2006), in which 

six genes (SSU, LSU, 5.8S rRNA, rpb1, rpb2 and tef1) for 

approximately 200 fungal taxa were used to present the first 

kingdom-level phylogeny, and a new classification for the Fungi 

(Hibbett et al. 2007). These studies also illustrated clearly that 

it was merely the ‘tip of the iceberg’, and that numerous genera 

must now be accommodated in this phylogenetic framework. 

A major problem encountered during the Assembling the 

Fungal Tree of Life (AFTOL, www.aftol.org) project, was that 

many genera are insufficiently known, and have never been 

cultured, or subjected to DNA analyses. This is especially true 

for the majority of apparently asexual microfungi, namely the 

1 

CBS Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The 

Netherlands; corresponding author e-mail: p.crous@cbs.knaw.nl. 

2 

Martin-Luther-Universität, Institut für Biologie, Bereich Geobotanik und 

Botanischer Garten, Herbarium, Neuwerk 21, D-06099 Halle (Saale), 

Germany. 

3 

Department of Genetics, Forestry and Agricultural Biotechnology Institute 

(FABI), University of Pretoria, Pretoria 0002, South Africa. 

4 

ARC – Plant Protection Research Institute, P. Bag X5017, Stellenbosch, 

7599, South Africa. 

5 

Division of Environmental Science & Ecological Engineering, Korea University, 

Seoul 136-701, Korea. 

6 

Royal Botanic Gardens and Domain Trust, Mrs. Macquaries Road, Sydney, 

NSW 2000, Australia. 

7 

Departamento de Fitopatologia, Universidade Federal de Viçosa, 36.570 

Viçosa, MG, Brazil. 

8 

Crop Protection Cluster, College of Agriculture, University of the Philippines, 

Los Baños College, Laguna 4031, Philippines. 

coelomycetes (Sutton 1980, Nag Raj 1993) and hyphomycetes 

(Ellis 1971, 1976, Carmichael et al. 1980). The only means to 

deal with this problem is, therefore, to encourage mycologists 

to recollect these genera and species, to establish cultures for 

them and to ultimately generate DNA sequence data (Shenoy 

et al. 2007), a process which can be described as ‘leafing out 

the fungal tree of life’. 

Ten genera of hyphomycetes not previously known from culture, 

or for which the phylogenetic classification is uncertain, 

are treated in the present study. These fungi were collected 

from diverse hosts from various continents, isolated in axenic 

culture, and subjected to DNA sequence analysis. They are 

shown to belong to the Chaetothyriales (Brycekendrickomyces, 

Cyphellophora, Vonarxia), Pleosporales (Chalastospora, Dictyosporium, 

Edenia, Xenostigmina), Helotiales (Thedgonia), and 

the Capnodiales, Mycosphaerellaceae (Trochophora, Verrucisporota). 

The present paper represents a further contribution in a series 

aiming to clarify the morphology and DNA phylogeny of obscure 

genera of microfungi. Other than resolving their phylogenetic 

relationships, several novelties are described, and keys are 

provided to the accepted species in these genera. 

Material and Methods 

Isolates 

Symptomatic leaves and leaf litter were collected on various 

continents, and sent to the Centraalbureau voor Schimmelcultures 

(CBS) for isolation of microfungi. Leaves with visible fruiting 

were immediately subjected to direct isolation of hyphomycetes, 

or alternatively were first incubated in moist chambers to 

© 2009 Nationaal Herbarium Nederland & Centraalbureau voor Schimmelcultures 

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140 Persoonia – Volume 22, 2009 

Table 1 Collection details and GenBank accession numbers for fungal species included in this study. 

Species Strain no. 1 Substrate Country Collector(s) GenBank Accession no. 2 

ITS LSU 

Brycekendrickomyces acaciae CBS 124104; CPC 15078 Acacia auriculiformis Indonesia M.J. Wingfield FJ839606 FJ839641 

Chalastospora cetera CBS 121340; E.G.S. 41.072 Elymus scabrus Australia R.G. Rees FJ839607 FJ839642 

Chalastospora ellipsoidea CBS 121331; E.G.S. 22.060 Triticum sp. Australia H.L. Harvey & S. Perth FJ839608 FJ839643 

Chalastospora gossypii CBS 112844; CPC 4571 Bromus tectorum USA F.M. Dugan AY251081 AY251081 

CBS 114005; CPC 4572 Festuca idahoensis USA F.M. Dugan AY251079 AY251079 

CBS 114809; MAF 943 Leaves of Anethum graveolens (dill) along with Itersonilia perplexans New Zealand J. Pike FJ839609 FJ839644 

CBS 114810; MAF 954 Quercus robur (oak) leaves in association with Tubakia dryina New Zealand H. Nettleton FJ839610 FJ839645 

CBS 148.66; CPC 3690; NRRL W 52-29 — USA C.W. Hesseltine FJ839611 FJ839646 

CBS 173.80; ATCC 200939; CPC 3685 Agricultural soil Syria M.I.A. Abdel-Kader — FJ839647 

CBS 216.65; NRRL A-13702 Triticum aestivum grain USA C.W. Hesseltine FJ839612 DQ008142 

CBS 266.75; ATCC 28332; CPC 3680; IMI 165252; PRE 44703 Wheat stubble South Africa W.F.O. Marasas FJ839613 FJ839648 

CBS 900.87; ATCC 200938 Soil Lebanon F. Seigle-Murandi FJ839614 FJ839649 

CPC 15567; C.F. Hill 2008/3899 Wood and wallpaper from inside walls of a dwelling New Zealand D. De Vanny FJ839615 FJ839650 

Chalastospora gossypii var. polymorpha CBS 112048; CPC 4570 Dormant buds (overwintered) of Vitis vinifera USA F.M. Dugan AY251080 AY251080 

Chalastospora obclavata CBS 124120; E.G.S. 12.128 Air USA C.T. Rogerson FJ839616 FJ839651 

Cyphellophora eugeniae CBS 124105; CPC 15172 Living leaves of Stenocalyx uniflorus Brazil A.C. Alfenas FJ839617 FJ839652 

Dictyosporium strelitziae CBS 123359; CPC 15359 Dead leaves of Strelitzia nicolai South Africa A. Wood FJ839618 FJ839653 

Edenia gomezpompae CBS 124106; CPC 15689 Senna alata Phillippines C.J.R. Cumagun FJ839619 FJ839654 

Haplographium catenatum CBS 196.73 Decaying wood Germany W. Gams FJ839620 FJ839655 

CBS 482.67; CMW 754 Decaying wood Germany W. Gams FJ839621 FJ839656 

CBS 739.68; CMW 755 Decaying wood Netherlands H.A. van der Aa FJ839622 FJ839657 

Lauriomyces bellulus CBS 517.93 Cupule of Castanea sativa Switzerland P.W. Crous FJ839623 FJ839658 

Lauriomyces heliocephalus CBS 112054; INIFAT CO2/59 Decaying leaf Brazil A. Stchigel & J. Guarro FJ839624 FJ839659 

Mycopappus aceris CBS 124109; CPC 14379 Fallen leaves of Acer macrophyllum Canada B. Callan FJ839625 FJ839660 

Mycosphaerella lupini CPC 1661 Lupinus sp. USA W. Kaiser AF362050 FJ839661 

Stenella anthuriicola CBS 118742 Anthurium leaf Thailand C. F. Hill FJ839626 FJ839662 

Stigmina platani CBS 110755; CPC 4299; IMI 136770 Platanus orientalis India — AY260090 FJ839663 

Thedgonia ligustrina CPC 10019 Ligustrum ovalifolium South Korea H.-D. Shin FJ839627 FJ839664 

CPC 10530 Ligustrum ovalifolium Netherlands P.W. Crous FJ839628 FJ839665 

CPC 10861 Ligustrum ovalifolium South Korea H.-D. Shin FJ839629 FJ839666 

CPC 14754 Ligustrum obtusifolium South Korea H.-D. Shin FJ839630 FJ839667 

CPC 4298; W1877 Ligustrum sp. Asia H. Evans EU040242 EU040242 

Trochophora fasciculata CPC 10281 Leaves of Daphniphyllum macropodum South Korea H.-D. Shin FJ839631 — 

CPC 10282 Leaves of Daphniphyllum macropodum South Korea H.-D. Shin FJ839632 FJ839668 

Verrucisporota daviesiae CBS 116002; VPRI 31767 Living leaves of Daviesia mimosoides Australia V. & R. Beilharz FJ839633 FJ839669 

Verrucisporota grevilleae CBS 124107; CPC 14761 Leaves of Grevillia decurrens Australia B. Summerell FJ839634 FJ839670 

Verrucisporota proteacearum CBS 116003; VPRI 31812 Grevillea sp. Australia V. Beilharz FJ839635 FJ839671 

Vonarxia vagans CBS 123533; CPC 15151 Stenocalyx uniflorus Brazil A.C. Alfenas FJ839636 FJ839672 

CPC 15152 Stenocalyx uniflorus Brazil A.C. Alfenas FJ839637 FJ839673 

Xenostigmina zilleri CBS 115685; CPC 4011 Living leaves of Acer sp. Canada K.A. Seifert FJ839638 FJ839674 

CBS 124108; CPC 14376 Fallen leaves of Acer macrophyllum Canada B. Callan FJ839639 FJ839675 

CBS 115686; CPC 4010 Living leaves of Acer sp. Canada K.A. Seifert FJ839640 FJ839676 

1 

ATCC: American Type Culture Collection, Virginia, USA; C.F. Hill: Culture collection of C.F. Hill, housed at MAF, New Zealand; CBS: CBS Fungal Biodiversity Centre, Utrecht, The Netherlands; CMW: Culture collection of M.J. Wingfield, housed at FABI, Pretoria, South Africa; CPC: 

Culture collection of P.W. Crous, housed at CBS; E.G.S.: Culture collection of E.G. Simmons, Indiana USA; IMI: International Mycological Institute, CABI-Bioscience, Egham, Bakeham Lane, U.K.; INIFAT: Alexander Humboldt Institute for Basic Research in Tropical Agriculture, 

Ciudad de La Habana, Cuba; MAF: Ministry of Agriculture and Forestry, New Zealand; NRRL: National Center for Agricultural Utilization Research, Peoria, USA; PRE: National collection of fungi, Pretoria, South Africa; VPRI: Victorian Department of Primary Industries, Knoxfield, 

Australia. 

2 

ITS: Internal transcribed spacers 1 and 2 together with 5.8S nrDNA; LSU: 28S nrDNA.

P.W. Crous et al.: Obscure genera of microfungi 

141 

stimulate sporulation. Single-conidial isolates were established 

on malt extract agar (MEA; 20 g/L Biolab malt extract, 15 g/L 

Biolab agar) using the technique outlined in Crous (1998). 

Cultures were later plated on fresh MEA, 2 % water agar (WA) 

supplemented with sterile pine needles, 2 % potato-dextrose 

agar (PDA), synthetic nutrient agar (SNA) and/or oatmeal agar 

(OA) (Crous et al. 2009), and subsequently incubated at 25 °C 

under near-ultraviolet light to promote sporulation. Reference 

strains are maintained in the culture collection of the CBS, 

Utrecht, the Netherlands (Table 1). Descriptions, nomenclature, 

and illustrations were deposited in MycoBank (www.mycobank. 

org, Crous et al. 2004b). 

DNA isolation, amplification and analyses 

Genomic DNA was isolated from fungal mycelium grown on 

MEA, using the UltraClean TM Microbial DNA Isolation Kit (Mo 

Bio Laboratories, Inc., Solana Beach, CA, USA) according to the 

manufacturer’s protocols. The Primers V9G (de Hoog & Gerrits 

van den Ende 1998) and LR5 (Vilgalys & Hester 1990) were 

used to amplify part of the nuclear rDNA operon spanning the 3’ 

end of the 18S rRNA gene (SSU), the first internal transcribed 

spacer (ITS1), the 5.8S rRNA gene, the second ITS region 

(ITS2) and the first 900 bases at the 5’ end of the 28S rRNA 

gene (LSU). The primers ITS4 (White et al. 1990) and LR0R 

(Rehner & Samuels 1994) were used as internal sequence 

primers to ensure good quality sequences over the entire length 

of the amplicon. The PCR conditions, sequence alignment 

and subsequent phylogenetic analysis followed the methods 

of Crous et al. (2006b). Alignment gaps were treated as new 

character states. Sequence data were deposited in GenBank 

(Table 1) and alignments in TreeBASE (www.treebase.org). 

The ITS sequences were compared with those sequences available 

in NCBI’s GenBank nucleotide database using a megablast 

search and the results are discussed where applicable under 

the taxonomic notes. Because the genus Chalastospora is relatively 

novel, species in this genus were supported by a separate 

phylogenetic tree. 

Morphology 

Fungal descriptions were based on cultures sporulating in 

vitro (media indicated). Wherever possible, 30 measurements 

(× 1 000 magnification) were made of structures mounted in 

lactic acid, with the extremes of spore measurements given in 

parentheses. Colony colours (surface and reverse) were assessed 

after 2–4 wk on different media at 25 °C in the dark, 

using the colour charts of Rayner (1970). 

Results 

Phylogenetic analysis 

Amplification products of approximately 1 700 bases were obtained 

for the isolates listed in Table 1. The LSU region of the 

sequences was used to obtain additional sequences from Gen- 

Bank, which were added to the alignment. Due to the inclusion 

of the shorter LSU sequences of Dictyosporium alatum (Gen- 

Bank accession DQ018101), Dictyosporium elegans (GenBank 

accession DQ018100) and Dictyosporium toruloides (GenBank 

accession DQ018104) in the alignment, it was not possible to 

subject the full length of the determined LSU sequences (Table 

1) to analyses. The manually adjusted LSU alignment contained 

115 sequences (including the two outgroup sequences) and, of 

the 568 characters used in the phylogenetic analyses, 267 were 

parsimony informative, 30 were variable and parsimony uninformative, 

and 271 were constant. Neighbour-joining analyses 

using three substitution models on the sequence data yielded 

trees supporting the same tree topology to one another but 

differed from the most parsimonious tree shown in Fig. 1 with 

regard to the placement of the clade containing Ochroconis and 

Fusicladium (in the distance analyses, this clade moves to a 

more basal position). Forty equally most parsimonious trees 

(TL = 1 039 steps; CI = 0.477; RI = 0.833; RC = 0.397), the first 

of which is shown in Fig. 1, were obtained from the parsimony 

analysis of the LSU alignment. 

The manually adjusted ITS alignment contained 28 sequences 

(including the outgroup sequence) and, of the 521 characters 

used in the phylogenetic analyses, 97 were parsimony informative, 

91 were variable and parsimony uninformative, and 333 

were constant. Neighbour-joining analyses using three substitution 

models on the sequence data yielded trees supporting the 

same tree topology to one another but differed from the most 

parsimonious tree shown in Fig. 2 with regard to the placement 

of Chalastospora ellipsoidea (in the distance analyses, this 

taxon moves to a more basal position in Chalastospora). Six 

equally most parsimonious trees (TL = 253 steps; CI = 0.913; 

RI = 0.938; RC = 0.856), the first of which is shown in Fig. 2, 

were obtained from the parsimony analysis of the ITS alignment. 

The results of the phylogenetic analyses are highlighted 

below under the taxonomic notes, or in the Discussion, where 

applicable. 

Taxonomy 

Brycekendrickomyces Crous & M.J. Wingf., gen. nov. — 

MycoBank MB509515 

Mycelium ex hyphis ramosis, septatis, laevibus, pallide brunneis, 1–2 µm 

latis compositum. Conidiophora solitaria, erecta, cylindrica, recta vel leviter 

flexuosa, cellula basali bulbosa, sine rhizoideis, stipite modice brunneo vel 

atro-brunneo, laevi, transverse euseptato, ad apicem cum (1–)2–4(–6) 

cellulis conidiogenis. Cellulae conidiogenae subcylindricae, allontoides vel 

doliiformes, rectae vel leviter curvatae, pallide brunneae, polyblasticae, 

sympodialiter proliferantes. Conidia hyalina, mucilagine aggregata (sed non 

catenata), ellipsoidea, apice subobtuso, basi subtruncata. 

Type species. Brycekendrickomyces acaciae Crous & M.J. Wingf. 

Etymology. Named for Bryce Kendrick, husband of Laurie Kendrick, for 

which Lauriomyces was named and that resembles the current genus. 

Mycelium consisting of branched, septate, smooth, pale brown, 

1–2 µm wide hyphae. Conidiophores solitary, erect, cylindrical, 

straight to somewhat flexuous, basal cell bulbous, without 

rhizoids; stalk medium to dark brown, smooth, transversely 

euseptate; upper cell giving rise to (1–)2–4(–6) conidiogenous 

cells. Conidiogenous cells subcylindrical to allantoid or doliiform, 

straight to gently curved, pale brown, polyblastic, proliferating 

sympodially. Conidia hyaline, aggregating in slimy mass (never 

in chains), ellipsoid, apex subobtuse, base subtruncate. 

Brycekendrickomyces acaciae Crous & M.J. Wingf., sp. nov. 

— MycoBank MB509517; Fig. 3 

Maculae modice brunneae vel atro-brunneae, margine elevato, rubro-purpureo, 

oblongae vel ellipticae, ad 7 mm diam, in consortione ‘Phaeotrichoconis’ 

crotalariae. In vitro (MEA): Mycelium ex hyphis ramosis, septatis, 

laevibus, pallide brunneis, 1–2 µm latis compositum. Conidiophora ex 

hyphis oriunda, solitaria, erecta, cylindrica, recta vel leviter flexuosa, cellula 

basali bulbosa, sine rhizoideis, 4–6 µm lata, ad basim 10–15 µm lata, 

stipite modice brunneo vel atro-brunneo, laevi, transverse 2–5-euseptato, 

(15–)30–50(–60) µm longo, (3–)4(–5) µm lato, ad apicem cum (1–)2–4(–6) 

cellulis conidiogenis. Cellulae conidiogenae subcylindricae, allontoides vel 

doliiformes, rectae vel leviter curvatae, pallide brunneae, 5–8 × 2–2.5 µm, 

polyblasticae, sympodialiter proliferantes. Conidia hyalina, mucilagine aggregata 

(sed non catenata), ellipsoidea, apice subobtuso, basi subtruncata, 

latitudine maxima in parte centrali vel in parte supra centrum, saepe leviter 

asymmetrica, (3.5–)4(–4.5) × 2(–2.5) µm. 

Etymology. Named after the host genus on which the fungus occurs, 

Acacia.


Kluyveromyces lodderae AY048161 

Saccharomyces cerevisiae Z73326 

82 

10 changes 

62 

1 

100 

2 

99 

Mycosphaerella punctiformis EU167569 

Ramularia pratensis var. pratensis EU019284 

100 

99 

74 

56 

64 

97 

76 

97 

51 

Trochophora fasciculata CPC 10282 

Mycosphaerella crystallina DQ204747 

Mycosphaerella waimeana AY260083 

Mycosphaerella heimii DQ204751 

Mycosphaerella heimioides DQ204753 

Pseudocercospora vitis DQ073923 

Pseudocercospora pseudoeucalyptorum DQ204766 

Pseudocercospora robusta DQ204767 

Stigmina platani CBS 110755 

Pseudocercospora natalensis DQ267576 

Pseudocercospora basitruncata DQ204759 

Mycosphaerella lupini CPC 1661 

Mycosphaerella marksii DQ246250 

Pseudocercospora epispermogonia DQ204758 

100 

Schizothyrium pomi EF134947 

Schizothyrium pomi EF134949 SCH 

69 Schizothyrium pomi EF134948 

Cibiessia minutispora EU019259 

82 100 Teratosphaeria readeriellophora DQ246238 

88 Batcheloromyces leucadendri EU019246 

96 Cibiessia nontingens EU019260 

Teratosphaeria nubilosa DQ246228 

95 Teratosphaeria fibrillosa EU019282 

Mycosphaerella madeirae DQ204756 

Mycosphaerella parkii DQ246245 

Ramichloridium biverticillatum EU041853 

Verrucisporota grevilleae CPC 14761 

97 99 Verrucisporota proteacearum CBS 116003 

83 Periconiella arcuata EU041836 

Verrucisporota daviesiae CBS 116002 

Periconiella velutina EU041840 

MYC (2) 

54 

Ramichloridium musae EU041857 

Ramichloridium strelitziae EU041860 

54 Stenella anthuriicola CBS 118742 

Ramichloridium cerophilum EU041855 

77 

85 

Zasmidium cellare EU041878 

Cyphellophora laciniata EU035416 HER (1) 

Vonarxia vagans CPC 15151 

80 100 Vonarxia vagans CPC 15152 

CHA 

Ceramothyrium carniolicum AY004339 

Cyphellophora Cyphellophora eugeniae hylomeconis CPC EU035415 

15172 

78 

Exophiala eucalyptorum EU035417 

71 Exophiala pisciphila DQ823101 

Capronia mansonii AY004338 

100 Capronia munkii EF413604 

97 Brycekendrickomyces acaciae CPC 15078 

63 Phaeococcomyces chersonesos AJ507323 

65 Cladophialophora proteae EU035411 

100 Exophiala placitae EU040215 

Phaeococcomyces catenatus AF050277 

88 

70 

MYC (1) 

Fig. 1 The first of 1 000 equally most parsimonious trees obtained from a heuristic search with 100 random taxon additions of the LSU sequence alignment. 

The scale bar shows 10 changes, and bootstrap support values from 1 000 replicates are shown at the nodes. Novel sequences generated for this study are 

shown in bold. Branches present in the strict consensus tree are thickened. Orders and families are coded as indicated in the legends. The tree was rooted 

to a sequence of Kluyveromyces lodderae (GenBank accession AY048161) and Saccharomyces cerevisiae (GenBank accession Z73326). 

Abbreviations used: Families: AMO = Amorphothecaceae, CHA = Chaetothyriaceae, HEL = Helotiaceae, HER = Herpotrichiellaceae, HYA = Hyaloscyphaceae, 

IC = Incertae cedis: LEP = Leptosphaeriaceae, LOP = Lophiostomataceae, MEL = Melanommataceae, MYC = Mycosphaerellaceae, PHA = Phaeosphaeriaceae, 

PLE = Pleosporaceae, PSE = Pseudeurotiaceae, RHY = Rhytismataceae, SCH = Schizothyriaceae, TER = Teratosphaeriaceae. Orders: 1 = Capnodiales, 

2 = Chaetothyriales, 3 = Incertae cedis, 4 = Rhytismatales, 5 = Helotiales, 6 = Hypocreales, 7 = Pezizales, 8 = Pleosporales. 

TER 

HER (2) 

Leaf spots medium to dark brown, margin raised, red-purple, 

oblong to ellipsoid, up to 7 mm diam, associated with ‘Phaeotrichoconis’ 

crotalariae. Description based on culture on MEA: 

Mycelium consisting of branched, septate, smooth, pale brown, 

1–2 µm wide hyphae. Conidiophores arising from mycelium, 

solitary, erect, cylindrical, straight to somewhat flexuous; basal 

cell bulbous, without rhizoids, 4–6 µm wide in upper part, but 

becoming 10–15 µm wide at basal part; stalk medium to dark 

brown, smooth, transversely 2–5-euseptate, (15–)30–50(–60) 

µm tall, (3–)4(–5) µm wide in the middle part; upper cell giving 

rise to (1–)2–4(–6) conidiogenous cells. Conidiogenous cells 

subcylindrical to allantoid or doliiform, straight to gently curved, 

pale brown, 5–8 × 2–2.5 µm; polyblastic, proliferating sympodially. 

Conidia hyaline, aggregating in slimy mass (never in 

chains), ellipsoid, apex subobtuse, base subtruncate, widest in 

the middle or upper third of the conidium, frequently somewhat 

asymmetrical, (3.5–)4(–4.5) × 2(–2.5) µm. 

Characteristics in culture — Colonies on MEA erumpent, 

spreading, with moderate aerial mycelium; surface folded, 

margin lobate, smooth; surface olivaceous-grey, outer margin 

iron-grey; reverse iron-grey; colonies reaching up to 20 mm 

after 1 mo. Colonies fertile on SNA, OA and MEA. 

Specimen examined. Indonesia, Pelalawan, living leaves of Acacia auriculiformis, 

Mar. 2008, leg. M.J. Wingfield, isol. P.W. Crous, holotype CBS 

H-20198, culture ex-type CPC 15078 = CBS 124104. 

Notes — Castañeda & Kendrick (1990) established the genus 

Lauriomyces, characterised by dark brown conidiophores,


143 

Fig. 1 (cont.) 

3 Lauriomyces bellulus CBS 517.93 

IC 

4 100 Lauriomyces heliocephalus CBS 112054 

Potebniamyces pyri DQ470949 

RHY 

5 Crinula caliciiformis AY544680 

HEL (1) 

100 

Holwaya mucida DQ257356 

Leuconeurospora pulcherrima AF096193 

61 6 

IC 

Pseudeurotium zonatum AF096198 

933 

PSE 

Pseudeurotium zonatum DQ470988 

3 97 

Amorphotheca resinae EU040230 

Amorphotheca resinae EU040231 

AMO 

100 Hyalodendriella betulae EU040232 

IC 

Phialea strobilina EF596821 

5 7 100 Haplographium catenatum CBS 482.67 

Haplographium catenatum CBS 196.73 

HYA (1) 

Haplographium catenatum CBS 739.68 

3 Clathrosporium intricatum AY616235 

66 Hydrocina chaetocladia AY789412 

Rhyzoscyphus ericae AM887699 

IC 

HEL (2) 

89 

Protoventuria alpina EU035444 

Thedgonia ligustrina CPC 10019 

100 

Thedgonia ligustrina EU040242 

Thedgonia ligustrina CPC 10861 IC 


3 


Trichoconis echinophila EU107315 

91 Trematosphaeria pertusa DQ678072 

10 changes 

8 

100 

100 

88 

Aposphaeria populina EU754130 

Herpotrichia juniperi DQ678080 

Wettsteinina macrotheca AY849969 

Seifertia azaleae EU030276 

Mycopappis aceris CPC 14379 

70 

Xenostigmina zilleri CBS 115685 

Xenostigmina zilleri CBS 115686 

Xenostigmina zilleri CPC 14376 

100 

Dictyosporium toruloides DQ018104 

Dictyosporium strelitziae CPC 15359 

Dictyosporium elegans DQ018100 

Dictyosporium alatum DQ018101 

91 

Pseudodiplodia sp. EU754201 

Coniothyrium concentricum EU754152 

Leptospora rubella DQ195792 

Phaeosphaeriopsis musae DQ885894 

HYA (2) 

MEL 

LOP 

PLE (1) 

60 

LEP 

IC 

PHA 

Phaeosphaeria nodorum EF590318 

Edenia gomezpompae CPC 15689 

Chalastospora gossypii CBS 112844 

94 Chalastospora gossypii DQ008142 



Chalastospora gossypii CBS 900.87 

Chalastospora gossypii var. polymorpha AY251080 

Chalastospora gossypii AY251081 

100 Chalastospora gossypii AY251079 

Chalastospora gossypii CPC 3685 


Chalastospora obclavata CBS 124120 




Chalastospora cetera CBS 121340 

64 

Chalastospora ellipsoidea CBS 121331 

IC 

PLE (2) 

and a series of branches, giving rise to chains of hyaline conidia 

via sympodial conidiogenesis. Brycekendrickomyces is morphologically 

similar to Lauriomyces, which in turn resembles 

Haplographium. The genus Haplographium is based on H. delicatum. 

Its confused history is discussed in detail by Zucconi 

& Pagano (1993). Haplographium delicatum was originally 

described by Berkeley & Broome as having conidia in chains 

(Mason 1933), which Saccardo (1886) also reported for the 

type species. Hughes (1958) noted that Stilbum catenatum 

was an older name for H. delicatum, which led Holubová-Jechová 

(1973) to place this species in Haplographium, while 

Castañeda & Kendrick (1990) placed it in Lauriomyces. If 

Haplographium and Lauriomyces are synonymous, the older 

name, Haplographium, would have preference. However, as 

shown here, ‘Lauriomyces’ catenatus is not congeneric with 

other species of Lauriomyces, such as L. heliocephalus (Rao 

& de Hoog 1986, Castañeda & Kendrick 1990) and L. bellulus 

(Crous & Wingfield 1994), suggesting that the two genera are 

distinct, and that the name Haplographium catenatum should 

be resurrected. Data from this study, furthermore, suggest that 

the strains of H. catenatum included here, probably represent 

a species complex. 

Brycekendrickomyces differs from Haplographium and Lauriomyces 

by the absence of an intricate conidiophore branching 

system, and in having conidia produced in slimy heads rather 

than in chains. Furthermore, it is not phylogenetically related 

to species of Lauriomyces or Haplographium presently known 

from culture (Fig. 1). Brycekendrickomyces is somewhat similar 

to Argopericonia (Sutton & Pascoe 1987), although the latter 

fungus produces hyaline, apical conidiogenous heads, and it 

has ellipsoidal, single to short catenate conidia, each with a 

prominent, globose guttule.


Phaeosphaeriopsis musae DQ885894 

100 

100 

Alternaria alternata ATCC 52170 FJ545250 

Alternaria alternata EGS 34-016 AF347031 

62 

10 changes 

99 

99 

100 

100 

75 

94 

59 

78 

100 

99 

77 

70 

82 

90 

62 

Alternaria triticina EGS 17-061 AY762948 

Alternaria triticina MUCL 44210 AY714476 

Lewia ethzedia AY278833 

Lewia ethzedia EGS 37-143 AF392987 

Alternaria conjuncta EGS 37-139 AF392988 

Alternaria conjuncta FJ266475 

Chalastospora cetera CBS 121340 

Chalastospora obclavata CBS 124120 

Chalastospora ellipsoidea CBS 121331 

Chalastospora gossypii STE-U 4571 AY251081 

Chalastospora gossypii var. polymorpha AY251080 

Chalastospora gossypii STE-U 4572 AY251079 

100 

86 

62 

Chalastospora gossypii ATCC 28332 AF393680 








Chalastospora 

Fig. 2 The first of 6 equally most parsimonious trees obtained from a heuristic 

search with 100 random taxon additions of the ITS sequence alignment. 

The scale bar shows 10 changes, and bootstrap support values (blue are 

from the parsimony analysis and red from the distance analysis using the 

HKY85 substitution model) from 1 000 replicates are shown at the nodes. 

Branches present in the strict consensus tree are thickened. The tree was 

rooted to a sequence of Phaeosphaeriopsis musae (GenBank accession 

DQ885894). 

65 

63 

61 

50 






Chalastospora E.G. Simmons, Alternaria. An identification 

manual: 668. 2007 

Type species. Chalastospora cetera (E.G. Simmons) E.G. Simmons. 

Conidiophores solitary, brown, smooth, arising from surface hyphae 

or as short, lateral branches from ropes of aerial hyphae; 

short, subcylindrical to flask-shaped, 0–2-transversely euseptate, 

seldom once geniculate or branched. Conidiogenous cells 

integrated, terminal or conidiophores reduced to conidiogenous 

cells, monotretic, determinate to polytretic, sympodial, conidiogenous 

loci visible as minute pores, without or with somewhat 

darkened and slightly thickened rim. Conidia in acropetal, 

branched chains, narrowly ellipsoid to narrowly ovoid, pale to 

medium brown, rarely 1–3 transversely euseptate, generally 

lacking longitudinal or oblique septa; conidial apex functioning 

as secondary conidiophore, proliferating laterally. 

Chalastospora gossypii (Jacz.) U. Braun & Crous, comb. nov. 

— MycoBank MB509518; Fig. 4 

Basionym. Cladosporium gossypii Jacz., Holopkovoe Delo 1929, 5–6: 

564. 1929 and Trudy Byuro Priklad. Bot. 24 (5): 181–182. 1931. 

= Cladosporium malorum Rühle, Phytopathology 21: 1146. 1931. 

≡ Alternaria malorum (Rühle) U. Braun, Crous & Dugan, Mycol. Progr. 2: 

5. 2003. 

= Phaeoramularia kellermaniana Marasas & I.H. Bredell, Bothalia 11: 

217. 1974. 

= Cladophialophora kellermaniana (Marasas & I.H. Bredell) U. Braun & 

Feiler, Microbiol. Res. 150: 83, 1995. 

≡ Pseudocladosporium kellermanianum (Marasas & I.H. Bredell) U. Braun, 

A monograph of Cercosporella, Ramularia and allied genera 2: 393. 1998. 

= Cladosporium porophorum Matsush., Icones Microfungorum a Matsushima 

lectorum: 36. 1975. 

Characteristics in culture — See Braun et al. (2003). 

Specimens examined. Canada, Saskatchewan, Matador, from grass litter, 

27 May 1968, G.C. Bhatt 255, IMI 144487 = ATCC 38025 = CBS 597.69; from 

(?) soil, 18 Sept. 1973, H.A.H. Wallace, IMI 179345; Alberta, from Bromus 

inermis, 1994, R.J. Howad 397, IMI 360655, HAL. – Central Asia (without 

detailed locality), on fibres of Gossypium sp., 1927 and 1928, V.S. Fedorov, 

LEP, syntypes of Cladosporium gossypii. – Lebanon, from soil, July 1987, 

F. Seigle-Murandi, ATCC 200938 = CBS 900.87. – Libya, from Prunus persica, 

April 1975, Casay, IMI 194863. – New Zealand, Wellington, 40 Epuni Street, 

Te Aro Valley, wood and wallpaper from inside walls of a dwelling, 5 Sept. 

2008, leg. D. De Vanny, isol. C.F. Hill 2008/3899, CPC 15567; Auckland, 

Henderson Valley Road, Henderson, leaves of Anethum graveolens (dill)


145 

a 

b 

c 

f 

d 

e 

Fig. 3 Brycekendrickomyces acaciae (CBS 124104). a. Colonies sporulating on MEA; b. colonies on SNA; c–e, g. conidiophores with conidiogenous apparatus; 

f. conidia. — Scale bars = 10 µm. 

g 

along with Itersonilia perplexans, 1 Dec. 2003, leg. J. Pike, isol. C.F. Hill, 

MAF 943 = CBS 114809; Auckland, 90 Aberdeen Road, Castor Bay, isolated 

from Quercus robur (oak) leaves in association with Tubakia dryina, 

5 Sept. 2008, leg. H. Nettleton, isol. C.F. Hill, MAF 954 = CPC 15567 = CBS 

114810. – Pakistan, Karachi, from stored grains, 5 Jan. 1967, S.S. Hussain, 

IMI 124270. – South Africa, Western Cape Province, Kopgat, Calvinia, 

from wheat stubble, Feb. 1972, W.F.O. Marasas OP-76, PREM 44703, IMI 

165252, cultures ATCC 28332 = IMI 165252 = PRE 44703 = CPC 3680 = 

CBS 266.75, ex-type cultures of Phaeoramularia kellermaniana. – Syria, 

from agricultural soil, Feb. 1980, M.I.A. Abdel-Kader, CPC 3685 = ATCC 

200939 = CBS 173.80. – Turkey, Manisa, from Hordeum sp., 16 June 1971, 

Maksu & Selçuc, IMI 159198; Gossypium seeds, M. Esentepe, CBS 540.75. 

– USA, New Mexico, Red River, from a polypore on Picea sp., 4 Sept. 1996, 

D. Wicklow, IMI 386094; Washington State, from Bing cherry fruit, June 1992, 

F.M. Dugan, ATCC 96020; from fruits of Malus domestica, F.D. Heald, ATCC 

36953; Washington State, Festuca idahoensis, F.M. Dugan, STE-U 4572 = 

CBS 114005; Pacific Northwest, Feb. 1966, C.W. Hesseltine, NRRL W 52- 

29 = CPC 3690 = CBS 148.66; Oregon, Portland, Triticum aestivum grain, 

June 1965, C.W. Hesseltine, NRRL A-13702 = CBS 216.65; Malus sylvestris 

fruit, Jan. 1931, F.D. Head, ATCC 36953 = MUCL 10096 = CBS 135.31; 

Washington State, Bromus tectorum, F.M. Dugan, CPC 4571 = CBS 112844; 

Washington State, Roza Canal near Prosser, isolated from dormant buds 

(overwintered) of Vitis vinifera, Mar. 2001, F.M. Dugan, holotype WSP 70286, 

cultures ex-type STE-U 4570 = CBS 112048 (var. polymorpha). – Uzbekistan, 

Bukhara, Experiment Station, on fibres of Gossypium hirsutum, 1928, V.S. 

Zelenetzi, LEP, lectotype of Cladosporium gossypii (selected here) (isolectotype 

in LEP); Bukhara, Shafrikanskoje, on fibres of Gossypium hirsutum, 

1928, V.S. Zelenetzi, LEP, syntype of Cladosporium gossypii. 

Notes — The genus Chalastospora appears to represent 

an anamorph lineage in the Pleosporales (Fig. 1). Chalastospora 

cetera and C. gossypii are clearly congeneric (Fig. 2). 

Based on the ITS data, there are some point mutations among 

strains of C. gossypii, suggesting that other genes need to be 

sequenced to fully elucidate the variation within this species 

(Fig. 2). On SNA, ramoconidia of CBS 114810 were 10–17 

× 3–5 µm, and conidia narrowly ellipsoid-ovoid, cylindrical 

to fusiform, 6–10 × 2–2.5 µm, thus much smaller than that 

reported by Braun et al. (2003) on PDA. Jaczewski introduced 

the name Cladosporium gossypii in 1929, and provided a brief 

Russian description, including shape and size of conidia. This 

description, published before 1935, is, however, valid. In his 

paper of 1931, he re-introduced C. gossypii together with a 

Latin description and a micrograph of conidia. Type material of 

C. gossypii was re-examined and it is identical to C. malorum. 

However, C. gossypii is an older name than C. malorum, which 

was published in 1931, and has priority. 

Chalastospora ellipsoidea Crous & U. Braun, sp. nov. — 

MycoBank MB509519; Fig. 5 

Chalastosporae gossypii similis, sed conidiis ellipsoideis, longioribus et leviter 

latioribus, (8–)10–15(–17) × 3(–3.5) µm. 

Etymology. Named after its ellipsoid conidia.


a 

b 

c 

d 

e 

f 

Fig. 4 Chalastospora gossypii (CBS 114810). a–f. Superficial mycelium on SNA showing conidiophores with branched conidial chains. — Scale bars = 10 µm. 

On SNA: Conidiophores arising singly from aerial and creeping 

hyphae; subcylindrical, erect, medium brown, smooth, up 

to 25 × 3 µm, frequently reduced to conidiogenous cells, 5–13 

× 3 µm; seldom once geniculate, mostly straight, with a slight 

swelling in the apical conidiogenous region; conidiogenous loci 

1–3 per conidiogenous cell, medium brown, slightly thickened, 

darkened, up to 1 µm wide. Ramoconidia (0–)1–3-septate, ellipsoid-ovoid, 

subcylindrical or fusiform, smooth, medium brown, 

(12–)15–18(–30) × 3(–4) µm; apex at times with short beak, 

giving rise to lateral branch. Conidia ellipsoid to fusoid, medium 

brown, smooth, in long acropetal chains, simple, or branched 

with short apical or basal, lateral branches, (8–)10–15(–17) 

× 3(–3.5) µm, 0–1(–2)-septate; hila thickened and darkened, 

0.5–1 µm wide. 

Characteristics in culture — Colonies on OA spreading, with 

moderate, flattened aerial mycelium, smoke-grey. On MEA cinnamon 

with patches of hazel on surface and reverse. On PDA 

olivaceous-grey, with moderate aerial mycelium; iron-grey in 

reverse. 

Specimen examined. Australia, on Triticum, H.L. Harvey & S. Perth, holotype 

CBS H-20199, culture ex-type E.G.S. 22.060 = CBS 121331. 

Notes — The most characteristic features of this species 

are its short lateral branches, and ellipsoid conidia. It is clearly 

distinct from C. cetera and C. gossypii based on ITS sequence 

data (Fig. 2). 

Chalastospora obclavata Crous & U. Braun, sp. nov. — 


Differt ab omnibus specibus Chalastosporae conidiis intercalaribus obclavatis. 

Etymology. Named after its obclavate conidia. 

Sporulating poorly on SNA. Conidiophores 17–30 × 3–4 µm, 

arising singly from aerial and creeping hyphae; subcylindrical, 

somewhat clavate near apex of conidiogenous region, erect, 

straight to once geniculate, medium brown, smooth, frequently 

reduced to conidiogenous cells, 5–10 × 3–4 µm; conidiogenous 

loci medium brown, slightly thickened, darkened, 1–1.5 µm 

wide. Ramoconidia medium brown, smooth, developing short 

lateral beaks at apex that give rise to lateral chains (verticillate-like 

appearance), obclavate, widest at base, 0–3-septate, 

(28–)30–35 × (3.5–)4–5(–6) µm. Conidia obclavate, widest 

at base, (23–)26–30(–35) × (3.5–)4 µm, 0–3-septate; hila 

thickened, darkened, 1–1.5 µm wide. 

Characteristics in culture — Colonies on OA spreading, with 

moderate, white aerial mycelium, grey-olivaceous to smoke 

grey; reverse grey-olivaceous. On MEA cream with dense aerial 

mycelial mat. 

Specimen examined. USA, Kansas, Manhattan, ex air, Jan. 1958, C.T. 

Rogerson, holotype CBS H-20200, culture ex-type E.G.S. 12.128 = CBS 

124120.


147 

a 

b 

c 

d 

e 

f 

g h i j 

Fig. 5 Chalastospora ellipsoidea (CBS 121331). a–f. Superficial mycelium on SNA showing conidiophores with conidial chains; g, h. microconidiophores; 

i, j. conidia in chains. — Scale bars = 10 µm. 

Notes — The most characteristic features of this species are 

its conidial branching pattern and conidial shape. This strain 

was discussed by Simmons under Alternaria cetera (Simmons 

1996), and under Chalastospora in Simmons (2007). It is clearly 

distinct from C. cetera (ex-type CBS 121340, Fig. 7), C. ellipsoidea 

and C. gossypii based on ITS sequence data (Table 1, 

Fig. 2). 

Key to species of Chalastospora 1 

1. Intercalary conidia usually longer than 20 µm . . . . . . . . . 2 

1. Intercalary conidia shorter than 20 µm . . . . . . . . . . . . . . 3 

2. Intercalary conidia narrowly ellipsoid to narrowly ovoid, widest 

in middle or lower third, (10–)19–24(–30) × 3(–4) µm, 

0–3-septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. cetera 

2. Intercalary conidia obclavate, widest at base, (23–)26–30 

(–35) × (3.5–)4 µm, 0–3-septate . . . . . . . . . C. obclavata 

3. Intercalary conidia narrowly ellipsoid-ovoid to cylindrical or 

fusiform, 6–10 × 2–2.5 µm, mostly aseptate. . C. gossypii 

3. Intercalary conidia ellipsoid, not cylindrical nor fusiform, 

(8–)10–15(–17) × 3(–3.5) µm, 0(–2)-septate C. ellipsoidea 

1 

Colonies cultivated on SNA. 

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

47. 1962 

Type species. Cyphellophora laciniata G.A. de Vries. 

Hyphae fertile, pale brown, 1.5–3 µm wide, at times constricted 

at septa. Conidiogenous cells phialidic, intercalary, at times on 

short lateral branches, with a prominent to indistinct collarette. 

Conidia sickle-shaped, brown, smooth-walled, 1–3-septate, 

adhering in bundles. 

Cyphellophora eugeniae Crous & Alfenas, sp. nov. — Myco- 

Bank MB509521; Fig. 8 

Cyphellophorae taiwanensis similis, sed conidiis valde longioribus, (40–)60– 

75(–90) × 2–2.5(–3) µm. 

Etymology. Named after the host on which it occurs, Eugenia. 

On PDA. Mycelium consisting of branched, greenish brown, 

septate, smooth, 3–5 µm wide hyphae, constricted at septa. 

Conidiogenous cells phialidic, intercalary, inconspicuous to subdenticulate, 

1 µm wide, with minute collarettes, with several loci 

aggregated at hyphal swellings. Conidia subcylindrical, tapering 

towards obtuse ends, curved, smooth, hyaline to olivaceous,


a 

b 

c 

d 

Fig. 6 Chalastospora obclavata (CBS 124120). a, b. Superficial mycelium on SNA showing conidiophores with branched conidial chains; c–e. conidia in 

chains. — Scale bar = 10 µm. 

e 

a 

b 

c 

d 

e 

f 

g 

Fig. 7 Chalastospora cetera (CBS 121340). a–g. Superficial mycelium on SNA showing conidiophores with conidial chains. — Scale bars = 10 µm.


149 

a 

b 

c 

d 

e 

f 

g h i j 

Fig. 8 Cyphellophora eugeniae (CBS 124105). a, b. Colonies sporulating on OA; c–e. conidia attached to conidiogenous cells (arrows denote loci); f–j. 

conidia. — Scale bars = 10 µm. 

finely guttulate, 4–6(–10)-septate, prominently constricted at 

septa, widest in the middle of conidium, (40–)60–75(–90) × 

2–2.5(–3) µm; conidia also anastomose and undergo microcyclic 

conidiation in culture. 

Characteristics in culture — Colonies on PDA erumpent, 

with sparse aerial mycelium and even margins; surface olivaceous-grey, 

with patches of iron-grey; reverse iron-grey. On 

MEA erumpent, with folded surface and smooth, lobate margin, 

and sparse aerial mycelium; surface pale olivaceous-grey to 

olivaceous-grey; reverse iron-grey. On OA spreading, flat, with 

even, smooth margins and sparse aerial mycelium, olivaceousgrey. 

Colonies reaching 15 mm diam after 1 mo at 25 °C, fertile, 

sporulating in slimy sporodochial masses. 

Specimen examined. Brazil, Rio Grande do Sul, Guaiba, living leaves of 

Stenocalyx uniflorus, 1 Apr. 2008, leg. A.C. Alfenas, isol. P.W. Crous, holotype 

CBS H-20201, culture ex-type CPC 15172 = CBS 124105. 

Notes — The indistinct conidiogenous loci of C. eugeniae are 

reminiscent of those of C. taiwanensis (Matsushima 1985). The 

two species can be distinguished by the much longer conidia in 

C. eugeniae. Based on the key provided by Decock et al. (2003), 

C. eugeniae appears to represent a new species. Further collections 

of this complex are required to confirm the synonymy 

of the genera Cyphellophora with Pseudomicrodochium and 

Kumbhayama (Decock et al. 2003, Crous et al. 2007b), which 

were originally distinguished based on the absence of conidial 

pigmentation. The ITS sequence of C. eugeniae has 89 % 

similarity to that of Cyphellophora hylomeconis (GenBank accession 

EU035415).


Key to species of Cyphellophora 

(adapted from Decock et al. 2003) 

1. Phialides intercalary, reduced to a sessile locus with collarette 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 

1. Phialides prominent, cylindrical, flask-shaped, sessile or with 

an elongated base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 

2. Conidia 1–3-septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 

2. Conidia usually more than 3-septate . . . . . . . . . . . . . . . . 4 

3. Conidia up to 2.5 µm wide (11–20 × 2–2.5 µm), 1(–2)-septate 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. fusarioides 

3. Conidia up to 5 µm wide (11–25 × 2–5 µm), 1–3-septate 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. laciniata 

4. Conidia up to 2 µm wide, 3–6-septate, sigmoid (16–35 × 

1.5–2 µm) . . . . . . . . . . . . . . . . . . . . . . . . . C. taiwanensis 

4. Conidia wider than 2 µm . . . . . . . . . . . . . . . . . . . . . . . . . 5 

5. Conidia subcylindrical, 4–6(–10)-septate, (40–)60–75(–90) 

× 2–2.5(–3) µm . . . . . . . . . . . . . . . . . . . . . . . C. eugeniae 

5. Conidia sigmoid, 1–5-septate, (15–)25–35(–55) × (2.5–) 

3(–4) µm . . . . . . . . . . . . . . . . . . . . . . . . . . C. hylomeconis 

6. Phialides short to long and cylindrical; conidia 1–1.2 µm 

wide, 2–3-septate . . . . . . . . . . . . . . . . . . . . . . . C. suttonii 

6. Phialides prominent, flask-shaped, sessile or with an elongated 

base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 

7. Conidia mainly straight, on average smaller than 20 µm, 1–5- 

septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. pluriseptata 

7. Conidia straight to more commonly falcate, curved, or sigmoid, 

on average longer than 20 µm . . . . . . . . . . . . . . . . 8 

8. Conidia (1–)3-septate, wider than 3 µm, 25–40 × 3.5–5.5 

µm; phialides commonly with an elongated base C. indica 

8. Conidia 2–8-septate, narrower than 3 µm; phialides without 

elongated base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 

9. Conidia vermiform, mostly curved, mostly 4–8-septate, 30– 

55 × 1.2–1.5 µm. . . . . . . . . . . . . . . . . . . . . . C. vermispora 

9. Conidia straight, falcate or slightly sigmoid, (2–)3–6-septate, 

(18–)19.5–28(–29) × 1.5–2 µm . . . . . . . . . C. guyanensis 

Dictyosporium Corda, in Weitenweber, Beitr. Gesammten 

Natur-Heilwiss., Prag 1: 87. 1836 

Type species. Dictyosporium elegans Corda. 

Conidiomata sporodochial, black, scattered. Mycelium predominantly 

immersed, consisting of branched, septate, smooth, thinwalled 

hyphae. Conidiophores micronematous, mononematous, 

pale brown, smooth to finely verruculose, thin-walled, septate, 

cylindrical. Conidiogenous cells monoblastic, integrated, pale 

to medium brown, smooth to finely verruculose, cylindrical, 

determinate; at times remaining attached to released conidium. 

Conidia cheiroid, medium to dark brown, smooth, euseptate, 

one cell-layer thick, cells arranged in 1–2 planes, fan-shaped; 

cell rows originating from a central basal cell; rows usually attached 

along their length; outer rows usually shorter than inner 

rows, at times paler in colour than central rows, and with or 

without hyaline, thin-walled, 1–2-celled appendages that are 

allantoid, clavate to globose, or fusoid to cylindrical. 

Dictyosporium strelitziae Crous & A.R. Wood, sp. nov. — 


Dictyosporii bulbosi valde simile, sed conidiis leviter longioribus, (30–)40– 

46(–55), et phylogenetice manifeste divergens. 

Etymology. Named after the host genus Strelitzia, on which it occurs. 

Leaf spots absent, colonies occurring on dead leaf tissue. 

Description based on colonies sporulating on WA with pine nee- 

a 

b 

c 

d 

e 

f 

Fig. 9 Dictyosporium strelitziae (CBS 123359). a. Colony sporulating on PDA; b, c. conidia attached to conidiogenous cells; d–h. conidia with hyaline, apical 

appendages. — Scale bars = 10 µm. 

g 

h


151 

dles (colonies also sporulate well on OA and MEA): Mycelium 

predominantly internal in host tissue, consisting of branched, 

septate, smooth, brown, 2–2.5 µm wide hyphae. Conidiomata 

sporodochial, scattered, black, up to 170 µm diam. Conidiophores 

subcylindrical, darker brown than hyphae, at times 

slightly verruculose, irregularly curved to geniculate-sinuous, 

1–3-septate, 10–25 × 2–2.5 µm; older conidiophores curved 

like sheperd’s crook. Conidiogenous cells terminal, medium 

brown, verruculose, subcylindrical, curved (semi-circular), 5–10 

× 2–2.5 µm. Conidia solitary, complanante, cheiroid, smoothwalled, 

uniformly pale brown, becoming uniformly medium 

brown at maturity; cells arranged in (4–)5(–6) rows, meeting at 

basal cell; outer rows with 8–10 cells, with a hyaline, globose, 

apical appendage, 5–10 µm diam; outer rows shorter than inner 

rows; inner rows with 7–11 cells; central row with 6–10 cells; 

conidia (30–)40–46(–55) × (20–)21–23(–25) µm. 

Characteristics in culture — Colonies on OA flat, spreading, 

without aerial mycelium, and with regular, even margin; on MEA 

flat, spreading, with moderate aerial mycelium and regular, 

smooth margin; surface buff, reverse cinnamon; colonies on 

both media reaching 30 mm diam after 1 mo at 25 °C. 

Specimen examined. South Africa, KwaZulu-Natal, Skyline Nature 

Reserve, Uvongo, on dead leaves of Strelitzia nicolai, 29 May 2008, leg. 

A. Wood, isol. P.W. Crous, CBS H-20202 holotype, cultures ex-type CPC 

15359–15361, CBS 123359. 

Notes — The genus Dictyosporium is well defined, and separated 

from similar genera by having smooth-walled, euseptate 

conidia produced from determinate conidiogenous cells (Sutton 

et al. 1996, Tsui et al. 2006). Based on the key provided by Cai 

et al. (2003b), D. strelitziae is morphologically most similar to 

D. bulbosum (conidia 27–46 × 11–30 µm), but its conidia are 

somewhat longer, and there is a 10 bp difference between the 

ITS sequences of D. strelitziae and D. bulbosum (DQ018086). 

Phylogenetically, D. strelitziae is closest to D. elegans (conidia 

44–80 × 24–36 µm; appendages absent) (5 bp difference in 

the ITS sequence, DQ018087), but it has smaller conidia than 

the latter species. Furthermore, it also appears distinct from all 

species not occurring in the key of Cai et al. (2003b) (Arambarri 

et al. 2001, Cai et al. 2003a, Zhao & Zhang 2003, Kodsueb et 

al. 2006, Cai & Hyde 2007, McKenzie 2008). 

Key to species of Dictyosporium 

(adapted from Cai et al. 2003b) 

1. Conidia with appendages. . . . . . . . . . . . . . . . . . . . . . . . . 2 

1. Conidia lacking appendages . . . . . . . . . . . . . . . . . . . . . 13 

2. Appendages apical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 

2. Appendages not apical . . . . . . . . . . . . . . . . . . . . . . . . . . 4 

3. Apical appendages aseptate . . . . . . . . . . . . . . . . . . . . . . 6 

3. Apical appendages frequently 1-septate, cylindrical, 24–51 

× 6–10.5 µm; conidia 27.5–47.5 × 20–25 µm, complanate, 

with 4–5 rows of cells . . . . . . . . . . . . . . . . D. canisporum 

4. Appendages subapical, cylindrical to clavate; conidia 52.5– 

72.5 × 18.5–26.5 µm, not complanate, with 5 rows of cells 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. tetraploides 

4. Appendages not subapical, but central or basal . . . . . . . 5 

5. Appendages central, hyaline, thin-walled, clavate to obovoid; 

conidia 36–45 × 16–21 µm, not complanate, mostly 7 rows 

of cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. musae 

5. Appendages basal, fusoid to cylindrical; conidia 22–28 × 12.5– 

18 µm, complanate, with 3 rows of cells . . D. manglietiae 

6. Conidia with 3 rows of cells, (27–)31–43 × 10–12 µm . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. freycinetiae 

6. Conidia with more than 3 rows of cells . . . . . . . . . . . . . . 7 

7. Conidia mostly with 4 rows of cells . . . . . . . . . . . . . . . . . 8 

7. Conidia with 5 or more rows of cells . . . . . . . . . . . . . . . 10 

8. Conidia with darker colour at apex of inner rows; apical 

cells of outer rows each bearing a hyaline, cylindrical appendage. 

. . . . . . . . . . . . . . . . . . . . . . . . . . D. nigroapice 

8. Conidia concolorous . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 

9. Conidia 24–40 × 14–20 µm; appendages clavate . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. tetraseriale 

9. Conidia 36–45 × 16–21 µm; appendages tapering . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. palmae 

10. Conidia mostly comprising 5 rows of cells. . . . . . . . . . 11 

10. Conidia mostly comprising 6–8 rows, 46–88 × 26–46 µm; 

appendages hyaline, curved . . . . . . . . . . . . D. digitatum 

11. Conidia longer than 32 µm, appendages globose to obovoid 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 

11. Conidia shorter than above, 26–32 × 15–24 µm; appendages 

cylindrical to clavate . . . . . . . . . . . . . . . . D. alatum 

12. Conidia up to 46 µm long, and 30 µm wide, 27–46 × 11–30 

µm; appendages globose to obovoid . . . . . D. bulbosum 

12. Conidia longer than 46 µm, but not wider than 25 µm, 

(30–)40–46(–55) × (20–)21–23(–25) µm; appendages 

globose . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. strelitziae 

13. Conidia complanate, one cell layer thick . . . . . . . . . . . 14 

13. Conidia not complanate, more than one cell layer thick 24 

14. Conidia regularly consisting of 3 rows of cells. . . . . . . 15 

14. Conidia consisting of at least 4 rows of cells . . . . . . . . 16 

15. Conidia 15–22.5 × 10–16.5 µm . . . . D. lakefuxianensis 

15. Conidia 26–32 × 16–18 µm . . . . . . . . . . . . . D. triseriale 

16. Conidia curved, with 5–7 rows of cells, each curving in the 

same direction, 34–56 × 20–38 µm . . . . . . . . D. foliicola 

16. Conidia not curved . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 

17. Conidia less than 25 µm long . . . . . . . . . . . . . . . . . . . 18 

17. Conidia more than 25 µm long . . . . . . . . . . . . . . . . . . 19 

18. Conidia 18–24 × 13–19 µm . . . . . . D. brahmaswaroopii 

18. Conidia 15–17 × 11–12 µm . . . . . . D. schizostachyfolium 

19. Conidia with paler outer rows . . . . . . . . . . . . . . . . . . . 20 

19. Conidia concolorous . . . . . . . . . . . . . . . . . . . . . . . . . . 21 

20. Conidia 25–45 × 22–38 µm, with (5–)6(–7) rows . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. yunnanensis 1 

20. Conidia 26–40 × 13–25 µm, mostly with 5 rows. . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. zeylanicum 

21. Conidia with 4 rows, 23.5–40 × 16–21.5 µm . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. tetrasporum 

21. Conidia with more than 4 rows . . . . . . . . . . . . . . . . . . 22 

22. Conidia 40–80 × 24–36 µm, mostly with 5 rows, slightly 

constricted at septa . . . . . . . . . . . . . . . . . . . . D. elegans 

22. Conidia mostly with more than 5 rows, strongly constricted 

at septa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 

23. Conidia 26–34 × 23–34 µm, mostly with 7–9 rows of cells; 

conidiomata sporodochial . . . . . . . . . . . . D. polystichum 

23. Conidia 38–56 × 25–32 µm, mostly 6–8 rows of cells; 

conidiomata not sporodochial . . . . . . . . . . . D. toruloides 

24. Conidia campaniform, with a darker base; with 12–16 rows 

of cells, 22–40 × 20–30 µm . . . . . . . . D. campaniforme 

24. Conidia more or less cylindrical, concolorous, comprising 

3–7 rows of cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 

25. Conidia regularly with 3 rows of cells; usually 13.5 µm or 

less wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 

25. Conidia mostly with 4–7 rows of cells; more than 13.5 µm 

wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 

1 

Appearing morphologically similar to D. taishanensis, also described 

from China; conidia with (3–)5(–7) cell layers, 27–43 × 15–30 µm (Zhao 

& Zhang 2003). Dictyosporium taishanensis (22 February 2003) is older 

than D. yunnanensis (March 2003), and would have priority if these fungi 

are shown to be synonymous.


26. Conidia 40–60 × 10–13.5 µm . . . . . . . . . . D. triramosum 

26. Conidia shorter than 43 µm . . . . . . . . . . . . . . . . . . . . . 27 

27. Conidia 36–43 × 11–12 µm; sporodochia usually covered 

with gelatinous matrix . . . . . . . . . . . . . . . D. australiense 

27. Conidia 20–30 × 10–12 µm; sporodochia not as above 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. micronesicum 

28. Conidia 40–50 × 18–25 µm, with 4–6 rows of cells, muriform, 

with hyaline, subglobose conidiogenous cell remaining 

attached as basal appendage . . . . . . . . . . D. gauntii 

28. Conidial morphology not as above . . . . . . . . . . . . . . . 29 

29. Conidia with rows of cells that are distinctly incurved or 

hook-like at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . 30 

29. Conidia with more or less straight rows of cells at the apex 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 

30. Conidia 105–121 × 25–32 µm . . . . . . . . . D. giganticum 

30. Conidia up to 80 µm long . . . . . . . . . . . . . . . . . . . . . . 31 

31. Conidia 50–80 × 20–30 µm . . . . . . . . . D. heptasporum 

31. Conidia 33–42 × 16–20 µm . . . . . . . . D. subramanianii 

32. Colonies effuse, not sporodochial; conidia irregularly cylindrical 

or oblong, strongly constricted at septa; 30–50 × 

12–30 µm . . . . . . . . . . . . . . . . . . . . . . . . . . D. oblongum 

32. Colonies sporodochial; conidia more or less cylindrical, 

slightly constricted at septa, 53–76 × 19–22 µm . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. cocophilum 

Edenia M.C. González, Anaya, Glenn, Saucedo & Hanlin, 

Mycotaxon 101: 254. 2007. 

Type species. Edenia gomezpompae M.C. González, Anaya, Glenn, 

Saucedo & Hanlin. 

Conidiophores fasciculate, subcylindrical, medium brown, 

finely roughened, 3–15-septate, straight to variously curved 

or geniculate-sinuous, irregular in width, constricted at some 

septa, with percurrent rejuvenation in upper part, situated on 

a submerged, brown stroma. Conidiogenous cells terminal, 

integrated, becoming paler brown towards apex, tapering to 

a subtruncate tip, with several lateral loci that are somewhat 

thickened and protruding (pimple-like), giving rise to conidia 

a 

b 

c 

d 

h 

e 

f 

g 

i 

j k l 

m 

Fig. 10 Edenia gomezpompae (CBS 124106). a. Hyphal tufts visible when cultivated on MEA; b. leaf spot with conidiophores; c. fasciculate conidiophores; 

d. conidiophores arising from conidioma; e–g. conidiophores and conidiogenous cells; h. conidia; i. conidiomata forming on OA; j. conidioma with ostiolar 

setae; k, l. conidiogenous cells; m. conidia. — Scale bars = 10 µm.


153 

via sympodial proliferation near apex. Conidia 11–16 × 3.5–6 

µm, subhyaline, smooth, thin-walled, finely guttulate, fusoidellipsoidal 

with obtuse apex and tapering from its widest point 

in the middle towards a subtruncate base, 1–1.5 µm wide. 

Edenia gomezpompae M.C. González, Anaya, Glenn, Saucedo 

& Hanlin, Mycotaxon 101: 254. 2007 — Fig. 10 

Leaf spots subcircular, 3–12 mm diam, grey-brown, with a 

dark brown, raised border, surrounded by a diffuse, black halo 

(absent in smaller spots). Conidiophores in fascicles of 5–30, 

subcylindrical, medium brown, finely roughened, 3–15-septate, 

straight to variously curved or geniculate-sinuous, 50–170 × 

4–6 µm, irregular in width, constricted at some septa, with percurrent 

rejuvenation in upper part; fascicles randomly distributed 

over lesion, amphigenous, visible as erect, dark brown to black 

tufts on lesions, situated on a submerged, brown stroma, up to 

60 µm wide and 40 µm high, intermingled among leaf trichomes 

(fruiting structures of a Ramularia sp. and ascomata of another 

fungus also present in some lesions). Conidiogenous cells 

15–30 × 3–4 µm, terminal, integrated, becoming paler brown 

towards apex, tapering to a subtruncate tip, with several lateral 

loci that are somewhat thickened and protruding (pimple-like), 

up to 1 µm diam, giving rise to conidia via sympodial proliferation 

near apex, but some conidiogenous cells also show 

signs of percurrent proliferation, but this appears to be linked 

to rejuvenation, not conidiogenesis. Conidia (11–)13–15(–16) 

× (3.5–)4.5–5.5(–6) µm, subhyaline, smooth, thin-walled, 

finely guttulate, fusoid-ellipsoidal with obtuse apex and tapering 

from its widest point in the middle towards a subtruncate 

base, 1–1.5 µm wide. 

Characteristics in culture — Colonies fluffy, with white hyphal 

strands that turn brown with age; surface woolly with abundant 

aerial mycelium; margins uneven. On MEA buff to rosy-buff 

(surface), brick to dark brick (reverse); on PDA fluffy, cream 

to buff (surface), dark brick to buff (reverse); on OA brick with 

patches of cream to buff. Colonies reaching 25 mm diam after 

2 wk at 25 °C, becoming fertile on OA. 

Specimens examined. Mexico, Quintana Roo, Isla Mujeres Municipality, El 

Eden Ecological reserve, from leaves of Callicarpa acuminata (Lamiaceae), 

May 2002, A. Saucedo-García & A.L. Anaya, holotype MEXU 25346. – Philippines, 

on Senna alata (≡ Cassia alata) (Caesalpiniaceae), Oct. 2008, leg. 

C.J.R. Cumagun, isol. P.W. Crous, epitype designated here CBS H-20203, 

cultures CPC 15689 = CBS 124106, CPC 15690, 15691. 

Notes — The genus Edenia was originally introduced for 

a sterile fungus (suspected to be a member of the Pleosporaceae), 

isolated as an endophyte from leaves of Callicarpa 

acuminata in Mexico (González et al. 2007). The genus was 

characterised by producing numerous sterile, whitish mycelial 

strands and coils on PDA. The present collection from Cassia 

alata in the Philippines has the same colony characteristics, 

and based on its identical DNA sequence data (GenBank 

EF565744.1), we believe that this is the same fungus. What 

is interesting, however, is the fact that the latter collection was 

made from conidia of a dematiaceous hyphomycete sporulating 

on leaf spots of C. alata. As other fungi were also present on 

these spots, its potential role as pathogen remains uncertain. 

On host tissue, however, some conidiophores were associated 

with a weakly developed layer of pale brown stromatic cells. On 

OA, cultures became fertile, and conidiophores were arranged 

around well-developed ostioles of submerged pycnidia (with a 

similar pale brown stromatic wall to that observed on the host). 

It is possible, therefore, that if the field material had been placed 

in moist chambers, the pycnidial state would have developed. 

The latter state resembles species that are pyronellea-like in 

morphology. 

Morphologically, the hyphomycete state of Edenia resembles 

genera such as Digitopodium, although species of this genus 

have rhizoids, and 1-septate, pale brown conidia that can also 

occur in short chains (Heuchert et al. 2005). It also shares some 

similarities with Blastophorum (Matsushima 1971), although 

the latter fungus is distinct in having solitary conidiophores with 

rhizoids, and a hyaline, upper conidiogenous region. 

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

Type species. Thedgonia ligustrina (Boerema) B. Sutton. 

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, 

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. 

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

Soc. 61: 428. 1973 — Fig. 11 

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. 

Characteristics in culture — On MEA erumpent, slow growing, 

5–8 mm after 2 wk, with moderate, white aerial mycelium 

and smooth, lobate margins; umber in reverse. On OA 5–8 mm 

diam after 2 wk, submerged to flattened on surface, sparse aerial 

mycelium, and smooth, even margins; umber on surface. 

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 — Kaiser & Crous (1998) linked ‘Thedgonia’ lupini as 

anamorph to Mycosphaerella lupini, and thus suggested that 

Thedgonia belongs in the Mycosphaerellaceae. Results of this 

study (Fig. 1), however, show that Thedgonia s.str. belongs to 

the Helotiales, and is unrelated to the Mycosphaerellaceae. 

Furthermore, there is presently no separate anamorph genus 

in the Mycosphaerellaceae to accommodate ‘T.’ lupini. Although 

‘T.’ lupini resembles species of Pseudocercosporella (Braun 

1995), it appears to represent a separate phylogenetic lineage. 

Trochophora R.T. Moore, Mycologia 47: 90. 1955 

Type species. Trochophora fasciculata (Berk. & M.A. Curtis) Goos (syn. 

T. simplex (Petch) R.T. Moore). 

Colonies hypophyllous, medium to dark brown, consisting of 

numerous synnemata. Stroma absent, but 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


a 

b 

c 

d 

e 

f 

g 

Fig. 11 Thedgonia ligustrina (CBS 124332). a, b. Leaf spots on Ligustrum; c. fasciculate conidiophores; d, e. conidiophores; f, g. conidia. — Scale bars = 

10 µm. 

unthickened scar, clavate. Conidia solitary, terminal or lateral 

on conidiogenous cells, prominently curved to helicoid, pale to 

medium brown, smooth, transversely septate with a darkened, 

thickened band at the septa. 

Trochophora fasciculata (Berk. & M.A. Curtis) Goos (as ‘fasciculatum’), 

Mycologia 78: 759. 1986 — Fig. 12 

Basionym. Helicoma fasciculatum Berk. & M.A. Curtis, U.S. North Pacific 

Exped.: 142. (1853–1856) 1853. 

≡ Helicosporium fasciculatum (Berk. & M.A. Curtis) Sacc., Syll. fung. 4: 

560. 1886. 

≡ Helicomyces fasciculatus (Berk. & M.A. Curtis) Pound & Clem., Minnesota 

Bot. Stud. 1: 658. 1896. 

= Helicosporium simplex Syd., Mém. Herb. Boissier 4: 7. 1900. 

≡ Helicoma simplex (Syd.) Linder, Ann. Missouri Bot. Gard. 16: 315. 

1929. 

= Helicostilbe simplex Petch, Ann. Roy. Bot. Gard. Peradeniya 7: 321. 

1922. 

≡ Trochophora simplex (Petch) R.T. Moore, Mycologia 47: 90. 1955. 

Specimen examined. Korea, Pusan, on leaves of Daphniphyllum macropodum, 

13 Nov. 2002, leg. H.D. Shin, isol. P.W. Crous, KUS-F19414, cultures 

CPC 10280–10282. 

Notes — Two species have been described in the genus, 

namely T. fasciculata and T. simplex; the latter recognised as 

a synonym of the former (Zhao et al. 2007). Within the Mycosphaerellaceae, 

pseudocercospora-like species cluster in two 

well-defined clades, namely the P. vitis clade (Pseudocercospora 

s.str.), and the P. heimii clade (pseudocercospora-like). 

Based on LSU DNA phylogeny (Fig. 1), Trochophora clusters 

basal to the pseudocercospora-like clade. Although it is tempting 

to use the name Trochophora for this clade, further collections 

of Trochophora are required to clarify the morphological 

variation among taxa with this unique conidial morphology. 

Using sequence data of the ITS gene, the closest taxa obtained 

from a BLAST search is the Mycosphaerella heimii species 

complex (96 % similarity). 

Zhao et al. (2007) consider T. fasciculata as a pathogen of Daphniphyllum, 

and report it from this host in several Asian countries, 

namely Sri Lanka, China (incl. Hong Kong and Taiwan) 

and India. 

Verrucisporota D.E. Shaw & Alcorn, Austral. Syst. Bot. 6: 273. 

1993 

≡ Verrucispora D.E. Shaw & Alcorn, Proc. Linn. Soc. New South Wales 

92: 171. 1967. (nom. illegit.). 

Type species. Verrucisporota proteacearum (D.E. Shaw & Alcorn) D.E. 

Shaw & Alcorn. 

Mycelium consisting of pale brown, septate, verrucose hyphae. 

Stroma forming in substomatal cavities, cells brown-walled, 

pseudoparenchymatous. Conidiophores macronematous, 

mononematous, simple, flexuous, often geniculate, septate, 

mainly smooth, pale to dark brown, tapering towards the apex, 

but often becoming more swollen, and also verruculose to verrucose 

at the apex. Conidiogenous cells cylindrical, becoming 

geniculate, integrated, terminal, becoming intercalary, polyblastic, 

proliferating sympodially, cicatrised; conidiogenous loci 

planate, conspicuous, protuberant, thickened and darkened. 

Conidia cylindrical, narrowing slightly to an obtuse apex and 

with a truncate base with a distinctly thickened hilum, medium 

brown, straight or curved, transversely septate, verrucose to 

verruculose.


155 

a 

b 

c 

d 

e 

f 

g 

Fig. 12 Trochophora fasciculata (CPC 10280). a. Leaf spots on Daphniphyllum; b. colony on MEA; c. fasciculate conidiophores; d. conidiophores and conidiogenous 

cells; e–g. conidia. — Scale bars = 10 µm. 

Verrucisporota daviesiae (Cooke & Massee) Beilharz & Pascoe, 

Mycotaxon 82: 360. 2002 

Basionym. Cercospora daviesiae Cooke & Massee, Grevillea 18: 7. 

1889. 

Teleomorph. Mycosphaerella daviesiicola Beilharz & Pascoe, Mycotaxon 

82: 364. 2002. 

Characteristics in culture — On MEA erumpent, spreading 

with folded surface, and sparse aerial mycelium and even, 

lobate margin; surface iron-grey to olivaceous-grey; reverse 

iron-grey; colonies reaching 7 mm diam after 2 wk. On PDA 

erumpent, spreading, with moderate aerial mycelium and 

uneven margins; surface white in middle, olivaceous-grey in 

outer region, iron-grey underneath; colonies reaching 8 mm 

diam after 2 wk. On OA erumpent, spreading, with moderate 

aerial mycelium and uneven margin; surface white in middle, 

olivaceous-grey in outer region; colonies reaching 8 mm diam 

after 2 wk. 

Specimen examined. Australia, Victoria, on living leaves of Daviesia 

mimosoides (≡ D. cormybosa var. mimosoides), V. & R. Beilharz, VPRI 31767 

= CBS 116002. 

Notes — The type species of the genus Stenella, S. araguata, 

clusters in the Teratosphaeriaceae (Crous et al. 2007a), 

and thus the majority of the stenella-like anamorphs in the 

Mycosphaerellaceae, will need to be placed in another genus. 

One option would be Zasmidium (Arzanlou et al. 2007), which 

clusters in the Mycosphaerellaceae, along with Verrucisporota 

(Fig. 1). This clade, however, is neither morphologically nor 

phylogenetically well resolved, and taxa need to be added to 

improve the phylogeny before a reasonable assessment can 

be made. The ITS sequence of this species is distinct from the 

other two species of this genus treated in this paper (Table 1). 

Verrucisporota grevilleae Crous & Summerell, sp. nov. — 


Differt a Verrucisporota protearum conidiis angustioribus et longioribus, (30–) 

50–65(–80) × (5–)6–7 µm, et conidiophoris brevioribus, (35–)80–120(–160) 

× (5–)6–7 µm. 

Etymology. Named after the host genus on which it occurs, Grevillea. 

Leaf spots angular, elongated, amphigenous, 1–2 mm wide, 

3–10 mm long, medium to dark brown to black, discrete. 

Mycelium immersed and superficial, hyphae medium brown, 

septate, verrucose, 1.5–3 µm wide. Stroma up to 60 µm wide 

and 40 µm high, forming in substomatal cavities, becoming 

erumpent, cells brown, thick-walled, pseudoparenchymatous. 

Conidiophores macronematous, mononematous, caespitose, 

emerging through the stomata, simple, flexuous, often geniculate-sinuous, 

4–7-septate, mainly smooth, dark brown, from 

a bulbous base tapering towards the apex, but often becoming 

more swollen, and also verrucose at the apex, (35–)80–120 

(–160) × (5–)6–7 µm. Conidiogenous cells cylindrical, becoming 

geniculate, integrated, terminal, polyblastic, proliferating 

sympodially, 20–45 × 5–7 µm, with conspicuous, cicatrised, 

protuberant, conidiogenous loci, 3 µm diam. Conidia subcylindrical, 

narrowing slightly to an obtuse apex (frequently 

swollen), and with a truncate base with a distinctly thickened, 

darkened, somewhat refractive hilum, 3 µm wide, red-brown, 

straight or curved, with 3–7(–12) mainly unconstricted eusepta, 

thick-walled, verrucose, (30–)50–65(–80) × (5–)6–7 µm. Conidiophores 

frequently arising from brown, erumpent spermatogonia, 

up to 150 µm wide. Spermatia hyaline, smooth, bacilliform, 

4–6 × 1–1.5 µm. 

Characteristics in culture — Colonies on MEA erumpent, 

with sparse aerial mycelium; margins feathery, crenate; surface


a 

b 

c 

d 

e 

f 

g 

h 

i 

Fig. 13 Verrucisporota grevilleae (CBS 124107). a. Leaf spots on Grevillea; b. conidiophores; c, d. conidiophores and conidiogenous cells; e–h. conidia; 

i. colony on PDA; j. colony on SNA. — Scale bars = 10 µm. 

j 

folded, with zones of salmon or smoke-grey mycelium; outer 

region and reverse olivaceous-grey; colonies reaching 10 mm 

diam after 1 mo. 

Specimen examined. Australia, Northern Territory, Emerald Springs 

(13°37'13.3" 131°36'40"), on leaves of Grevillea decurrens, 22 Sept. 2007, 

leg. B. Summerell, isol. P.W. Crous, CBS H-20205, cultures CPC 14761 = 

CBS 124107, CPC 14762, 14763. 

Notes — Conidia of V. grevilleae are narrower and longer, 

and conidiophores shorter than those of V. protearum (conidia 

23–51 × 5.6–10.5 µm, conidiophores up to 290 µm long, 4.5– 

8.5 µm wide; Shaw & Alcorn 1967). South African specimens 

from the genus Protea have conidia that are (20–)31–36(–49) 

× (7–)8.5–9.5(–12) µm (Crous et al. 2004a). These findings 

suggest that the fungus treated as V. protearum on Proteaceae 

(Shaw & Alcorn 1967, 1993, Beilharz & Pascoe 2002, Crous et 

al. 2004a), probably represents a complex of several taxa.


157 

Verrucisporota proteacearum (D.E. Shaw & Alcorn) 

D.E. Shaw & Alcorn, Austral. Syst. Bot. 6: 273. 1993 

Basionym. Verrucispora proteacearum D.E. Shaw & Alcorn, Proc. Linn. 

Soc. New South Wales 92: 171. 1967. 

Characteristics in culture — On MEA erumpent with sparse 

aerial mycelium; surface cream to pale olivaceous-grey, folded, 

with smooth, even margin; reverse brown-vinaceous; reaching 

8 mm diam after 2 wk. On PDA erumpent with sparse aerial 

mycelium and smooth to feathery margin; surface cream to pale 

olivaceous-grey; reverse olivaceous-grey, reaching 8 mm diam 

after 2 wk. On OA erumpent, with moderate aerial mycelium 

and uneven margin, pale white in middle, pale olivaceous-grey 

in outer region; reaching 10 mm diam after 2 wk. 

Specimen examined. Australia, Grevillea sp., V. Beilharz, VPRI31812 = 

CBS 116003. 

Notes — Because V. proteacearum was originally described 

from Finschia (conidia 23–51 × 5.6–10.5 µm; Shaw & Alcorn 

1967), there is a strong possibility that the strain listed here from 

Grevillea (conidia 30–45 × 10–12 µm on OA) may represent a 

different taxon to the one occurring on Finschia. Although apparently 

identical based on the LSU phylogeny (see Fig. 1), the 

ITS sequence of this isolate is different to that of V. grevilleae 

(95 % similarity and 4 % gaps). 

Key to species of Verrucisporota 

1. Conidia wider than 4.5 µm . . . . . . . . . . . . . . . . . . . . . . . . 2 

1. Conidia narrower than 4.5 µm . . . . . . . . . . . . . . . . . . . . . 3 

2. Conidia up to 56 µm long. . . . . . . . . . . . . . . . . . . . . . . . . 4 

2. Conidia longer than 56 µm, 3–7(–12)-septate, (30–)50– 

65(–80) × (5–)6–7 µm; on Grevillea . . . . . . . V. grevilleae 

3. Conidia mostly up to 30 µm long, (0–)2–3(–7)-septate, 

13–30(–70) × 2.75–4 µm; on Capparis . V. kimberleyana 

3. Conidia longer, mostly up to 77 µm long, 1–11-septate, 

(10–)27–77(–108) × 3–4.5 μm; on Struthanthus . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. struthanthicola 

4. Conidia up to 3-septate, obclavate, 1–3-septate, 32.5–55 

× 7–10.5 µm; on Celastrus . . . . . . . . . . . . . . . . . V. indica 

4. Conidia more than 3 septa . . . . . . . . . . . . . . . . . . . . . . . 5 

5. Conidia up to 32 µm long; (1–)3–4(–5)-septate, 20–32 × 

6–10 µm; on Bridelia . . . . . . . . . . . . . . . . . . . . V. brideliae 

5. Conidia frequently longer than above . . . . . . . . . . . . . . . 6 

6. Conidia 0–6-septate, 18–56 × 4.5–7 µm; on Daviesia (Beilharz 

& Pascoe 2002) . . . . . . . . . . . . . . . . . . . .V. daviesiae 

6. Conidia 3–7-septate, 23–51 × 5.6–10.5 µm; on Finschia 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. proteacearum 

Vonarxia Bat., Publ. Inst. Micol. Univ. Fed. Pernambuco 283: 

5. 1960 

Type species. Vonarxia anacardii Bat. & J.L. Bezerra. 

Mycelium immersed and superficial, composed of branched, 

septate, pale to medium brown, smooth to finely roughened 

hyphae. Conidiomata sporodochial; basal stroma composed 

of globose-ellipsoidal, brown, slightly roughened cells. Setae 

irregularly scattered throughout colony, simple, subulate with a 

bulbous base, straight to slightly curved, dark brown, smooth to 

slightly roughened, thick-walled, 5–16-euseptate, septa rather 

thick, but becoming thinner towards apex. Conidiogenous cells 

arise from upper cells of the stroma, tightly aggregated, doliiform 

to ellipsoid, pale brown to subhyaline or hyaline, smooth, giving 

rise to a cluster of conidia by means of sympodial proliferation, 

with successive conidia forming at a higher level. Conidia 

hyaline, smooth-walled, tetraradiate, basal cell subcylindrical to 

clavate to doliiform, 0–1-septate; upper three arms arise from 

the apical part of the basal cell, 3–10-septate, subcylindrical 

to cylindrical, apex subobtuse. 

Vonarxia vagans (Speg.) Aa, Persoonia 13: 128. 1986 — Fig. 

14 

Basionym. Ypsilonia vagans Speg., Revista Mus. La Plata, Secc. Bot. 

15: 35. 1908. 

≡ Kazulia vagans (Speg.) Nag Raj, Canad. J. Bot. 55: 1621. 1977. 

On PDA. Mycelium immersed and superficial, composed of 

branched, septate, pale to medium brown, smooth to finely 

roughened, 3–5 µm wide hyphae. Conidiomata sporodochial, 

flattened to erect and globose (especially on WA, not so on MEA 

or PDA, tending to be more flattened, and more hemispherical 

on OA), up to 300 µm diam; basal stroma up to 70 µm thick, 

composed of globose-ellipsoidal, brown, slightly roughened 

cells, 5–10 µm diam. Setae irregularly scattered throughout 

colony, simple, subulate with a bulbous base, straight to 

slightly curved, dark brown, smooth to slightly roughened, 

thick-walled (1–1.5 µm diam), (5–)10–12(–16)-septate, septa 

rather thick, but becoming thinner towards apex, basal cell 

10–13 µm wide, with slight taper towards bluntly rounded, obtuse 

apex, (120–)150–200(–220) µm; width at basal septum 

(5–)6(–7) µm; width at apical septum, 2–3(–5) µm; apical 

two cells frequently pale brown; individual cells 10–25 µm 

long. Conidiogenous cells arise from upper cells of the stroma, 

tightly aggregated, doliiform to ellipsoid, pale brown to subhyaline 

or hyaline, smooth, 8–10 × 3–5 µm, giving rise to a 

cluster of conidia by means of sympodial proliferation, with 

successive conidia forming at a higher level. Conidia hyaline, 

smooth-walled, tetraradiate, basal cell subcylindrical to clavate 

to doliiform, 0–1-septate, 10–15 × (1.5–)2–3 µm (10–18 µm 

long on OA); upper three arms arise from the apical part of the 

basal cell, 3–5-septate (prominently constricted at septa on WA 

and MEA, up to 10-septate on these media), subcylindrical to 

cylindrical, apex subobtuse, arms 20–55 µm long (20–90 µm 

on OA), 1.5–2 µm wide (2–3 µm wide on OA). 

Characteristics in culture — Colonies on OA spreading, 

with sparse aerial mycelium, and uneven, striate surface, with 

crenate margin; surface black, with patches of mouse-grey, 

reaching up to 25 mm diam after 1 mo; on PDA spreading, 

with sparse aerial mycelium and crenate margins; surface 

pale mouse-grey, outer region grey-olivaceous; reverse greyolivaceous, 

reaching up to 25 mm diam after 1 mo; on MEA 

spreading, erumpent with sparse aerial mycelium; surface 

prominently striate, margin crenate; centre black, outer region 

mouse-grey; reverse black; colonies reaching up to 20 mm 

diam after 1 mo. 

Specimens examined. Brazil, São Paulo Horto Botanico, leaves of Spiraea 

cantoniensis, Sept. 1905, leg. Usteri no. 15 bis, holotype LPS 12280; 

Rio Grande do Sul, Guaiba, living leaves of Stenocalyx uniflorus, 1 Apr. 2008, 

leg. A.C. Alfenas, isol. P.W. Crous, epitype designated here CBS H-20206, 

culture ex-type CPC 15151 = CBS 123533, CPC 15152. 

Notes — The holotype specimen (LPS 12280) was described 

and illustrated in detail by Nag Raj (1977). The species was 

originally described from leaves of Spiraea cantoniensis collected 

in the São Paulo Botanical Garden, where it occurred 

on leaves of several tree species, suggesting that it is not host 

specific. The present collection was obtained by incubating 

Eugenia leaves with leaf spots of Phaeophleospora eugeniae 

in moist chambers, which resulted in a few conidiophores of 

Vonarxia vegans developing. 

Nag Raj (1977) erected Kazulia for a genus of hyphomycetes 

with dark brown, septate setae, and tetraradiate conidia, 

which he regarded as morphologically distinct, and a probable 

anamorph of the Chaetothyriaceae. The fact that he did 

not compare Kazulia with Vonarxia is not surprising, because


a 

b 

c 

d 

e 

f 

g 

h 

j 

k 

l 

i 

m 

Fig. 14 Vonarxia vagans (CBS 123533). a, b. Colony on PDA; c. colony with setae; d, e. setae with rounded apices and swollen bases, lacking rhizoids; f–i. 

conidiogenous cells giving rise to conidia; j–o. conidia. — Scale bars = 10 µm. 

n 

o 

Batista et al. (1960) who initially described Vonarxia, showed 

setae on the outside of the pycnidia, and thus this fungus was 

regarded as a coelomycete. Later comments from Nag Raj 

(1977) (as Kazulia) suggest, however, that these bodies are 

perithecia of a probable teleomorph. In a subsequent study 

Van der Aa & Van Oorschot (1985) and Van der Aa & Von Arx 

(1986) showed that Kazulia is a synonym of Vonarxia. Wu & 

Sutton (1995) were not convinced of the distinction between 

Vonarxia and another hyphomycete genus, Fumagopsis, due 

to insufficient material, and chose to use the name Fumagopsis 

for F. complexa, which they described from Eugenia leaves collected 

in India. Based on the present collection of V. vagans, it


159 

is apparent, that these are two distinct genera. In Fumagopsis 

the setae are aseptate, arranged around the sporodochium, 

and taxa have rhizoid-like structures. In contrast, the setae of 

Vonarxia are septate, irregularly distributed and do not surround 

the sporodochium, and have a simple, bulbous base. 

Key to species of Vonarxia 

1. Setae 87–155 µm long; apical conidial arms 12–35 µm 

long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V. anacardii 

1. Setae and conidial arms longer; setae 120–220 µm long; 

apical conidial arms 20–55(–90) µm long . . . . V. vagans 

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

Type species. Xenostigmina zilleri (A. Funk) Crous. 

Associated with 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, 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. 

Xenostigmina zilleri (A. Funk) Crous, Mycol. Mem. 21: 155. 

1998 — Fig 15 

Basionym. Stigmina zilleri A. Funk, Canad. J. Bot. 65: 482. 1987. 

Synanamorph. Mycopappus aceris (Dearn. & Barthol.) Redhead & G.P. 

White, Canad. J. Bot. 63: 1430. 1985. 

Basionym. Cercosporella aceris Dearn. & Barthol., Mycologia 9: 362. 

1917. 

Teleomorph. ? Didymella mycopappi (A. Funk & Dorworth) Crous, Mycol. 

Mem. 21: 152. 1998. 

Basionym. Mycosphaerella mycopappi A. Funk & Dorworth, Canad. J. 

Bot. 66: 295. 1988. 

Characteristics in culture — Colonies spreading on PDA with 

moderate to abundant aerial mycelium, and feathery margins; 

olivaceous-grey with patches of iron-grey and pale olivaceousgrey; 

iron-grey in reverse. On OA spreading, with abundant 

aerial mycelium, olivaceous-grey with patches of pale olivaceous-grey. 

On MEA erumpent, spreading, with abundant aerial 

mycelium, pale olivaceous-grey with patches of olivaceous-grey 

and iron-grey; reverse iron-grey. 

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 Acer macrophyllum, 6 Sept. 2007, leg. B. Callan, isol. P.W. Crous, CBS 

H-20208, CPC 14376 = CBS 124108, CPC 14377, 14378 (Xenostigmina zilleri), 

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

Notes — Although Stigmina s.str. has been shown to reside in 

Pseudocercospora s.str. (Crous et al. 2006a, Braun & Crous 2006, 

2007), this is not the case for Xenostigmina (Crous 1998), which 

appears to be related to Seifertia (Seifert et al. 2007) in the 

Dothideomycetes. Isolates of the Xenostigmina state are shown 

here (Fig. 1) to be identical to those of the Mycopappus state, 

which proves that these two genera are indeed synanamorphs. 

No ascospore isolates were obtained, however, to confirm their 

relationship to ‘Mycosphaerella’ mycopappi, though this species 

is clearly not a member of the Mycosphaerellaceae. Xenostigmina 

wolfii (Crous & Corlett 1998), which is the anamorph of 

Mycosphaerella stigmina-platani, and a Pseudocercospora 

synanamorph, is not congeneric with X. zilleri, and would be 

better accommodated in Pseudocercospora (Crous et al. 2006a) 

than in Xenostigmina. 

a 

b 

c 

d 

e 

f 

g 

h 

Fig. 15 Xenostigmina zilleri (CBS 124108). a–c. Conidial propagules of Mycopappus aceris; d. setae on the surface of conidial propagules; e. colony of 

Xenostigmina zilleri; f, g. fasciculate conidiophores; h. conidia. — Scale bars = 10 µm.


Acknowledgements We acknowledge Drs B.E. Callan (Canadian Forest 

Service, Natural Resources Canada), F. Dugan (Washington State University, 

Pullman), H. Evans (CABI, UK) and V. Beilharz (Department of Primary Industries, 

Knoxfield, Victoria, Australia), who provided valuable specimens for 

study. We are grateful to the technical staff including A. van Iperen, M. Vermaas, 

and M. Starink for providing assistance with cultures, photoplates and DNA 

sequencing, respectively. Dr E.G. Simmons (Crawfordsville, Indiana, USA) is 

acknowledged for bringing to our attention the problem of Alternaria malorum 

and for providing strains representing related novel taxa known to him, for 

inclusion in this study. 

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Phylogeny and taxonomy of obscure genera of microfungi - Persoonia

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