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Mycosphaerella is polyphyletic

¹ CBS Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD, Utrecht, The Netherlands
² Martin-Luther-Universität, Institut für Biologie, Geobotanik und Botanischer Garten, Herbarium, Neuwerk 21, D-06099 Halle, Germany

^* Correspondence: Pedro W. Crous, p.crous{at}cbs.knaw.nl

	Abstract

TOP Abstract INTRODUCTION MATERIALS AND METHODS RESULTS DISCUSSION References

 
Mycosphaerella, one of the largest genera of ascomycetes, encompasses several thousand species and has anamorphs residing in more than 30 form genera. Although previous phylogenetic studies based on the ITS rDNA locus supported the monophyly of the genus, DNA sequence data derived from the LSU gene distinguish several clades and families in what has hitherto been considered to represent the Mycosphaerellaceae. Several important leaf spotting and extremotolerant species need to be disposed to the genus Teratosphaeria, for which a new family, the Teratosphaeriaceae, is introduced. Other distinct clades represent the Schizothyriaceae, Davidiellaceae, Capnodiaceae, and the Mycosphaerellaceae. Within the two major clades, namely Teratosphaeriaceae and Mycosphaerellaceae, most anamorph genera are polyphyletic, and new anamorph concepts need to be derived to cope with dual nomenclature within the Mycosphaerella complex.

Taxonomic novelties: Batcheloromyces eucalypti (Alcorn) Crous & U. Braun, comb. nov., Catenulostroma Crous & U. Braun, gen. nov., Catenulostroma abietis (Butin & Pehl) Crous & U. Braun, comb. nov., Catenulostroma chromoblastomycosum Crous & U. Braun, sp. nov., Catenulostroma elginense (Joanne E. Taylor & Crous) Crous & U. Braun, comb. nov., Catenulostroma excentricum (B. Sutton & Ganap.) Crous & U. Braun, comb. nov., Catenulostroma germanicum Crous & U. Braun, sp. nov., Catenulostroma macowanii (Sacc.) Crous & U. Braun, comb. nov., Catenulostroma microsporum (Joanne E. Taylor & Crous) Crous & U. Braun, comb. nov., Catenulostroma protearum (Crous & M.E. Palm) Crous & U. Braun, comb. nov., Penidiella Crous & U. Braun, gen. nov., Penidiella columbiana Crous & U. Braun, sp. nov., Penidiella cubensis (R.F. Castañeda) U. Braun, Crous & R.F. Castañeda, comb. nov., Penidiella nectandrae Crous, U. Braun & R.F. Castañeda, nom. nov., Penidiella rigidophora Crous, R.F. Castañeda & U. Braun, sp. nov., Penidiella strumelloidea (Milko & Dunaev) Crous & U. Braun, comb. nov., Penidiella venezuelensis Crous & U. Braun, sp. nov., Readeriella blakelyi (Crous & Summerell) Crous & U. Braun, comb. nov., Readeriella brunneotingens Crous & Summerell, sp. nov., Readeriella considenianae (Crous & Summerell) Crous & U. Braun, comb. nov., Readeriella destructans (M.J. Wingf. & Crous) Crous & U. Braun, comb. nov., Readeriella dimorpha (Crous & Carnegie) Crous & U. Braun, comb. nov., Readeriella epicoccoides (Cooke & Massee) Crous & U. Braun, comb. nov., Readeriella gauchensis (M.-N. Cortinas, Crous & M.J. Wingf.) Crous & U. Braun, comb. nov., Readeriella molleriana (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov., Readeriella nubilosa (Ganap. & Corbin) Crous & U. Braun, comb. nov., Readeriella pulcherrima (Gadgil & M. Dick) Crous & U. Braun, comb. nov., Readeriella stellenboschiana (Crous) Crous & U. Braun, comb. nov., Readeriella toledana (Crous & Bills) Crous & U. Braun, comb. nov., Readeriella zuluensis (M.J. Wingf., Crous & T.A. Cout.) Crous & U. Braun, comb. nov., Teratosphaeria africana (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov., Teratosphaeria alistairii (Crous) Crous & U. Braun, comb. nov., Teratosphaeria associata (Crous & Carnegie) Crous & U. Braun, comb. nov., Teratosphaeria bellula (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov., Teratosphaeria cryptica (Cooke) Crous & U. Braun, comb. nov., Teratosphaeria dentritica (Crous & Summerell) Crous & U. Braun, comb. nov., Teratosphaeria excentrica (Crous & Carnegie) Crous & U. Braun, comb. nov., Teratosphaeria fimbriata (Crous & Summerell) Crous & U. Braun, comb. nov., Teratosphaeria flexuosa (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov., Teratosphaeria gamsii (Crous) Crous & U. Braun, comb. nov., Teratosphaeria jonkershoekensis (P.S. van Wyk, Marasas & Knox-Dav.) Crous & U. Braun, comb. nov., Teratosphaeria maxii (Crous) Crous & U. Braun, comb. nov., Teratosphaeria mexicana (Crous) Crous & U. Braun, comb. nov., Teratosphaeria molleriana (Thüm.) Crous & U. Braun, comb. nov., Teratosphaeria nubilosa (Cooke) Crous & U. Braun, comb. nov., Teratosphaeria ohnowa (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov., Teratosphaeria parkiiaffinis (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov., Teratosphaeria parva (R.F. Park & Keane) Crous & U. Braun, comb. nov., Teratosphaeria perpendicularis (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov., Teratosphaeria pluritubularis (Crous & Mansilla) Crous & U. Braun, comb. nov., Teratosphaeria pseudafricana (Crous & T.A. Cout.) Crous & U. Braun, comb. nov., Teratosphaeria pseudocryptica (Crous) Crous & U. Braun, comb. nov., Teratosphaeria pseudosuberosa (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov., Teratosphaeria quasicercospora (Crous & T.A. Cout.) Crous & U. Braun, comb. nov., Teratosphaeria readeriellophora (Crous & Mansilla) Crous & U. Braun, comb. nov., Teratosphaeria secundaria (Crous & Alfenas) Crous & U. Braun, comb. nov., Teratosphaeria stramenticola (Crous & Alfenas) Crous & U. Braun, comb. nov., Teratosphaeria suberosa (Crous, F.A. Ferreira, Alfenas & M.J. Wingf.) Crous & U. Braun, comb. nov., Teratosphaeria suttonii (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov., Teratosphaeria toledana (Crous & Bills) Crous & U. Braun, comb. nov., Teratosphaeriaceae Crous & U. Braun, fam. nov.

Keywords Ascomycetes / Batcheloromyces / Colletogloeopsis / Readeriella / Teratosphaeria / Trimmatostroma / DNA sequence comparisons / systematics

	INTRODUCTION

TOP Abstract INTRODUCTION MATERIALS AND METHODS RESULTS DISCUSSION References

 
The genus Mycosphaerella Johanson as presently circumscribed contains close to 3 000 species (Aptroot 2006), excluding its anamorphs, which represent thousands of additional species (Crous et al. 2000, 2001, 2004a, b, 2006a, b, 2007b, Crous & Braun 2003). Crous (1998) predicted that Mycosphaerella would eventually be split according to its anamorph genera, and Crous et al. (2000) recognised six sections, as originally defined by Barr (1972). This was followed by a set of papers (Crous et al. 2001, Goodwin et al. 2001), where it was concluded, based on ITS DNA sequence data, that Mycosphaerella was monophyletic. A revision of the various coelomycete and hyphomycete anamorph concepts led Crous & Braun (2003) to propose a system whereby the asexual morphs could be allocated to various form genera affiliated with Mycosphaerella holomorphs.

In a recent study that formed part of the US "Assembling the Fungal Tree of Life" project, Schoch et al. (2006) were able to show that the Mycosphaerellaceae represents a family within Capnodiales. Furthermore, some variation was also depicted within the family, which supported similar findings in other recent papers employing LSU sequence data, such as Hunter et al. (2006), and Batzer et al. (2007). To further elucidate the phylogenetic variation observed within the Mycosphaerellaceae in these studies, a subset of isolates was selected for the present study, representing the various species recognised as morphologically distinct from Mycosphaerella s. str.

The genus Mycosphaerella has in recent years been linked to approximately 30 anamorph genera (Crous & Braun 2003, Crous et al. 2007b). Many of these anamorph genera resulted from a reassessment of cercosporoid forms. Chupp (1954) was of the opinion that they all represented species of the genus Cercospora Fresen., although he clearly recognised differences in their morphology. In a series of papers by Deighton, as well as others such as Sutton, Braun and Crous, the genus Cercospora was delimited based on its type species, Cercospora penicillata (Ces.) Fresen., while taxa formerly included in the genus by Chupp (1954) but differing in conidiophore arrangement, conidiogenesis, pigmentation, conidial catenulation, septation, and scar/hilum structure were allocated to other genera. Similar studies in which the type species were recollected and subjected to DNA sequence analysis were undertaken to characterise Mycosphaerella (Verkley et al. 2004), and anamorph genera such as Pseudocercospora Speg., Stigmina Sacc., Phaeoisariopsis Ferraris (Crous et al. 2006a), Ramulispora Miura (Crous et al. 2003), Batcheloromyces Marasas, P.S. van Wyk & Knox-Dav. (Taylor et al. 2003), Phaeophleospora Rangel and Dothistroma Hulbary (Crous et al. 2000, 2001, Barnes et al. 2004).

To assess the phylogeny of the species selected for the present study, DNA sequences were generated of the 28S rRNA (LSU) gene. In a further attempt to address monophyletic groups within this complex, these data were integrated with their morphological characteristics. To further resolve pleomorphism among the species studied, isolates were examined on a range of cultural media to induce possible synanamorphs.

	MATERIALS AND METHODS

TOP Abstract INTRODUCTION MATERIALS AND METHODS RESULTS DISCUSSION References

 
Isolates
Chosen isolates represent various species previously observed to be morphologically distinct from Mycosphaerella s. str. (Crous 1998, Crous et al. 2004a, b, 2006a, b, 2007b). In a few cases, specifically Teratosphaeria fibrillosa Syd. & P. Syd. and Coccodinium bartschii A. Massal., fresh material had to be collected from South Africa and Canada, respectively. Excised tissue pieces bearing ascomata were soaked in water for approximately 2 h, after which they were placed in the bottom of Petri dish lids, with the top half of the dish containing 2 % malt extract agar (MEA) (Gams et al. 2007). Ascospore germination patterns were examined after 24 h, and single-ascospore and conidial cultures established as described by Crous (1998). Colonies were sub-cultured onto synthetic nutrient-poor agar (SNA), potato-dextrose agar (PDA), oatmeal agar (OA), MEA (Gams et al. 2007), and incubated at 25 °C under continuous near-ultraviolet light to promote sporulation.

DNA phylogeny
Fungal colonies were established on agar plates, and genomic DNA was isolated following the CTAB-based protocol described in Gams et al. (2007). 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 5' end of the 28S rRNA gene (LSU). The primers ITS4 (White et al. 1990), LR0R (Rehner & Samuels 1994), LR3R (www.biology.duke.edu/fungi/mycolab/primers.htm), and LR16 (Moncalvo et al. 1993), were used as internal sequence primers to ensure good quality sequences over the entire length of the amplicon. The ITS1, ITS2 and 5.8S rRNA gene (ITS) were only sequenced for isolates of which these data were not available. The ITS data were not included in the analyses but deposited in GenBank where applicable. The PCR conditions, sequence alignment and subsequent phylogenetic analysis using parsimony, distance and Bayesian analyses followed the methods of Crous et al. (2006c). Gaps longer than 10 bases were coded as single events for the phylogenetic analyses; the remaining gaps were treated as new character states. Sequence data were deposited in GenBank (Table 1) and alignments in TreeBASE (www.treebase.org).

Taxonomy
Wherever possible, 30 measurements (x 1 000 magnification) were made of structures mounted in lactic acid, with the extremes of spore measurements given in parentheses. Ascospores were frequently also mounted in water to observe mucoid appendages and sheaths. Colony colours (surface and reverse) were assessed after 1-2 mo on MEA at 25 °C in the dark, using the colour charts of Rayner (1970). All cultures obtained in this study are maintained in the culture collection of the Centraalbureau voor Schimmelcultures (CBS) in Utrecht, the Netherlands (Table 1). Nomenclatural novelties and descriptions were deposited in MycoBank (www.MycoBank.org).

	RESULTS

TOP Abstract INTRODUCTION MATERIALS AND METHODS RESULTS DISCUSSION References

 
DNA phylogeny
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 GenBank which were added to the alignment. The manually adjusted alignment contained 97 sequences (including the two outgroup sequences) and 844 characters including alignment gaps. Of the 844 characters used in the phylogenetic analysis, 308 were parsimony-informative, 105 were variable and parsimony-uninformative, and 431 were constant.

The parsimony analysis of the LSU region yielded 1 135 equally most parsimonious trees (TL = 1 502 steps; CI = 0.446; RI = 0.787; RC = 0.351), one of which is shown in Fig. 1. Three orders are represented by the ingroup isolates, namely Chaetothyriales (100 % bootstrap support), Helotiales (100 % bootstrap support) and Capnodiales (100 % bootstrap support). These are discussed in detail in the Taxonomy and Discussion sections. A new collection of Coccodinium bartschii A. Massal clusters (100 % bootstrap support) with members of the Herpotrichiellaceae (Chaetothyriales), whereas the type species of the genus Trimmatostroma Corda, namely T. salicis Corda, as well as T. betulinum (Corda) S. Hughes, are allied (99 % bootstrap support) with the Dermateaceae (Helotiales). The Capnodiales encompasses members of the Capnodiaceae, Trichosphaeriaceae, Davidiellaceae, Schizothyriaceae and taxa traditionally placed in the Mycosphaerellaceae, which is divided here into the Teratosphaeriaceae, (65 % bootstrap support), and the Mycosphaerellaceae (76 % bootstrap support), which contains several subclades. Also included in the Capnodiales are Devriesia staurophora (W.B. Kendr.) Seifert & N.L. Nick., Staninwardia suttonii Crous & Summerell and Capnobotryella renispora Sugiy. as sister taxa to Teratosphaeriaceae s. str. Neighbour-joining analysis using three substitution models on the sequence data yielded trees supporting the same topologies, but differed from the parsimony tree presented with regard to the order of the families and orders at the deeper nodes, e.g., the Helotiales and Chaetothyriales are swapped around, as are the Capnodiaceae and the Trichosphaeriaceae/Davidiellaceae (data not shown). Using neighbour-joining analyses, the Mycosphaerellaceae s. str. clade obtained 71 %, 70 % and 70 % bootstrap support respectively with the uncorrected "p", Kimura 2-parameter and HKY85 substitution models wherease the Teratosphaeriaceae clade obtained 74 %, 79 % and 78 % bootstrap support respectively with the same models. The Schizothyriaceae clade appeared basal in the Capnodiales, irrespective of which substitution model was used.

View larger version (33K): [in this window] [in a new window]   Fig. 1. One of 1 135 equally most parsimonious trees obtained from a heuristic search with 100 random taxon additions of the LSU sequence alignment using PAUP v. 4.0b 10. The scale bar shows 10 changes, and bootstrap support values from 1000 replicates are shown at the nodes. Thickened lines indicate the strict consensus branches and ex-type sequences are printed in bold face. The tree was rooted to two sequences obtained from GenBank (Athelia epiphylla AY586633 and Paullicorticium ansatum AY586693).

View larger version (33K): [in this window] [in a new window]   Fig. 2. Consensus phylogram (50 % majority rule) of 18 815 trees resulting from a Bayesian analysis of the LSU sequence alignment using MRBAYES v. 3.1.2. Bayesian posterior probabilities are indicated at the nodes. Ex-type sequences are printed in bold face. The tree was rooted to two sequences obtained from GenBank (Athelia epiphylla AY586633 and Paullicorticium ansatum AY586693).

Key to Mycosphaerella, and Mycosphaerella-like genera treated

• 1. Ascomata thyrothecial; anamorph Zygosporium................................................................................................................ Schizothyrium.
  
• 1. Ascomata pseudothecial........................................................................................................................................................................... 2
  
• 2. Ascospores with irregular, angular lumens typical of Davidiella; anamorph Cladosporium s. str............................................... Davidiella
  
• 2. Ascospores guttulate or not, lacking angular lumens; anamorph other than Cladosporium..................................................................... 3
  
• 3. Ascomata frequently linked by superficial stroma; hamathecial tissue, ascospore sheath, multi-layered endotunica, prominent periphysoids, and ascospores turning brown in asci frequently observed........................................................................ Teratosphaeria
  
• 3. Ascomata not linked by superficial stroma; hamathecial tissue, ascospore sheath, multi-layered endotunica, prominent periphysoids, ascospores turning brown in asci not observed........................................................................................................................................ 4
  
• 4. Conidiophores solitary, pale brown, giving rise to primary and secondary, actively discharged conidia; anamorph Dissoconium................................................................................................................................................................... teleomorph Mycosphaerella-like
  
• 4. Conidiomata variable from solitary conidiophores to sporodochia, fascicles to pycnidia, but conidia not actively discharged..................................................................................................................................................................................................... Mycosphaerella s. str.
  

Treatment of phylogenetic clades
Davidiellaceae clade
Davidiella Crous & U. Braun, Mycol. Progr. 2: 8. 2003.

Type species: Davidiella tassiana (De Not.) Crous & U. Braun, Mycol. Progr. 2: 8. 2003.

Basionym: Sphaerella tassiana De Not., Sferiacei Italici 1: 87. 1863.

Description: Schubert et al. (2007 - this volume).

Anamorph: Cladosporium Link, Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 7: 37. 1816.

Type species: Cladosporium herbarum (Pers.: Fr.) Link, Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 7: 37. 1816.

Basionym: Dematium herbarum Pers., Ann. Bot. (Usteri), 11 Stück: 32. 1794: Fr., Syst. Mycol. 3: 370. 1832.

Description: Schubert et al. (2007 - this volume).

Notes: The genus Davidiella (Davidiellaceae) was recently introduced for teleomorphs of Cladosporium s. str. (Braun et al. 2003). The genus Cladosporium is well-established, and contains around 772 names (Dugan et al. 2004), while Davidiella presently has 33 names (www.MycoBank.org), of which only around five have acknowledged Cladosporium states.

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

Type species: Teratosphaeria fibrillosa Syd. & P. Syd., Ann. Mycol. 10: 40. 1912. Fig. 3.

View larger version (143K): [in this window] [in a new window]   Fig. 3. Teratosphaeria fibrillosa (epitype material). A. Leaf spots. B. Subepidermal ascomata linked by means of stromatic tissue. C. Paraphyses among asci. D. Periphysoids. E. Ascospores becoming brown in asci. F-G. Multi-layered endotunica. H-K. Ascospores, becoming brown and verruculose. L-M. Germinating ascospores. Scale bars = 10 µm.

Notes: Although similar in morphology, the genus Teratosphaeria was separated from Mycosphaerella based on its ascomatal arrangement, and periphysate ostioles (Müller & Oehrens 1982). It was later synonymised under Mycosphaerella by Taylor et al. (2003), who showed that the type species clustered within Mycosphaerella based on ITS DNA sequence data. The LSU sequence data generated in the present study, has clearly shown that Mycosphaerella is polyphyletic, thus contradicting earlier reports of monophyly by Crous et al. (2000) and Goodwin et al. (2001), which were based on ITS data.

A re-examination of T. fibrillosa, the type species of Teratosphaeria, revealed several morphological features that characterise the majority of the taxa clustering in the clade, though several characters have been lost in some of the small-spored species. These characters are discussed below:

1. Teratosphaeria fibrillosa has a superficial stroma linking ascomata together, almost appearing like a spider's web on the leaf surface. Although this feature is not seen in other taxa in this clade, some species, such as M. suberosa Crous, F.A. Ferreira, Alfenas & M.J. Wingf. and M. pseudosuberosa Crous & M.J. Wingf. have a superficial stroma, into which the ascomata are inbedded (Crous 1998, Crous et al. 2006b).
   
2. Ascospores of Teratosphaeria become brown and verruculose while still in their asci. This feature is commonly observed in species such as M. jonkershoekensis P.S. van Wyk, Marasas & Knox-Dav., M. alistairii Crous, M. mexicana Crous, M. maxii Crous and M. excentricum Crous & Carnegie (Crous 1998, Crous & Groenewald 2006a, b, Crous et al. 2007b).
   
3. A few ascomata of T. fibrillosa were found to have some pseudoparaphysoidal remnants (cells to distinguish pseudoparaphyses), though they mostly disappear with age. This feature is rather uncommon, though pseudoparaphyses were observed in ascomata of M. eucalypti (Wakef.) Hansf.
   
4. Ascospores of Teratosphaeria were found to be covered in a mucous sheath, which is commonly observed in other taxa in this clade, such as M. bellula Crous & M.J. Wingf., M. pseudocryptica Crous, M. suberosa, M. pseudosuberosa, M. associata Crous & Carnegie, M. dendritica Crous & Summerell and M. fimbriata Crous & Summerell (Crous et al. 2004b, 2006b, 2007b). Re-examination of fresh collections also revealed ascospores of M. cryptica (Cooke) Hansf. and M. nubilosa (Cooke) Hansf. to have a weakly definable sheath. Germinating ascospores of species in this clade all exhibit a prominent mucoid sheath.
   
5. Asci of T. fibrillosa were observed to have a multi-layered endotunica, which, although not common, can be seen in species such as M. excentrica, M. maxii, M. alistairii, M. pseudosuberosa, M. fimbriata (Crous et al. 2006b, 2007b, Crous & Groenewald 2006a, b), and also M. nubilosa.
   
6. Finally, ascomata of T. fibrillosa and T. proteae-arboreae P.S. van Wyk, Marasas & Knox-Dav. have well-developed ostiolar periphyses, which are also present in species such as M. suberosa, M. pseudosuberosa, M. maxii and T. microspora Joanne E. Taylor & Crous (Crous 1998, Crous et al. 2004a, b, 2006b). Morphologically thus, the Teratosphaeria clade is distinguishable from Mycosphaerella s. str., though these differences are less pronounced in some of the smaller-spored species. Based on these distinct morphological features, as well as its phylogenetic position within the Capnodiales, a new family is herewith proposed to accommodate species of Teratosphaeria:
   

Teratosphaeriaceae Crous & U. Braun, fam. nov. MycoBank MB504464.

Ascomata pseudotheciales, superficiales vel immersa, saepe in stromate ex cellulis brunneis pseudoparenchymatibus disposita, globulares, uniloculares, papillata, apice ostiolato, periphysata, saepe cum periphysoidibus; tunica multistratosa, ex cellulis brunneis angularibus composita, strato interiore ex cellulis applanatis hyalinis; saepe cum pseudoparaphysibus subcylindricis, ramosis, septatis, anastomosibus. Asci fasciculati, octospori, bitunicati, saepe cum endotunica multistratosa. Ascosporae ellipsoideae-fusiformes vel obovoideae, 1-septatae, hyalinae, deinde pallide brunneae et verruculosae, saepe mucosae.

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.

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

Teratosphaeria africana (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov. MycoBank MB504466.

Basionym: Mycosphaerella africana Crous & M.J. Wingf., Mycologia 88: 450. 1996.

Teratosphaeria associata (Crous & Carnegie) Crous & U. Braun, comb. nov. MycoBank MB504467.

Basionym: Mycosphaerella associata Crous & Carnegie, Fungal Diversity 26: 159. 2007.

Teratosphaeria alistairii (Crous) Crous & U. Braun, comb. nov. MycoBank MB504468.

Basionym: Mycosphaerella alistairii Crous, in Crous & Groenewald, Fungal Planet, No. 4. 2006.

Anamorph: Batcheloromyces sp.

Teratosphaeria bellula (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov. MycoBank MB504469.

Basionym: Mycosphaerella bellula Crous & M.J. Wingf., Mycotaxon 46: 20. 1993.

Teratosphaeria cryptica (Cooke) Crous & U. Braun, comb. nov. MycoBank MB504470.

Basionym: Sphaerella cryptica Cooke, Grevillea 20: 5. 1891.

Mycosphaerella cryptica (Cooke) Hansf., Proc. Linn. Soc. New South Wales 81: 35. 1956.

Anamorph: Readeriella nubilosa (Ganap. & Corbin) Crous & U. Braun, comb. nov. MycoBank MB504471.

Basionym: Colletogloeum nubilosum Ganap. & Corbin, Trans. Brit. Mycol. Soc. 72: 237. 1979.

Colletogloeopsis nubilosum (Ganap. & Corbin) Crous & M.J. Wingf., Canad. J. Bot. 75: 668. 1997.

Teratosphaeria dendritica (Crous & Summerell) Crous & U. Braun, comb. nov. MycoBank MB504472.

Basionym: Mycosphaerella dendritica Crous & Summerell, Fungal Diversity 26: 161. 2007.

Anamorph: Nothostrasseria dendritica (Hansf.) Nag Raj, Canad. J. Bot. 61: 25. 1983.

Basionym: Spilomyces dendriticus Hansf., Proc. Linn. Soc. New South Wales 81: 32. 1956.

Teratosphaeria excentrica (Crous & Carnegie) Crous & U. Braun, comb. nov. MycoBank MB504473.

Basionym: Mycosphaerella excentrica Crous & Carnegie, Fungal Diversity 26: 164. 2007.

Anamorph: Catenulostroma excentricum (B. Sutton & Ganap.) Crous & U. Braun, comb. nov. MycoBank MB504475.

Basionym: Trimmatostroma excentricum B. Sutton & Ganap., New Zealand J. Bot. 16: 529. 1978.

Teratosphaeria fibrillosa Syd. & P. Syd., Ann. Mycol. 10: 40. 1912.

Mycosphaerella fibrillosa (Syd. & P. Syd.) Joanne E. Taylor & Crous, Mycol. Res. 107: 657. 2003.

Specimens examined: South Africa, Western Cape Province, Bains Kloof near Wellington, on living leaves of Protea grandiflora, 26 Feb. 1911, E.M. Doidge, holotype PREM; Stellenbosch, Jonkershoek valley, S33° 59' 44.7", E18° 58' 50.6", 1 Apr. 2007, on leaves of Protea sp., P.W. Crous & L. Mostert, epitype designated here CBS H-19913, culture ex-epitype CBS 121707 = CPC 13960.

Teratosphaeria fimbriata (Crous & Summerell) Crous & U. Braun, comb. nov. MycoBank MB504476.

Basionym: Mycosphaerella fimbriata Crous & Summerell, Fungal Diversity 26: 166. 2007.

Teratosphaeria flexuosa (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov. MycoBank MB504477.

Basionym: Mycosphaerella flexuosa Crous & M.J. Wingf., Mycol. Mem. 21: 58. 1998.

Teratosphaeria gamsii (Crous) Crous & U. Braun, comb. nov. MycoBank MB504478.

Basionym: Mycosphaerella gamsii Crous, Stud. Mycol. 55: 113. 2006.

Teratosphaeria jonkershoekensis (P.S. van Wyk, Marasas & Knox-Dav.) Crous & U. Braun, comb. nov. MycoBank MB504479.

Basionym: Mycosphaerella jonkershoekensis P.S. van Wyk, Marasas & Knox-Dav., J. S. African Bot. 41: 234. 1975.

Teratosphaeria maxii (Crous) Crous & U. Braun, comb. nov. MycoBank MB504480.

Basionym: Mycosphaerella maxii Crous, in Crous & Groenewald, Fungal Planet No. 6. 2006.

Teratosphaeria mexicana (Crous) Crous & U. Braun, comb. nov. MycoBank MB504481.

Basionym: Mycosphaerella mexicana Crous, Mycol. Mem. 21: 81. 1998.

Teratosphaeria microspora Joanne E. Taylor & Crous, Mycol. Res. 104: 631. 2000.

Mycosphaerella microspora (Joanne E. Taylor & Crous) Joanne E. Taylor & Crous, Mycol. Res. 107: 657. 2003.

Anamorph: Catenulostroma microsporum (Joanne E. Taylor & Crous) Crous & U. Braun, comb. nov. MycoBank MB504482.

Basionym: Trimmatostroma microsporum Joanne E. Taylor & Crous, Mycol. Res. 104: 631. 2000.

Teratosphaeria molleriana (Thüm.) Crous & U. Braun, comb. nov. MycoBank MB504483.

Basionym: Sphaerella molleriana Thüm., Revista Inst. Sci. Lit. Coimbra 28: 31. 1881.

Mycosphaerella molleriana (Thüm) Lindau, Nat. Pfanzenfam. 1: 424. 1897.

= Mycosphaerella vespa Carnegie & Keane, Mycol. Res. 102: 1275. 1998.

= Mycosphaerella ambiphylla A. Maxwell, Mycol. Res. 107: 354. 2003.

Anamorph: Readeriella molleriana (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov. MycoBank MB504484.

Basionym: Colletogloeopsis molleriana Crous & M.J. Wingf., Canad. J. Bot. 75: 670. 1997.

Teratosphaeria nubilosa (Cooke) Crous & U. Braun, comb. nov. MycoBank MB504485.

Basionym: Sphaerella nubilosa Cooke, Grevillea 19: 61. 1892.

Mycosphaerella nubilosa (Cooke) Hansf., Proc. Linn. Soc. New South Wales 81: 36. 1965.

= Mycosphaerella juvenis Crous & M.J. Wingf., Mycologia 88: 453. 1996.

Teratosphaeria ohnowa (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov. MycoBank MB504486.

Basionym: Mycosphaerella ohnowa Crous & M.J. Wingf., Stud. Mycol. 50: 206. 2004.

Teratosphaeria parkiiaffinis (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov. MycoBank MB504487.

Basionym: Mycosphaerella parkiiaffinis Crous & M.J. Wingf., Fungal Diversity 26:168. 2007.

Teratosphaeria parva (R.F. Park & Keane) Crous & U. Braun, comb. nov. MycoBank MB504488.

Basionym: Mycosphaerella parva R.F. Park & Keane, Trans. Brit. Mycol. Soc. 79: 99. 1982.

= Mycosphaerella grandis Carnegie & Keane, Mycol. Res. 98: 414. 1994.

Teratosphaeria perpendicularis (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov. MycoBank MB504489.

Basionym: Mycosphaerella perpendicularis Crous & M.J. Wingf., Stud. Mycol. 55: 113. 2006.

Teratosphaeria pluritubularis (Crous & Mansilla) Crous & U. Braun, comb. nov. MycoBank MB504490.

Basionym: Mycosphaerella pluritubularis Crous & Mansilla, Stud. Mycol. 55: 114. 2006.

Teratosphaeria pseudafricana (Crous & T.A. Cout.) Crous & U. Braun, comb. nov. MycoBank MB504491.

Basionym: Mycosphaerella pseudafricana Crous & T.A. Cout., Stud. Mycol. 55: 115. 2006.

Teratosphaeria pseudocryptica (Crous) Crous & U. Braun, comb. nov. MycoBank MB504492.

Basionym: Mycosphaerella pseudocryptica Crous, Stud. Mycol. 55: 116. 2006.

Anamorph: Readeriella sp.

Teratosphaeria pseudosuberosa (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov. MycoBank MB504493.

Basionym: Mycosphaerella pseudosuberosa Crous & M.J. Wingf., Stud. Mycol. 55: 118. 2006.

Anamorph: Catenulostroma sp.

Teratosphaeria quasicercospora (Crous & T.A. Cout.) Crous & U. Braun, comb. nov. MycoBank MB504494.

Basionym: Mycosphaerella quasicercospora Crous & T.A. Cout., Stud. Mycol. 55: 119. 2006.

Teratosphaeria readeriellophora (Crous & Mansilla) Crous & U. Braun, comb. nov. MycoBank MB504495.

Basionym: Mycosphaerella readeriellophora Crous & Mansilla, Stud. Mycol. 50: 207. 2004.

Anamorph: Readeriella readeriellophora Crous & Mansilla, Stud. Mycol. 50: 207. 2004. Fig. 18.

View larger version (152K): [in this window] [in a new window]   Fig. 18. A-E. Readeriella mirabilis. A. Conidium with conidial cirrus. B. Conidiogenous cells with percurrent proliferation. C. Macroconidia. D. Slightly pigmented, verruculose conidiogenous cell. E. Macro- and microconidia. F-I. Readeriella readeriellophora (type material). F. Colony on OA. G. Central stromatal tissue giving rise to conidiophores. H. Conidiogenous cells. I. Conidia. Scale bars = 10 µm.

Basionym: Mycosphaerella secundaria Crous & Alfenas, Stud. Mycol. 55: 122. 2006.

Teratosphaeria stramenticola (Crous & Alfenas) Crous & U. Braun, comb. nov. MycoBank MB504497.

Basionym: Mycosphaerella stramenticola Crous & Alfenas, Stud. Mycol. 55: 123. 2006.

Teratosphaeria suberosa (Crous, F.A. Ferreira, Alfenas & M.J. Wingf.) Crous & U. Braun, comb. nov. MycoBank MB504498.

Basionym: Mycosphaerella suberosa Crous, F.A. Ferreira, Alfenas & M.J. Wingf., Mycologia 85: 707. 1993.

Teratosphaeria suttonii (Crous & M.J. Wingf.) Crous & U. Braun, comb. nov. MycoBank MB504499.

Basionym: Mycosphaerella suttonii Crous & M.J. Wingf. (suttoniae), Canad. J. Bot. 75: 783. 1997.

Anamorph: Readeriella epicoccoides (Cooke & Massee) Crous & U. Braun, comb. nov. MycoBank MB504500.

Basionym: Cercospora epicoccoides Cooke & Massee apud Cooke, Grevillea 19: 91. 1891.

Phaeophleospora epicoccoides (Cooke & Massee) Crous, F.A. Ferreira & B. Sutton, S. African J. Bot. 63: 113. 1997.

Kirramyces epicoccoides (Cooke & Massee) J. Walker, B. Sutton & Pascoe, Mycol. Res. 96: 919. 1992.

= Hendersonia grandispora McAlp., Proc. Linn. Soc. New South Wales 28: 99. 1903.

= Phaeoseptoria eucalypti Hansf., Proc. Linn. Soc. New South Wales 82: 225. 1957.

= Phaeoseptoria luzonensis T. Kobayashi, Trans. Mycol. Soc. Japan 19: 377. 1978.

Synanamorph: Pseudocercospora sp.

Teratosphaeria toledana (Crous & Bills) Crous & U. Braun, comb. nov. MycoBank MB504501.

Basionym: Mycosphaerella toledana Crous & Bills, Stud. Mycol. 50: 208. 2004.

Anamorph: Readeriella toledana (Crous & Bills) Crous & U. Braun, comb. nov. MycoBank MB504502.

Basionym: Phaeophleospora toledana Crous & Bills, Stud. Mycol. 50: 208. 2004.

Key to treated anamorph genera of Teratosphaeria (Teratosphaeriaceae)

• 1. Hyphae submerged to superficial, disarticulating into arthroconidia......................................................................................................... 2
  
• 1. Hyphae not disarticulating into arthroconidia............................................................................................................................................ 3
  
• 2. Mature, brown hyphae disarticulating into thick-walled, spherical, smooth to verruculose 0(-2) transversely septate, brown conidia........................................................................................................................... Pseudotaeniolina (= Friedmanniomyces)
  
• 2. Hyphae superficial, brown to green-brown, smooth, disarticulating to form pale brown, cylindrical, 0-3-septate conidia with subtruncate ends, frequently with a Readeriella synanamorph...................................................................................................................... Cibiessia
  
• 3. Hyphal ends forming endoconidia; hyphae pale to medium brown, verruculose, end cells dividing into several brown, verruculose, thick-walled, ellipsoid to obovoid endoconidia................................................................................................................. Phaeothecoidea
  
• 3. Endoconidia absent.................................................................................................................................................................................... 4
  
• 4. Conidiogenous cells integrated in hyphae; well-developed conidiomata or long, solitary, macronematous, terminally penicillate conidiophores absent................................................................................................................................................................................ 5
  
• 4. Conidiomata well-developed or with long, solitary, terminally penicillate conidiophores........................................................................... 7
  
• 5. Conidia in chains, holoblastic, pseudocladosporium-like in morphology, but scars and hila not excessively thickened, nor refractive, producing chlamydospores in culture; species are mostly heat resistant................................................................................... Devriesia
  
• 5. Conidia solitary on indistict to well defined phialides on hyphae............................................................................................................... 6
  
• 6. Conidiogenous cells integrated in the distal ends of hyphae; conidia thick-walled, brown, smooth, 1-septate......................................................................................................................................................................... Capnobotryella
  
• 6. Conidiophores short and frequently reduced to conidiogenous cells that proliferate percurrently via wide necks, giving rise to hyaline, 0(-2)-septate, broadly ellipsoidal conidia................................................................................................................................................ Hortaea
  
• 7. Conidia brown, with hyaline basal appendages; conidiomata pycnidial, conidiogenous cells phialidic, but also percurrent, subhyaline....................................................................................................................................................................... Nothostrasseria
  
• 7. Conidia brown, but basal appendages lacking, amero- to scolecospores................................................................................................ 8
  
• 8. Conidiomata pycnidial to acervular........................................................................................................................................................... 9
  
• 8. Conidiomata not enclosed by host tissue, fasciculate to sporodochial or solitary, hyphomycetous........................................................ 10
  
• 9. Conidia solitary, dry, without mucilaginous sheath................................................................................................................... Readeriella
  
• 9. Conidia catenulate, with persistant mucilaginous sheath..................................................................................................... Staninwardia
  
• 10. Conidiophores usually solitary, rarely densely fasciculate to synnematous (in vivo), penicillate, with a branched, apical conidiogenous apparatus giving rise to ramoconidia and branched chains of secondary conidia; scars not to slightly thickened and darkened-refractive.................................................................................................................................................................................................... Penidiella
  
• 10. Conidiophores not penicillate, without a branched conidiogenous apparatus, in vivo fasciculate to sporodochial................................. 11
  
• 11. Biotrophic; fruiting composed of sporodochia and radiating layers of hyphae arising from the stromata, conidiophores arising from superficial sporodochia and radiating hyphae, conidiogenous cells unilocal, with conspicuous annellations, conidia solitary or in fragile disarticulating chains, aseptate or transversely 1-3-septate, usually with distinct frills, secession rhexolytic.............. Batcheloromyces
  
• 11. Biotrophic, leaf-inhabiting, with distinct, subepidermal to erumpent, well-developed sporodochia, or saxicolous, saprobic, sometimes causing opportunistic human infections; radiating layers of hyphae arising from sporodochia; conidiogenous cells without annellations; conidia in true simple or branched basipetal chains, transversely 1- to pluriseptate or with longitudinal and oblique septa (dictyosporous), occasionally distoseptate...................................................................................................................... Catenulostroma
  

To explain the arguments behind the selection and synonymies of some of these anamorphic genera, they are briefly discussed below:

Acidomyces Baker et al., Appl. Environ. Microbiol. 70: 6270. 2004. (nom. inval.)

Type species: Acidomyces richmondensis Baker et al., Appl. Environ. Microbiol. 70: 6270. 2004. (nom. inval.)

Notes: The genus presently clusters among isolates in the Teratosphaeria clade based on sequences deposited in GenBank. Acidomyces lacks a Latin description and holotype specimen, and is thus invalidly described. The genus, which was distinguished from other taxa based on its DNA phylogeny (Dothideomycetes), forms filamentous hyphae with disarticulating cells. It is unclear how it differs from Friedmanniomyces Onofri and Pseudotaeniolina J.L. Crane & Schokn.

Batcheloromyces Marasas, P.S. van Wyk & Knox-Dav., J. S. African Bot. 41: 41. 1975.

Type species: Batcheloromyces proteae Marasas, P.S. van Wyk & Knox-Dav., J. S. African Bot. 41: 43. 1975.

Description: Crous et al. (2004a; figs 4-26).

View larger version (90K): [in this window] [in a new window]   Fig. 4. Batcheloromyces leucadendri in vitro. A-B. Batcheloromyces state with synanamorph (arrows). C-D. Conidia occurring solitary or in short chains. Scale bar = 10 µm.

The finding that Stigmina s. str. [based on S. platani (Fuckel) Sacc., the type species] is a generic synonym of Pseudocercospora Speg. (Crous et al. 2006a), and that the type species of Trimmatostroma (T. salicis, Fig. 5) belongs to the Helotiales (Fig. 1), raises the question of where to place stigmina- and trimmatostroma-like anamorphs that reside in the Teratosphaeria clade. Although the stigmina-like species can be accommodated in Batcheloromyces (see Sutton & Pascoe 1989), a new genus is required for Teratosphaeria anamorphs that have a trimmatostroma-like morphology. The recognition of Batcheloromyces and the introduction of a new anamorph genus for trimmatostroma-like anamorphs of Teratosphaeria are also morphologically justified. Batcheloromyces is easily distinguishable from Stigmina s. str. by its special structure of the fruiting body, composed of sporodochia and radiating layers of hyphae arising from the sporodochia and the conidia often formed in delicate disarticulating chains. Trimmatostroma-like anamorphs of Teratosphaeria are morphologically also sufficently distinct from Trimmatostroma s. str. (see notes under Catenulostroma Crous & U. Braun) as well as Batcheloromyces (see key above).

View larger version (115K): [in this window] [in a new window]   Fig. 5. Trimmatostroma salicis. A. Sporodochia on twig. B-E. Chains of disarticulating conidia. Scale bars = 10 µm.

Basionym: Stigmina eucalypti Alcorn, Trans. Brit. Mycol. Soc. 60: 151. 1973.

Capnobotryella Sugiy., in Sugiyama, Pleomorphic Fungi: The Diversity and its Taxonomic Implications (Tokyo): 148. 1987.

Type species: Capnobotryella renispora Sugiy., in Sugiyama, Pleomorphic Fungi: The Diversity and its Taxonomic Implications (Tokyo): 148. 1987.

Description: Sugiyama & Amano (1987, figs 7.5-7.8).

Notes: The genus forms brown, septate, thick-walled hyphae, with ellipsoidal, 0-1-septate conidia forming directly on the hyphae, via minute phialides. Hambleton et al. (2003) also noted the occurrence of endoconidiation.

Catenulostroma Crous & U. Braun, gen. nov. MycoBank MB504474.

Etymology: Named after its catenulate conidia, and stromata giving rise to sporodochia.

Hyphomycetes. Differt a Trimmatostromate habitu phytoparasitico, maculis formantibus, conidiophoris saepe fasciculatis, per stoma emergentibus vel habitu saxiphilo-saprophytico, interdum sejunctis ex mycosibus humanis.

Habit plant pathogenic, leaf-spotting or saxicolous-saprobic, occasionally isolated from opportunistic human mycoses. Mycelium internal and external; hyphae dark brown, septate, branched. Conidiomata in vivo vary from acervuli to sporodochia or fascicles of conidiophores arising from well-developed or reduced, pseudoparenchymatal stromata. Setae and hyphopodia absent. Conidiophores arising from hyphae or stromata, solitary, fasciculate to sporodochial, in biotrophic, plant pathogenic species emerging through stomata, little differentiated, semimacronematous, branched or not, continuous to septate, brown, smooth to verruculose. Conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, holoblastic-thalloblastic, meristematic, unilocal, delimitation of conidium by a single septum with retrogressive delimitation of next conidium giving an unconnected chain of conidia, brown, smooth to verruculose, conidiogenous scars (conidiogenous loci) inconspicuous, truncate, neither thickened nor darkened. Conidia solitary or usually forming simple to branched basipetal chains of transversely to muriformly eu- or distoseptate, 1- to multiseptate, brown, smooth, verruculose to verrucose conidia, conidial secession schizolytic.

Type species: Catenulostroma protearum (Crous & M.E. Palm) Crous & U. Braun, comb. nov.

Description: Crous & Palm (1999), Crous et al. (2004a; figs 364-365).

Notes: Catenulostroma contains several plant pathogenic species previously placed in Trimmatostroma, a morphologically similar but, based on its type species, phylogenetically distinct genus belonging to Helotiales (Fig. 1). Trimmatostroma s. str. is well-distinguished from most Catenulostroma species by being saprobic, living on twigs and branches of woody plants, or occasionally isolated from leaf litter, i.e., they are not associated with leaf spots. The conidiomata of Trimmatostroma species are subepidermal, acervular-sporodochial with a well defined wall of textura angularis, little differentiated, micronematous conidiophores giving rise to long chains of conidia that disarticulate at the surface to form a grey-black to brown powdery mass. The generic affinity of other species assigned to Trimmatostroma, e.g. those having a lichenicolous habit, is unresolved.

Trimmatostroma abietis Butin & Pehl (Butin et al. 1996) clusters together with the plant pathogenic Catenulostroma species, but differs from these species in having a more complex ecology. Trimmatostroma abietis is usually foliicolous on living or necrotic conifer needles on which characteristic acervuli to sporodochia with densely arranged, fasciculate fertile hyphae are formed, comparable to the fasciculate conidiomata of the plant pathogenic species of Catenulostroma (Butin et al. 1996: 205, fig. 1). Although not discussed by Butin et al. (1996), T. abietis needs to be compared to T. abietina Doherty, which was orginally described from Abies balsamea needles collected in Guelph, Canada (Doherty 1900). Morphologically the two species appear to be synonymous, except for reference to muriformly septate conidia, which is a feature not seen in vivo in the type of T. abietis. Furthermore, as this is clearly a species complex, this matter can only be resolved once fresh Canadian material has been collected to serve as epitype for T. abietina.

Isolates from stone, agreeing with T. abietis in cultural, morphological and physiological characteristics, have frequently been found (Wollenzien et al. 1995, Butin et al. 1996, Gorbushina et al. 1996, Kogej et al. 2006, Krumbein et al. 1996). Furthermore, isolates from humans (ex skin lesions and ex chronic osteomycelitis of human patients) and Ilex leaves are known (Butin et al. 1996). De Hoog et al. (1999) included strains of T. abietis from stone, man and Ilex leaves in molecular sequence analyses and demonstrated their genetical identity based on 5.8S rDNA and ITS2 data, but strains from conifer needles were not included. Furthermore, we consider T. abietis, as presently defined, to represent a species complex, with Dutch isolates from Pinus again appearing distinct from German Abies isolates, suggesting that different conifer genera could harbour different Catenulostroma species. Isolates from stone form stromatic, durable microcolonies, which are able to grow under extreme xerophilic environmental conditions. Cultural growth resembles that of other meristematic black yeasts (Butin et al. 1996, Kogej et al. 2006). Another fungus isolated from stone in Germany is in vitro morphologically close to C. abietis, but differs in forming conidia with oblique septa. Furthermore, a human pathogenic isolate from Africa clusters together with other Catenulostroma species. The habit and origin of this human pathogenic fungus in nature and its potential morphology on "natural" substrates, which typically deviates strongly from the growth in vitro, are still unknown. However, C. abietis, usually growing as a foliicolous and saxicolous fungus, has already shown the potential ability of Catenulostroma species to cause opportunistic human infections.

Key to Catenulostroma species

• 1. Conidia formed in basipetal chains, smooth, 4-celled, consisting of two basal cells with truncate lateral sides, each giving rise to a secondary globose apical cell, that can extend and develop additional septa, appearing as two lateral arms................. C. excentricum
  
• 1. Conidia variable in shape, but without two basal cells giving rise to two lateral arms............................................................................... 2
  
• 2. Conidia smooth or almost so, at most very faintly rough-walled; usually foliicolous on conifer needles or saxicolous, forming stromatic, xerophilic durable microcolonies on stone, occasionally causing opportunistic human infections............................................................ 3
  
• 2. Conidia distinctly verruculose to verrucose; plant pathogenic, forming leaf spots.................................................................................... 5
  
• 3. Conidia (8-)20-35(-60) x 4-5(-7) µm, 1-10-septate................................................................................... C. chromoblastomycosum
  
• 3. Conidia much shorter, 8-20 µm long, 0-5-septate................................................................................................................................... 4
  
• 4. Conidia 0-5 times transversely septate, mostly two-celled; usually foliicolous on conifer needles or saxicolous...................... C. abietis
  
• 4. Conidia 2-4 times transversely septate and often with 1-2 oblique septa; isolated from stone....................................... C. germanicum
  
• 5. Conidia rather broad, usually wider than 10 µm....................................................................................................................................... 6
  
• 5. Conidia narrower, width below 10 µm....................................................................................................................................................... 7
  
• 6. Conidia distoseptate, rather long, (12-)25-35(-45) x (7-)10-15(-25) µm; conidiomata large, up to 250 µm diam, on Protea anceps.............................................................................................................................................................................................. C. protearum
  
• 6. Conidia euseptate, shorter, (9-)16-20(-36) x (10-)14-18(-27) µm; sporodochia 90-100 x 40-80 µm; on Protea grandiceps................................................................................................................................................................................................ C. elginense
  
• 7. Conidia 1- to multiseptate, (10-)15-17(-23) x (5-)6.5-7(-9) µm; on various Proteaceae................................................. C. macowanii
  
• 7. Conidia in vivo predominantly 1-septate, (8-)13-15(-21) x (3.5-)5.5-6(-8) µm; on Protea cynaroides........................................................................................................................................ C. microsporum (Teratosphaeria microspora)
  

Catenulostroma abietis (Butin & Pehl) Crous & U. Braun, comb. nov. MycoBank MB504504.

View larger version (115K): [in this window] [in a new window]   Fig. 6. Catenulostroma chromoblastomycosum (type material). A. Sporodochium on pine needle in vitro. B-H. Chains of disarticulating conidia. Scale bars: A = 350, B, E, G, H = 10 µm.

Notes: Catenulostroma abietis needs to be compared to Trimmatostroma abietina Doherty (Abies balsamea needles Canada), which is either an older name for this species, or a closely related taxon. Presently T. abietina is not known from culture, and needs to be recollected.

Catenulostroma chromoblastomycosum Crous & U. Braun, sp. nov. MycoBank MB504505. Fig. 6.

Etymology: Named after the disease symptoms observed due to opportunistic human infection.

Differt a C. abieti et C. germanico conidiis longioribus, (8-)20-35(-60) x 4-5(-7) µm, 1-10-septatis.

Description based on cultures sporulating on WA supplemented with sterile pine needles. Mycelium consisting of branched, septate, smooth to finely verruculose, medium to dark brown, thick-walled, 3-4 µm wide hyphae. Conidiomata brown, superficial, sporodochial, up to 350 µm diam. Conidiophores reduced to inconspicuous conidiogenous loci on hyphae, 2-4 µm wide, neither darkened nor thickened or refractive. Conidia occurring in branched chains, that tend to remain attached to each other, subcylindrical with subtruncate ends, straight to slightly curved, (8-)20-35(-60) x 4-5(-7) µm, 1-10-septate, medium brown, smooth to finely verruculose.

View larger version (158K): [in this window] [in a new window]   Fig. 7. Catenulostroma germanicum (type material). A-D. Chains of disarticulating conidia in vitro. Scale bars = 10 µm.

Specimen examined: Zaire, Pawa, isolated from man with chromoblastomycosis, Mar. 1997, V. de Brouwere, holotype CBS H-19935, culture ex-type CBS 597.97.

Notes: Catenulostroma chromoblastomycosum was originally identified as an isolate of Stenella araguata Syd. The latter fungus is morphologically distinct, however, having much shorter and narrower conidia, formed in acropetal chains, as well as quite different conidiogenous loci and conidial hila which are small, thickened and darkened.

Catenulostroma elginense (Joanne E. Taylor & Crous) Crous & U. Braun, comb. nov. MycoBank MB504506.

Basionym: Trimmatostroma elginense Joanne E. Taylor & Crous, Mycol. Res.104: 633. 2000.

Catenulostroma excentricum, see Teratosphaeria excentrica.

Catenulostroma germanicum Crous & U. Braun, sp. nov. MycoBank MB504507. Fig. 7.

Etymology: Named after the geographic location of its type strain in Germany.

Differt a C. abieti conidiis 1-2 oblique septatis.

Mycelium consisting of branched, septate, smooth, pale to medium brown, 2-4 µm wide hyphae, giving rise to conidial chains. Conidiophores integrated, subcylindrical, branched or not, septate, little differentiated, micronematous, 3-5 µm wide, 3- to multiseptate, medium brown, thick-walled; conidiogenous cells integrated, terminal, inconspicuous, unilocal, conidiogenous loci inconspicuous. Conidia in simple or branched basipetal chains, subcylindrical, straight to flexuous, (8-)10-15(-20) x 4-5(-6) µm, 2-4 transversely septate or with 1-2 oblique septa, medium to dark brown, thick-walled, smooth.

Cultural characteristics: Colonies on OA erumpent, spreading, with even, smooth margins and sparse to moderate aerial mycelium; olivaceous-grey, with iron-grey margins (surface). Colonies reaching 12 mm diam after 1 mo at 25 °C in the dark; colonies fertile.

Specimen examined: Germany (former West-Germany), isolated from stone, Oct. 1988, J. Kuroczkin, holotype CBS H-19936, culture ex-type CBS 539.88.

Notes: Catenulostroma germanicum was originally deposited as Taeniolina scripta (P. Karst.) P.M. Kirk. It is clearly distinct, however, as the latter fungus forms intricate, branched, brown conidia (Kirk 1981), unlike those of C. germanicum. Phylogenetically C. germanicum forms part of the C. abietis species complex.

Catenulostroma macowanii (Sacc.) Crous & U. Braun, comb. nov. MycoBank MB504508.

Basionym: Coniothecium macowanii Sacc., Syll. Fung. 4: 512. 1886.

Coniothecium punctiforme G. Winter, Hedwigia 24: 33. 1885, non C. punctiforme Corda, Icones Fungorum (Prague) 1: 2. 1837.

Trimmatostroma macowanii (Sacc.) M.B. Ellis, More Dematiacous Hyphomycetes: 29. 1976.

Catenulostroma microsporum, see Teratosphaeria microspora.

Catenulostroma protearum (Crous & M.E. Palm) Crous & U. Braun, comb. nov. MycoBank MB504509.

Basionym: Trimmatostroma protearum Crous & M.E. Palm, Mycol. Res. 103: 1303. 1999.

Cibiessia Crous, Fungal Diversity 26: 151. 2007.

Type species: Cibiessia dimorphospora Crous & C. Mohammed, Fungal Diversity 26: 151. 2007.

Description: Crous et al. (2007b; figs 3-5).

Notes: The genus Cibiessia was introduced to accommodate species with chains of disarticulating conidia (arthroconidia). Some species have been shown to have a Readeriella synanamorph.

Devriesia Seifert & N.L. Nick., Can. J. Bot. 82: 919. 2004.

Type species: Devriesia staurophora (W.B. Kendr.) Seifert & N.L. Nick., Canad. J. Bot. 82: 919. 2004.

Description: Seifert et al. (2004; figs 2-42).

Notes: The genus is characterised by producing chains of pale brown, subcylindrical to fusiform, 0-1-septate conidia with somewhat thickened, darkened hila, forming chlamydospores in culture, and being heat resistant. Morphologically they resemble taxa placed in Pseudocladosporium U. Braun (= Fusicladium Bonord.; Venturiaceae), though phylogenetically Devriesia is not allied to this family.

Hortaea Nishim. & Miyaji, Jap. J. Med. Mycol. 26: 145. 1984.

Type species: Hortaea werneckii (Horta) Nishim. & Miyaji, Jap. J. Med. Mycol. 26: 145. 1984.

Description: de Hoog et al. (2000, illust. p. 721).

Notes: The genus forms brown, septate, thick-walled hyphae, with ellipsoidal, 0-1-septate (becoming muriformly septate), hyaline to pale brown conidia forming directly on the hyphae, via phialides with percurrent proliferation. Isolates of H. werneckii are restricted to tropical or subtropical areas, where they occur as halophilic saprobes, frequently being associated with tinea nigra of humans (de Hoog et al. 2000). The generic distinction with Capnobotryella is less clear, except that the latter tends to have darker, thick-walled conidia, and reduced, less prominent phialides.

Penidiella Crous & U. Braun, gen. nov. MycoBank MB504463.

Etymology: Named after its penicillate conidiophores.

Differt a Periconiellae conidiophoris apice penicillato ex cellulis conidiogenis et ramoconidiis compositis, cellulis conidiogenis saepe 1-3(-4) locis conidiogenis, terminalibus vel subterminalibus, subdenticulatis, non vel subincrassatis, non vel leviter fuscatis-refractivis, ramoconidiis praesentibus, saepe numerosis, conidiis ramicatenatis.

Mycelium consisting of branched, septate, smooth to verruculose, subhyaline to pale brown hyphae. Conidiophores macronematous, occasionally also with some micronematous conidiophores; macronematous conidiophores arising from superficial mycelium or stromata, solitary, fasciculate or in synnemata, erect, brown, thin- to thick-walled, smooth to finely verruculose; terminally penicillate, branched terminal part consisting of a conidiogenous apparatus composed of a series of conidiogenous cells and/or ramoconidia. Conidiogenous cells integrated, terminal, intercalary or pleurogenous, unbranched, pale to medium brown, smooth to finely verruculose, tapering to a flattened or rounded apical region or tips slightly inflated, polyblastic, sympodial, giving rise to a single or several sets of ramoconidia on different levels; with relatively few conidiogenous loci, 1-3(-4), terminal or subterminal, subdenticulate, denticle-like loci usually conical, terminally truncate, usually unthickened or at most very slightly thickened, not to slightly darkened or somewhat refractive. Conidia in branched acropetal chains. Ramoconidia 0-1-septate, pale to medium brown, smooth to verruculose, thin-walled, ellipsoidal, obovoid, fusiform, subcylindrical to obclavate; conidia subcylindrical, fusiform to ellipsoid-ovoid, 0-1-septate, pale olivaceous to brown, smooth to verruculose, thin-walled, catenate; hila truncate, unthickened or almost so, barely to somewhat darkened-refractive.

Type species: Penidiella columbiana Crous & U. Braun, sp. nov.

Notes: Three ramichloridium-like genera cluster within Capnodiales, namely Periconiella Sacc. [type: P. velutina (G. Winter) Sacc.], Ramichloridium Stahel ex de Hoog [type: R. apiculatum (J.H. Mill., Giddens & A.A. Foster) de Hoog] and Penidiella [type: P. columbiana Crous & U. Braun]. All three genera have brown, macronematous conidiophores with similar conidial scars. Within this complex, Ramichloridium is distinct in having a prominent rachis giving rise to solitary conidia. Periconiella and Penidiella are branched in the apical part of their conidiophores, and lack a rachis. In Periconiella conidia are solitary or formed in short, mostly simple chains, ramoconidia are lacking. The apical conidiogenous apparatus is composed of conidiogenous cells or branches with integrated, usually terminal conidiogenous cells, which are persistent. The conidiogenous cells are subcylindrical to somewhat clavate, usually not distinctly attenuated towards the tip, and have several, often numerous loci, aggregated or spread over the whole cell, terminal to usually lateral, flat, non-protuberant, not denticle-like, usually distinctly thickened and darkened, at least at the rim. In contrast, Penidiella has a quite distinct branching system, consisting of a single terminal conidiogenous cell giving rise to several ramoconidia that form secondary ramoconidia, etc., or the branched apparatus is composed of several terminal and sometimes lateral conidiogenous cells giving rise to sequences of ramoconidia (conidiogenous cells and ramoconidia are often barely distinguishable, with conidiogenous cells disarticulating, becoming ramoconidia). The branched apparatus may be loose to dense, metula-like. The conidiogenous cells have only few, usually 1-3 (-4), terminal or subterminal subdenticulate loci, and ramoconidia are prominent and numerous, giving rise to branched chains of secondary conidia with flat-tipped hila. Some species of Penidiella with compact, metula-like branched apices are morphologically close to Metulocladosporiella Crous, Schroers, J.Z. Groenew., U. Braun & K. Schub. (Crous et al. 2006d). This genus encompasses two species of banana diseases belonging to Herpotrichiellaceae (Chaetothyriales), characterised by having conidiophore bases with rhizoid hyphal appendages and abundant micronematous conidiophores. Penidiella species with less pronounced penicillate apices, e.g. P. strumelloidea (Milko & Dunaev) Crous & U. Braun, are comparable with species of the genus Pleurotheciopsis B. Sutton (see Ellis 1976). The latter genus is distinct in having unbranched, often percurrently proliferating conidiophores, lacking ramoconidia and colourless conidia formed in simple chains.

Cladosporium helicosporum R.F. Castañeda & W.B. Kendr. (Castañeda et al. 1997) is another penidiella-like fungus with terminally branched conidiophores, subdenticulate conidiogenous loci and conidia in long acropetal chains, but its affinity to Penidiella has still to be proven.

Key to Penidiella species

• 1. Conidiophores in vivo in well-developed, dense fascicles and distinct synnemata arising from a basal stroma; on fallen leaves of Ficus sp., Cuba............................................................................................................................................................................... P. cubensis
  
• 1. Conidiophores solitary, at most loosely aggregated.................................................................................................................................. 2
  
• 2. Conidiophores with a terminal conidiogenous cell, often somewhat swollen, giving rise to several ramoconidia (on one level) that form chains of straight to distinctly curved conidia; isolated from leaf of Carex sp., Russia.................................................... P. strumelloidea
  
• 2. Penicillate apex of the conidiophores composed of a system of true branchlets, conidiogenous cells and ramoconidia or at least a sequence of ramoconidia on several levels; conidia usually straight......................................................................................................................... 3
  
• 3. Mycelium verruculose; long filiform conidiophores ending with a subdenticulate cell giving rise to sets of penicillate conidiogenous cells or ramoconidia which are barely distinguishable and turn into each other; ramoconidia and conidia consistently narrow, (1.5-)2(-2.5) µm wide, and aseptate, ramoconidia sometimes heterochromous; on living leaves of Nectandra coriacea, Cuba................... P. nectandrae
  
• 3. Mycelium more or less smooth; penicillate apex at least partly with true branchlets; conidia wider, 2-5 µm, at least partly septate, uniformly pigmented................................................................................................................................................................................................. 4
  
• 4. Hyphae, conidiophores and conidia frequently distinctly constricted at the septa; penicillate apex of the conidiophores sparingly developed, branchlets more or less divergent; isolated from leaf litter of Smilax sp., Cuba.................................................................. P. rigidophora
  
• 4. Hyphae and conidia without distinct constrictions at the septa; penicillate apex of the conidiophores usually well-developed, with abundant branchings................................................................................................................................................................................................. 5
  
• 5. Conidiophores short, up to 120 x 3-4 µm, frequently with intercalary conidiogenous cell, swollen at the conidiogenous portion just below the upper septum which render the conidiophores subnodulose to distinctly nodulose, apex ± loosely penicillate; conidia (4-)5-7(-8) µm long; occasionally with micronematous conidiophores; isolated from man with tinea nigra, Venezuela......................... P. venezuelensis
  
• 5. Conidiophores much longer, up to 800 µm, 7-9 µm wide at the base, not distinctly nodulose, penicillate apex loose to often more compact, tight, metula-like; conidia longer, 7-25 x 2-5 µm; micronematous conidiophores lacking; isolated from dead leaf of Paepalanthus columbianus, Colombia....................................................................................................................................................... P. columbiana
  

Penidiella columbiana Crous & U. Braun, sp. nov. MycoBank MB504510. Figs 8-9.

View larger version (126K): [in this window] [in a new window]   Fig. 8. Penidiella columbiana (type material). A. Conidiophores on pine needle in vitro. B-H. Conidiophores with chains of disarticulating conidia. Scale bars: A = 450, B-C = 10 µm.

View larger version (16K): [in this window] [in a new window]   Fig. 9. Penidiella columbiana (type material). A. Conidiophores. B. Ramoconidia. C. Secondary conidia. Scale bar = 10 µm. U. Braun del.

Mycelium ex hyphis ramosis, septatis, levibus, pallide brunneis, 2-3 µm latis compositum. Conidiophora ex hyphis superficialibus oriunda, penicillata, erecta, brunnea, crassitunicata, minute verruculosa, ad 800 µm longa, ad basim 7-9 µm lata, ad apicem pluriramosa, ex ramibus diversibus et cellulis conidiogenis composita, ramibus primariis (-2) subcylindraceis, 1-7-septatis, 50-120 x 4-6 µm; ramibus secundariis (-2) subcylindraceis, 1-5-septatis, 40-60 x 4-6 µm; ramibus tertiariis et subsequentibus 1-4-septatis, 10-30 x 3-5 µm. Cellulae conidiogenae terminales vel laterales, non ramosae, 5-15 x 3-5 µm, modice brunneae, minute verruculosae, apicem versus attenuatae, truncatae vel rotundatae, polyblasticae, sympodiales, cicatrices conidiales incrassatae, sed leviter fuscatae et non refractivae. Ramoconidia 0-1-septata, modice brunnea, levia, ellipsoidea, obclavata vel obovoidea, cum 1-3 hilis terminalibus, 10-20 x 3-5 µm; conidia subcylindrica vel ellipsoidea, 0-1-septata, pallide brunnea, catenata (-10), hila truncata, non incrassata, vix vel leviter fuscata.

Mycelium consisting of branched, septate, smooth, pale brown, 2-3 µm wide hyphae. Conidiophores arising from superficial mycelium, terminally penicillate, erect, brown, wall up to 1 µm wide, almost smooth to finely verruculose, up to 800 µm tall, 7-9 µm wide at the base; conidiogenous region consisting of a series of branches composed of true branchlets, conidiogenous cells and ramoconidia, branched portion usually rather compact, even metula-like, but also looser, with divergent branches; primary branches (-2), subcylindrical, 1-7-septate, 50-120 x 4-6 µm; secondary branches (-2), subcylindrical, 1-5-septate, 40-60 x 4-6 µm; tertiary and additional branches 1-4-septate, 10-30 x 3-5 µm. Conidiogenous cells terminal, intercalary or lateral, unbranched, 5-15 x 3-5 µm, medium brown, finely verruculose, tapering to a flattened or rounded (frequently swollen) apical region, scars thickened, but only somewhat darkened, not refractive. Ramoconidia 0-1-septate, medium brown, smooth, wall 1 µm wide, ellipsoidal to obclavate or obovoid, with 1-3 apical hila, 10-25 x 3-5 µm, ramoconidia with broadly truncate base, not or barely attenuated, up to 4 µm wide, or at least somewhat attenuated at the base, hila 1.5-3 µm wide. Conidia subcylindrical to ellipsoid, 0(-1)-septate, pale brown, in chains of up to 10, 7-15 x 2-3 µm, hila truncate, unthickened, barely to somewhat darkened, 1-2 µm wide.

Cultural characteristics: Colonies on PDA erumpent, spreading, with moderate aerial mycelium and smooth, even, submerged margins; olivaceous-grey in central part, iron-grey in outer region (surface); colonies fertile.

Specimen examined: Colombia, Páramo de Guasca, 3400 m alt., isolated from dead leaf of Paepalanthus columbianus (Eriocaulaceae), Aug. 1980, W. Gams, holotype CBS H-19937, culture ex-type CBS 486.80.

Notes: This isolate was originally identified as belonging to the Stenella araguata species complex. The latter name has been somewhat confused in the literature, and has been incorrectly applied to isolates associated with opportunistic human infections (de Hoog et al. 2000). The "araguata" species complex is treated elsewhere in the volume (see Crous et al. 2007a - this volume).

Penidiella cubensis (R.F. Castañeda) U. Braun, Crous & R.F. Castañeda, comb. nov. MycoBank MB504511. Fig. 10.

View larger version (26K): [in this window] [in a new window]   Fig. 10. Penidiella cubensis (type material). A. Swollen stromatic base of synnema. B. Conidiophores. C. Ramoconidia. D. Secondary conidia. Scale bar = 10 µm. U. Braun del.

In vivo: Colonies on fallen leaves, amphigenous, effuse, pilose, brown. Mycelium usually external, superficial, but also internal, composed of branched, septate, brown, thin-walled, smooth to rough-walled hyphae, 2-3 µm wide. Stromata present, 40-80 µm diam, brown, immersed. Conidiophores densely fasciculate or in distinct synnemata, arising from stromata, erect, synnemata up to about 1000 µm long and (10-)20-40(-50) µm wide, individual threads filiform, pluriseptate throughout, brown, thin-walled ( 0.5 µm), smooth or almost so to distinctly verruculose, apically penicillate. Conidiogenous cells integrated, terminal and intercalary, 10-30 µm long, subcylindrical, terminal conidiogenous cells often slightly enlarged at the tip, with (1-)2-3(-4) terminal or subterminal subdenticulate conidiogenous loci, short conically truncate, 1-2 µm diam, unthickened or almost so, but often slightly refractive or darkened-refractive, intercalary conidiogenous cells usually with a single lateral locus just below the upper septum, conidiogenous cells giving rise to a single set of primary ramoconidia, or a sequence of ramoconidia at different levels. Ramoconidia cylindrical to ellipsoid-fusoid, 8-18(-25) x 2-3 µm, aseptate, pale olivaceous, olivaceous-brown to brown, thin-walled, smooth or almost so to faintly verruculose, ramoconidia with broadly truncate base, barely narrowed, or ramoconidia more or less attenuated at the base, hila 1-2 µm wide, unthickened or almost so, but often slightly refractive or darkened-refractive. Conidia in long acropetal chains, narrowly ellipsoid-ovoid, fusiform, 5-12(-15) x (1-)1.5-3 µm, aseptate, pale olivaceous to brownish, thin-walled, smooth to faintly rough-walled, ends attenuated, hila 1-1.5 µm wide, unthickened, not darkened, at most somewhat refractive.

Specimen examined: Cuba, Guantánamo, Maisí, on fallen leaves of Ficus sp., 24 Apr. 1986, M. Camino, holotype INIFAT C86/134 (HAL 2019 F, ex holotype).

Notes: Cladosporium cubense was not available in culture and molecular sequence data are not available, but type material could be re-examined and revealed that this species is quite distinct from Cladosporium s. str., but agreeing with the concept of the genus Penidiella. Penidiella cubensis differs from all other species of this genus in having densely fasciculate conidiophores to synnematous conidiomata, arising from stromata.

Penidiella nectandrae Crous, U. Braun & R.F. Castañeda, nom. nov. MycoBank MB504512. Fig. 11.

View larger version (14K): [in this window] [in a new window]   Fig. 11. Penidiella nectandrae (type material). A. Conidiophores. B. Ramoconidia. C. Secondary conidia. Scale bar = 10 µm. U. Braun del.

In vivo: Colonies amphigenous, brown. Mycelium internal and external, superficial, composed of sparingly branched hyphae, septate, 1-3 µm wide, pale olivaceous-brown or brown, thin-walled ( 0.5 µm), smooth or almost so to distinctly verruculose, fertile cells giving rise to conidiophores somewhat swollen at the branching point, up to 5 µm diam, and somewhat darker. Stromata lacking. Conidiophores erect, straight, filiform, up to 350 µm long, 2.5-4 µm wide, pluriseptate throughout, brown, darker below and paler above, thin-walled, smooth, apex penicillate, terminal cell of the conidiophore with 2-4 short denticle-like loci giving rise to sets of conidiogenous cells or ramoconidia that then form a sequence of new sets of ramoconidia on different levels, i.e., the loose to dense, metula-like branching system is composed of conidiogenous cells and ramoconidia which are often barely distinguishable and turn into each other; conidiogenous loci terminal or subterminal, usually 1-3(-4), subdenticulate, 1-2 µm diam, conical, apically truncate, unthickened or almost so, not to somewhat darkened-refractive. Ramoconidia with truncate base, barely attenuated, or ramoconidia distinctly attenuated at the truncate base, up to 20 x 2 µm, aseptate, at the apex with 2-3(-4) subdenticulate hila, subcylindrical, very pale olivaceous, olivaceous-brown to brown, sometimes with different shades of brown (heterochromatous), thin-walled ( 0.5 µm), smooth to faintly verruculose. Conidia in long acropetal chains, narrowly ellipsoid-ovoid, fusiform to cylindrical, 5-16 x (1.5-)2(-2.5) µm, aseptate, very pale olivaceous, olivaceous-brown to brown, thin-walled, smooth to very faintly rough-walled, primary conidia with rounded apex and trunacte base, somewhat attenuated, secondary conidia truncate at both ends, hila 1-1.5 µm diam, unthickened or almost so, at most slightly darkened-refractive.

Cultural characteristics: Colonies on PDA slimy, smooth, spreading; aerial mycelium absent, margins smooth, irregular; surface black with patches of cream. Colonies reaching 20 mm diam after 1 mo at 25 °C in the dark; colonies sterile on PDA, SNA and OA.

Specimen examined: Cuba, Matanzas, San Miguel de los Baños, isolated from living leaves of Nectandra coriacea (Lauraceae), 24 Jan. 1987, R.F. Castañeda and G. Arnold, holotype INIFAT C87/45, culture ex-type CBS 734.87, and HAL 2018 F (ex-holotype).

Notes: Although the ex-type strain of Cladosporium ferrugineum is sterile, its LSU DNA phylogeny reveals it to be unrelated to Cladosporium s. str. (see Fig. 1 in Crous et al. 2007a - this volume). Based on a re-examination of the type material it could clearly be shown that the morphology of this species fully agrees with the concept of the new genus Penidiella, which is supported by its phylogenetic position within Capnodiales.

Penidiella rigidophora Crous, R.F. Castañeda & U. Braun, sp. nov. MycoBank MB504513. Figs 12-13.

View larger version (131K): [in this window] [in a new window]   Fig. 12. Penidiella rigidophora (type material). A-F. Micronematous conidiophores giving rise to chains of conidia. G-H. Macronematous conidiophores (note base in G, and apex in H). I. Conidia. Scale bars = 10 µm.

View larger version (17K): [in this window] [in a new window]   Fig. 13. Penidiella rigidophora (type material). A. Hyphae. B. Conidiophores. C. Ramoconidia. D. Secondary conidia. Scale bar = 10 µm. U. Braun del.

Cladosporium rigidophorum R.F. Castañeda, nom. nud. (herbarium name).

Differt a specibus Penidiellae conidiophoris dimorphosis, hyphis et conidiis ad septa saepe distincte constrictis.

Mycelium consisting of strongly branched, septate, smooth or almost so, pale olivaceous to medium brown, guttulate, commonly constricted at septa, 2-6 µm wide hyphae, swollen cells up to 8 µm wide, wall up to 1(-1.5) µm wide. Conidiophores dimorphic. Macronematous conidiophores separate, erect, subcylindrical, predominantly straight to slightly curved, terminally loosely penicillate, up to 120 µm long, and 4-5 µm wide at the base, which is neither lobed nor swollen, and lacks rhizoids, up to 10-septate, medium to dark brown, wall up to 1(-1.5) µm wide. Micronematous conidiophores erect, subcylindrical, up to 40 µm tall, 3-4 µm wide, 1-3-septate, pale to medium brown (concolorous with hyphae). Conidiogenous cells predominantly terminal, rarely intercalary, medium brown, smooth, subcylindrical, but frequently swollen at apex, 10-20 x 5-6 µm, loci (predominantly single in micronematous conidiophores, but up to 4 in macronematous conidiophores) flat-tipped, sub-denticulate or not, 1-1.5 µm wide, barely to slightly darkened and thickened-refractive. Conidia in branched chains, medium brown, verruculose, (appearing like small spines under light microscope), ellipsoid to cylindrical-oblong, up to 1(-1.5) µm wide, frequently constricted at septa, which turn dark with age; ramoconidia (10-)13-17(-25) x 3-4(-5) µm, 1(-3)-septate; secondary conidia (7-)8-10(-12) x 3-4(-5); hila unthickened to very slightly thickened and darkened, not refractive, (0.5-)1(-1.5) µm.

Cultural characteristics: Colonies on PDA erumpent, spreading, with lobate margins and moderate aerial mycelium; iron-grey (surface), with a greenish black margin; reverse greenish black. Colonies reaching 20 mm diam after 1 mo at 25 °C in the dark; colonies fertile.

Specimen examined: Cuba, isolated from leaf litter of Smilax sp. (Smilacaceae), 6. Nov. 1994, R.F. Castañeda, holotype CBS H-19938, culture ex-type CBS 314.95.

Notes: Cladosporium rigidophorum is a herbarium name, which was never validly published. The ex-type strain, however, represents a new species of Penidiella, for which a valid name with Latin diagnosis is herewith provided. This species is easily distinguishable from all other taxa of Penidiella by forming distinct constrictions at hyphal and conidial septa as well as micronematous conidiophores (except for P. venezuelensis in which a few micronematous conidiophores have been observed). It is also phylogenetically distinct from the other taxa of Penidiella (see Fig. 1 in Crous et al. 2007a - this volume).

Penidiella strumelloidea (Milko & Dunaev) Crous & U. Braun, comb. nov. MycoBank MB504514. Figs 14-15.

View larger version (144K): [in this window] [in a new window]   Fig. 14. Penidiella strumelloidea (type material). A-B. Micronematous conidiophores. C-D. Macronematous conidiophores. E-G. Conidia. Scale bars = 10 µm.

View larger version (11K): [in this window] [in a new window]   Fig. 15. Penidiella strumelloidea (type material). A. Hyphae. B. Conidiophores. C. Ramoconidia. D. Secondary conidia. Scale bars = 10 µm. U. Braun del.

Basionym: Cladosporium strumelloideum Milko & Dunaev, Novosti Sist. Nizsh. Rast. 23: 134. 1986.

Mycelium consisting of branched, septate, smooth, hyaline to pale olivaceous, 1-4 µm wide hyphae, sometimes constricted at somewhat darker septa. Conidiophores solitary, erect, arising from superficial mycelium, micronematous, i.e., reduced to conidiogenous cells, or macronematous, subcylindrical, straight to slightly curved, subcylindrical throughout or often somewhat attenuated towards the apex, 12-80 x (2-)2.5-4 µm, 0-6-septate, medium brown, smooth, wall 0.75 µm, penicillate apex formed by a terminal conidiogenous cell giving rise to a single set of ramoconidia. Conidiogenous cells terminal, integrated, subcylindrical, straight, 8-12 x 1.5-2(-2.5) µm, pale brown, thin-walled, smooth, apex obtusely rounded to somewhat clavate; loci terminal, occasionally subterminal or lateral, unthickened or almost so to slightly thickened and darkened, not refractive, 1-1.5 µm wide. Conidia in branched chains; ramoconidia subcylindrical, with 1-3 terminal loci, olivaceous-brown, smooth; secondary conidia ellipsoidal, with one side frequently straight and the other convex, straight to slightly curved, (8-)10-12(-20) x 2(-3) µm, subhyaline to olivaceous-brown, smooth, thin-walled; hila unthickened or almost so to somewhat thickened and darkened, not refractive, 1 µm wide.

Cultural characteristics: Colonies on PDA erumpent, spreading, with abundant, dense to woolly aerial mycelium, and uneven, feathery margins; surface pale olivaceous grey, reverse iron-grey. Colonies reaching 25 mm diam after 1 mo at 25 °C in the dark; colonies fertile.

Specimen examined: Russia, Yaroslavl Region, Rybinsk Reservoir, mouth of Sutka River, isolated from leaf of Carex sp. (Cyperaceae), from stagnant water, S. Ozerskaya, holotype BKMF-2534, culture ex-type CBS 114484.

Notes: Penidiella strumelloidea resembles other species of Penidiella by having penicillate conidiophores with a conidiogenous apparatus giving rise to branched conidial chains. It differs, however, from all other species of this genus in having a rather simple penicillate apex composed of a single terminal conidiogenous cell giving rise to one set of ramoconidia which form frequently somewhat curved conidia. It is also phylogenetically distinct from the other taxa of Penidiella (see Fig. 1 in Crous et al. 2007a - this volume).

Penidiella venezuelensis Crous & U. Braun, sp. nov. MycoBank MB504515. Figs 16-17.

View larger version (136K): [in this window] [in a new window]   Fig. 16. Penidiella venezuelensis (type material). A. Microconidiophore. B. Apical part of macroconidiophore. C-F. Chains of conidia. Scale bars = 10 µm.

View larger version (17K): [in this window] [in a new window]   Fig. 17. Penidiella venezuelensis (type material). A. Hypha. B. Micronematous conidiophores. C. Macronematous conidiophores. D-E. Ramoconidia. F. Secondary conidia. Scale bar = 10 µm. U. Braun del.

Differt a P. columbiana conidiophoris bevioribus et angustioribus, ad 20 x 3-4 µm, subnodulosis, apice plus minusve laxe penicillatis et conidiis brevioribus, (4-)5-7(-8) µm longis.

Mycelium consisting of branched, septate, smooth to faintly rough-walled, thin-walled, subhyaline, pale olivaceous to medium brown, (1.5-)2-3 µm wide hyphae. Conidiophores solitary, erect, macronematous, subcylindrical, straight to flexuous to once geniculate, up to 120 µm long, 3-4 µm wide, 1-12-septate, pale to medium olivaceous-brown or brown, thin-walled (up to about 1 µm), terminally penicillate, branched portion composed of true branchlets and/or a single set or several sets of ramoconidia, branchlets up to 50 µm long; occasionally with a few additional micronematous conidiophores, about 10-15 x 2-3 µm. Conidiogenous cells terminal and intercalary, unbranched, subcylindrical, 5-12 x 3-4 µm, medium brown, smooth or almost so to finely verruculose, apex of conidiogenous cells frequently swollen, up to 6 µm diam, with 1-3(-4) flat-tipped, non to slightly thickened, non to slightly darkened-refractive loci, 1-1.5 µm wide, frequently appearing subdenticulate, up to 1.5 µm long, intercalary conidiogenous cells also slightly swollen at the conidiogenous portion just below the upper septum, which render the conidiophores subnodulose to nodulose, swellings round about the conidiophore axis or unilateral. Conidia ellipsoid-ovoid, subcylindrical, pale to medium olivaceous-brown or brown, finely verruculose, wall 0.5 µm wide, guttulate or not, occurring in branched chains. Ramoconidia 0-1(-3)-septate, 5-15(-22) x 3-4(-5) µm, with 1-3 subdenticulate apical hila; secondary conidia 0(-1)-septate, ellipsoid, obovoid to irregular, (4-)5-7(-8) x (2-)2.5-3(-4) µm; hila non to slightly thickened, non to slightly darkened-refractive, (0.5-)1(-1.5) µm wide.

Cultural characteristics: Colonies on OA erumpent, spreading, with dense, compact aerial mycelium, and even, smooth margins; olivaceous-grey (surface), margins iron-grey. Colonies reaching 22 mm diam after 1 mo at 25 °C in the dark.

Specimen examined: Venezuela, isolated from man with tinea nigra, Jan. 1975, D. Borelli, holotype CBS H-19934, culture ex-type CBS 106.75.

Notes: The type culture of Penidiella venezuelensis was originally determined as Stenella araguata from which it is, however, quite distinct by having smooth mycelium, long penicillate conidiophores with subdenticulate conidiogenous loci, smaller conidia, and agreeing with the concept of the genus Penidiella. It is phylogenetically distinct from the other taxa of Penidiella (see Fig. 1 in Crous et al. 2007a - this volume).

Pseudotaeniolina J.L. Crane & Schokn., Mycologia 78: 88. 1986.

? = Friedmanniomyces Onofri, Nova Hedwigia 68: 176. 1999.

Type species: Pseudotaeniolina convolvuli (Esfand.) J.L. Crane & Schokn., Mycologia 78: 88. 1986.

Description: Crane & Schoknecht (1986, figs 3-19).

View larger version (100K): [in this window] [in a new window]   Fig. 19. Readeriella brunneotingens (type material). A. Leaf spot. B. Colony on MEA. C-D. Conidia. Scale bar = 10 µm.

Readeriella Syd. & P. Syd., Ann. Mycol. 6: 484. 1908.

= Kirramyces J. Walker, B. Sutton & Pascoe, Mycol. Res. 96: 919. 1992.

= Colletogloeopsis Crous & M.J. Wingf., Canad. J. Bot. 75: 668. 1997.

Synanamorphs: Cibiessia Crous, Fungal Diversity 26: 151. 2007; also pseudocercospora-like, see Crous (1998).

Type species: Readeriella mirabilis Syd. & P. Syd., Ann. Mycol. 6: 484. 1908.

Description: Crous et al. (2004b; figs 36-38).

Notes: Several coelomycete genera are presently available to accommodate anamorphs of Capnodiales that reside in Teratosphaeriaceae, for which Readeriella is the oldest name. Other genera such as Phaeophleospora Rangel, Sonderhenia H.J. Swart & J. Walker and Lecanosticta Syd. belong to Mycosphaerellaceae.

Readeriella is polyphyletic within Teratosphaeriaceae. The recognition and circumscription (synonymy) of this genus follows the principles for anamorph genera within Capnodiales as outlined in the introduction to this volume. The only unifying character is conidial pigmentation, and the mode of conidiogenesis. Conidiogenous cells range from mono- to polyphialides with periclinal thickening, to phialides with percurrent proliferation, as observed in the type species, R. mirabilis (Fig. 18). Within the form genus conidia vary from aseptate to multiseptate, smooth to rough, and have a range of synanamorphs. Readeriella mirabilis has a synanamorph with cylindrical, aseptate conidia, while other species of Readeriella again have Cibiessia synanamorphs (scytalidium-like, with chains of dry, disarticulating conidia), suggesting the conidial morphology to be quite plastic. A re-examination of R. readeriellophora Crous & Mansilla revealed pycnidia to form a central cushion on which the conidiogenous cells are arranged (Fig. 18). This unique feature is commonly known in genera such as Coniella Höhn. and Pilidiella Petr. & Syd. (Diaporthales) (Van Niekerk et al. 2004), and has never been observed among anamorphs of the Capnodiales. Another species of Readeriella, namely "Phaeophleospora" toledana Crous & Bills, again forms paraphyses interspersed among conidiogenous cells, a rare feature in this group of fungi, while several species have conidiomata ranging from acervuli to pycnidia (Cortinas et al. 2006). Phylogenetically this coelomycete morphology, with its characteristic conidiogenesis, has evolved several times in Teratosphaeriaceae.

Readeriella blakelyi (Crous & Summerell) Crous & U. Braun, comb. nov. MycoBank MB504516.

Basionym: Colletogloeopsis blakelyi Crous & Summerell, Fungal Diversity 23: 342. 2006.

Readeriella brunneotingens Crous & Summerell, sp. nov. MycoBank MB504517. Fig. 19.

Etymology: Named after the diffuse brown pigment visible in agar when cultivated on MEA.

Readeriellae gauchensi similis, sed coloniis viridi-atris et pigmento brunneo in agaro diffundente distinguenda.

Leaf spots amphigenous, irregular specks up to 3 mm diam, medium brown with a thin, raised, concolorous border. Conidiomata amphigenous, substomatal, exuding conidia in black masses; conidiomata pycnidial in vivo and in vitro, globose, brown to black, up to 120 µm diam; wall consisting of 3-4 cell layers of brown cells of textura angularis. Conidiogenous cells brown, verruculose, aseptate, doliiform to ampulliform, or reduced to inconspicuous loci on hyphae (in vitro), proliferating percurrently near the apex, 5-7 x 3-5 µm; sympodial proliferation also observed in culture. Conidia brown, smooth to finely verruculose, ellipsoidal to subcylindrical, apex obtuse to subobtuse, tapering to a subtruncate or truncate base (1-1.5 µm wide) with inconspicuous, minute marginal frill, (5-)6-7(-8) x 2-3(-3.5) µm in vitro, becoming 1-septate; in older cultures becoming swollen, and up to 2-septate, 15 µm long and 5 µm wide.

Cultural characteristics: Colonies on MEA reaching 20 mm diam after 2 mo at 25 °C; colonies erumpent, aerial mycelium sparse to absent, margins smooth but irregularly lobate; surface irregularly folded, greenish black, with profuse sporulation, visible as oozing black conidial masses; a diffuse dark-brown pigment is also produced, resulting in inoculated MEA plates appearing dark-brown.

Specimen examined: Australia, Queensland, Cairns, Eureka Creek, 48 km from Mareeba, S 17° 11' 13.2", E 145° 02' 27.4", 468 m, on leaves of Eucalyptus tereticornis, 26 Aug. 2006, P.W. Crous, CBS-H 19838 holotype, culture ex-type CPC 13303 = CBS 120747.

Notes: Conidial dimensions of R. brunneotingens closely match those of Readeriella gauchensis (M.-N. Cortinas, Crous & M.J. Wingf.) Crous (Cortinas et al. 2006). The two species can be distinguished in culture, however, in that colonies of R. brunneotingens are greenish black in colour, sporulate profusely, and exude a diffuse, brown pigment into the agar, whereas colonies of R. gauchensis are more greenish olivaceous, and exude a yellow pigment into the agar (Cortinas et al. 2006).

Readeriella considenianae (Crous & Summerell) Crous & U. Braun, comb. nov. MycoBank MB504518.

Basionym: Colletogloeopsis considenianae Crous & Summerell, Fungal Diversity 23: 343. 2006.

Readeriella destructans (M.J. Wingf. & Crous) Crous & U. Braun, comb. nov. MycoBank MB504519.

Basionym: Kirramyces destructans M.J. Wingf. & Crous, S. African J. Bot. 62: 325. 1996.

Phaeophleospora destructans (M.J. Wingf. & Crous) Crous, F.A. Ferreira & B. Sutton, S. African J. Bot. 63: 113. 1997.

Readeriella dimorpha (Crous & Carnegie) Crous & U. Braun, comb. nov. MycoBank MB504520.

Basionym: Colletogloeopsis dimorpha Crous & Carnegie, Fungal Diversity 23: 345. 2006.

Readeriella gauchensis (M.-N. Cortinas, Crous & M.J. Wingf.) Crous & U. Braun, comb. nov. MycoBank MB504521.

Basionym: Colletogloeopsis gauchensis M.-N. Cortinas, Crous & M.J. Wingf., Stud. Mycol. 55: 143. 2006.

Readeriella pulcherrima (Gadgil & M. Dick) Crous & U. Braun, comb. nov. MycoBank MB504522.

Basionym: Septoria pulcherrima Gadgil & M. Dick, New Zealand J. Bot. 21: 49. 1983.

Stagonospora pulcherrima (Gadgil & M. Dick) H.J. Swart, Trans. Brit. Mycol. Soc. 90: 285. 1988.

= Cercospora eucalypti Cooke & Massee, Grevillea 18: 7. 1889.

Kirramyces eucalypti (Cooke & Massee) J. Walker, B. Sutton & Pascoe, Mycol. Res. 96: 920. 1992.

Phaeophleospora eucalypti (Cooke & Massee) Crous, F.A. Ferreira & B. Sutton, S. African J. Bot. 63: 113. 1997.

Notes: The epithet "eucalypti" is preoccupied by Readeriella eucalypti (Gonz. Frag.) Crous (Summerell et al., 2006), and thus the synonym "pulcherrima" becomes the next available name for this species.

Readeriella readeriellophora, see Teratosphaeria readeriellophora. Fig. 18.

Readeriella stellenboschiana (Crous) Crous & U. Braun, comb. nov. MycoBank MB504523.

Basionym: Colletogloeopsis stellenboschiana Crous, Stud. Mycol. 55: 110. 2006.

Readeriella zuluensis (M.J. Wingf., Crous & T.A. Cout.) Crous & U. Braun, comb. nov. MycoBank MB504524.

Basionym: Coniothyrium zuluense M.J. Wingf., Crous & T.A. Cout., Mycopathologia 136: 142. 1997.

Colletogloeopsis zuluensis (M.J. Wingf., Crous & T.A. Cout.) M.-N. Cortinas, M.J. Wingf. & Crous (zuluense), Mycol. Res. 110: 235. 2006.

Staninwardia B. Sutton, Trans. Br. Mycol. Soc. 57: 540. 1971.

Type species: Staninwardia breviuscula B. Sutton, Trans. Br. Mycol. Soc. 57: 540. 1971.

Description: Sutton (1971; fig. 1).

Notes: The genus Staninwardia presently contains two species, namely S. breviuscula and Staninwardia suttonii Crous & Summerell (Summerell et al. 2006), though its placement in Capnodiales was less well resolved. The genus forms acervuli on brown leaf spots, with brown, catenulate conidia covered in a mucilaginous sheath.

Schizothyriaceae clade
Schizothyrium Desm., Ann. Sci. Nat., Bot., sér. 3:11. 1849.

Type species: Schizothyrium acerinum Desm., Ann. Sci. Nat., Bot., sér. 3:11. 1849.

Description: Batzer et al. (2007; figs 3-7).

Notes: Species of Schizothyrium (Schizothyriaceae) have Zygophiala E.W. Mason anamorphs, and were recently shown to be allied to Mycosphaerellaceae (Batzer et al. 2007). Although species of Schizothyrium have thyrothecia, they cluster among genera with pseudothecial ascomata, questioning the value of this character at the family level. Based on its bitunicate asci and 1-septate ascospores, the teleomorph is comparable to others in the Capnodiales.

Mycosphaerellaceae clade
Mycosphaerella subclade
Mycosphaerella Johanson, Öfvers. Förh. Kongl. Svenska Vetensk.-Akad. 41(9): 163. 1884.

Type species: Mycosphaerella punctiformis (Pers.: Fr.) Starbäck, Bih. Kongl. Svenska Vetensk.-Akad. Handl. 15(3, 2): 9. 1889.

Anamorph: Ramularia endophylla Verkley & U. Braun, Mycol. Res. 108: 1276. 2004.

Description: Verkley et al. (2004; figs 3-16).

Notes: The genus Mycosphaerella has in the past been linked to 23 anamorph genera (Crous et al. 2000), while additional genera have been linked via DNA-based studies, bringing the total to at least 30 genera (Crous & Braun 2003, Crous et al. 2007b). However, based on ITS and SSU DNA phylogenetic studies and a reassessment of morphological characters and conidiogenesis, several anamorph genera have recently been reduced to synonymy (Crous & Braun 2003, Crous et al. 2006a). Furthermore, the DNA sequence data generated to date clearly illustrate that the anamorph genera in Mycosphaerella are polyphyletic, residing in several clades within Mycosphaerella. If future collections not known from culture or DNA sequences are to be described in form genera, we recommend that the concepts as explained in Crous & Braun (2003) be used until such stage as they can be placed in Mycosphaerella, pending a modification of Art. 59 of the International Code of Botanical Nomenclature. The genus Mycosphaerella and its anamorphs represent a future topical issue of the Studies in Mycology, and will thus be treated separately.

Dissoconium subclade
Dissoconium de Hoog, Oorschot & Hijwegen, Proc. K. Ned. Akad. Wet., Ser. C, Biol. Med. Sci. 86(2): 198. 1983.

Type species: Dissoconium aciculare de Hoog, Oorschot & Hijwegen, Proc. K. Ned. Akad. Wet., Ser. C, Biol. Med. Sci. 86(2): 198. 1983.

? = Uwebraunia Crous & M.J. Wingf., Mycologia 88: 446. 1996.

Teleomorph: Mycosphaerella-like.

Description: de Hoog et al. (1983), Crous (1998), Crous et al. (2004b; figs 3-10).

Notes: The genus Dissoconium presently encompasses six species (Crous et al. 2007b), of which two, M. lateralis Crous & M.J. Wingf. (D. dekkeri de Hoog & Hijwegen), and M. communis Crous & Mansilla (D. commune Crous & Mansilla) are also known from their Mycosphaerella-like teleomorphs. No teleomorph genus will be introduced for this clade, however, until more sexual species have been collected to help clarify the morphological features of this genus. A further complication lies in the fact that yet other species, morphologically distinct from Dissoconium, also cluster in this clade (Crous, unpubl. data).

"Passalora" zambiae subclade
"Passalora" zambiae Crous & T.A. Cout., Stud. Mycol. 50: 209. 2004.

Description: Crous et al. (2004b; figs 32-33).

Notes: This fungus was placed in the form genus "Passalora" based on its smooth mycelium, giving rise to conidiophores forming branched chains of brown conidia with thickened, darkened, refractive hila. Although derived from single ascospores, the teleomorph material was lost, and thus it needs to be recollected before the relavance of its phylogenetic position can be fully understood.

Additional teleomorph genera considered
Coccodinium A. Massal., Atti Inst. Veneto Sci. Lett. Arti, Série 2, 5: 336. 1860. (Fig. 20).

View larger version (130K): [in this window] [in a new window]   Fig. 20. Coccodinium bartschii. A. Ascomata on host. B. Ostiolar area. C. Periphysoids. D-E. Ascospores shot onto agar. F-I. Asci with thick ectotunica. J-K. Young ascospores. L-M. Mature ascospores. N. Colony on MEA. O-Q. Conidiogenous cells giving rise to conidia. R-S. Conidia. Scale bars: A, N = 250, B, D, F-G, I, L-M, O = 10 µm.

Description: Eriksson (1981, figs 34-35).

Notes: The genus Coccodinium (Coccodiniaceae) is characterised by having ascomata that are sessile on a subiculum, or somewhat immersed, semiglobose, collapsed when dry, brownish, uniloculate, with a centrum that stains blue in IKI (iodine potassium iodide). Asci are bitunicate, stalked, 8-spored, saccate, and have a thick, undifferentiated endotunica. Periphyses and periphysoids are well-developed and numerous. Ascospores are elongate, fusiform, ellipsoidal or clavate, transversely septate or muriform, hyaline or brownish (Eriksson 1981), and lack a mucous sheath. Based on a SSU sequence (GenBank accession U77668 [GenBank] ) derived from a strain identified as C. bartschii (Winka et al. 1998), Coccodinium appears to be allied to the taxa treated here in Teratosphaeria. Freshly collected cultures are relatively slow growing, and on MEA they form erumpent round, black colonies with sparse hyphal growth. On the surface of these colonies hyphal strands, consisting of brown, globose cells, give rise to conidia. Older cells (up to 15 µm diam) become fertile, giving rise to 1-3 conidia via inconspicuous phialidic loci. Conidia are fusoid-ellipsoidal to clavate, 3-5-septate, becoming constricted at the transverse septa, apex obtuse, base subtruncate, guttulate, smooth, widest in the upper third of the conidium, 15-40 x 4-7 µm. Phylogenetically Coccodinium is thus allied to the Chaetothyriales (Fig. 1), and not the Teratosphaeriaceae.

Stigmidium Trevis., Consp. Verruc.: 17. 1860. (Fig. 21).

View larger version (134K): [in this window] [in a new window]   Fig. 21. Stigmidium schaereri. A. Lichenicolous habit on Dacampia hookeri. B. Vertical section through an ascoma. C-D. Asci. E-G. Ascospores. H. Older, brown ascospores. Scale bars = 10 µm.

Type species: Stigmidium schaereri (A. Massal.) Trevis., Consp. Verruc.: 17. 1860.

Description: Roux & Triebel (1994, figs 47-50).

Notes: The type species of the genus is lichenicolous, characterised by semi-immersed, black, globose ascomata with ostiolar periphyses and periphysoids. Asci are 8-spored, fasciculate, bitunicate, (endotunica not giving a special reaction in Congo red or toluidine blue). Ascospores are fusoid-ellipsoidal, medianly 1-septate, guttulate, thin-walled, lacking a sheath. Presently no culture is available, and thus the placement of Stigmidium remains unresolved.

	DISCUSSION

TOP Abstract INTRODUCTION MATERIALS AND METHODS RESULTS DISCUSSION References

 
From the LSU sequence data presented here, it is clear that Mycosphaerella is not monophyletic as previously suggested (Crous et al. 2001, Goodwin et al. 2001). The first step to circumscribe natural genera within this complex was taken by Braun et al. (2003), who separated Cladosporium anamorphs from this complex, and erected Davidiella (Davidiellaceae; Schoch et al. 2006) to accommodate their teleomorphs. The present study reinstates the genus Teratosphaeria for a clade of largely extremotolerant fungi (Selbmann et al. 2005) and foliar pathogens of Myrtaceae and Proteaceae (Crous et al. 2004a, b, 2006b, 2007b), and further separates generic subclades within the Mycosphaerellaceae, while Batzer et al. (2007) again revealed Schizothyrium Desm. (Schizothyriaceae) to cluster within the Mycosphaerellaceae. Our results, however, provide support for recognition of Schizothyrium as a distinct genus, although Schizothyriaceae was less well supported as being separate from Mycosphaerellaceae (Capnodiales).

Although pleomorphism represents a rather unstudied phenomenon in this group of fungi, it has been observed in several species. Within the Teratosphaeria clade, Crous et al. (2007b) recently demonstrated teleomorphs to have Readeriella and Cibiessia synanamorphs, while the black yeast genera that belong to this clade, commonly have more than one anamorph state in culture. The present study also revealed Readeriella mirabilis to have two conidial types in culture, and to be highly plastic regarding its mode of conidiogenesis, and Readeriella to be the oldest generic name available for a large group of leaf-spotting coelomycetes in the Teratosphaeriaceae (Capnodiales).

Although not commonly documented, there are ample examples of synanamorphs in Capnodiales. Within Mycosphaerella, Beilharz et al. (2004) described Passalora perplexa Beilharz, Pascoe, M.J. Wingf. & Crous as a species with a coelomycete and yeast synanamorph, while Crous & Corlett (1998) described Mycosphaerella stigmina-platani F.A. Wolf to have a Cercostigmina U. Braun and Xenostigmina Crous synanamorph, and recent collections also revealed the presence of a similar species that has typical "Stigmina" (distoseptate conidia) and Pseudocercospora (euseptate conidia) synanamorphs (Crous, unpubl. data), and Crous (1998) reported Readeriella epicoccoides (coelomycete) to have a Cercostigmina (hyphomycete) synanamorph in culture.

Although the Mycosphaerella complex encompasses thousands of names, it may appear strange that it is only now that more clarity is obtained regarding the phylogenetic relationships among taxa in this group. This is partly due to the fact that these organisms are cultivated with difficulty, and also that the first paper to address the taxonomy of this complex based on DNA sequence data was only relatively recently published (Stewart et al. 1999). In the latter study, the genus Paracercospora Deighton (scars minutely thickened along the rim), was shown to be synonymous with the older genus Pseudocercospora. Similarily, Crous et al. (2001) showed that Cercostigmina (rough, irregular percurrent proliferations) was also synonymous with Pseudocercospora. This led Crous & Braun (2003) to conclude that conidiomatal type, conidial catenulation, septation and proliferation of conidiogenous cells were of less importance in separating species at the generic level. Mycovellosiella Rangel and Phaeoramularia Munt.-Cvetk. were subsequently reduced to synonymy with the older name, Passalora Fr., and characters identified as significant at the generic level were pigmentation (Cercospora vs. Passalora), scar structure (Passalora vs. Pseudocercospora), and verruculose superficial hyphae (Stenella vs. Passalora). Due to the unavailability of cultures, no decision was made regarding Stenella (verrucose conidia and mycelium), Stigmina (distoseptate conidia), and several other, less well-known genera such as Asperisporium Maubl., Denticularia Deighton, Distocercospora N. Pons & B. Sutton, Prathigada Subram., Ramulispora, Pseudocercosporidium Deighton, Stenellopsis B. Huguenin and Verrucisporota D.E. Shaw & Alcorn. In a recent study, however, Crous et al. (2006a) were able to show that Phaeoisariopsis (synnemata, conidia with slightly thickened hila) and Stigmina (distoseptate conidia) were also synonyms of Pseudocercospora.

The present study shows that most anamorph genera are polyphyletic within Teratosphaeria, and paraphyletic within Capnodiales. In some cases, generic concepts of anamorphs based on morphology and conidium ontogeny conform well with phylogenetic relationships, though this is not true in all cases due to convergence. Nevertheless, anamorphs still convey valuable morphological information that is contained in the anamorph name, and naming anamorphs continue to provide a practical system to identify the various asexual taxa encountered.

	Acknowledgments

 
We gratefully acknowledge several colleagues in different countries who have collected the material studied here, without which this work would not have been possible. We thank M. Vermaas for preparing the photographic plates. M. Grube is gratefully acknowledged for sending us fresh material of Stigmidium to include in this study, and D. Triebel and M. Grube are thanked for advice regarding the strain of hamathecial tissue in Stigmidium. K.A. Seifert is thanked for numerous discussions about anamorph concepts and advice pertaining to this paper, and for recollecting Coccodinium bartschii to enable us to clarify its phylogeny. J.K. Stone is thanked for commenting on an earlier draft version of this paper.

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