The genus Cladosporium and similar dematiaceous ... - CBS - KNAW

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available online at www.studiesinmycology.org doi:10.3114/sim.2007.58.02 Studies in Mycology 58: 33–56. 2007. Delimiting Cladosporium from morphologically similar genera P.W. Crous 1* , U. Braun 2 , K. Schubert 3 and J.Z. Groenewald 1 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 (Saale), Germany; 3 Botanische Staatssammlung München, Menzinger Straße 67, D-80638 München, Germany *Correspondence: Pedro Crous, p.crous@cbs.knaw.nl Abstract: The genus Cladosporium is restricted to dematiaceous hyphomycetes with a coronate scar type, and Davidiella teleomorphs. In the present study numerous cladosporium-like taxa are treated, and allocated to different genera based on their morphology and DNA phylogeny derived from the LSU nrRNA gene. Several species are introduced in new genera such as Hyalodendriella, Ochrocladosporium, Rachicladosporium, Rhizocladosporium, Toxicocladosporium and Verrucocladosporium. A further new taxon is described in Devriesia (Teratosphaeriaceae). Furthermore, Cladosporium castellanii, the etiological agent of tinea nigra in humans, is confirmed as synonym of Stenella araguata, while the type species of Stenella is shown to be linked to the Teratosphaeriaceae (Capnodiales), and not the Mycosphaerellaceae as formerly presumed. Taxonomic novelties: Devriesia americana Crous & Dugan, sp. nov., Hyalodendriella Crous, gen. nov., Hyalodendriella betulae Crous sp. nov., Ochrocladosporium Crous & U. Braun, gen. nov., Ochrocladosporium elatum (Harz) Crous & U. Braun, comb. nov., Ochrocladosporium frigidarii Crous & U. Braun, sp. nov., Rachicladosporium Crous, U. Braun & Hill, gen. nov., Rachicladosporium luculiae Crous, U. Braun & Hill, sp. nov., Rhizocladosporium Crous & U. Braun, gen. nov., Rhizocladosporium argillaceum (Minoura) Crous & U. Braun, comb. nov., Toxicocladosporium Crous & U. Braun, gen. nov., Toxicocladosporium irritans Crous & U. Braun, sp. nov., Verrucocladosporium K. Schub., Aptroot & Crous, gen. nov., Verrucocladosporium dirinae K. Schub., Aptroot & Crous, sp. nov. Key words: Cladosporium, Davidiella, food spoilage, hyphomycetes, indoor air, LSU phylogeny, taxonomy. Introduction Cladosporioid hyphomycetes are common, widespread fungi. The genus Cladosporium Link is based on the type species, Cladosporium herbarum (Pers. : Fr.) Link, which in turn has been linked to Davidiella Crous & U. Braun teleomorphs (Braun et al. 2003, Schubert et al. 2007b – this volume). Cladosporium is one of the largest, most heterogeneous genera of hyphomycetes, comprising more than 772 names (Dugan et al. 2004), and including endophytic, fungicolous, human pathogenic, phytopathogenic and saprobic species. Species of this genus affect daily human life in various ways. The common saprobic members of Cladosporium occur on all kinds of senescing and dead leaves and stems of herbaceous and woody plants, as secondary invaders on necrotic leaf lesions caused by other fungi, are frequently isolated from air, soil, food stuffs, paint, textiles and other organic matters, are also known to be common endophytes (Riesen & Sieber 1985, Brown et al. 1998, El-Morsy 2000) as well as phylloplane fungi (Islam & Hasin 2000, De Jager et al. 2001, Inacio et al. 2002, Stohr & Dighton 2004, Levetin & Dorsey 2006). Furthermore, some Cladosporium species are known to be potential agents of medical relevance. Cladosporium herbarum is, for instance, a common contaminant in clinical laboratories and causes allergic lung mycoses (de Hoog et al. 2000, Schubert et al. 2007b – this volume). In spite of the enormous relevance of this genus, there is no comprehensive modern revision of Cladosporium, but some attempts to revise and monograph parts of it have been initiated during the last decade (David 1997, Partridge & Morgan-Jones 2002, Wirsel et al. 2002, Braun et al. 2003, Dugan et al. 2004, Park et al. 2004, Seifert et al. 2004, Schubert & Braun 2004, 2005a, b, 2007, Heuchert et al. 2005, Schubert 2005a, b, Schubert et al. 2006). Previous molecular studies employing rDNA ITS sequence data (Crous et al. 2001) have shown Cladosporium spp. to cluster adjacent to the main monophyletic Mycosphaerella Johanson cluster, suggesting a position apart from the latter genus. Braun et al. (2003) carried out more comprehensive sequence analyses, based on ITS (ITS-1, 5.8S and ITS-2) and 18S rDNA data, providing further evidence that Cladosporium s. str. represents a sister clade of Mycosphaerella. Various authors discussed the taxonomy and circumscription of Cladosporium (von Arx 1981, 1983, McKemy & Morgan-Jones 1990, Braun 1995), reaching different conclusions. However, a first decisive revision of Cladosporium, leading to a more natural concept of this genus, was published by David (1997), who carried out comprehensive scanning electron microscopic examinations of the scar and hilum structure in Cladosporium and Heterosporium Klotzsch ex Cook. The first Scanning Electron Micrograph (SEM) studies of these structures, published by Roquebert (1981), indicated that the conidiogenous loci and conidial hila in Cladosporium are characterised by having a unique structure. David (1997) confirmed these observations, based on a wide range of Cladosporium and Heterosporium species, and demonstrated that the structures of the conidiogenous loci and hila in the latter genus fully agree with those of Cladosporium, proving that Heterosporium was indeed a synonym of Cladosporium s. str. He introduced the term “coronate” for the Cladosporium scar type, which is characterised by having a central convex part (dome), surrounded by a raised periclinal rim (David 1997), and showed that this type is confined to anamorphs, as far as experimentally proven, connected with teleomorphs belonging in “Mycosphaerella” s. lat. These results were confirmed in a later phylogenetic study by Braun et al. (2003). Cladosporium s. str. was shown to be a sister clade to Mycosphaerella s. str., for which the new teleomorph genus Davidiella was proposed. Although no clear morphological differences were reported between Davidiella and Mycosphaerella, a further study by Aptroot (2006) found ascospores of Davidiella to have characteristic irregular cellular inclusions (lumina), which are absent in species of Mycosphaerella, along with periphysoids and pseudoparaphyses (Schubert et al. 2007b – this volume). Furthermore, a higher order phylogeny study by Schoch et al. (2006), which employed DNA sequence data of four loci (SSU nrDNA, LSU nrDNA, EF-1α, RPB2), revealed species of Davidiella to cluster in a separate family (Davidiellaceae) from species of Mycosphaerella (Mycosphaerellaceae), with both families residing in the Capnodiales (Dothideomycetes), and not Dothideales as always presumed. 33
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Page 7 and 8: CONTENTS P.W. Crous, U. Braun and J
Page 9 and 10: lectotype for the genus by Clements
Page 11: Schubert K (2005a). Morphotaxonomic
Page 14 and 15: Crous et al. Table 1. Isolates for
Page 16 and 17: Crous et al. Table 1. (Continued).
Page 18 and 19: Crous et al. 100 10 changes 65 100
Page 20 and 21: Crous et al. Treatment of phylogene
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Page 60 and 61: Crous et al. Fig. 10. Ochrocladospo
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Page 70 and 71: Arzanlou et al. To date 26 species
Page 72 and 73: Arzanlou et al. Table 1. (Continued
Page 74 and 75: Arzanlou et al. 10 changes Athelia
Page 76 and 77: Arzanlou et al. Athelia epiphylla A
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Arzanlou et al. Fig. 26. Veronaea c
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Arzanlou et al. Cultural characteri
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Arzanlou et al. Fig. 30. Myrmecridi
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Arzanlou et al. Fig. 32. Radulidium
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Arzanlou et al. Fig. 34. Rhodoveron
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Arzanlou et al. even further, thoug
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available online at www.studiesinmy
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Dichocladosporium gen. nov. 10 chan
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Dichocladosporium gen. nov. Fig. 3.
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Dichocladosporium gen. nov. to intr
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Dichocladosporium gen. nov. Czechos
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Cladosporium herbarum species compl
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Cladosporium sphaerospermum species
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Crous et al. The aim of the present
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Crous et al. 10 changes Athelia epi
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Crous et al. Table 1. Isolates used
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Crous et al. Table 1. (Continued).
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Crous et al. 0.1 expected changes p
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Crous et al. Fig. 6. Cladophialopho
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Crous et al. Fig. 11. Cladophialoph
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Crous et al. Fig. 13. Cladophialoph
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Crous et al. Fig. 15. Exophiala sp.
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Crous et al. Fig. 18. Cylindrosympo
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Crous et al. Fig. 19. Fusicladium a
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Crous et al. Fig. 22. Fusicladium f
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Crous et al. Fig. 24. Fusicladium p
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Crous et al. Fusicladium rhodense C
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Crous et al. Excluded taxa Polyscyt
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Crous et al. the conidial tips are
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Cladophialophora carrionii complex
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Hormoconis resinae and morphologica
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Fig. 6 Hormoconis resinae and morph
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Cladophialophora, 52 k , 54 k -55,
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Fusicladium phillyreae, 189 c , 191
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Ramichloridium epichloës, 60, 89 P
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Trimmatostroma salicis, 3 t , 5, 6
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