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

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Cladosporium sphaerospermum species complex EXF-322 CBS 119413 EXF-604 60 100 C. salinae CBS 189.54 71 CBS 601.84 CBS 101880 dH 12459 dH 13833 96 CBS 109868 dH 11736 62 EXF-332 EXF-391 77 EXF-714 CBS 177.71 C. bruhnei 66 89 ATCC 66670 Davidiella tassiana 67 EXF-343 C. subinflatum 85 C. psychrotolerans 87 EXF-696 EXF-718 100 EXF-732 C. dominicanum CPC 11683 EXF-598 100 CBS 102045 EXF-458 EXF-1069 EXF-455 72 EXF-1061 77EXF-739 76 EXF-962 C. sphaerospermum 81 EXF-738 76 CBS 109.14 CBS 193.54 CBS 300.96 Cladosporium sp. EXF-449 100 90 EXF-397 C. fusiforme CBS 452.71 100 EXF-466 EXF-471 CBS 119417 C. velox 84 98 EXF-334 EXF-382 C. spinulosum CBS 170.54 C. cladosporioides 89 CBS 125.80 ATCC 66669 C. oxysporum dH 12862 dH 13911 EXF-646 EXF-703 EXF-228 EXF-1072 C. halotolerans EXF-572 EXF-977 EXF-564 EXF-380 CBS 280.49 “Mycosphaerella” hyperici C. langeronii 10 steps Fig. 4. One of 32 equally most parsimonious and equally looking phylogenetic trees based on a heuristic tree search using aligned exons and introns of the partial actin gene. The tree was randomly selected. Support based on 10 000 replicates of a fast step-wise addition bootstrap analysis is indicated near the branches. Trees were rooted with the strains of Cladosporium salinae. Most monophyletic species clades received high, but deeper branches weak or no, bootstrap support (CI = 0.586, RI = 0.885, PIC = 103). growing colonies at all tested temperatures and relatively large, oblong conidia. However, phylogenetic analyses of the β-tubulin and actin gene indicate that C. langeronii presents two cryptic species (Figs 3–4). The species clade of C. psychrotolerans is moderately supported in analyses of the actin gene but highly by means of the β-tubulin gene. It is evident from all three analyses (Figs 2– 4) that C. langeronii and C. psychrotolerans are closely related species. The species node of Cladosporium spinulosum, which is morphologically clearly distinguished from all other species by its conspicuous ornamentation consisting of digitate projections (Fig. 5), is supported by β-tubulin (Fig. 3) and actin (Fig. 4) sequence data but not by those of the ITS rDNA (Fig. 2). Analyses of all loci, however, indicate that it is a member of the C. herbarum complex. The analyses of sequences of the ITS and the β-tubulin and actin gene introns and exons (Figs 2–4) do not allow the full elucidation of phylogenetic relationships among these Cladosporium species. Statistical support of the interior tree branches resulting from analyses of the β-tubulin and actin genes is low (bootstrap values mostly < 50 %). While the sister group relationship of C. sphaerospermum and C. fusiforme is highly supported in the analysis based on the β-tubulin gene, analysis of the ITS rDNA indicate that these two species are unrelated, and that C. sphaerospermum is closely related to C. dominicanum. It is clear from the data that the species morphologically resembling C. sphaerospermum are not phylogenetically closely related and that the data we present here do not allow their classification in natural subgroups of the genus Cladosporium. Only C. spinulosum was placed in all analyses among species of the C. herbarum complex www.studiesinmycology.org 165
Zalar et al. Fig. 5. Conidial scars and surface ornamentation of ramoconidia and conidia (SEM). A. C. dominicanum (strain EXF-732). B. C. fusiforme (strain EXF-449). C. C. halotolerans (strain EXF-572). D. C. langeronii (strain CBS 189.54). E. C. psychrotolerans (strain EXF-391). F. C. salinae (strain EXF-335 = CBS 119413). G. C. sphaerospermum (strain CBS 193.54). H. C. spinulosum (strain EXF-334). I. C. velox (strain CBS 119417). Scale bars = 5 µm. (Photos: K. Drašlar). and all analyses supported close relatedness of C. langeronii and C. psychrotolerans. The majority of species described here have slightly ornamented conidia ranging from minutely verruculose (C. fusiforme, C. langeronii, C. psychrotolerans, C. sphaerospermum, C. velox) to verrucose (C. halotolerans) (Fig. 5). The verrucose conidia of C. halotolerans can be recognised also under the light microscope and used as a distinguishing character. Almost smooth to minutely verruculose conidia are encountered in C. dominicanum and C. salinae (Fig. 5). Cladosporium spinulosum, a member of the C. herbarum species complex, has conidia with a digitate ornamentation that can appear spinulose under the light microscope; however, when using the SEM it became clear that its projections have parallel sides and a blunt end (Fig. 5). DISCUSSION The genus Cladosporium was established by Link (1816) who originally included four species, of which C. herbarum is the type species of the genus (Clements & Shear 1931). In 1950, von Arx reported a teleomorph connection for this species with Mycosphaerella tassiana (De Not.) Johanson. Based on SSU rDNA data the majority of Mycosphaerella species, including the type species of the genus, M. punctiformis (Pers.) Starbäck, clustered within the Mycosphaerellaceae, a family separated from M. tassiana (Braun et al. 2003). Therefore, Mycosphaerella tassiana was reclassified as Davidiella tassiana (De Not.) Crous & U. Braun, the type of the new genus Davidiella. All anamorphs with a cladosporium- and heterosporium-like appearance and with a supposed Dothideomycetes relationship were maintained under the anamorph name Cladosporium, morphologically characterised by scars with a protuberant hilum consisting of a central dome surrounded by a raised rim (David 1997). The concept of distinguishing ramoconidia from secondary ramoconidia has been adopted from Schubert et al. (2007). In the species described here, ramoconidia have been observed often in C. sphaerospermum, sometimes in C. psychrotolerans, C. langeronii and C. spinulosum, and only sporadically in all other species. Therefore, ramoconidia can be seen as important for distinguishing species although sometimes, they can be observed only with difficulty. When using ramoconidia as a diagnostic criterion, colonies only from SNA and not older than 7 d should be taken into account. Cladosporium sphaerospermum was described by Penzig (1882) from decaying Citrus leaves and branches in Italy. He described C. sphaerospermum as a species with (i) branched, septate and dark conidiophores having a length of 150–300 µm and a width of the main conidiophore stipe of 3.5–4 µm, (ii) spherical to ellipsoid, acrogenously formed conidia of 3.4–4 µm diam, and (iii) ramoconidia of 6–14 × 3.5–4 µm. Penzig’s original material is not known to be preserved. Later, a culture derived from CBS 193.54, originating from a human nail, was accepted as typical of C. sphaerospermum. However, de Vries (1952), incorrectly cited it as “lectotype”, and thus the same specimen is designated as neotype in this study (see below), with the derived culture (CBS 193.54) 166
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Studies in Mycology 58 (2007) The g
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Studies in Mycology The Studies in
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CONTENTS P.W. Crous, U. Braun and J
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lectotype for the genus by Clements
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Schubert K (2005a). Morphotaxonomic
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Crous et al. Table 1. Isolates for
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Crous et al. Table 1. (Continued).
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Crous et al. 100 10 changes 65 100
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Crous et al. Treatment of phylogene
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Crous et al. Teratosphaeria bellula
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Crous et al. 6. Conidiophores short
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Crous et al. Habit plant pathogenic
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Crous et al. Fig. 7. Catenulostroma
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Crous et al. system, consisting of
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Crous et al. Fig. 9. Penidiella col
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Crous et al. Fig. 12. Penidiella ri
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Crous et al. conidiophores, about 1
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Crous et al. have conidiomata rangi
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Crous et al. Schizothyriaceae clade
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Crous et al. Fig. 21. Stigmidium sc
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Crous et al. Gams W, Verkley GJM, C
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Crous et al. Table 1. Isolates for
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Crous et al. 100 61 100 52 100 10 c
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Crous et al. Fig. 3. Rachicladospor
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Crous et al. Fig. 4. Toxicocladospo
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Crous et al. Fig. 5. Verrucocladosp
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Crous et al. Fig. 7. Stenella aragu
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Crous et al. Helotiales, incertae s
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Crous et al. Fig. 10. Ochrocladospo
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Crous et al. Fig. 12. Rhizocladospo
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Crous et al. 7. Conidiophores unbra
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Crous et al. 33. Terminal conidioge
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Crous et al. Crous PW, Kang JC, Bra
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Arzanlou et al. To date 26 species
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Arzanlou et al. Table 1. (Continued
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Arzanlou et al. 10 changes Athelia
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Arzanlou et al. Athelia epiphylla A
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Arzanlou et al. Fig. 3. Periconiell
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Arzanlou et al. Cultural characteri
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Arzanlou et al. Fig. 10. A. Ramichl
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Arzanlou et al. Ramichloridium musa
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Arzanlou et al. Fig. 20. Rhinocladi
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Arzanlou et al. Fig. 22. Rhinocladi
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Arzanlou et al. Rhinocladiella mack
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Arzanlou et al. Fig. 26. Veronaea c
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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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Page 126 and 127: Cladosporium herbarum species compl
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Page 170 and 171: Cladosporium sphaerospermum species
Page 196 and 197: Crous et al. The aim of the present
Page 198 and 199: Crous et al. 10 changes Athelia epi
Page 200 and 201: Crous et al. Table 1. Isolates used
Page 202 and 203: Crous et al. Table 1. (Continued).
Page 204 and 205: Crous et al. 0.1 expected changes p
Page 206 and 207: Crous et al. Fig. 6. Cladophialopho
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Page 212 and 213: Crous et al. Fig. 15. Exophiala sp.
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Page 216 and 217: Crous et al. Fig. 19. Fusicladium a
Page 218 and 219: Crous et al. Fig. 22. Fusicladium f
Page 220 and 221: Crous et al. Fig. 24. Fusicladium p
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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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The genus Cladosporium and similar dematiaceous ... - CBS - KNAW

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