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

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Cladosporium sphaerospermum species complex 69 EXF-322 CBS 119413 EXF-604 10 steps 100 85 C. salinae 62 64 79 98 EXF-780 EXF-321 EXF-946 CBS 170.54 ATCC 76499 ATCC 66669 EXF-699 EXF-710 EXF-697 EXF-711 CBS 125.80 EXF-563 EXF-452 EXF-371 Moricca et al. ATCC 38027 EXF-714 EXF-332 EXF-391 EXF-326 87 CBS 109868 dH 13833 CBS 601.84 dH 11736 CBS 101880 CBS 189.54 dH 12459 100 EXF-449 CBS 452.71 EXF-397 EXF-466 CBS 119417 EXF-471 EXF-739 CBS 102045 CBS 109.14 EXF-455 EXF-738 EXF-962 EXF-1061 CBS 193.54 EXF-458 CBS 300.96 Cladosporium sp. EXF-732 80 90 62 100 CPC 11683 EXF-718 EXF-720 EXF-727 72 63 79 99 EXF-977 EXF-1072 dH 12991 dH 12862 dH 13911 EXF-564 C. halotolerans EXF-380 EXF-703 EXF-646 EXF-228 EXF-572 CBS 280.49 “Mycosphaerella” hyperici CBS 177.71 C. bruhnei CBS 812.71 EXF-595 Cladosporium sp. EXF-454 C. ramotenellum ATCC 66670 Davidiella tassiana EXF-333 EXF-382 EXF-334 EXF-343 C. subinflatum C. spinulosum C. cladosporioides C. oxysporum C. tenuissimum C. sphaerospermum C. dominicanum C. psychrotolerans C. langeronii C. fusiforme C. velox Fig. 2. One of 600 equally most parsimonious and equally looking phylogenetic trees based on a heuristic tree search using aligned sequences of the internal transcribed spacer regions 1 and 2 and the 5.8S rDNA. 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 some deeper branches moderate, bootstrap support (CI = 0.804, RI = 0.975, PIC = 68). RESULTS Descriptive statistical parameters of phylogenetic analyses and calculated tree scores for each analysed sequence locus are summarised in Table 2. Mainly reference material such as extype or ex-neotype strains was analysed on the level of SSU rDNA sequences. Downloaded and newly generated SSU rDNA sequences of members of Cladosporium s. str. were compared with related taxa of the Mycosphaerellaceae, Dothioraceae and Dothideaceae. The somewhat more distantly related Fusicladium effusum (Venturiaceae) (Braun et al. 2003: Fig. 2) was selected as outgroup. Anungitopsis amoena R.F. Castañeda & Dugan (now placed in Fusicladium Bonord., see Crous et al. 2007b), also a member of the Venturiaceae, was included in the analyses. All taxa included in the SSU rDNA analysis belong to the Dothideomycetes www.studiesinmycology.org (Schoch et al. 2006), within which the ingroup is represented by the orders Capnodiales (Davidiellaceae, Mycosphaerellaceae, Teratosphaeriaceae) and Dothideales (Dothioraceae, Dothideaceae) (see also Schoch et al. 2006). The genus Cladosporium, of which some species are linked to Davidiella Crous & U. Braun teleomorphs (Braun et al. 2003), forms a statistically strongly supported monophyletic group (Davidiellaceae). It also accommodates species newly described in this paper, namely, C. halotolerans Zalar, de Hoog & Gunde-Cimerman, C. fusiforme Zalar, de Hoog & Gunde- Cimerman, C. dominicanum Zalar, de Hoog & Gunde-Cimerman, C. salinae, C. psychrotolerans Zalar, de Hoog & Gunde-Cimerman, C. velox Zalar, de Hoog & Gunde-Cimerman and C. spinulosum Zalar, de Hoog & Gunde-Cimerman (Fig. 1). A sister group relationship of Cladosporium s. str. with a clade of taxa characterised, among others, by Mycosphaerella Johanson teleomorphs, containing various anamorphic genera such as Septoria Sacc., 163
Zalar et al. 97 EXF-322 CBS 119413 EXF-604 10 steps 89 C. salinae 96 61 87 93 91 98 98 CBS 601.84 CBS 101880 dH 12459 CBS 189.54 CBS 109868 100 dH 13833 dH 11736 EXF-332 EXF-326 EXF-391 EXF-714 100 CBS 177.71 C. bruhnei ATCC 66670 Davidiella tassiana CBS 170.54 C. cladosporioides EXF-343 C. subinflatum EXF-382 C. spinulosum EXF-334 ATCC 66669 CBS 125.80 CBS 119417 C. velox C. oxysporum EXF-572 EXF-564 EXF-1072 EXF-380 C. halotolerans EXF-703 EXF-228 EXF-646 EXF-977 CBS 280.49 “Mycosphaerella” hyperici 98 dH 12862 100 63 84 100 95 93 100 100 dH 12941 dH 13911 dH 12991 EXF-727 90 8862 100 99 EXF-458 98 EXF-455 CBS 102045 EXF-1061 CBS 193.54 EXF-738 CBS 109.14 EXF-739 EXF-962 EXF-397 EXF-449 CBS 452.71 78 99 EXF-720 EXF-732 EXF-718 CPC 11683 EXF-696 C. halotolerans C. sphaerospermum C. fusiforme C. langeronii C. psychrotolerans C. dominicanum Fig. 3. One of 90 equally most parsimonious and equally looking phylogenetic trees based on a heuristic tree search using aligned exons and introns of a part of the β-tubulin 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 = 0.538, RI = 0.883, PIC = 220). Ramularia Unger, Cercospora Fresen., Pseudocercospora Speg., “Trimmatostroma” Corda (now Catenulostroma Crous & U. Braun) (see Crous et al. 2004, 2007a – this volume) and the somewhat cladosporium-like genus Devriesia Seifert & N.L. Nick. (Seifert et al. 2004), was statistically only moderately supported (Fig. 1), whereas in an analogous analysis by Braun et al. (2003: Fig. 2) it was highly supported. These data also support the conclusion by Braun et al. (2003) and Crous et al. (2006) that Cladosporium is not a member of the distantly related Herpotrichiellaceae (Chaetothyriomycetes), which is also rich in cladosporium-like taxa (Crous et al. 2006). None of the fungi isolated from hypersaline environments belonged to the Herpotrichiellaceae. The SSU rDNA sequences do not resolve a phylogenetic structure within Cladosporium s. str. Only a moderately supported clade comprising C. halotolerans, C. dominicanum, C. velox, C. sphaerospermum and C. fusiforme is somewhat distinguished from a statistically unsupported clade with C. herbarum (Pers. : Fr.) Link, C. cladosporioides (Fresen.) G.A. de Vries, C. oxysporum Berk. & Broome, C. spinulosum, and C. psychrotolerans, etc. Because C. salinae appeared most distinct within the genus Cladosporium in analyses of the SSU rDNA (Fig. 1), it was used as outgroup in analyses of the ITS rDNA and the β-tubulin and actin genes. Analyses of the more variable ITS rDNA and partial β-tubulin and actin gene introns and exons supported the species clades of C. halotolerans, C. dominicanum, C. sphaerospermum, C. fusiforme and C. velox (Figs 2–4), of which C. velox was distinguished in the β-tubulin tree by a particular long terminal branch of the only sequenced strain (Fig. 3). Cladosporium salinae also clustered as a well-supported species clade in preliminary analyses using various Mycosphaerella species as outgroup (not shown). All strains of C. langeronii (Fonseca, Leão & Nogueira) Vuill. are particularly well distinguishable from other Cladosporium species by strikingly slow- 164
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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 124 and 125: 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).
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Page 206 and 207: Crous et al. Fig. 6. Cladophialopho
Page 208 and 209: Crous et al. Fig. 11. Cladophialoph
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Page 212 and 213: Crous et al. Fig. 15. Exophiala sp.
Page 214 and 215: Crous et al. Fig. 18. Cylindrosympo
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
Page 222 and 223: 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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