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

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Crous et al. Table 1. (Continued). Anamorph Teleomorph Accession number 1 Host Country Collector GenBank Accession number Teratosphaeria ohnowa CBS 112896*; CMW 4937; CPC 1004 Eucalyptus grandis South Africa M.J. Wingfield EU019305 Teratosphaeria secundaria CBS 115608; CPC 504 Eucalyptus grandis Brazil A.C. Alfenas EU019306 Teratosphaeria sp. CBS 208.94; CPC 727 Eucalyptus grandis Indonesia A.C. Alfenas EU019307 1 ATCC: American Type Culture Collection, Virginia, U.S.A.; CBS: Centraalbureau voor Schimmelcultures, Utrecht, The Netherlands; CPC: Culture collection of Pedro Crous, housed at CBS; CMW: Culture collection of Mike Wingfield, housed at FABI, Pretoria, South Africa; IAM: Institute of Applied Microbiology, University of Tokyo, Institute of molecular and cellular bioscience, Tokyo, Japan; IFO: Institute For Fermentation, Osaka, Japan; IMI: International Mycological Institute, CABI-Bioscience, Egham, Bakeham Lane, U.K.; INIFAT: Alexander Humboldt Institute for Basic Research in Tropical Agriculture, Ciudad de La Habana, Cuba; JCM: Japan Collection Of Microorganisms, RIKEN BioResource Center, Japan; VKM: All-Russian Collection of Microorganisms, Institute of Biochemistry and Physiology of Microorganisms, Russian Academy of Sciences, Pushchino, Russia. *Ex-type cultures. 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 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 nutrientpoor 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 (× 1 000 magnification)
Phylogenetic lineages in the Capnodiales 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 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. Bayesian analysis was conducted on the same aligned LSU dataset using a general time-reversible (GTR) substitution model with inverse gamma rates and dirichlet base frequencies. The Markov Chain Monte Carlo (MCMC) analysis of 4 chains started from a random tree topology and lasted 23 881 500 generations. Trees were saved each 100 generations, resulting in 238 815 saved trees. Burn-in was set at 22 000 000 generations after which the likelihood values were stationary, leaving 18 815 trees from which the consensus tree (Fig. 2) and posterior probabilities (PP’s) were calculated. The average standard deviation of split frequencies was 0.011508 at the end of the run. The same overall topology as that observed using parsimony was obtained, with the exception of the inclusion of Staninwardia suttonii in the Mycosphaerellaceae (PP value of 0.74) and not in the Teratosphaeriaceae. The Mycosphaerellaceae s. str. clade, as well as the Teratosphaeriaceae clade, obtained a PP value of 1.00. Taxonomy Based on the dataset generated in this study, several well-supported genera could be distinguished in the Mycosphaerella complex (Figs 1–2), for which we have identified morphological characters. These genera, and a selection of their species, are treated below. 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. www.studiesinmycology.org
Page 1: Studies in Mycology 58 (2007) The g
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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
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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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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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The genus Cladosporium and similar dematiaceous ... - CBS - KNAW

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