Colletotrichum: complex species or species ... - CBS - KNAW

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Damm et al.1.001 change971.000.991.00831.00791.000.98991.000.971.00931.0099 100CBS 133.44 Godetia DenmarkCBS 285.50 Malus unknownCBS 131331 Juglans AustriaCBS 126527 Prunus UKIMI 376331 Prunus NorwayCBS 155.25 nut shell unknownCBS 131332 Agrimonia AustriaCBS 126522 Prunus NLCBS 126512 Bonzai NLCBS 125972 Fragaria NLCBS 126516 Fragaria NLCBS 126503 Fragaria UKIMI 351589 Fragaria IrelandCBS 125974 Fragaria UKIMI 351253 Fragaria UKIMI 345026 Fragaria SpainCBS 126520 Parthenocissus NLIMI 362149b Laurus UKCBS 129942 Mahonia GermanyCBS 129951 Vitis UKCBS 130251 Olea ItalyCBS 193.32 Olea GreeceCBS 171.59 Juglans unknownCBS 160.50 Citrus unknownCBS 796.72 Aeschynomene USACBS 126376 Fragaria NLIMI 345035 Fragaria FranceCBS 130252 Olea ItalyIMI 351262 Fragaria BelgiumCBS 198.53 Malus NLIMI 351248 Ceanothus UKCBS 129911 Podocarpus ZACBS 129912 Podocarpus ZACBS 129913 Podocarpus ZACBS 129934 Prunus IsraelCBS 862.70 Sambucus NLIMI 381927 Rubus TurkeyCBS 129809 Solanum ColombiaCBS 129816 Solanum ColombiaCBS 129917 Schinus MexicoCBS 127561 Ugni ChileIMI 357027 Citrus NZCBS 128532 Solanum NZ1.00 1.00CBS 128531 Pyrus NZCBS 118194 Phormium GermanyCBS 199.35 Phormium UKCBS 102054 Phormium NZCBS 118201 Phormium NZCBS 118197 Phormium NZCBS 483.82 Phormium NZCBS 124953 Phormium NLCBS 118191 Phormium ZA911.00100 721.001.000.990.98100100871.00 1.001.00CBS 198.35 Phormium UK100 CBS 116478 Trachycarpus ZA1.00 CBS 131325 Hakea Australia1.00 CBS 607.94 Salix NL1.00 CBS 191.56 Salix Germany75CBS 159.27 unknown unknownCBS 115.14 Solanum Germany0.96CBS 113.14 unknown GermanyCBS 128558 Salix NZCBS 223.36 Populus NLCBS 129972 Acer USACBS 129356 Rhododendron Latvia100 CBS 128559 Pyrus NZ1.00IMI 345581 Fragaria NZIMI 345585 Fragaria NZCBS 129973 Acer USACBS 128557 Fragaria NZCBS 192.56 Salix GermanyCBS 465.83 Araucaria USACBS 128556 Fragaria NZCBS 239.49 unknown unknownCBS 180.97 Populus NLIMI 385055 Malus NZCBS 632.80 Dendrobium USACBS 631.80 Ascocenda USA73 IMI 309357 Phalaenopsis UKCBS 119291 Cygnoches PanamaCBS 129953 Olea PortugalCBS 131322 Vaccinium USACBS 128530 Malus NZC. godetiaeC. johnstoniiC. pyricolaC. phormiiC. rhombiformeC. acerbumC. kinghorniiC. australeC. salicisC. orchidophilumA4A6A7F1*F3F7F8F2F1*K1Clade 5Fig. 1. (Continued).first clade is well supported with a bootstrap support of 100 %and a Bayesian posterior probability value of 1.00. It consists oftwo frequently isolated, well-supported clades (bootstrap support/Bayesian posterior probability value of both 100/1.00) comprisingseveral strains each, representing C. lupini (A1, J2/J6) and C.tamarilloi (A8). Other less frequently encountered subclades in thefirst clade include C. costaricense (94/1.00) with two strains, C.cuscutae and C. melonis both represented by single-strain cladeson long branches, and several short-branched single-strain clades,including the known species C. limetticola (J3) and six furtherunnamed strains. The majority of strains in clade 2 (A2, 86/1.00)belong to C. nymphaeae (98/1.00, D2/D4), while most of the other11 subclades of this clade are occupied by only one or few strains.The clade representing C. scovillei (99/1.00) consists of threestrains and groups (98/1.00) with a single-strain clade formed byC. guajavae. These two adjacent clades probably correspond toclade D3 in Guerber et al. (2003). The other sister clades representC. simmondsii (99/1.00), C. chrysanthemi (100/1.00), C. paxtonii(99/1.00), C. laticiphilum (99/1.00), C. cosmi, C. walleri, C. sloanei,C. indonesiense and C. brisbanense, the last five of which consistof single-strain clades. Clades 3 and 4 are well-supported (100/1.00and 100/0.87) and on long branches; they represent C. fioriniae(A3, C1/C2/C3) and C. acutatum (A5, J4). Clade 5 consists of twosister clades. Colletotrichum godetiae (A4, F1*, 99/1.00), formedby a large number of strains, belongs to the first sister clade andgroups (97/1.00) with C. johnstonii (F8, 100/1.00) and a single-strainclade representing C. pyricola (F2). The other sister clade (75/1.00)consists of six subclades: a large, long-branched and almosthomogenous subclade representing C. salicis (A7, K1, 100/1.00);a short-branched subclade representing C. phormii (91/1.00); C.rhombiforme (A6, 100/1.00), which groups with a single-strainclade representing C. acerbum on a long branch (F1*, 100/1.00);plus C. australe (100/1.00) and C. kinghornii on long branches.Strains named F1 appear in the phylogeny of Guerber et al. (2003)in different subclades, corresponding to C. acerbum, C. godetiaeand probably also C. rhombiforme. Colletotrichum pseudoacutatum42
The Colletotrichum acutatum species complexFig. 2. Colletotrichum acerbum (from ex-holotype strain CBS 128530). A–B. Conidiomata. C. Tip of a seta. D. Basis of a seta. E–J. Conidiophores. K–O. Appressoria. P–Q.Conidia. A, C–G, P. from Anthriscus stem. B, H–O, Q. from SNA. A–B. DM, C–Q. DIC, Scale bars: A = 100 µm, E = 10 µm. Scale bar of A applies to A–B. Scale bar of E appliesto C–Q.is only distantly related to the C. acutatum complex and is thereforenot included in the phylogeny, while C. orchidophilum was found tobe more closely related and was therefore used as outgroup. Thephylogenetic position of these and all other species included here isexhibited in fig. 1 and 2 of Cannon et al. (2012, this issue).The individual alignments and maximum parsimony analysesof the six single genes were compared with respect to theirperformance in species recognition. With ITS and CHS-1, only 11and 13 species, respectively, can be recognised. All subclades arerecognised with TUB2 and GAPDH. TUB2 performs better thanGAPDH due to higher numbers of base pair differences, but evenwith TUB2 there are clades with differerences of only 1 bp, whichsuggests that both genes should be used for identification. Theperformance of the other two genes is intermediate between ITSand TUB2/GAPDH.TaxonomyBased on DNA sequence data and morphology, the 331 strainsstudied (Table 1) are assigned to 31 species, of which 29 speciesare within the C. acutatum species complex and two outside thisgroup, including 21 species that proved to be new to science. Twospecies formed sexual morphs in vitro. All species studied in cultureare characterised below.Colletotrichum acerbum Damm, P.F. Cannon & Crous, sp.nov. MycoBank MB800494. Fig. 2.Etymology: acerbus = Latin for bitter; referring to bitter rot, thevernacular name for Colletotrichum disease of apple.Sexual morph not observed. Asexual morph on SNA. Vegetativehyphae 1–6 µm diam, hyaline, smooth-walled, septate,branched. Chlamydospores not observed. Conidiomata absent,conidiophores formed directly on hyphae. Setae not observed.Conidiophores hyaline, smooth-walled, septate, branched, to 30µm long. Conidiogenous cells hyaline, smooth-walled, cylindrical toampulliform, sometimes lacking a basal septum and continuous withthe conidiophore, sometimes extending to form new conidiogenousloci, polyphialides sometimes observed, discrete phialides measure7–18 × 3–4.5 µm, opening 1.5–2 µm diam, collarette 0.5–1.5 µmlong, periclinal thickening distinct. Conidia hyaline, smooth-walled,aseptate, straight, cylindrical to clavate with one end round andone end slightly acute or both ends round, 15.5–20.5(–29) × (4–)4.5–5 µm, mean ± SD = 17.9 ± 2.4 × 4.7 ± 0.2 µm, L/W ratio =3.8. Appressoria single or in loose groups, medium to dark brown,smooth-walled, clavate, ovate or irregular outline, the edge entireor undulate, sometimes lobate, (8–)9–14(–16.5) × (4–)5–7.5(–9.5)µm, mean ± SD = 11.3 ± 2.4 × 6.2 ± 1.2 µm, L/W ratio = 1.8.Asexual morph on Anthriscus stem. Conidiomata acervular,conidiophores formed on a cushion of pale brown angular cells,www.studiesinmycology.org43
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Damm et al.Table 3. Appressoria mea
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Damm et al.Fig. 25. Colletotrichum
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Damm et al.to olivaceous grey, filt
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Damm et al.are formed directly from
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Damm et al.Arx JA von (1957). Die A
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Damm et al.Masaba DM, Waller JM (19
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114
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Weir et al.grass-inhabiting species
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Weir et al.Table 1. A list of strai
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Weir et al.Table 1. (Continued).Spe
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Weir et al.Table 1. (Continued).Spe
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Weir et al.Table 2. Primers used in
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Weir et al.11ICMP 17789 Malus USA1I
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Weir et al.Kahawae cladeC. alataeC.
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Weir et al.A0.991ICMP 17905 Coffea
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Weir et al.Conidial width variation
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Weir et al.G. c.“f.sp. camelliae
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Weir et al.Fig. 10. Colletotrichum
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Weir et al.Fig. 18. Glomerella cing
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Weir et al.images under C. fructico
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Weir et al.Specimen examined: Thail
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Weir et al.Notes: First described f
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Weir et al.gossypii var. cephalospo
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Weir et al.to utilise either citrat
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Weir et al.(ex-epitype culture CBS
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Weir et al.of this species to C. ps
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Weir et al.Table 4. Performance of
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Weir et al.Crous PW, Gams W, Stalpe
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Weir et al.Simmonds JH (1968). Type
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Cannon et al.182
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Cannon et al.In rare instances, Col
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Cannon et al.Table 1. Summary of pr
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Cannon et al.Table 1. (Continued).P
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Cannon et al.Table 2. Colletotrichu
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Cannon et al.as did those for Colle
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Cannon et al.Table 3. Authentic seq
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Cannon et al.Table 3. (Continued).S
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Cannon et al.110.74 C. cuscutae IMI
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Cannon et al.Boninense cladeThe bon
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Cannon et al.C. lindemuthianum AB08
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Cannon et al.A further important ac
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Cannon et al.Hawksworth DL (2011).
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Cannon et al.Rodríguez-Guerra R, R
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Studies in MycologyStudies in Mycol
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CBS Biodiversity SeriesNo. 11:Taxon
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Studies in Mycology (ISSN 0166-0616
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