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Damm et al.Sutton BC (1980). The Coelomycetes. Fungi Imperfecti with Pycnidia, Acervuli andStromata. Commonwealth Mycological Institute, Kew, UK.Sutton BC (1992). The genus Glomerella and its anamorph Colletotrichum. In:Colletotrichum: Biology, Pathogenicity, and Control (Bailey JA , Jeger MJ, eds).CAB International, Wallingford, UK: 1–26.Swofford DL (2000). PAUP* 4.0: phylogenetic analysis using parsimony (* and othermethods). Sinauer Associates, Sunderland, MA.Tarnowski TLB, Ploetz RC (2010). First report of Colletotrichum boninense, C.capsici, and a Glomerella sp. as causes of postharvest anthracnose of passionfruit in Florida. Plant Disease 94: 786.Thaung MM (2008). Biodiversity survey of coelomycetes in Burma. AustralasianMycologist 27: 74–110.Tozze Jr HJ, Massola Jr NM, Camara MPS, Gioria R, Suzuki O, Brunelli KR,Braga RS, Kobori RF (2009). First report of Colletotrichum boninense causinganthracnose on pepper in Brazil. Plant Disease 93: 106.Tozze Jr HJ, Fischer IH, Camara MPS, Massola Jr NS (2010). First report ofColletotrichum boninense infecting yellow passion fruit (Passiflora edulis f.flavicarpa) in Brazil. Australasian Plant Disease Notes 5: 70–72Uecker FA (1994). Ontogeny of the ascoma of Glomerella cingulata. Mycologia 86:82–88.Vassiljevski NI, Karakulin BP (1950). Fungi imperfecti Parasitici. Pars II.Melanconiales. Academiae Scientiarum URSS, Moscow and Leningrad.Vega FE, Simpkins A, Aime MC, Posada F, Peterson SW, et al. (2010). Fungalendophyte diversity in coffee plants from Colombia, Hawai’i, Mexico and PuertoRico. Fungal Ecology 3: 122–138.Weir BS, Johnston PR, Damm U (2012). The Colletotrichum gloeosporioidesspecies complex. Studies in Mycology 73: 115–180.White TJ, Bruns T, Lee S, Taylor J (1990). Amplification and direct sequencing offungal ribosomal RNA genes for phylogenetics. In: PCR Protocols: a guide tomethods and applications (Innis MA, Gelfand DH, Sninsky JJ, White TJ, eds),Academic Press, San Diego: 315–322.Woudenberg JHC, Aveskamp MM, Gruyter J de, Spiers AG, Crous PW (2009).Multiple Didymella teleomorphs are linked to the Phoma clematidinamorphotype. Persoonia 22: 56–62.Yang YL, Liu ZY, Cai L, Hyde KD, Yu ZN, McKenzie EHC (2009). Colletotrichumanthracnose of Amaryllidaceae. Fungal Diversity 39: 123–146.Yang YL, Cai L, Yu ZN, Liu ZY, Hyde KD (2011). Colletotrichum species onOrchidaceae in southwest China. Cryptogamie Mycologie 32: 229–253.Yuan ZL, Su ZZ, Mao LJ, Peng YQ, Yang GM (2011). Distinctive endophytic fungalassemblage in stems of wild rice (Oryza granulata) in China with specialreference to two species of Muscodor (Xylariaceae). Journal of Microbiology49: 15–23.Zhuang W-Y, Ed. (2001). Higher Fungi of Tropical China. Mycotaxon, Ltd., Ithaca,NY.36
available online at www.studiesinmycology.orgStudies in Mycology 73: 37–113.The Colletotrichum acutatum species complexU. Damm 1* , P.F. Cannon 2 , J.H.C. Woudenberg 1 , and P.W. Crous 1,3,41CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands; 2 CABI Europe-UK, Bakeham Lane, Egham, Surrey TW20 9TY, UK and RoyalBotanic Gardens, Kew, Richmond TW9 3AB, UK; 3 Microbiology, Department of Biology, Utrecht University, Padualaan 8, 3584 CH Utrecht, The Netherlands; 4 WageningenUniversity and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands*Correspondence: Ulrike Damm, u.damm@cbs.knaw.nlAbstract: Colletotrichum acutatum is known as an important anthracnose pathogen of a wide range of host plants worldwide. Numerous studies have reported subgroups withinthe C. acutatum species complex. Multilocus molecular phylogenetic analysis (ITS, ACT, TUB2, CHS-1, GAPDH, HIS3) of 331 strains previously identified as C. acutatum andother related taxa, including strains from numerous hosts with wide geographic distributions, confirmed the molecular groups previously recognised and identified a series ofnovel taxa. Thirty-one species are accepted, of which 21 have not previously been recognised. Colletotrichum orchidophilum clusters basal to the C. acutatum species complex.There is a high phenotypic diversity within this complex, and some of the species appear to have preferences to specific hosts or geographical regions. Others appear to beplurivorous and are present in multiple regions. In this study, only C. salicis and C. rhombiforme formed sexual morphs in culture, although sexual morphs have been describedfrom other taxa (especially as laboratory crosses), and there is evidence of hybridisation between different species. One species with similar morphology to C. acutatum but notbelonging to this species complex was also described here as new, namely C. pseudoacutatum.Key words: anthracnose, Ascomycota, Colletotrichum acutatum, Gloeosporium, Glomerella, phylogeny, systematics.Taxonomic novelties: New combinations - Colletotrichum limetticola (R.E. Clausen) Damm, P.F. Cannon & Crous, C. lupini (Bondar) Damm, P.F. Cannon & Crous, C. salicis(Fuckel) Damm, P.F. Cannon & Crous. New species - C. acerbum Damm, P.F. Cannon & Crous, C. australe Damm, P.F. Cannon & Crous, C. brisbanense Damm, P.F. Cannon& Crous, C. cosmi Damm, P.F. Cannon & Crous, C. costaricense Damm, P.F. Cannon & Crous, C. cuscutae Damm, P.F. Cannon & Crous, C. guajavae Damm, P.F. Cannon &Crous, C. indonesiense Damm, P.F. Cannon & Crous, C. johnstonii Damm, P.F. Cannon & Crous, C. kinghornii Damm, P.F. Cannon & Crous, C. laticiphilum Damm, P.F. Cannon& Crous, C. melonis Damm, P.F. Cannon & Crous, C. orchidophilum Damm, P.F. Cannon & Crous, C. paxtonii Damm, P.F. Cannon & Crous, C. pseudoacutatum Damm, P.F.Cannon & Crous C. pyricola Damm, P.F. Cannon & Crous, C. rhombiforme Damm, P.F. Cannon & Crous, C. scovillei Damm, P.F. Cannon & Crous, C. sloanei Damm, P.F. Cannon& Crous, C. tamarilloi Damm, P.F. Cannon & Crous, C. walleri Damm, P.F. Cannon & Crous. Typifications: Epitypifications - C. acutatum J.H. Simmonds, C. limetticola (R.E.Clausen) Damm, P.F. Cannon & Crous, C. nymphaeae (Pass.) Aa, C. phormii (Henn.) D.F. Farr & Rossman, C. salicis (Fuckel) Damm, P.F. Cannon & Crous. Lectotypifications- C. nymphaeae (Pass.) Aa, C. orchidearum Allesch.Published online: 16 August 2012; doi:10.3114/sim0010. Hard copy: September 2012.Studies in MycologyINTRODUCTIONColletotrichum acutatum is one of the most frequently reportedspecies of the genus and causes diseases commonly known asanthracnose on numerous host plants worldwide (Farr & Rossman2012). Originally described from diseased tissues of Caricapapaya, Capsicum frutescens and Delphinium ajacis in Australiaby Simmonds (1965), the C. acutatum species complex is todayknown as especially destructive on fruits like strawberry (Garridoet al. 2009), citrus (Peres et al. 2008), apple (Lee et al. 2007),olive (Talhinhas et al. 2011), cranberry (Polashock et al. 2009) andblueberry (Wharton & Schilder 2008). It is also implicated in the“terminal crook” disease of pine (Dingley & Gilmour 1972) and inthe anthracnose of leather leaf fern (Schiller et al. 2006). There arealso reports of a disseminated infection of a sea turtle (Manire etal. 2002) and the infection of a scale insect (Marcelino et al. 2008).Reviews of the species in its broad sense and its pathology werepublished by Wharton & Diéguez-Uribeondo (2004) and Peres etal. (2005).On strawberry, C. acutatum mainly causes black spot of fruitbut can also attack crowns, roots and leaves (Freeman & Katan1997), and is one of the most serious diseases in commercial fruitproduction. Largely due to its economic importance as a strawberryCopyright CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.pathogen, C. acutatum was treated for many years as a regulatedplant quarantine pest by the European and Mediterranean PlantProtection Organization (EPPO), though it is absent from thecurrent list (EPPO 2011) – presumably due to its now widespreaddistribution in Europe. Inoculum sources are frequently transplantmaterial, mostly with quiescent infections (Rahman & Louws 2008),infected plants, weeds and other hosts (McInnes et al. 1992,Parikka et al. 2006), while the survival rate of conidia in natural fieldsoil is low (Freeman et al. 2002).The most well-known morphological feature of C. acutatum (s.lat.) is the shape of its conidia, which have acute ends (Simmonds1965). However, other conidial shapes, especially ± cylindricalwith only one acute end, are frequently encountered, especially instrains that have been repeatedly subcultured, but these conidialshapes can also occur in species outside the C. acutatum speciescomplex. Even the differentiation between C. acutatum (s. lat.) andC. gloeosporioides (s. lat.) is difficult, because many intermediatestrains exist with a restricted number of typical fusiform conidia andmany cylindrical ones (Van der Aa et al. 1990). On the host, conidiaare formed in acervuli; in culture, conidia are often also producedin the aerial mycelium (Johnston & Jones 1997). Colletotrichumacutatum has also been observed to form secondary conidia on thesurface of living strawberry leaves (Leandro et al. 2001) that wereYou are free to share - to copy, distribute and transmit the work, under the following conditions:Attribution:You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work).Non-commercial: You may not use this work for commercial purposes.No derivative works: You may not alter, transform, or build upon this work.For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you getpermission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights.37
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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 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.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.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.as did those for Colle
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Cannon et al.110.74 C. cuscutae IMI
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Cannon et al.Hawksworth DL (2011).
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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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