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

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Damm et al.to olivaceous grey, filter paper Anthriscus stem and medium partlycovered with felty white aerial mycelium (and salmon acervuli),reverse same colours; growth rate 17.5–21.5 mm in 7 d (28.5–31.5mm in 10 d). Colonies on OA flat with entire margin; surface honey,isabelline to olivaceous, almost entirely covered by felty white topale olivaceous grey aerial mycelium, reverse buff, olivaceous,pale olivaceous grey, olivaceous grey to iron-grey, growth rate16–18 mm in 7 d (26–29 mm in 10 d). Conidia in mass salmon.Material examined: Colombia, Cundinamarca, from fruit anthracnose of Solanumbetaceum, 13 Aug. 2010, J. Molina, (CBS H-20726 holotype, culture ex-type CBS129814 = T.A.6); Cundinamarca, from anthracnose on a fruit of Solanum betaceum,13 Aug. 2010, J. Molina, culture CBS 129811 = T.A.3; Antioquia, Santa Rosa, from aflower of Solanum betaceum, 1998, collector unknown, CBS H-20728, culture CBS129955 = Tom-12.Notes: Afanador-Kafuri et al. (2003) identified several strains fromtamarillo in Colombia as C. acutatum, three of which are included inthis study. Sreenivasaprasad & Talhinhas (2005) recognised thesestrains as a separate molecular group, A8, closely related to A1 (C.lupini).Colletotrichum tamarilloi can be separated from other speciesusing CHS-1, HIS3, TUB2 and GAPDH sequences, most effectivelywith GAPDH, and forms a uniform cluster even with six genes (Fig.1). Afanador-Kafuri et al. (2003) observed uniformity of bandingpatterns with apPCR, RAPD-PCR and A+T-rich DNA analyses ofthe strains they studied. They speculated that selection for clonalityand homogeneity had occurred among the isolates, all of whichwere collected in one region in Colombia where only one cultivarof the host was cultivated. Conidia of C. tamarilloi are uniformlyfusiform on SNA, and almost so on Anthriscus stem, while C. lupiniforms conidia that are usually clavate on SNA and cylindrical onthe stems. Additionally, we found that appressoria of C. lupini havean undulate to lobate margin, while those of C. tamarilloi have anentire or rarely slightly undulate edge.This species is only known on Solanum betaceum in Colombia.There are no previously described species associated with thishost. Three Colletotrichum species are reported from tamarillo inthe USDA fungal databases (Farr & Rossman 2012): C. acutatum(Guerber et al. 2003, Gadgil 2005) and C. gloeosporioides (Gadgil2005) in New Zealand and C. simmondsii in Australia (Shivas & Tan2009). None of these species/groups is identical with C. tamarilloi.While C. lupini and C. tamarilloi form well-supported clusters, thereare several additional species and unnamed strains from varioushosts in Central and South America, as well as in Florida that areclosely related to C. lupini and C. tamarilloi. One of these is fromtamarillo in the same locality in Colombia (CBS 129810).A recently reported anthracnose pathogen of tamarillo inthe USA (Jones & Perez 2012) probably belongs to C. fioriniaeaccording to its ITS sequence (JN863589). The Colletotrichumstrains available to us from tamarillo in Colombia and New Zealandbelong to C. godetiae, C. tamarilloi and an unnamed strain relatedto C. tamarilloi (this study), as well as C. boninense, C. constrictumand C. karstii belonging to the C. boninense species complex(Damm et al. 2012, this issue). Yearsley et al. (1988) report C.acutatum (s. lat.) infections of tamarillo in New Zealand; howevernone of our tamarillo strains isolated from New Zealand belongsto the C. acutatum group. The strains from this host included inGuerber et al. (2003) and assigned to group F2 formed a clade withstrains described as C. johnstonii in this study. We did not find anyspecies on tamarillo occurring in both Colombia and New Zealand.Falconi & van Heusden (2011) studied Colletotrichum isolatescollected from Lupinus mutabilis and tamarillo in the EcuadorianAndes. They formed two different subgroups within C. acutatumbased on ITS sequence data. The isolates from lupins werepathogenic to tamarillo and vice versa, but lupin and tamarilloisolates were each more virulent to their own hosts. ITS sequenceof the ex-type strain of C. tamarilloi, CBS 129814, matched with100 % identity with JN543070 from isolate Tam7 from tamarillo, aswell as JN543066 from isolate Lup28 from L. mutabilis in Ecuador(Falconi et al. 2012).The closest TUB2 blastn matches for CBS 129814 (with 99% identity, 4 bp differences) were FN611029 and FN611028 fromisolates DPI and CS-1 from Citrus aurantifolia and Citrus sinensisfrom USA, Florida (Ramos et al. 2006). The closest GAPDHmatches (with 97 % identity) were EU647323 from leatherleaf fernand EU168905, EU647318 and EU647319 from sweet orangeisolates, all from Florida, USA (Peres et al. 2008, MacKenzie etal. 2009).Colletotrichum walleri Damm, P.F. Cannon & Crous, sp.nov. MycoBank MB800517. Fig. 32.Etymology: Named after J.M. Waller, tropical pathologist extraordinaireand a key worker on the most important Colletotrichumpathogen of coffee.Sexual morph not observed. Asexual morph on SNA. Vegetativehyphae 1–6 µm diam, hyaline, smooth-walled, septate, branched.Chlamydospores not observed. Conidiomata not developed,conidiophores formed directly on hyphae. Setae not observed.Conidiophores hyaline, smooth-walled, septate, branched, to 70µm long. Conidiogenous cells hyaline, smooth-walled, cylindrical toampulliform, 10–14 × 3–4 µm, opening 1–1.5 µm diam, collarette0.5–1 µm long, periclinal thickening distinct. Conidia hyaline,smooth-walled, aseptate, straight, cylindrical to fusiform with bothends slightly acute or one end round, (6–10.5)15.5–(–19.5) ×(3–)3.5–4.5(–5.5) µm, mean ± SD = 13.0 ± 2.7 × 4.0 ± 0.5 µm,L/W ratio = 3.3. Appressoria single, medium brown, smooth-walled,elliptical, clavate, sometimes irregularly shaped, the edge entire orundulate, (4.5–)5.5–12.5(–18.5) × (3.5–)4.5–7.5(–10.5) µm, mean± SD = 9.0 ± 3.3 × 5.9 ± 1.4 µm, L/W ratio = 1.5.Asexual morph on Anthriscus stem. Conidiomata either notdeveloped, conidiophores formed directly on hyphae, or acervular,conidiophores formed on pale brown, angular, basal cells 3.5–7 µmdiam. Setae not observed. Conidiophores hyaline to pale brown,smooth-walled, septate, branched, to 70 µm long. Conidiogenouscells hyaline to pale brown, smooth-walled, cylindrical, 12–23× 2.5–3 µm, opening 1–1.5 µm diam, collarette 0.5–1 µm long,periclinal thickening visible to distinct. Conidia hyaline, smoothwalled,aseptate, straight, sometimes slightly curved, cylindrical tofusiform with both ends ± acute or one end round, (10.5–)12–16(–18.5) × 3.5–4(–4.5) µm, mean ± SD = 13.9 ± 1.8 × 4.0 ± 0.3 µm,L/W ratio = 3.5.Culture characteristics: Colonies on SNA flat with entire margin,hyaline, filter paper pale olivaceous grey, medium, filter paper andAnthriscus stem covert with fely white aerial mycelium, reversesame colours; 21–24 mm in 7 d (31–34 mm in 10 d). Colonies onOA flat with entire margin; surface covert with felty or short floccosewhite to pale olivaceous grey aerial mycelium, reverse olivaceousgrey to iron grey, olivaceous in the centre and white towards themargin; 20–26 mm in 7 d (30.5–37.5 mm in 10 d). Conidia in masssalmon.106
The Colletotrichum acutatum species complexFig. 32. Colletotrichum walleri (from ex-holotype strain CBS 125472). A–B. Conidiomata. C–H. Conidiophores. I–N. Appressoria. O–P. Conidia. A, C–E, O. from Anthriscus stem.B, F–N, P. from SNA. A–B. DM, C–P. DIC, Scale bars: A = 100 µm, C = 10 µm. Scale bar of A applies to A–B. Scale bar of C applies to C–P.Material examined: Vietnam, Buon Ma Thuot-Dak Lac, from leaf tissue of Coffeaarabica, unknown collection date, H. Nguyen, (CBS H-20795 holotype, culture extypeCBS 125472 = BMT(HL)19).Notes: Species of the C. gloeosporioides species complex arewell-known as pathogens of Coffea, especially the African coffeeberry disease pathogen C. kahawae (Waller et al. 1993). AdditionalCoffea-associated components of this species complex fromVietnam and Thailand have been studied by Nguyen et al. (2009)and Prihastuti et al. (2009); see Weir et al. (2012, this issue) forfurther review.Masaba & Waller (1992) commented that strains identified asC. acutatum may cause minor disease of ripening coffee berries.Kenny et al. (2006) and Nguyen et al. (2010) respectively isolated,in Papua New Guinea and Vietnam, taxa in this species complexfrom coffee leaves, twigs and fruits. None of the Vietnameseisolates could infect undamaged coffee berries (Nguyen et al.2010). One of the C. acutatum cultures studied by Nguyen et al.(BMT(HL)19) was sent to CBS and a dried sample of this strain ishere designated as holotype of C. walleri. In this study, this is theonly coffee isolate from Asia, while six other isolates from coffee,originating from Africa and Central America, belong to three otherspecies within the C. acutatum species complex (C. fioriniae, C.acutatum s. str. and C. costaricense). Two of these strains wereincluded in the study by Waller et al. (1993).Colletotrichum walleri is separated from other species byalmost all genes. It is most easily distinguished using HIS3 and ITSsequences, while sequences of other genes differ by only one bpfrom those of other species. The CHS-1 sequence is the same asthat of C. sloanei. The closest TUB2 blastn match for CBS 125472(with 99 % identity, 5 bp differences) was GU246633 from isolateR14 from Capsicum annuum from South Korea (Sang et al. 2011).The closest GAPDH match for a sequence covering ± the fullgene length (with 98 % identity, 4 bp differences) was HQ846724from isolate OBP6 from an unnamed plant, probably from India(Chowdappa P, Chethana CS, Madhura S, unpubl. data). The only100 % match with the ITS sequence was FJ968601, the sequenceof the same isolate previously sequenced by Nguyen et al. (2009).DISCUSSIONColletotrichum acutatum (in the broad sense) was originallydistinguished using morphological characteristics. The primarydiagnostic feature was given as the possession of fusiform conidiawith acute ends (Simmonds 1965). More detailed research hashowever shown that this characteristic is not absolute; while moststrains of species within the C. acutatum complex have at least aproportion of conidia with at least one acute end, it is common tofind significant variation in conidial shape within species and evenwithin individual strains. Conidia that are more or less cylindricalare frequently encountered. The variation may have multiplecauses; in some circumstances it seems that secondary conidiawww.studiesinmycology.org107
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Colletotrichum: complex species or
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Cannon et al.In rare instances, Col
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Studies in Mycology (ISSN 0166-0616
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