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

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Damm et al.Notes: Von Arx (in litt.) identified the strain CBS 330.75 as C.acutatum but with the remark “deviating by lack of pigment andless fusiform conidia”. While the main causal agent of coffee berrydisease (CBD) is C. kahawae (Waller et al. 1993) that belongsto the C. gloeosporioides species complex (Weir et al. 2012, thisissue), strains from the C. acutatum aggregate are not frequentlyencountered associated with coffee. Hindorf (1973) studiedColletotrichum populations from Coffea arabica in Kenya andillustrated conidia or ascospores of some strains diverging fromeach other in morphology and culture appearance, including astrain identified as C. acutatum and another as C. gloeosporioideswith conidia some of which are ellipsoidal and acute-ended. Oneof the two strains from western Kenya that are assigned to C.acutatum s. str. is derived from a suspected disease symptomon a coffee berry from Kenya that did not cause CBD (Gielink& Vermeulen, 1983). One of the endophytic strains from Coffearobusta in Brazil studied by Sette et al. (2006) showing antimicrobialactivity against Staphylococcus aureus belongs to the C. acutatumspecies complex; since only a short ITS sequence of this strainwas generated (DQ123614), the species cannot be identified.Colletotrichum walleri (clade 2) is known from a single strain fromcoffee, from Vietnam. Colletotrichum costaricense is quite distinctfrom either of these taxa based on molecular sequence data.Two Colletotrichum species have previously been describedfrom leaves of Coffea sp. in Costa Rica, C. brachysporum andC. coffeophilum. Conidia of the first are smaller than those of C.costaricense and have a different shape; they are subgloboseovoidand measure 7–8 × 4–6 µm (Saccardo et al. 1931), whilethose of C. costaricense measure on average 14.6 × 3.7 µm or 14.8× 3.8 µm depending on the medium. Conidia of C. coffeophilum arewider than those of C. costaricense, being ellipsoidal and straightor slightly curved (navicular), and measuring 13–15 × 6–8 µm(Saccardo et al. 1931).Colletotrichum costaricense may be differentiated from the otherspecies accepted here by TUB2, GAPDH and ACT sequences,and most effectively with TUB2. The ACT sequences of the twostrains differ by 2 bp, but have only 1 bp in common to separatethem from C. lupini and some of the unnamed single strains. Theclosest match in a blastn search with the TUB2 sequence of strainCBS 330.75 with 99 % identity (3 bp differences) was FN611028from a Citrus sinensis isolate (Ramos et al. 2006), while theclosest matches with the GAPDH sequence with 99 % identity (2differences) were EU647322 and EU647324 from leatherleaf fernisolates (MacKenzie et al. 2009). All isolates were from Florida,USA. The closest matches with the ITS sequence with 100 %identity were FN566877 from isolate DPI from Citrus aurantifolia inFlorida, USA (Ramos et al. 2006) and isolate c2 from Citrus sp. inBrazil (Giaretta et al. 2010).Colletotrichum cuscutae Damm, P.F. Cannon & Crous, sp.nov. MycoBank MB800500. Fig. 9.Etymology: Named after the host plant, Cuscuta.Sexual morph not observed. Asexual morph on SNA. Vegetativehyphae 1–5.5 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, simple or septate andbranched, to 35 µm long. Conidiogenous cells hyaline, smoothwalled,cylindrical to ampulliform, often integrated, polyphialidesoccationally observed, discrete phialides measuring 4–14.5 ×2.5–4.5 µm, opening 1.5–2 µm diam, collarette 0.5–1.5 µm long,periclinal thickening conspicuous. Conidia hyaline, smooth-walled,aseptate, straight, cylindrical with both ends acute, (15.5–)17.5–21(–27) × (3–)3.5–4.5 µm, mean ± SD = 19.2 ± 1.7 × 4.0 ± 0.3µm, L/W ratio = 4.8. Appressoria single or in loose clusters, palebrown, smooth-walled, elliptical to clavate, entire edge (3.5–)5.5–11.5(–15.5) × (2–)3.5–5.5(–6.5) µm, mean ± SD = 8.5 ± 3.2 × 4.6± 0.9 µm, L/W ratio = 1.8.Asexual morph on Anthriscus stem. Conidiomata acervular,conidiophores formed on pale brown angular basal cells, 3–8 µmdiam. Setae not observed. Conidiophores hyaline to pale brown,smooth-walled, septate, branched, to 40 µm long. Conidiogenouscells hyaline, smooth-walled, cylindrical to fusiform with bothends acute, 8–21 × 2–3.5 µm, opening 1–2 µm diam, collarette0.5–1 µm long, periclinal thickening conspicuous. Conidia hyaline,smooth-walled, aseptate, straight, cylindrical with both ends acute,(15–)17–20(–21) × (3.5–)4–4.5 µm, mean ± SD = 18.6 ± 1.5 × 4.2± 0.2 µm, L/W ratio = 4.5.Culture characteristics: Colonies on SNA flat with entire margin,hyaline to buff, on filter paper and Anthriscus stem partly coveredwith woolly to felty white to pale grey aerial mycelium and orangeacervuli, reverse hyaline to buff, under filter paper pale olivaceousgrey; growth 20 mm in 7 d (30 mm in 10 d). Colonies on OA flat toraised with entire margin; surface partly covered with woolly whiteto pale olivaceous grey aerial mycelium and olivaceous grey toorange acervuli appearing in rings, reverse buff, pale olivaceousgrey to olivaceous grey with orange sectors; growth 19–21 mm in 7d (27.5–31 mm in 10 d). Conidia in mass orange.Material examined: Dominica, Castle Comfort, from Cuscuta sp., 1986, C. Prior (IMI304802 holotype, CBS H-20784 isotype, culture ex-type IMI 304802).Notes: Colletotrichum cuscutae is known from a single strain,reported from Dominica. The multigene analysis indicates thatit occupies a single subclade within clade 1, quite distinct fromthe principal subclade of C. lupini. Its conidia are substantiallylonger than is typical for C. lupini (mean length 18.6 µm asopposed to 12 µm for C. lupini), though the length range for thelatter species is considerable. The appressoria of C. cuscutaeare narrower than those of C. lupini and also greater in length/width ratio.Colletotrichum species have been reported previouslyas parasitising Cuscuta species, which are themselves nonphotosyntheticparasites of other plants. Colletotrichumdestructivum was found to affect Cuscuta campestris parasitisingalfalfa crops in NW USA (Leach 1958). A strain identified as C.gloeosporioides f. sp. cuscutae was apparently used widely as abiological control agent “Lu Bao no. 1” of Cuscuta in China afterits adoption in the 1960s (Zhang 1985, Gao & Gan 1992), but itscurrent commercial status is unknown and it may no longer be inproduction (Watson et al. 2000). According to Watson et al. (2000)and Dinoor et al. (2009) the Lu Bao strain belongs to C. acutatumrather than C. gloeosporioides. However, no detailed morphologicaldata are available and the identification as C. acutatum was madeby means of primers that at that time were considered to bespecies-specific for the two species that are both now recognisedas species complexes.Guerber et al. (2003) studied isolates from Cuscuta in the USAand China that belong to two different species, neither of which64
The Colletotrichum acutatum species complexFig. 9. Colletotrichum cuscutae (from ex-holotype strain IMI 304802). A–B. Conidiomata. C–J. Conidiophores. K–P. Appressoria. Q–R. Conidia. A, C–E, Q. from Anthriscus stem.B, F–P, R. from SNA. A–B. DM, C–R. DIC, Scale bars: A = 100 µm, C = 10 µm. Scale bar of A applies to A–B. Scale bar of C applies to C–R.is conspecific with C. cuscutae. Based on GAPDH sequences(Guerber et al. 2003), isolates FRC2 and FRC7 from dodder inthe USA are C. fioriniae, while strain 783 from China (apparentlyidentical with strain Lu Bao no. 1) belongs to a subclade of clade2 that is not included in this study. Strain 783 was found to havea haplotype of MspI mtDNA identical to those of two Australianstrains causing terminal crook disease of pine, and distinct fromthose of other Cuscuta strains.In an attempt to compare endophytes of a Cuscuta parasite andits hosts in India, Suryanarayanan et al. (2000) isolated 44 fungalendophytes from Cuscuta reflexa, including C. gloeosporioides,C. truncatum and a “Colletotrichum sp.” that was not furthercharacterised. None of the strains is included in this study, andthere are no corresponding sequences available on GenBank.Colletotrichum cuscutae is separated from other species by allgenes studied except for ITS, most effectively by TUB2 and ACT.In blastn searches with the ITS, TUB2 and GAPDH sequences ofthe ex-type strain IMI 304802, no sequence matched with 100 %homology. Closest matches with the TUB2 sequence (with 98 %identity, 8 bp differences) were FN611029 and FN611028 fromCitrus aurantifolia and Citrus sinensis from Florida, USA (Ramos etal. 2006) and the closest matches with the GAPDH sequence (with98 % identity and 4 bp differences) were EU168905, EU647318 andEU647319 from sweet orange (Peres et al. 2008, MacKenzie et al.2009). In a blastn search with the ITS sequence a large number ofstrains were 99 % identical with that of strain IMI 304802 includingseveral ITS sequences from Key lime isolates, e.g. EU647307 andEU647308 (MacKenzie et al. 2009).Colletotrichum fioriniae (Marcelino & Gouli) R.G. Shivas &Y.P. Tan, Fungal Diversity 39: 117. 2009. Fig. 10.Basionym: Colletotrichum acutatum var. fioriniae Marcelino &Gouli, Mycologia 100: 362. 2008.≡ Glomerella fioriniae (Marcelino & Gouli) R.G. Shivas & Y.P. Tan, FungalDiversity 39: 117. 2009.≡ Glomerella acutata var. fioriniae Marcelino & Gouli, Mycologia 100: 362.2008.Sexual morph not observed. Asexual morph on SNA. Vegetativehyphae 1.5–7.5 µm diam, hyaline to pale brown, smooth-walled,septate, branched. Chlamydospores not observed. Conidiomataconidiophores formed directly on hyphae. Setae not observed.Conidiophores hyaline to pale brown, smooth-walled, septate,branched, up to 35 µm long. Conidiogenous cells hyaline to palebrown, smooth-walled, cylindrical to ampulliform, sometimes lackinga basal septum and continuous with the conidiophore, sometimescovert with a mucous coating, discrete phialides measure 4–12 ×2.5–3.5 µm, opening 1–2 µm diam, collarette 1 µm long, periclinalthickening distinct. Conidia hyaline, smooth-walled, aseptate,straight, fusiform to cylindrical with both ends acute, (10–)13.5–16.5(–19.5) × 4–5(–5.5) µm, mean ± SD = 15.0 ± 1.6 × 4.5 ± 0.3µm, L/W ratio = 3.3 µm, conidia of strain CBS 129947 are smaller,measuring (10.5–)12–15(–17) × 3.5–5(–6) µm, mean ± SD = 13.5± 1.7 × 4.1 ± 0.8 µm, L/W ratio = 3.3 µm. Appressoria solitary orin loose groups, pale to medium brown, smooth-walled, ellipsoidal,www.studiesinmycology.org65
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Studies in Mycology 73 (September 2
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Colletotrichum: complex species or
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Damm et al.Table 1. Strains of Coll
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Damm et al.Table 1. (Continued).Gen
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Damm et al.combined sequence datase
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Damm et al.Fig. 2. Colletotrichum a
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The Colletotrichum boninense specie
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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.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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