Damm et al.belonging to the C. crassipes group as accepted by Sutton (1980).No type details were given in the <strong>or</strong>iginal description. There isa probable type specimen of C. heveae in K(M), collected fromleaves of Hevea (Petch 2228) on 7 Oct. 1905, presumably from SriLanka. It is fragmentary and also contains a fungus identified onthe label as Gloeosp<strong>or</strong>ium brunneum. Acc<strong>or</strong>ding to Petch (1927),C. heveae causes an indeterminate leaf spot, and is perhapsan invader following mechanical damage; it was not consideredto be a significant disease of rubber at that time. No fungusc<strong>or</strong>responding to the description of C. heveae was found on thetype specimen, and a slide previously made from this material(IMI 80135) also does not contain this <strong>species</strong>. Glomerellaphyllanthi (from the related plant Phyllanthus acidus) was initiallyregarded as the sexual m<strong>or</strong>ph of C. heveae (Pai 1970), but waslater revealed to belong to the C. boninense <strong>species</strong> <strong>complex</strong>,as was another <strong>species</strong> on Hevea, C. annellatum (Damm et al.2012, this issue).The conidia of Gm. heveae are about the same size as those ofC. laticiphilum (12–17 × 3.5–5 µm); however the sp<strong>or</strong>es extrude ina pale brown mass, which would be unusual f<strong>or</strong> a <strong>Colletotrichum</strong>.Also the size range of the “basidia” (= conidiogenous cells) is givenas 20–34 × 2 µm; c<strong>or</strong>responding structures of C. laticiphilum aresh<strong>or</strong>ter and much wider. There is no material in K(M) identifiedas Gm. heveae. It is possible that the fungus identified as Gm.brunneum in the type collection of C. heveae is actually Gm.heveae, as Gm. brunneum is a completely unrelated fungus<strong>or</strong>iginating from Populus leaves in the USA (Ellis & Everhart1889). Petch could have realised after writing the packet label, butbef<strong>or</strong>e publication, that naming his fungus Gm. brunneum wouldcreate a later homonym. Petch (1927) indicated that Gm. heveaewas found only in one isolated instance in 1905, when it causedleaf fall in young nursery-grown plants and resulted in generaldiscol<strong>or</strong>ation and death of the whole leaf blade. The disease wassuccessfully controlled by reducing exposure to shade. Synonymyof Gm. heveae with our fungus would not affect the naming of C.laticiphilum as a combination into <strong>Colletotrichum</strong> based on Gm.heveae would be a later homonym of C. heveae.It seems possible that Gm. alb<strong>or</strong>ubrum might be referableto the <strong>species</strong> described here. Acc<strong>or</strong>ding to Saha et al. (2002) asymptom consisting of raised spots had been attributed to this<strong>species</strong>. The fungus was <strong>or</strong>iginally described from green stems ofHevea brasiliensis, but Petch (1927) stated that it caused abn<strong>or</strong>malleaf fall and appeared to spread to green ends of the branches tocause dieback. He thought that it might be a secondary invaderfollowing Phytophth<strong>or</strong>a infection. These symptoms do not seemto c<strong>or</strong>respond well with those described by Saha and colleagues.The conidia of Gm. alb<strong>or</strong>ubrum were measured as 15–20 × 3–4µm, and described as oblong with rounded ends, straight <strong>or</strong> slightlycurved, issuing in thick pink <strong>or</strong> white tendrils (Petch 1906). Thesize is similar to C. laticiphilum; we also observed slightly curvedconidia, especially in the isolate from Colombia (<strong>CBS</strong> 129827). Theconidial shape of both C. laticiphilum isolates on Anthriscus stemis not fusif<strong>or</strong>m, but cylindrical with one end round and one endonly slightly acute. Therer are three specimens in K(M) identifiedby Petch as belonging to Gm. alb<strong>or</strong>ubrum, but none can be typematerial as they were all collected after publication of the name.Bearing in mind that definite type material of all three namesis either missing <strong>or</strong> fragmentary and that none of the authenticmaterial would be likely to yield good sequences, we think that itis m<strong>or</strong>e practical to publish a new taxon rather than to epitypify <strong>or</strong>neotypify one of the earlier names with a specimen that we are notconfident is conspecific with the type.<strong>Colletotrichum</strong> laticiphilum is separated from other <strong>species</strong> byits TUB2, GAPDH and CHS-1 sequences, and most differentiallywith TUB2. With CHS-1 there is only one bp difference from C.indonesiense, while the HIS3 sequence is the same as that ofthat <strong>species</strong>. The closest match with the GAPDH sequence (with99 % identity, 1 bp difference) was HQ846719 from an unnamedplant, probably from India (P. Chowdappa, C.S. Chethana, S.Madhura, unpubl. data). The ITS sequence of strain <strong>CBS</strong> 112989matches 100 % with AB042306 and AB042307 from isolates fromCarthamus and Glebionis from Japan (J. M<strong>or</strong>iwaki, T. Tsukiboshi,T. Sato, S. Uematsu, unpubl. data), with AJ749675 from isolatePD85/694 (= <strong>CBS</strong> 126519, C. chrysanthemi), and with AB219024from strawberry in Japan (Chung et al. 2006).<strong>Colletotrichum</strong> limetticola (R.E. Clausen) Damm, P.F.Cannon & Crous, comb. nov. MycoBank MB455483. Fig. 17.Basionym: Gloeosp<strong>or</strong>ium limetticola [as Gm. limetticolum] R.E.Clausen, Phytopathology 2: 231. 1912.Sexual m<strong>or</strong>ph not observed. Asexual m<strong>or</strong>ph on leaf of Citrusaurantifolia (BPI 394978). Conidiomata conidioph<strong>or</strong>es f<strong>or</strong>med ona cushion of pale brown angular cells 3–6 µm diam. Setae notobserved. Conidioph<strong>or</strong>es hyaline, smooth-walled, septate andbranched, up to 75 µm. Conidiogenous cells hyaline, smoothwalled,cylindrical, sometimes slightly inflated, 10–18 × 2.5–4µm, opening 1–1.5 µm diam, collarette 0.5–1 µm long, periclinalthickening visible, sometimes distinct. Conidia hyaline, smoothwalled,aseptate, straight, sometimes slightly flexuous, cylindricalwith one end round and one end slightly acute to truncate, <strong>or</strong> bothends slightly acute, (10–)12.5–17.5(–20) × (3.5–)4–4.5(–4.5) µm,mean ± SD = 15.1 ± 2.4 × 4.1 ± 0.3 µm, L/W ratio = 3.7. Appress<strong>or</strong>iafew observed on specimen, pale to medium brown, smooth-walled,subglobose, ovoid to ellipsoidal outline, entire edge.Asexual m<strong>or</strong>ph on SNA (<strong>CBS</strong> 114.14). Vegetative hyphae1–8.5 µm diam, hyaline, smooth-walled, septate, branched.Chlamydosp<strong>or</strong>es not observed. Conidiomata not developed,conidioph<strong>or</strong>es f<strong>or</strong>med directly on hyphae. Setae not observed.Conidioph<strong>or</strong>es hyaline, smooth-walled, simple <strong>or</strong> septate andbranched, up to 45 µm. Conidiogenous cells hyaline, smoothwalled,cylindrical to ampullif<strong>or</strong>m, sometimes integrated (notseparated from fertile hyphae by a septum, polyphialides rarelyobserved, 8.5–20 × 3–5.5 µm, opening 1–1.5 µm diam, collarette0.5–1 µm long, periclinal thickening visible, sometimes distinct.Conidia hyaline, smooth-walled, aseptate, straight, sometimesslightly curved, cylindrical to clavate with one end round andone end slightly acute to truncate, <strong>or</strong> both ends slightly acute,sometimes slightly constricted in the middle, (9–)12–20.5(–29) ×(3–)4–5(–6) µm, mean ± SD = 16.3 ± 4.2 × 4.5 ± 0.6 µm, L/W ratio= 3.6. Appress<strong>or</strong>ia single <strong>or</strong> in loose groups, pale to medium brown,smooth-walled, subglobose, ovoid to ellipsoidal outline, entire <strong>or</strong>undulate edge (5–)6–8.5(–11) × (4–)4.5–6(–7) µm, mean ± SD =7.4 ± 1.3 × 5.3 ± 0.7 µm, L/W ratio = 1.4.Asexual m<strong>or</strong>ph on Anthriscus stem (<strong>CBS</strong> 114.14). Conidiomataconidioph<strong>or</strong>es f<strong>or</strong>med directly on hyphae <strong>or</strong> on a cushion ofpale brown angular cells 3.5–6.5 µm diam. Setae not observed.Conidioph<strong>or</strong>es hyaline, smooth-walled, septate, branched, to 80µm long. Conidiogenous cells hyaline, smooth-walled, cylindricalslightly inflated, 6–13 × 2.5–4.5 µm, opening 1–1.5 µm diam,collarette 0.5–1 µm long, periclinal thickening visible. Conidiahyaline, smooth-walled, aseptate, straight, cylindrical, clavate,cylindrical to fusif<strong>or</strong>m with one end round and one end (often only76
The <strong>Colletotrichum</strong> acutatum <strong>species</strong> <strong>complex</strong>Fig. 17. <strong>Colletotrichum</strong> limetticola (from ex-epitype strain <strong>CBS</strong> 114.14). A–B. Conidiomata. C–K. Conidioph<strong>or</strong>es. L–Q. Appress<strong>or</strong>ia. R–S. Conidia. A, C–G, R. from Anthriscusstem. B, H–Q, S. from SNA. A–B. DM, C–S. DIC, Scale bars: A = 100 µm, F = 10 µm. Scale bar of A applies to A–B. Scale bar of F applies to C–S.slightly) acute <strong>or</strong> both ends acute, (12–)13–18(–24) × (3.5–)4–4.5(–5.5) µm, mean ± SD = 15.5 ± 2.3 × 4.3 ± 0.4 µm, L/W ratio = 3.6.Culture characteristics: Colonies on SNA flat to low convex withentire margin, hyaline, filter paper partly pale salmon to straw, partlycovered with felty white aerial mycelium, reverse hyaline to paleochreous, filter paper partly straw; 18.5–20 mm in 7 d (26–30.5 mmin 10 d). Colonies on OA flat with entire margin; surface moist, whiteto pale luteous, saffron towards the centre due to sp<strong>or</strong>ulation, aerialmycelium lacking, reverse whitish, buff to rosy buff, 18–21.5 mm in7 d (26–29 mm in 10 d). Conidia in mass salmon.Material examined: Cuba, Herradura, inoculation experiment XV in Berkeley,Alameda Co., Calif<strong>or</strong>nia, from twig of Citrus medica var. acida (= Citrus aurantifolia),unknown collection date, (inoculated 30 Jan. 1912, photographed 20 Mar. 1912 byR.E. Clausen), Earle (UC 302386 lectotype [not seen], BPI 394978 isolectotype).USA, Fl<strong>or</strong>ida, from young twig of Citrus aurantifolia, collection date and collect<strong>or</strong>unknown (deposited in <strong>CBS</strong> collection Feb. 1914 by R.E. Clausen as Gloeosp<strong>or</strong>iumlimetticola), (<strong>CBS</strong> H-20910 epitype, here designated, culture ex-epitype <strong>CBS</strong>114.14).Notes: Gloeosp<strong>or</strong>ium limetticola was described by Clausen (1912)following pathogenicity trials in Calif<strong>or</strong>nia on young sour lime (Citrusmedica var. acida = Citrus aurantifolia, Key lime) trees inoculatedwith strains from sour lime from Cuba and with strains from <strong>or</strong>ange,lemon, pomelo and tangerine from Cuba, Calif<strong>or</strong>nia, and Fl<strong>or</strong>ida.The Cuban sour lime strain from Herradura consistently causedwither tip disease symptoms on tester plants from that <strong>species</strong>, andanother Cuban strain (from Santiago de las Vegas) caused broadlysimilar symptoms on both sour lime and lemon (Citrus limon) trees.Clausen stated that a virulent f<strong>or</strong>m of wither tip occurred in Fl<strong>or</strong>idaalso, but this auth<strong>or</strong> was unable to access diseased material tocompare with the Cuban pathogen.Type material of Gm. limetticola was deposited by Clausen inthe dried fungus collections at the University of Calif<strong>or</strong>nia (UC) andWashington DC (BPI). However, its identity (in particular its localgeographical <strong>or</strong>igin, i.e. from Herradura <strong>or</strong> Santiago de las Vegas)was not specified in the <strong>or</strong>iginal paper. The <strong>species</strong> was described[translated from the Latin] as occurring “in young leaves and stemsof Citrus medica var. acida, acting as a pathogen naturally in Cuba,and also artificially inoculated in greenhouses in Calif<strong>or</strong>nia onleaves and stems of C. medica var. acida, C. limetta and C. limon)”.The relevant accession at UC consists of a single packet (UC302386) containing three further packets. One is from Clausen’sExperiment XV and is marked “lime type”; another is from lemon(Experiment XXVII) and Cuban lime material (presumably the<strong>or</strong>iginal diseased sample) and is marked “type material”. Thelemon sample is definitely from a genetic source different from thatof the lime collections, and, it was not marked as type material. Thetwo lime samples may well be genetically identical and could beregarded collectively as the holotype, but on balance we feel thattreating them as two syntypes is m<strong>or</strong>e reasonable. That conclusionwas also reached by Tavares et al. (1997), who designated thecollection from Experiment XV in UC as lectotype of Gm. limetticola.www.studiesinmycology.<strong>or</strong>g77
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