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Damm et al. Fig. 12. Colletotrichum guajavae (from ex-holotype strain IMI 350839). A–B. Conidiomata. C. Seta. D–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, D = 10 µm. Scale bar of A applies to A–B. Scale bar of D applies to C–P. agar and resembled Glomerella. The Gloeosporium stage it links to may well fall within the C. acutatum complex: Gm. psidii, described from Psidium pomiferi (= Psidium guajava) in Mexico, forms ellipsoidal-ovoid conidia measuring 10–16 × 4–6 µm (Saccardo 1906); its conidia are thus broader than those of C. guajavae. Gloeosporium fructus-psidii was found on fruits of Psidium in Sao Paulo, Brazil, and was described as forming oblong, subfusoid to clavate, hyaline conidia, measuring 14–20 × 5–6 µm (Saccardo et al. 1931). The shape of the conidia of that species points also at the C. acutatum complex, however, there is no species in this complex with conidia on average generally wider than 5 µm. Conidia of C. guajavae are substantially smaller, measuring on average 13.4 × 3.5 µm on SNA and 14.6 × 3.8 µm on Anthriscus stem. Peres et al. (2002) isolated C. acutatum (s. lat.) from a guava fruit in Brazil. It caused lesions on guava fruits that were slightly larger than those caused by a C. acutatum (s. lat.) isolate from strawberry. Based on the ITS sequences they generated, the isolates from guava and strawberry from Brazil belong to the same major clade as C. guajavae; the ITS sequence is in fact identical to that of C. guajavae, but also the same as a number of other species in this complex, making an identification to species level impossible without additional information. Based on a phylogeny from combined GAPDH and GS sequences in the study by Guerber et al. (2003), both strains belong to clade D (= clade 2 in this study), but not to the same subclade. The GAPDH sequence generated in Guerber et al. (2003) differs in 5 bp from that of C. guajavae ex-holotype strain IMI 350839. A strain from guava from New Zealand, included in the same study, belongs to clade J3 sensu Guerber et al. (2003) (= C. acutatum s. str.). Apart from C. acutatum s. lat. and C. psidii, Farr & Rossman (2012) list reports from Psidium for C. coccodes in Myanmar, C. gloeosporioides in Brazil, China, Cuba, India, Mexico, Puerto Rico, South Africa, USA, Virgin Islands and Mexico, and Colletotrichum sp. in Brazil, Jamaica and Mexico; it is possible that some of these reports should be referred to C. guajavae. Colletotrichum guajavae can be distinguished from other species of clade 2 of the C. acutatum complex using TUB2, GAPDH and ACT sequences, most effectively with GAPDH. With data from GAPDH alone the species sits within the very variable C. nymphaeae cluster. With TUB2 and ACT there is only 1 bp difference between C. guajavae and C. scovillei, while CHS- 1 and HIS3 sequences are the same as those of C. scovillei. Colletotrichum guajavae is not reliably distinguishable from these species using morphological characteristics. Blastn searches with the GAPDH sequence of strain CBS 853.73 shows 100 % identity with HM038337 from Colletotrichum sp. isolate MFU 09 0624 from Ziziphus mauritiana (jujube) from Laos (Phoulivong et al. 2010), and it is therefore probable that this strain also belongs to C. guajavae. The closest match with the TUB2 sequence of strain CBS 853.73, with 100 % identity, was GU246633 from isolate R14 from Capsicum annuum from South Korea (Sang et al. 2011). We identify that isolate as C. scovillei; the available sequence does not include the region containing the single nucleotide polymorphism that distinguishes TUB2 sequences of C. guajavae and C. scovillei. 70
The Colletotrichum acutatum species complex Fig. 13. Colletotrichum indonesiense (from ex-holotype strain CBS 127551). A–B. Conidiomata. C–G. Conidiophores. H–M. Appressoria. N–O. Conidia. A, C–E, N. from Anthriscus stem. B, F–M, O. from SNA. A–B. DM, C–O. DIC, Scale bars: A = 100 µm, C = 10 µm. Scale bar of A applies to A–B. Scale bar of C applies to C–O. Colletotrichum indonesiense Damm, P.F. Cannon & Crous, sp. nov. MycoBank MB800502. Fig. 13. Etymology: Named after the country of origin, Indonesia. Sexual morph not observed. Asexual morph on SNA. Vegetative hyphae 1–7 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 and branched, to 40 µm long. Conidiogenous cells hyaline, smoothwalled, cylindrical to ± inflated, 8–21 × 2–3.5 µm, opening 1–1.5 µm diam, collarette 1–1.5 µm long, periclinal thickening distinct. Conidia hyaline, smooth-walled, aseptate, straight, cylindrical with one end round and one end acute, (8–)10–14.5(–18) × (2.5–)3.5– 4(–4.5) µm, mean ± SD = 12.3 ± 2.4 × 3.8 ± 0.3 µm, L/W ratio = 3.2. Appressoria single, pale to medium brown, smooth-walled, elliptical, to subglobose in outline, the edge entire, sometimes undulate, 5.5–9(–14.5) × (5–)5.5–7.5(–9) µm, mean ± SD = 7.5 ± 1.8 × 6.3 ± 1.0 µm, L/W ratio = 1.2. Asexual morph on Anthriscus stem. Conidiomata acervular, conidiophores formed on pale brown, angular, basal cells 2.5–6 µm diam. Setae not observed. Conidiophores hyaline to pale brown, smooth-walled, septate, branched, to 60 µm long. Conidiogenous cells hyaline to pale brown, smooth-walled, cylindrical, sometimes ± inflated, 9–25 × 2–4 µm, opening 1–1.5 µm diam, collarette 1–1.5 µm long, periclinal thickening visible. Conidia hyaline, smooth-walled, aseptate, straight, cylindrical to fusiform with both ends acute, (10.5–)13–17.5(–19) × (3–)3.5–4 µm, mean ± SD = 15.4 ± 2.2 × 3.7 ± 0.2 µm, L/W ratio = 4.1. Culture characteristics: Colonies on SNA flat, partly raised with entire margin, hyaline, buff to pale honey, on medium, filter paper and Anthriscus stem partly covered with irregular white aerial mycelium, Anthriscus stem partly covered with orange acervuli, reverse hyaline, white, buff to cinnamon, filter paper partly olivaceous grey; growth rate 18.5–20 mm in 7 d (30–31 mm in 10 d). Colonies on OA flat, partly raised with entire margin; surface covered with irregular floccose to woolly white to olivaceous grey aerial mycelium and few orange acervuli, reverse buff, cinnamon to dark purplish grey; growth rate 22.5–24 mm in 7 d (32.5–34 mm in 10 d). Conidia in mass orange. Material examined: Indonesia: Sumatra, Tele, from leaf spots developing after herbicide treatment of Eucalyptus sp., 1 Jan. 2008, M.J. Wingfield, (CBS H-20798 holotype, culture ex-type CBS 127551 = CPC 14986). Notes: Eucalyptus is not a well-known disease reservoir for Colletotrichum species. Colletotrichum eucalypti was described from Brazil by Bitancourt (1927) and noted again by Viégas (1946) from the Campinas region, causing anthracnose of Eucalyptus leaves. Viégas described the species as having conidia that are elongate-fusiform to oblong and 10–20 × 3–5 µm in size; the www.studiesinmycology.org 71
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