Mycosphere Doi 10.5943/mycosphere/3/3/5 New species and new records of cercosporoid from Cuba and Venezuela (Part 1)

Braun U1* and Urtiaga R2

1Martin-Luther-Universität, Institut für Biologie, Bereich Geobotanik und Botanischer Garten, Herbarium, Neuwerk 21, 06099 Halle (Saale), Germany 2Apartado 546, Barquisimeto, Lara, Venezuela.

Braun U, Urtiaga R 2012 – New species and new records of cercosporoid hyphomycetes from Cuba and Venezuela (Part 1). Mycosphere 3(3), 301–329, Doi 10.5943 /mycosphere/3/3/5

Numerous cercosporoid leaf-spotting hyphomycetes have been continuously collected in Venezuela and several new species and records have been published. Additional specimens, including various collections made between 1966 and 1970 in Cuba and Venezuela, are treated in this paper. The latter material is now housed at K (previously deposited at IMI as “ sp.”). Venezuelan collections made between about 1990 and 2012 (most of them since 2006) are now deposited at HAL. Several species are new to Venezuela, some new host are included, and the following new species and new varieties are introduced: Cercospora hadroanthi, Passalora emmeorhizae, P. melochiae, Pseudocercospora andirae, P. cordiae-alliodorae, P. cordiigena, P. crescentiae, P. gonolobicola, P. jahnii var. amaculata, P. pehriicola, P. rauvolfiae-tetraphyllae, P. trichophila var. punctata, Zasmidium asclepiadis.The new combinations Pseudocercospora trichophila var. solani- asperi and Zasmidium gongronematis are proposed.

Key words – – Cercospora – – Pseudocercospora – South America

Article Information Received 5 April 2012 Accepted 26 April 2012 Published online 13 May 2012 *Corresponding author: U. Braun – e-mail – [email protected]

Introduction analyses, based on increasing data sets and a Cercosporoid fungi are anamorphic asco- broader sampling, indicate that generic mycetes [Ascomycota, Pezizomycotina, Dothi- delimitations within cercosporoid genera are deomycetidae, , Mycosphaerella- not yet fully perceived and far from being ceae (Schoch et al. 2006)] and represent one of finally established. In the present paper we the largest and most diverse groups of hypho- follow generic concepts outlined in Crous & mycetes causing a wide range of diseases of Braun (2003). wild as well as numerous cultivated plants. Cercosporoid hyphomycetes are wide- Most of them were previously assigned to a spread, almost cosmopolitan fungi with an single genus, Cercospora Fresen., which was exceptional diversity in tropical and subtropical later variously split into smaller units (e.g., regions. Venezuela is a tropical country with a Deighton 1967, 1973, 1974, 1976, 1979, Braun great biodiversity of vascular , and 1993). Several of the segregated genera gained accordingly a similarly high diversity of foli- wide acceptance, e.g. Pseudocercospora Speg., icolous fungi. The exploration of this fungal whereas the circumscription and application of group is, however, far from being complete, other genera, e.g. Passalora Fr., remains debat- i.e., the cercosporoid hyphomycetes from able up to now. Recent molecular sequence Venezuela are insufficiently known. A first

301 Mycosphere Doi 10.5943/mycosphere/3/3/5 contribution to the knowledge of cercosporoid Cercospora apii Fresen. s. lat. (C. apii complex, fungi of Venezuela was published by Chupp sensu Crous & Braun 2003) (1934), and descriptions and records from this Material examined – VENEZUELA, La- work were also used for his monograph of ra, Barquisimeto, on leaves of Plumeria rubra Cercospora (Chupp 1954). Pons (1984, 1988, L. (Apocynaceae), Sep. 2010, R. Urtiaga 398 1993, 2004, 2007) and García et al. (1996) (HAL 2491 F); l.c., on leaves of Spondias added further Venezuelan records of cerco- mombin L. (Anacardiaceae), Dec. 2011, R. sporoids and a few new taxa. These data were Urtiaga 457 (HAL 2470 F). also used for the preparation of the annotated Notes – The collection on Plumeria world checklist of Cercospora species rubra is a sparse sample with little fructi- published by Crous & Braun (2003), which is, fication (stromata intraepidermal, brown, 10– therefore, also an important reference list for 50 µm diam., conidiophores fasciculate, 4–7 this fungal group in Venezuela. The second µm wide, conidiogenous loci 2.5–3.5 µm diam., author of the present paper has collected conidia acicular, colourless, 3–4 µm wide). A cercosporoid anamorphs since about 1966. specific Cercospora on Plumeria spp. has not Early collections from Cuba and Venezuela been described. The Cercospora on Plumeria were deposited at IMI as Cercospora sp. rubra is associated with an Asteromella state (recently completely transferred to K). A first (pycnidia 30–75 µm diam., with an irregular set out of these specimens has been sent on terminal porus, conidia bacilliform, 2–4 × 0.8– loan to the first author to be determined and for 1.2 µm). There is no specific Cercospora further treatment. Venezuelan collections made decribed on Spondias and this specimen between about 1990 and 2012 (most of them belongs to the morphological C. apii complex. since 2006) have been directly sent to the first C. verniciferae Chupp & Viégas on Rhus author and are now deposited at HAL. First vernicifera DC. in Brazil coincides morpho- results of examinations of the collections logically with the on Spondias. concerned have already been published by Braun & Urtiaga (2008) and Braun et al. Cercospora beticola Sacc. (2010). Additional results, including new Material examined – VENEZUELA, La- species and new varieties, new records for ra, Barquisimeto, market, on leaves of Beta Venezuela and new host species, are included vulgaris var. cicla L. (Chenopodiaceae), Jan. in the present publication, which will be 2011, R. Urtiaga 430 (HAL 2481 F). continued as numerous additional collections Notes – Known from Venezuela (Crous have not yet been examined. & Braun 2003).

Methods Cercospora brachiata Ellis & Everh. Sporulating structures were mounted in Material examined – VENEZUELA, Mi- distilled water without any staining, and randa, Carabobo, on leaves of Amaranthus examined using oil immersion (bright field and viridis L. (Amaranthaceae), Apr. 2011, R. Urti- phase contrast), with standard light microscopy aga 435 (HAL 2482 F). (Olympus BX 50, Hamburg, Germany). Thirty Known from Venezuela (Crous & Braun measurements ( 1000 magnification) of 2003). conidia and other structures were made, with the extremes given in parentheses. Cercospora coffeicola Berk. & Cooke Material examined – VENEZUELA, La- Results and discussion ra, Sanare, Sabana Redonda Arriba, on leaves New records of cercosporoid hyphomy- of Coffea arabica L. (Rubiaceae), Sep. 2010, R. cetes from Cuba and Venezuela and descrip- Urtiaga 423 (HAL 2495 F). tions of new species and new varieties are Notes – Known from Venezuela (Crous listed in alphabetical order by genus and & Braun 2003). species. Discussion and comments are added to each taxon. Cercospora cyperigena U. Braun & Crous

302 Mycosphere Doi 10.5943/mycosphere/3/3/5 Material examined – VENEZUELA, La- basal hilum 1–3 µm diam., thickened and ra, Barquisimeto, on leaves of Cyperus rotundi- darkened. folius L. (Cyperaceae), Nov. 2010, R. Urtiaga Material examined – CUBA, Bayamo, on 427 (HAL 2472 F); l.c., May 2011, R. Urtiaga leaves of Hadroanthus serratifolius (Vahl) 441 (HAL 2471 F). S.O. Grose [≡ Tabebuia serratifolia (Vahl) G. Notes – This species, hitherto only Nicholson] (Bignoniaceae), 28 Sep. 1967, R. known from the type collection, was described Urtiaga (IMI 129442b = K(M) 173059), holo- from Africa (Tanzania), on Cyperus sp. (Crous type). & Braun 2003: 151). This species is new to Notes – Hadroanthus is not yet known as Venezuela and was found on a new host. The a host genus of Cercospora species, but several current material agrees perfectly with the type species of the closely related genus Tabebuia material. The conidiophores are very short, 5– have been recorded as a host. Tabebuia sp. has 15 × 3–5 µm, 0(–1)-septate, densely fasciculate, been listed as host of C. apii s. lat. (Crous & and emerge through stomata. The conidia are Braun 2003). These authors recommended to long and narrow, 1.5–2.5 µm, and scars and assign Cercospora collections on hosts of new hila are small, 1–2 µm diam. families, genera or species, which are morphologically indistinguishable from C. apii, Cercospora hadroanthi U. Braun & Urtiaga, to C. apii s. lat. (= C. apii complex) since sp. nov. Fig. 1 specialized as well as plurivorous taxa are MycoBank, MB 800018. involved in this complex. In such cases, the Etymology – epithet derived from the can only be elucidated on the base of host genus, Hadroanthus. cultures and molecular sequence analyses. True Cercosporae tabebuiae-impetiginosae si- collections of C. apii s. str. as well as s. lat. are milis, sed laesionibus distinctis, stromatibus characterized by having acicular conidia with distincte minoribus, 10–50 µm diam., conidio- truncate base. However, C. hadroanthi is easily phoris longioribus, ad 200 µm, cicatricibus distinguishable from this complex by having conspicuis et conidiis longioribus, ad 100 µm. conidia with obconically truncate base. In this Leaf spots large, amphigenous, subcircu- respect, C. hadroanthi resembles C. tabebuiae- lar to irregular, up to 25 mm diam., grey to impetiginosae Inácio & Dianese (Inácio & greyish white, with narrow dark brown margin. Dianese 1998), described from Brazil on Caespituli amphigenous, punctiform, dark Hadroanthus impetiginosus (Mart. ex DC.) brown. Mycelium internal; stromata substoma- Mattos ( Tabebuia impetiginosa (Mart. ex tal, 10–50 µm diam., brown, cells 2–6 µm DC.) Standl.), but the latter species is easily diam. Conidiophores in small to usually distinguishable by its very large stromata, up to moderately large, divergent to moderately 107 µm diam., rather short conidiophores, up dense fascicles (5–30), arising from stromata, to 60 µm, in large, dense fascicles, rather in- erect, straight, subcylindrical to moderately conspicuous conidiogenous loci and shorter geniculate, unbranched, 20–90 µm long and conidia. Tabebuia spp. have also been recorded 2.5–4 µm wide, medium brown throughout or as hosts of C. tecomae Viégas & Chupp, tips somewhat paler, wall thin to slightly described on Tecoma sp. from Brazil (Viégas thickened, smooth, pluriseptate; conidiogenous 1945, Chupp 1954, Crous & Braun 2003). The cells integrated, terminal and intercalary, 10– latter species is, however, part of the C. apii 25 µm long, almost straight to distinctly complex with acicular conidia (base truncate) geniculate, with 1–6 conspicuous conidio- and differs additionally in having broader genous loci (scars), circular in front view, conidiophores (4–6 µm) and much shorter brown, 1.5–3.5 µm diam., thickened and conidia (usually 25–50 µm long). darkened. Conidia formed singly, long conidia acicular to obclavate-oblong, short conidia Cercospora lactucae-sativae Sawada fusiform-subcylindrical, 20–90 × 2.5–4 µm, Material examined – VENEZUELA, La- usually 3–8-septate, colourless to faintly ra, Barquisimeto, on leaves of Lactuca sativ L. greenish, smooth, apex acute or subacute, base (Asteraceae), Jan. 2011, R. Urtiaga 433 (HAL truncate to usually short obconically truncate, 2480 F).

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Fig. 1 – Cercospora hadroanthi. Based on type material. a Conidiophore fascicle. b Conidiophores. c Conidia. – Bar = 10 µm.

Notes – Known from Venezuela (Crous (Solanaceae), Jun. 2000, R. Urtiaga (HAL & Braun 2003). 2468 F). Notes – New to Venezuela (not listed in Cercospora mikaniicola F. Stevens Crous & Braun 2003). Material examined – VENEZUELA, La- ra, Sanare, Sabana Redonda Arriba, on leaves Passalora emmeorhizae U. Braun & Urtiaga, of Mikania sp. (Asteraceae), Sep. 2010, R. sp. nov. Fig. 2 Urtiaga 402 (HAL 2493 F). MycoBank, MB 800006. Notes – New to Venezuela (not listed in Etymology – epithet derived from the Crous & Braun 2003). host genus, Emmeorhiza. Differt a Passalora cephalanthi et a P. Passalora capsicicola (Vassiljevsky) U. Braun oldenlandiae conidiis valde brevioribus et & F. Freire latioribus.  Cercospora capsicicola Vassiljevsky. Leaf spots almost indistinct, or diffuse to  Phaeoramularia capsicicola (Vassil- subcircular-irregular, yellowish, olivaceous to jevsky) Deighton. brownish, about 2–8 mm diam., margin Material examined – VENEZUELA, La- indefinite. Caespituli amphigenous, but mainly ra, Barquisimeto, El Trompillo, on leaves of hypophyllous, punctiform to subeffuse, oli- Capsicum annuum L. [= C. frutescens L.] vaceous-brown to medium brown. Mycelium

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Fig. 2 – Passalora emmeorhizae. Based on type material. a Conidiophore fascicles. b Conidiophores. c Conidia. – Bar = 10 µm. internal; stromata lacking or only with small of Emmeorhiza umbellata (Spreng.) K. Schum. substomatal hyphal aggregations, 10–30 µm (Rubiaceae), May 2009, R. Urtiaga (HAL 2465 diam., brown. Conidiophores in small, loose to F, holotype). moderately large and dense fascicles, arising Notes – Passalora emmeorhizae is char- from stromata, emerging through stomata, acterized by lacking superficial mycelium, erect, straight, subcylindrical-conical to dis- fasciculate conidiophores and catenate conidia, tinctly geniculate-sinuous, unbranched, 10–60 i.e. it belongs to a group of species previously × 3–7 µm, 0–3-septate, pale olivaceous to assigned to the genus Phaeoramularia Munt.- olivaceous-brown, medium olivaceous-brown Cvetk., which is currently considered a syno- in mass, thin-walled, smooth; conidiogenous nym of Passalora. Species of the latter genus cells integrated, terminal or conidiophores on Emmeorhiza are unknown, but three com- reduced to conidiogenous cells, 10–25 µm parable Phaeoramularia-like species have been long, conidiogenous loci (scars) conspicuous, described from other hosts of the Rubiaceae, somewhat thickened and darkened, 1.5–2 µm viz. P. cephalanthi (Ellis & Kellerm.) U. Braun diam. Conidia in chains, ellipsoid-ovoid to & Crous on Cephalanthus in North America cylindrical, 15–50 × 4–7 µm, 0–4-septate, sub- (Chupp 1954, Crous & Braun 2003), and P. hyaline pale olivaceous to olivaceous-brown, oldenlandiae (Hansf.) U. Braun on Borreria thin-walled, smooth, apex rounded to obco- and Oldenlandia in Africa (Chupp 1954, Ellis nically truncate, base obconically truncate, hila 1976, Crous & Braun 2003), which are mor- 1.5–2 µm wide, slightly thickened and some- phologically quite distinct by their very long what darkened. and narrow, pluriseptate conidia (10–100 × 2– Material examined – VENEZUELA, La- 4.5 µm in P. cephalanthi and 22–90 × 3–5 µm ra, Sanare, Sabana Redonda Arriba, on leaves in P. oldenlandiae), as well as P. pseudo-

305 Mycosphere Doi 10.5943/mycosphere/3/3/5 capnoides O.L. Pereira & R.W. Barreto on terized by forming catenate conidia, i.e. this Mitracarpus hirtus (L.) DC. in Brazil, which species belongs to a group of Passalora species differs in having much narrower conidia (2–4 previously assigned to Phaeoramularia. P. µm wide, Pereira & Barreto 2005). meridiana (Chupp) U. Braun & Crous on Helicteres spp. (Chupp 1954, Crous & Braun Passalora melochiae U. Braun & Urtiaga, sp. 2003) is the only comparable species on a host nov. Fig. 3 of the Sterculioideae (= Sterculiaceae), but MycoBank, MB 800007. differs in having large, dense, coremoid fas- Etymology – epithet derived from the cicles, much longer conidiophores, up to at host genus, Melochia. least 120 µm, and much longer and narrower Passalorae meridianae similis, sed fasci- conidia, up to 125 × 2.5–4 µm with up to 11 culis conidiophorum minoribus, laxioribus, septa. P. helicteris-viscidae Phengsintham, conidiophoris brevioribus, ad 50 µm, conidiis Chukeatirole, K.D. Hyde & U. Braun (Pheng- brevioribus et latioribus, ad 70 × 6 µm, saepe sintham et al. 2010) is also characterized by 2–7-septatis. catenate conidia, which are, however, quite Leaf spots amphigenous, subcircular to distinct by being short and narrow, 8–44 × 1–3 somewhat irregular, 1–4 mm diam., centre µm, 0–4-septate and hyaline. Other species are ochraceous to greyish white, margin narrow, Mycovellosiella-like, i.e. with superficial hy- purple-red to brown, sometimes with diffuse phae and solitary conidiophores [Passalora purple halo. Caespituli amphigenous, puncti- sterculiacearum U. Braun & Crous (Braun & form, dark brown to blackish, scattered. Myce- Crous 2007), P. dombayae (Crous & U. Braun) lium internal; epiphyllous stromata intraepi- Crous & U. Braun (Crous & Braun 2003)], or dermal, hypophyllous stromata substomatal, the conidia are consistently formed singly [P. 10–40 µm diam., subcircular, elliptical-oval to helicteris (Soni, Dadwal & Jamaluddin) Poo- somewhat irregular in outline, medium brown, nam Srivast. (Soni et al. 1984)]. cells 2–6 µm diam. Conidiophores in small to moderately large fascicles, arising from stro- Pseudocercospora andirae U. Braun & Urti- mata, erumpent or emerging through stomata, aga, sp. nov. Fig. 4 erect, subcylindrical-conical, straight to MycoBank, MB 800008. slightly geniculate-sinuous, unbranched, 10–50 Etymology – epithet derived from the × 2.5–6 µm, 0–2-septate, thin-walled, smooth, host genus, Andira. sub-hyaline, pale olivaceous, light brown, Pseudocercosporae vataireae similis, sed medium olivaceous-brown in mass; stromatibus minoribus, 10–50 µm diam., coni- conidiogenous cells integrated, terminal or diophoris laevibus, saepe geniculatis-sinuosis. conidiophores reduced to conidiogenous cells, Differt a P. stevensii ad species Andirae laesi- 10–35 µm long, coni-diogenous loci (scars) onibus plene distinctis, hyphis superficialibus conspicuous, circular, 1–2 µm diam., slightly cum conidiophoris solitariis formantibus et thickened and darkened. Conidia formed singly conidiophoris valde brevioribus, 10–70(–80) or in short chains, cylindrical or subcylindrical, µm. 20–70 × 3–5.5 µm, (0–)2–7(–8)-septate, Leaf spots amphigenous, conspicuous, subhyaline, pale oli-vaceous, olivaceous to subcircular to somewhat angular-irregular, 2– light brown in mass, apex obtuse to truncate in 10 mm diam. or confluent and larger, occa- catenate conidia, base short obconically sionally somewhat zonate, brown, ranging truncate, hila 1–2 µm wide, somewhat from pale to medium dark brown, later grayish thickened and darkened. brown to dingy grey or greyish white, margin Material examined – CUBA, Bayamo, on indefinite or with a narrow darker marginal leaves of Melochia pyramidata L. (Malvaceae, line, often with narrow to broad, brown, Sterculioideae), 8 May 1967, R. Urtiaga (IMI necrotic halo. Caespituli amphigenous, puncti- 127445 = K(M) 173070, holotype); Bayamo, form to confluent and dense, dark brown to on leaves of M. pyramidata, 29 May 1967, R. blackish. Mycelium internal and external; Urtiaga (IMI 126779 = K(M) 173071). superficial hyphae only on the lower leaf Notes – Passalora melochiae is charac- surface, branched, straight to sinuous, 1–4 µm

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Fig. 3 – Passalora melochiae. Based on type material. a Conidiophore fascicles. b Coni-diophores. c – conidia. – Bar = 10 µm. wide, septate, subhyaline to pale olivaceous or olivaceous-brown, thin-walled, smooth to olivaceous-brown, thin-walled, smooth; stro- faintly rough-walled, apex subobtuse, base mata substomatal or intraepidermal, subcircular short obconically truncate, 1–2 µm wide, hilum to elongated or somewhat irregular in outline, neither thickened nor darkened. medium to dark olivaceous-brown, hypo- Material examined – VENEZUELA, La- phyllous stromata 10–50 µm diam., epi- ra, Barquisimeto, Bosque Macuto, on leaves of phyllous ones up to 80 µm diam., cells 2.5–7 Andira surinamensis (Bondt) Splitg. ex Pulle µm diam. Conidiophores in small to mode- (Fabaceae), Mar. 2010, R. Urtiaga (HAL 2466 rately large, loose to dense fascicles, arising F, holotype); Barquisimeto, Zoo, on leaves of from stromata, through stomata or solitary, Andira inermis (W. Wright) Kunth ex DC. arising from superficial hyphae, lateral, rarely (Fabaceae), Jan. 2011, R. Urtiaga 432 (HAL single conidiophores emerging through stoma- 2479 F, paratype). ta, erect, straight, subcylindrical-conical to Notes – Pseudocercospora stevensii (E. geniculate-sinuous, simple or occasionally Yong) U. Braun & Crous (Chupp 1954, Crous branched, 10–70(–80) × 3–5 µm, 0–5-septate, & Braun 2003) is known from South America pale to medium olivaceous or olivaceous- on Andira spp., including A. surinamensis. brown, thin-walled, smooth or almost so; Type material of this species has been conidiogenous cells integrated, terminal or examined (on Andira sp., Puerto Rico, Dos conidiophores reduced to conidiogenous cells, Bocas, below Utuado, 30 Dec. 1913, F.L. 10–25 µm long, conidiogenous loci incon- Stevens, ILL 6008 and PC). This species is, spicuous to denticle-like, but always un- however, quite distinct from P. andirae (leaf thickened and not darkened. Conidia formed spots lacking or only with indistinct small singly, obclavate-subcylindrical, short conidia reddish brown spots, mycelium internal, occasionally fusiform, (15–)25–80 × (2.5–)3– superficial hyphae with solitary conidiophores 5(–6) µm, (1–)3–8-septate, pale olivaceous to not formed and conidiophores very long, about

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Fig. 4 – Pseudocercospora andirae. Based on type material. a Superficial hyphae with solitary conidiophores. b Conidiophore fascicles. c Conidiophores emerging through a stoma. d Coni- diophores. e Conidia. f Germinating conidia – Bar = 10 µm.

50–200 µm, mostly in dense, often almost carpus spp. in Asia and Oceania (Yen & Lim coremioid fascicles). In phylogenetic analyses, 1980, Braun et al. 1999, Crous & Braun 2003), the genus Andira clustered in a basal un- superficial hyphae and solitary conidiophores resolved position within the aeschynomenoid are lacking. In addition, the conidia in P. group (Dalbergieae and Aeschynomeneae, see dalbergiae and P. pongamiae-pinnatae are Doyle et al. 2000). Among Pseudocercospora much narrower, 2–3 µm, and in P. ptero- spp. on hosts of closely as well as distantly carpicola they are much wider, (4–)5–8(–10) allied genera of the Dalbergieae and Aeschy- µm. The South American P. lonchocarpi (J.A. nomeneae, there are only few comparable Stev.) Crous & M.P.S Câmara (Crous & species. P. vataireae (Henn.) U. Braun & Câmara 1998) forms superficial mycelium, but Freire (Braun & Freire 2002) on Derris spp. in without any solitary conidiophores. Further- Brazil is morphologically similar by forming more, the conidiogenous cells are often solitary conidiophores arising from superficial percurrent and the verruculose conidia of this hyphae, but this species forms much larger species are much longer and narrower, (30–) stromata, up to 150 µm diam., and the 50–100(–120) × (2–)3–3.5(–4.5) µm (Chupp conidiophores are usually subcylindrical and 1954, Crous & Câmara 1998). straight, i.e. non-geniculate, verruculose to- wards the tip and often percurrently proli- Pseudocercospora atromarginalis (G.F. Atk.) ferating. In other species, viz. P. dalbergiae Deighton (S.H. Sun) J.M. Yen on Dalbergia spp. in Asia  Cercospora atromarginalis G.F. Atk. (Hsieh & Goh 1990, Guo et al. 1995), P. Material examined – VENEZUELA, La- pongamiae-pinnatae Raghu Ram & Mallaiah ra, Sanare, Sabana Redonda Arriba, on leaves on Pongamia sp. in India (Raghu Ram & of Solanum nigrum L. (Solanaceae), Jun. 2010, Mallaiah 1993) and P. pterocarpicola (J.M. R. Urtiaga 389 (HAL 2497 F). Yen) J.M. Yen var. pterocarpicola and var. Notes – New to Venezuela (not listed in guzmanii (Tak. Kobay.) U. Braun on Ptero- Crous & Braun 2003). 308 Mycosphere Doi 10.5943/mycosphere/3/3/5 Pseudocercospora cordiae-alliodorae U. septatis. Braun & Urtiaga, sp. nov. Fig. 5 Leaf spots amphigenous, angular-irre- MycoBank, MB 800009. gular, often vein-limited, darker brown on the Etymology – epithet derived from the upper leaf surface, paler brown below, 1–8 mm host species, Cordia alliodora. diam. or confluent and larger, margin not Pseudocercosporae cordiicolae similis, differentiated, but often with yellowish to sed conidiis valde brevioribus et latioribus, 12– ochraceous halo. Caespituli amphigenous, usu- 40 × 2–4 µm, modo 1–4-septatis. ally epiphyllous, rather inconspicuous. Myce- Leaf spots lacking or only with diffuse lium internal; stromata small to well de- greyish brown discolorations, rather incon- veloped, 15–45 µm, intraepidermal on the spicuous. Colonies hypophyllous, effuse, oli- upper side, hypophyllous stromata also sub- vaceous-brown, but rather inconspicuous. stomatal, subcircular to somewhat irregular in Mycelium internal and external, hyphae super- outline, medium to dark olivaceous-brown, ficial, emerging through stomata, also climbing cells circular to angular in outline, 3–8 µm leaf hairs, branched, sometimes anastomosing, diam. Conidiophores in small to moderately 1–4 µm wide, septate, subhyaline to pale large fascicles, arising from stromata, erect, olivaceous or olivaceous-brown, thin-walled, straight, unbranched to somewhat curved or smooth; stromata lacking. Conidiophores soli- slightly geniculate-sinuous, almost cylindrical, tary, arising from superficial hyphae, lateral, somewhat attenuated towards the tip or sub- erect, straight, subcylindrical to conical or clavate, 8–25 × 3–6 µm, up to 55 µm long with somewhat geniculate or curved-sinuous, un- still attached young conidia, pale to medium branched, 4–25 × 2–4 µm, 0(–1)-septate, sub- olivaceous-brown throughout or paler towards hyaline, pale olivaceous to olivaceous-brown, the tip, subhyaline to very pale olivaceous, thin-walled, smooth; conidiophores usually young conidiophores sometimes subhyaline, reduced to conidiogenous cells, conidiogenous wall thin to slightly thickened (up to 0.75 µm), loci inconspicuous. Conidia formed singly, smooth; conidiogenous cells integrated, termi- obclavate-cylindrical, short conidia sometimes nal or conidiophores one-celled, i.e. reduced to ellipsoid-fusiform, 12–40 × 2–4 µm, 1–4- conidiogenous cells, 8–20 µm long, conidio- septate, subhyaline to pale olivaceous or genous loci inconspicuous. Conidia formed olivaceous-brown, thin-walled, smooth, apex singly, obclavate, young conidia subcylin- obtuse to subacute, base short obconically drical, broadly fusiform or subclavate, often truncate, 1–1.5(–2) µm wide, hila unthickened, long attached at conidiogenous cells, 25–70(– not darkened. 90) × 4–8 µm, 0–7-septate, pale to medium Material examined – VENEZUELA, La- olivaceous or olivaceous-brown, wall up to ra, Barquisimeto, zoological garden, on leaves 0.75 µm wide, smooth, apex obtuse, rounded, of Cordia alliodora (Ruiz & Pav.) Oken base short obconically truncate, 2–3.5 µm (Boraginaceae), May 2011, R. Urtiaga (HAL wide, hila neither thickened nor darkened. 2464 F, holotype). Material examined – CUBA, Bayamo, on Notes – The Asian Pseudocercospora leaves of Cordia dentata Poir. (Boraginaceae), cordiicola (J.M. Yen) J.M. Yen (Yen & Lim 21 Jan. 1967, R. Urtiaga (IMI 124810 = K(M) 1980) is a comparable species, but differs from 173055, holotype). the new South American P. cordiae-alliodorae Notes – Crous et al. (2000) described in having much longer and narrower conidia, Pseudocercospora cordiana Crous & Bench. about 80–125 × 1.5–2.5 µm, with 4–8 septa. from Brazil on Cordia goeldiana Huber. This species is superficially similar, but has Pseudocercospora cordiigena U. Braun & narrower conidiophores, only 2.5–4 µm wide, Urtiaga, sp. nov. Fig. 6 and narrower conidia, (30–)40–46 µm long and MycoBank, MB 800010. only 2–3 µm wide and 1–3(–5)-septate. Two Etymology – epithet derived from the other Pseudocercospora species described on host genus, Cordia. Pseudocercosporae Cordia spp. are morphologically distinct. P. cordianae similis, sed conidiophoris latioribus cordiae Kamal & R.P. Singh (Kamal & Singh (3–6 µm) et conidiis latioribus, 4–8 µm, 0–7- 1980) differs in having longer, pluriseptate

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Fig. 5 – Pseudocercospora cordiae-alliodorae. Based on type material. a Superficial hyphae. b Hyphae emerging through a stoma. c Superficial hyphae with solitary conidiophores. d Conidiophores. e Conidia – Bar = 10 µm. conidiophores, 54–90 µm, and smaller caespitulis epiphyllis et hypophyllis distincte conidia,21.5–30 × 3.5–4.5 µm, and P. dimorphis, conidiophoris interdum ramosis et cordiicola (J.M. Yen) J. M. Yen (Yen & Lim conidiis ad basim breviter obconice truncatis. 1980) is characterized by lacking stromata, Leaf spots amphigenous, subcircular to superficial hyphae with solitary conidiophores angular-irregular, 1–10 mm diam., rarely and very long and narrow conidia, 80–125 × larger, brown, greyish brown, later becoming 1.5–2.5 µm. very pale, grey-brown to greyish white, with narrow brown margin. Caespituli amphi- Pseudocercospora costi (F. Stevens) U. Braun ogenous, on the upper leaf surface con- & Crous spicuously punctiform, dark brown to blackish,  Cercospora costi F. Stevens. on the lower side punctiform to subeffuse. Material examined – VENEZUELA, La- Mycelium internal; epiphyllous stromata large, ra, Sanare, Sabana Redonda Arriba, on leaves 30–70 µm diam., intraepidermal, dark olivace- of Costus sp. (Zingiberaceae), Sep. 2010, R. ous-brown to brown, cells rounded to angular Urtiaga 424 (HAL 2489 F). in outline, 2–7 µm diam., stromata on the lower Notes – This species was described from side lacking or smaller, 10–50 µm diam., Panama and is known from Venezuela on mostly substomatal. Conidiophores fasciculate, Costus sp. (Crous & Braun 2003). fascicles dimorphic, on the upper leaf surface always in large, dense, erumpent sporodochial Pseudocercospora crescentiae U. Braun & conidiomata, straight to slightly geniculate- Urtiaga, sp. nov. Fig. 7 sinuous, unbranched, subcylindrical-conical, 5– MycoBank, MB 800011. 25 × 2–4.5 µm, 0–1-septate, on the lower side Etymology – epithet derived from the in smaller, mostly loose fascicles, usually host genus, Crescentia. Pseudocercosporae emerging through stomata, erect to decumbent, tabebuiae-roseoalbae valde similis, sed almost straight, cylindrical to mostly distinctly 310 Mycosphere Doi 10.5943/mycosphere/3/3/5

Fig. 6 – Pseudocercospora cordiigena. Based on type material. a Conidiophore fascicles. b Conidiophores. c Conidia. – Bar = 10 µm. geniculate-sinuous, simple or often branched, K(M) 173062). 10–60 × 2.5–6 µm, 0–4-septate, pale to some- Notes – Many Pseudocercospora species what darker olivaceous-brown, thin-walled, have been described on hosts belonging to the smooth or almost so; conidiogenous cells Bignoniaceae. P. crescentiae differs from all integrated, terminal or occasionally intercalary, known species in forming distinctly dimorphic 10–30 µm long, conidiogenous loci (scars) caespituli, with obvious differences between inconspicuous. Conidia solitary, obclavate- epiphyllous and hypophyllous conidiophore subcylindrical, short conidia sometimes cylin- fascicles. On the upper leaf surface, the coni- drical-fusiform, 12–60 × (1.5–)2–4(–4.5 µm), diophores form sporodochial conidiomata with 1–5-septate, subhyaline to pale olivaceous, large stromata. Numerous Pseudocercospora thin-walled, smooth, apex obtuse to subacute, species on hosts of the Bignoniaceae are quite base short obconically truncate, in short co- distinct from P. crescentiae by lacking or nidia sometimes truncate, hila 1–2 µm diam., possessing very small stromata [viz., P. neither thickened nor darkened. arrabidaeae R. Kirschner (Kirschner & Material examined – CUBA, Media Lu- Piepenbring 2006), P. bignoniacearum B.K. na, on leaves of Crescentia cujete L. (Bigno- Gupta & Kamal (Gupta & Kamal 1987), P. niaceae), 12 Jul. 1967, R. Urtiaga 799 (IMI brasiliensis U. Braun & F.O. Freire (Braun & 129036 = K(M) 173063); Bayamo, on leaves Freire 2004), P. dolichandrones (Chupp) of Crescentia cujete, 18 Mar. 1968, R. Urtiaga Deighton (Chupp 1954), P. hansfordii (Chupp) 1210 (IMI 132556 = K(M) 173060). VENE- Deighton (Chupp 1954), P. millingtoniae ZUELA, Sucre State, Cumanacoa, on leaves of Raghu Ram & Mallaiah (Raghu Ram & Crescentia cujete, 15 Jan. 1971, R. Urtiaga Mallaiah 1996), P. oroxyligena J.M. Yen, A.K. 1345 (IMI 156326 = K(M) 173061, holotype); Kar & B.K. Das (Yen et al. 1982a), P. without locality, on leaves of Crescentia cujete, pandoreae U. Braun & C.F. Hill (Braun et al. 23 Jun. 1970, R. Urtiaga 1237 (IMI 149973 = 2006), P. sordida (Sacc.) Deighton (Chupp

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Fig. 7 – Pseudocercospora crescentiae. Based on type material. a Conidiophore fascicles. b Conidiophores. c Conidia. – Bar = 10 µm.

1954), P. tecomae-heterophyllae (J.M. Yen) superficial hyphae with solitary conidiophores Y.L. Guo & X.J. Liu (Guo & Hsieh 1995)]. developed; Dianese et al. 1999). Among species with well-developed stromata, Pseudocercospora crescentiae is close to P. Pseudocercospora cruenta (Sacc.) Deighton jahnii (Syd.) U. Braun & Crous (Chupp 1954,  Cercospora cruenta Sacc. Material Crous & Braun 2003) and P. tabebuiae- examined – VENEZUELA, La-ra, roseoalbae Inácio & Dianese (Inácio & Barquisimeto, on leaves of Vigna ungu-iculata Dianese 1998). However, the latter two species (Fabaceae), Sep. 2010, R. Urtiaga 422 (HAL do not form comparable dimorphic coni- 2493 F). diophore fascicles. The unbranched coni- Notes – Known from Venezuela (Crous diophores always arise from stromata. Other & Braun 2003). This species is widespread and species with larger stromata are easily common on various legumes. distinguishable: P. catalpigena U. Braun & Crous (conidia cylindrical; Braun et al. 2003), Pseudocercospora durantae N. Pons, U. Braun P. catalpicola U. Braun (superficial hyphae & Crous with solitary conidiophores developed; Braun 1999), P. stereospermicola Srisk. & Sivan.  Cercospora durantae Chupp & A.S. (longer conidia, 50–110 µm, with up to 10 Mull., nom. inval. septa; Sriskantha & Sivanesan 1980), P. Material examined – VENEZUELA, La- tabebuiae-caraibae Inácio & Dianese (with ra, Barquisimeto, on leaves of erecta large lesions, up to 30 µm diam., conidiophores L. [= D. repens L.] (), Jul. 2011, 4–6 µm wide, conidia up to 100 µm long, with R. Urtiaga 449 (HAL 2477 F). up to eight septa; Inácio & Dianese 2006), P. Notes – Pons et al. (in Crous & Braun zeyheriae (Henn.) Dianese, Furlanetto & L.T.P. 2003: 168) validated Cercospora durantae. Santos (stromata large, 60–240 µm diam., Type material of this species is from Venezuela conidia up to 100 µm long, with up to 13 septa, on Duranta mutisii. D. erecta is known as host

312 Mycosphere Doi 10.5943/mycosphere/3/3/5 of this species from Florida, USA. This is the without locality, on leaves of Gonolobus first record on the latter host from Venezuela. rostratus (Vahl) R. Br. ex Schult. (Asclepia- Morphological characters of the present daceae), 14 Mar. 1969, R. Urtiaga 247 (IMI material agrees perfectly with the type of this 139317 = K(M) 173064, holotype). species. Notes – Cercospora gonolobi W.W. Ray has been described from North America (USA, Pseudocercospora gonolobicola U. Braun & Oklahoma) on Gonolobus laevis Michx. ( Urtiaga, sp. nov. Fig. 8 Cynanchum leave (Michx.) Pers.). MycoBank, MB 800012. Etymology – epithet derived from the The generic affinity of this species is still host genus, Gonolobus. unresolved, but due to pigmented conidia (Chupp 1954) this species undoubtedly does Differt a Cercospora gonolobi hyphis not belong to Cercospora s. str. In any case, C. superficialibus cum conidiophoris solitariis, gonolobi is morphologically quite distinct by conidiophoris valde brevioribus, 3–25 µm, 0– lacking superficial mycelium and by its much 1(–2)-septatis et conidiis angustioribus, 2–4 longer, pluriseptate conidia (up to 80 µm) µm. which are also wider (4–5 µm). There is no Lesions diffuse to angular-irregular, up to comparable species among numerous 20 mm diam., sometimes vein-limited, yellow- Pseudocercospora species described from ish, olivaceous to brownish, margin indefinite. hosts of the Asclepiadaceae. Guo & Hsieh Colonies amphigenous, epiphyllous caespituli (1995) recorded, described and illustrated P. punctiform, dark brown, scattered, hypo- marsdeniae (Hansf.) Deighton from China on phyllous colonies effuse to aggregated, dingy Dregea sinensis Hemsl., characterized by greyish to olivaceous-brown. Mycelium in- having fasciculate conidiophores arising from ternal and external, superficial hyphae mainly stromata as well as solitary conidiophores hypophyllous, emerging through stomata, arising from superficial hyphae, but sometimes forming ropes, sparingly branched, conidiophores and conidia are much longer and 1.5–4 µm wide, subhyaline to pale olivaceous, above all broader (3–6.5 µm) than those of P. later olivaceous-brown, septate, thin-walled, gonolobicola. However, the Chinese collection smooth; stromata mainly epiphyllous, intra- on Dregea sinensis is undoubtedly not epidermal, 15–50 µm diam., dark brown, conspecific with the African P. marsdeniae, composed of medium brown cells, subcircular which does not form any superficial hyphae to somewhat angular-irregular in outline, 2.5–7 (Chupp 1954, Deighton 1976), and probably µm diam. Conidiophores in small to mode- represents a separate undescribed species. rately large fascicles, arising from stromata, erumpent, or conidiophores solitary, arising Pseudocercospora jahnii (Syd.) U. Braun & from superficial hyphae, lateral, erect, straight, Crous var. jahnii Fig. 9 subcylindrical-conical to moderately genicu- late-sinuous, unbranched, 3–25 × 2–5 µm, 0–  Cercospora jahnii Syd. 1(–2)-septate, subhyaline to pale olivaceous or = Cercoseptoria tabebuiicola Kamal, olivaceous-brown, thin-walled, smooth; coni- Narayan & R.P. Verma. diogenous cells integrated, terminal or conidio- = Pseudocercospora tabebuiae-roseo- phores often aseptate, i.e. reduced to coni- albae Inácio & Dianese. diogenous cells, 2–20 µm long, conidiogenous Leaf spots amphigenous, subcircular to loci inconspicuous. Conidia formed singly, angular-irregular, 2–15 mm diam., pale or dull narrowly obclavate-cylindrical, short conidia brown, greyish brown, finally greyish white, sometimes ellipsoid-fusiform, 15–70 × 2–4 with darker margin, brown to reddish brown. µm, (1–)2–6(–7)-septate, subhyaline to pale Caespituli hypophyllous, rarely amphigenous, olivaceous, thin-walled, smooth, apex subacute delicately to distinctly punctiform, scattered to or subobtuse, base short obconically truncate, aggregated, blackish brown, greyish black. (1–)1.5–2 µm wide, hila neither thickened nor Mycelium internal and external; superficial darkened. hyphae branched, septate, thin-walled, smooth, Material examined – VENEZUELA, 1.5–3 µm wide, pale olivaceous to olivaceous-

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Fig. 8 – Pseudocercospora gonolobicola. Based on type material. a Conidiophore fascicles. b Superficial hyphae with solitary conidiophores. c Conidiophores. d Conidia. – Bar = 10 µm. brown; stromata lacking or small to moderately serratifolia (Vahl) G. Nicholson] (Bignoni- large, 10–50 µm diam., substomatal or intra- aceae), 10 Oct. 2002, F. Freire [U. Braun, epidermal, brown, subglobose to irregularly Fungi sel. exs. 18] (HAL). INDIA, UP, North shaped, cells subcircular to angular in outline, Gorakhpur forest Division, on leaves of 2–6 µm diam. Conidiophores in small, loose to Tabebuia rosea (Bertol.) DC. (Bignoniaceae), moderately large, dense fascicles, arising from Nov. 1980, R.P. Verma (IMI 257246), isotype internal hyphae or stromata, emerging through of Cercoseptoria tabebuiicola. VENEZELA, stomata or erumpent through the cuticle, erect, without locality, on leaves of Spathodea occasionally decumbent, almost straight, sub- campanulata P. Beauv. (Bignoniaceae), 14 cylindrical to strongly geniculate-sinuous, Mar. 1969, R. Urtiaga 229 (IMI 139309 = usually simple or branched, 5–40(–60) × 2–5 K(M) 179309); Aragua, La Victoria, on leaves µm, 0–3-septate, pale olivaceous to medium of Tabebuia rosea (Bertol.) DC. (Bignoni- olivaceous-brown, thin-walled, smooth; coni- aceae), 4 Feb. 1928, H. Sydow (BPI 437408, diogenous cells integrated, usually terminal, 4– lectotype of Cercospora jahnii, designated 30 µm long, conidiogenous loci (scars) incon- here). spicuous or visible as truncate tips, but neither Notes – Pseudocercospora jahnii occurs thickened nor darkened. Conidia solitary, on several species of Tabebuia and related obclavate-cylindrical, (10–)15–65 × 2.5–5 µm, genera and is rather widespread in Central and 1–7(–8)-septate, pale olivaceous to olivace- South America, and also known from Asia ous-brown, thin-walled, smooth, apex obtuse or (India). P. tabebuiae-roseoalbae (Inácio & only slightly pointed, base obconically trun- Dianese 1998: 703) and Cercoseptoria tabe- cate, hila 1–2 µm wide, unthickened, not buiicola (Kamal et al. 1986: 456) are darkened. morphologically not clearly distinguishhable Material examined – BRAZIL, State of from P. jahnii. Collections on Hadroanthus Ceará, Ubajara City, on leaves of Hadroanthus serratifolius and Spathodea campanulata are serratifolius (Vahl) S.O. Grose [≡ Tabebuia also morphologically indistinguishable from P.

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Fig. 9 – Pseudocercospora jahnii var. jahnii on Spathodea campanulata. Based on K(M) 179309. a Conidiophore fascicles. b Superficial hyphae with solitary conidiophores. c Conidiophores. d Conidia. – Bar = 10 µm. jahnii. Due to its wide host range and Pseudocercospora marcelliana (Chupp) U. distribution, it is possible that this species Braun & Crous Fig. 11 represents a complex of closely allied, morpho-  Cercospora marcelliana Chupp. logically barely distinguishable cryptic species. Material examined – VENEZUELA, La- However, this problem can only be solved on ra, Sanare, Sabana Redonda Arriba, on leaves the base of cultures and molecular sequence of Solanum torvum L. s. lat. [probably var. analyses as well as inoculation experiment. hartwegianum Sendtner, = var. ochraceo-ferru- gineum Dunal, S. rudepannum Dunal] (Sola- Pseudocercospora jahnii var. amaculata U. naceae), Sep. 2010, R. Urtiaga 413 (HAL 2486 Braun & Urtiaga, var. nov. Fig. 10 F). MycoBank, MB 800019. Notes – This species, described from Etymology – derived from the lack of Venezuela on Solanum nudum Dunal [= S. leaf spots. micranthum Willd. ex Roem. & Schult. (Chupp Differt a var. jahnii maculis foliorum 1954)], is Cercoseptoria-like, i.e. with sporo- nullis. dochial conidiomata, numerous densely ar- Distinct from var. jahnii by lacking leaf ranged very short conidiophores, and hyaline spots. or very pale, narrowly cylindrical-filiform to Material examined – VENEZUELA, acicular, pluriseptate conidia. In the present without locality, on leaves of Tabebuia shaferi collection, the conidiophores are 5–25 × 1.5– Britton (Bignoniaceae), 14 Mar. 1969, R. Urti- 3.5 µm, and the conidia are 30–110 × 2–3.5 aga 230 (IMI 139210 = K(M) 173057, holo- µm, 3–11-septate. S. torvum is a new host for type). this species. Notes – Conidiophores and conidia in the collections on Tabebuia shaferi agree well Pseudocercospora mikaniigena J.M. Yen & with Pseudocercospora jahnii, but typical Lim Fig. 12 collections of this species form distinct leaf  Asperisporium mikaniigena (J.M. Yen spots. We prefer to introduce a variety for & Lim) R.W. Barreto collections without distinct lesions. Leaf spots amphigenous, subcircular to

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Fig. 10 – Pseudocercospora jahnii var. amaculata. Based on type material. a Superficial hyphae. b Superficial hyphae with solitary conidiophores. c Conidiophore fascicles d Conidiophores. e Conidia. – Bar = 10 µm. angular-irregular, brownish to greyish brown or Material examined – VENEZUELA, La- dingy grey, 1–6 mm diam., margin indefinite or ra, Sanare, Sabana Redonda Arriba, on leaves with narrow darker marginal line. Caespituli of Mikania cordifolia (L. f.) Willd. (Astera- hypophyllous, punctiform, medium to dark ceae), Sep. 2010, R. Urtiaga 409 (HAL 2492 brown or blackish brown. Mycelium internal. F). Stromata lacking or small, substomatal, brown, Notes – Pseudocercospora mikaniigena, 10–25 µm diam. Conidiophores in small to described from Malaysia on Mikania cordata moderately large fascicles, divergent to dense, (Burm. f.) B.L. Rob., and later recorded from very dense fascicles sometimes subcoremioid, Brazil (Yen & Lim 1983, Barreto & Evans arising from substomatal hyphae or stromata, 1995), is close to P. plunkettii (Chupp) R.F. emerging through stomata, erect, straight, Castañeda & U. Braun (lectotype material subcylindrical-filiform to moderately geni- examined – on Mikania cordifolia, Mexico, culate-sinuous, unbranched, 30–110 × 2.5–5 CUP 40596), but the conidiophores of the latter µm, continuous to pluriseptate, pale to medium species are much shorter, 10–40 µm long, brown or olivaceous-brown, paler towards the never coremioid, and the conidia are somewhat tip, thin-walled, smooth; conidiogenous cells narrower. The collection from Venezuela is integrated, terminal, 10–50 µm long, conidio- morphologically close to the original des- genous loci inconspicuous to subdenticulate, cription, except for lacking superficial hyphae but always unthickened and not darkened. with solitary conidiophores. This species is Conidia solitary, obclavate-subcylindrical, 30– new to Venezuela. 75 × 3–5 µm, 3–7-septate, pale olivaceous to olivaceous-brown, thin-walled, smooth, apex Pseudocercospora mirandensis (Chupp) R.F. obtuse to subacute, base short to longer Castañeda & U. Braun obconically truncate, 1.5–2 µm wide, hila  Cercospora mirandensis Chupp. neither thickened nor darkened. Material examined – VENEZUELA, La-

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Fig. 11 – Pseudocercospora marcelliana. Based on HAL 2486 F. a Conidiophore fas-cicle. b Conidiophores. c Conidia. – Bar = 10 µm. Fig. 11 – Pseudocercospora marcelliana. Based on HAL 2486 F. a Conidiophore fas-cicle. b Conidiophores. c Conidia. – Bar = 10 µm. ra, Sanare, Sabana Redonda Arriba, on leaves = C. hyptidicola Chupp & A.S. Mull., of Miconia sp. (Melastomataceae), Sep. 2010, nom. inval. R. Urtiaga 396 (HAL 2484 F). Material examined – VENEZUELA, La- Notes – This species was described ra, Barquisimeto, market, on leaves of Ocimum from Venezuela on Miconia ibaguensis sanctum L. (Lamiaceae), Jul. 2011, R. Urtiaga (Bonpl.) Triana. Type material has been 447 (HAL 2475 F). examined (Edo Miranda, Santa Lucia, 13 Apr. Notes – This species is known from 1939, Whetzel & Muller, CUP 3093). Braun & Venezuela on Hyptis spp. (Braun & Urtiaga Urtiaga (2008) recorded this species from 2008). Ocimum sanctum is a new host for this Venezuela on Clidemia hirta (L.) D. Don. The country. present collection on Miconia sp. differs from other samples in almost lacking a lesion. Pseudocercospora palicoureina (Petr. & Cif.) U. Braun Pseudocercospora ocimicola (Petr. & Cif.)  Cercospora palicoureina Petr. & Cif. Deighton Material examined – VENEZUELA, La-  Cercospora ocimicola Petr. & Cif. ra, Sanare, Sabana Redonda Arriba, on leaves

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Fig. 12 – Pseudocercospora mikaniigena. Based on HAL 2484 F. a Conidiophore fas-cicles. b Conidiophores. c Conidia. – Bar = 10 µm. of Palicourea perquandrangularis Wernham longer conidia, up to 115 µm. (Rubiaceae), Sep. 2010, R. Urtiaga 416 (HAL 2488 F). Pseudocercospora pancratii (Ellis & Everh.) Notes – New host species and new to U. Braun & R.F. Castañeda Venezuela. This collection agrees well with P.  Cercospora pancratii Ellis & Everh. palicoureina (conidiophores in fascicles, ari- = Cercospora hymenocallidis Pat. sing from stromata, 10–40 µm diam., 10–60 × Material examined – VENEZUELA, La- 2.5–5 µm, conidia obclavate-subcylindrical, ra, Barquisimeto, on leaves of Hymenocallis 30–65 × 3–4.5 µm, 3–6-septate). P. palicou- tubiflora Salisb. (Amaryllidaceae), Nov. 2010, reae O.L. Pereira & R.W. Barreto (Pereira & R. Urtiaga 428 (HAL 2474 F); l.c., on leaves of Barreto 2006), described from Brazil on Hymenocallis sp., Sep. 2010, R. Urtiaga 419 Palicourea marcgravii A. St.-Hill, is charac- (HAL 2473 F). terized by its distinct lesions, lacking stromata, Notes – New to Venezuela and new host conidiophores solitary or formed in small species (Crous & Braun 2003). P. pancratii is fascicles emerging through stomata and much widespread on various hosts of the Amarylli-

318 Mycosphere Doi 10.5943/mycosphere/3/3/5 daceae (Crous & Braun 2003). darkened. Material examined – VENEZUELA, Ca- Pseudocercospora pehriicola U. Braun & rabobo State, Miranda, on leaves of Pehria Urtiaga, sp. nov. Fig. 13 compacta (Rusby) Sprague (Lythraceae), Apr. MycoBank, MB 800013. 2011, R. Urtiaga 434 (HAL 2483 F, holotype). Etymology – epithet derived from the Notes – Braun & Urtiaga (2008) des- host genus, Pehria. cribed Pseudocercospora pehriae on Pehria Pseudocercosporae lagerstroemiigenae compacta from Venezuela. This species is, valde similis, sed hospite distincto, maculis however, quite distinct from P. pehriicola by differentibus, hyphis superficialibus sparse lacking superficial mycelium and stromata, evolutis, laxis. Differt a P. pehriae hyphis very long conidiophores, up to 150 µm, and superficialibus cum conidiophoris solitariis et broadly obclavate-cylindrical conidia, 20–70 × stromatibus formantibus, conidiophoris brevio- 4–7 µm. Several additional Pseudocercospora ribus, 10–50 µm, et conidiis angustioribus, 2– species on hosts belonging to diverse genera of 3.5 µm. the Lythraceae are known. The Asian P. Leaf spots amphigenous, circular to lagerstroemiigena Goh & W.H. Hsieh on somewhat irregular, 2–10 mm diam., occa- Lagerstroemia speciosa (Hsieh & Goh 1990) is sionally zonate, brownish, greyish brown to very similar, but differs in forming different grey or finally greyish white, with a very lesions and well-developed superficial hyphae, narrow dark marginal line, occasionally purple partly dense and aggregated in ropes. P. and slightly raised. Caespituli amphigenous, lagerstroemiae-lanceolatae U. Braun & Crous delicately punctiform on the upper side, (Crous & Braun 2003) and P. lythri H.D. Shin scattered to aggregated, less conspicuous on & U. Braun (Shin & Kim 2001) are two the lower surface, dark brown to blackish. species that form superficial mycelium, but the Mycelium internal and partly external; super- former differs in having very long, pluriseptate ficial hyphae emerging through stomata, spar- conidiophores, up to 100 µm, and the latter has ingly developed, more abundant on the lower wider conidia, 3–5 µm. In other species with surface, sparingly branched, septate, sub- similarly narrow conidia, the conidiophores are hyaline to pale olivaceous or olivaceous- consistently fasciculate, i.e. solitary conidio- brown, smooth, 1–4 µm wide. Stromata lacking phores and superficial hyphae are lacking, viz. to well-developed, 10–60 µm diam., on the P. cupheae (Syd.) U. Braun, P. lythracearum upper side intraepidermal and larger, sub- (Heald & F.A. Wolf) X.J. Liu & Y.L. Guo stomatal and smaller below, medium to dark (incl. P. lagerstroemiae-subcostatae (Sawada) brown or olivaceous-brown, cells 3–6 µm Goh & W.H. Hsieh), P. neseae (Ellis & Everh.) diam. Conidiophores in fascicles, epiphyllous U. Braun, P. woodfordiigena U. Braun & fascicles rather large and dense, hypophyllous Crous (Chupp 1954, Hsieh & Goh 1990, Guo ones smaller and looser, individual conidi- & Hsieh 1995). P. sydowiana (Chupp) U. ophores almost straight to usually distinctly Braun & Crous is characterized by its very and strongly geniculate-sinuous, unbranched, long conidiophores, up to 260 µm, and broad 10–50 × 1.5–4 µm, aseptate to sparingly conidia 20–70 × 4–7 µm, and P. lagerstroe- septate, pale to medium olivaceous or oliva- miae-parviflorae H.S.G. Rao, S. Narayan & ceous-brown, thin-walled, smooth; conidio- Bhartiya has very long, pluriseptate conidio- genous cells integrated, terminal or coni- phores, up to 204 µm, and somewhat wider diophores reduced to conidiogenous cells, 10– conidia, 3–5 µm (Chupp 1954, Goh & Hsieh 25 µm long, conidiogenous loci inconspicuous. 1990, Rao et al. 1996). Conidia formed singly, narrowly obclavate- filiform, occasionally subacicular, 35–80 × 2– Pseudocercospora punicae (Henn.) Deighton 3.5 µm, indistinctly 3–8-septate, hyaline, sub-  Cercospora punicae Henn. hyaline to very pale olivaceous, thin-walled, Material examined – VENEZUELA, La- smooth, apex acute to subobtuse, base usually ra, Barquisimeto, on leaves of Punica grana- obconically truncate, occasionally truncate, 1– tum L. (Punicaceae), Sep. 2010, R. Urtiaga 1.5 µm wide, hila neither thickened nor 404 (HAL 2485 F).

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Fig. 13 – Pseudocercospora pehriicola. Based on type material. a Superficial hyphae. b Superficial hyphae with solitary conidiophores. c Conidiophore fascicle. d Conidiophores and hyphae emerging through a stoma. e Conidiophores, f Conidia – Bar = 10 µm.

Notes – Known from Venezuela (Crous ramosis, hyphis superficialibus cum conidio- & Braun 2003). phoris solitariis formantibus). Leaf spots amphigenous, angular-irre- Pseudocercospora rauvolfiae-tetraphyllae U. gular, 1–10 mm diam., on the upper leaf sur- Braun & Urtiaga, sp. nov. Fig. 14 face at first brown to dark brown, later with MycoBank, MB 800014. pale centre, greyish brown to dingy grey, with Etymology – epithet derived from the a narrow to moderately broad darker margin, host species, Rauvolfia tetraphylla. on the lower side paler brown and margin Differt ab omnibus speciebus Pseudo- indefinite. Caespituli amphigenous, on the cercosporae ad Bignoniaceas (P. liebenbergii, upper side punctiform, dark brown to blackish, P. rauvolfiae, P. rauwolfiae-serpentinae et P. scattered to aggregated, below rather incon- serpentinae) caespitulis epiphyllis et hypo- spicuous. Mycelium internal and external on phyllis distincte dimorphis (caespitulis epi- the lower surface; superficial hyphae emerging phyllis punctiformibus, stromatibus intraepi- through stomata, straight to sinuous, branched, dermalibus bene evolutis, 15–60 µm diam., 1–4 µm wide, septate, subhyaline to pale conidiophoris numerosis, dense fasciculatis; olivaceous-brown, smooth; stromata on the caespitulis hypophyllis indistinctis, stromatibus upper side well-developed, intraepidermal, nullis vel parvis, substomatalibus, conidiopho- brown, 15–60 µm diam., erumpent, subcircular ris parvis, laxe fasciculatis, interdum valde to somewhat irregular in outline, lacking or

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Fig. 14 – Pseudocercospora rauvolfiae-tetraphyllae. Based on type material. a Conidiophore fascicles. b Hypha with solitary conidiophore. c Conidiophores and hyphae emerging through a stoma. d Conidiophores. e Conidia – Bar = 10 µm. only small below, substomatal, cells 2–5 µm tary, arising from superficial hyphae, lateral, diam. Conidiophores on the upper leaf surface straight, subcylindrical, subclavate or slightly in small to moderately large, often dense attenuated towards the tip, unbranched or fascicles, arising from stromata, erumpent, on hypophyllously sometimes slightly to strongly the lower side in smaller and usually looser branched, (5–)10–40 × 2–5 µm, 0–2-septate, fascicles, arising from internal hyphae or small thin-walled, smooth, pale to medium oli- stromata, emerging through stomata, or soli- vaceous or olivaceous-brown; conidiogenous

321 Mycosphere Doi 10.5943/mycosphere/3/3/5 cells integrated, terminal or conidiophores ficial hyphae lacking) and was reduced to reduced to conidiogenous cells, 5–30 µm long, synonymy with the latter species in Kamal conidiogenous loci inconspicuous. Conidia (2010). However, P. rauwolfiae-serpentinae formed singly, obclavate-cylindrical, short should rather be separated from the African conidia sometimes ellipsoid-subcylindrical or species due to its much shorter conidiophores fusoid, 15–80 × 2–4.5 µm, 1–7-septate, sub- and conidia. P. liebenbergii (Syd.) Deighton, hyaline to pale olivaceous or olivaceous- confined to R. caffra Sond. in South Africa, is brown, smooth, apex obtuse to subacute, base quite distinct from P. rauvolfiae-tetraphyllae. short obconically truncate, 1.5–2 µm wide, hila The conidiophores often proliferate percur- neither thickened nor darkened. rently, the conidia become rough-walled, and Material examined – CUBA, Bayamo, on superficial hyphae are lacking (Chupp 1954, leaves of Rauvolfia tetraphylla L. (Apocy- Deighton 1976, Crous & Braun 1996). Chupp naceae), 27 Feb. 1967, R. Urtiaga (IMI 126169 (1954) recorded this species from Venezuela = K(M) 173068, holotype); CUBA, Bayamo, on Rauvolfia sp. The material concerned has on leaves of R. tetraphylla, 12 Nov. 1966, R. not been seen, but it is possible that this record Urtiaga (IMI 123577 = K(M) 173066); CUBA, refers to P. rauvolfiae-tetraphyllae. Bayamo, on leaves of R. tetraphylla, 12 Feb. 1967, R. Urtiaga (IMI 126081 = K(M) Pseudocercospora tibouchinae (Viégas) 173067). VENEZUELA, without locality, on Deighton Fig. 15 leaves of R. tetraphylla, 11 Jul. 1969, R.  Cercospora tibouchinae Viégas. Urtiaga 389 (IMI 141512 = K(M) 173072).  Cercoseptoria tibouchinae (Viégas) Notes – This species, known from Cuba Deighton and Venezuela on Rauvolfia tetraphylla, differs Material examined – VENEZUELA, La- from all other species of Pseudocercospora on ra, Sanare, Sabana Redonda Arriba, on leaves Rauvolfia spp. in forming characteristically of Tibouchina longifolia (Vahl) Baill. dimorphic fructification, i.e. with obvious dif- (Melastomataceae), Sep. 2010, R. Urtiaga 407 ferences between epiphyllous and hypophyll- (HAL 2490 F). ous caespituli. On the upper leaf surface, well- Notes – This species has been described developed intraepidermal stromata with large, from Brazil on Tibouchina sp. Type material dense fascicles of conidiophores are developed, was re-examined and redescribed by Crous et whereas on the lower side stromata are lacking al. (1997). This is the first record of this or small, substomatal, and the conidiophores, species from Venezuela, and T. longifolia is a sometimes distinctly branched, are formed in new host species (Crous & Braun 2003). The small, divergent fascicles and superficial material from Venezuela agrees well with type hyphae with solitary conidiophores are also material of this species (conidiophores arising developed. In P. rauvolfiae Deighton from well-developed stromata, short and (Deighton 1983), described from Africa aseptate, 5–20 × 2–3 µm, forming sporodochial (Guinea and Sierra Leone) on Rauvolfia conidiomata; conidia solitary, narrowly cylin- vomitoria Afzel., superficial hyphae with drical-filiform to somewhat acicular, up to 90 × solitary conidiophores are lacking and the 2–3.5 µm, pluriseptate, apex obtuse to sub- conidiophores are longer, up to 85 µm, mostly acute, base truncate to short obconically 30–50 µm. The Indian P. serpentinae truncate). (Pandotra & Husain) Deighton (Deighton 1976, 1983) on R. serpentina (L.) Benth. ex Kurz is Pseudocercospora trichophila var. punctata characterized by lacking stromata, effuse U. Braun & Urtiaga, var. nov. Fig. 16 hypophyllous colonies, frequently branched MycoBank, MB 800018. conidiophores and abundant superficial hyphae Etymology: the name of the variety with solitary conidiophores. P. rauwolfiae- refers to the presence of punctiform serpentinae H.S. Rao, Arch. Singh & Kamal epiphyllous caes-pituli. (Rao et al. 1995), also described on R. Differt a var. trichophila caespitulis am- serpentina from India, is close to P. rauvolfiae phigenis, in epiphyllo punctiformibus, e stro- (conidiophores arising from stromata, super- matibus, 10–50 µm diam., et conidiophoris fas-

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Fig. 15 – Pseudocercospora tibouchinae. Based on HAL 2490 F. a Conidiophore fascicle. b Conidiophores. c Conidia. – Bar = 10 µm. ciculatis compositis. brown, thin-walled, smooth; stromata small to Leaf spots amphigenous, 2–8 mm diam., well-developed on the upper side, intra- subcircular to angular-irregular,brownish, epidermal, 10–50 µm diam., medium to dark greyish brown, dingy grey, with narrow darker olivaceous-brown, lacking or very small below. border or halo on the upper leaf surface, Conidiophores on the upper side in small to hypophyllous leaf spots less conspicuous, with moderately large fascicles, loose to dense, indistinct margin. Caespituli amphigenous, on arising from stromata, erect, straight, sub- the upper leaf surface punctiform, dark brown, cylindrical to geniculate-sinuous, unbranched, effuse and less conspicuous below. Mycelium 10–70 × 3–5 µm, pale olivaceous to medium internal and external, superficial hyphae olivaceous-brown, 0–4-septate, thin-walled, usually lacking on the upper side, abundant smooth, on the lower side solitary, arising from below, emerging through stomata, sparingly superficial hyphae, lateral, occasionally ter- branched, 1–3.5 µm wide, septate, hyaline, minal, short, 3–25 × 2.5–5 µm, conical-sub- subhyaline to pale olivaceous or olivaceous- cylindrical, 0–1-septate; conidiogenous cells

323 Mycosphere Doi 10.5943/mycosphere/3/3/5 integrated, terminal or conidiophores reduced asperi and tentatively maintained it as separate to conidiogenous cells, 3–20 µm long, conidio- species. However, the latter is barely more than genous loci inconspicuous or subdenticulate, an additional variety of the morphologically but always unthickened and not darkened. variable P. trichophila.: Conidia solitary, subcylindrical or oblavate- cylindrical, (10–)25–90(–110) × (2.5–)3–5 µm, Pseudocercospora trichophila var. solani- (1–)3–10(–12)-septate, distance between septa asperi (R.E.D. Baker & W.T. Dale) U. Braun, 6–18 µm, subhyaline to pale olivaceous, apex comb. et stat. nov. obtuse, rounded to somewhat pointed, base MycoBank, MB 800044. obconically truncate, 1–1.5(–2) µm wide, hila Basionym: Cercospora solani-asperi R. neither thickened nor darkened. E.D. Baker & W.T. Dale, Mycol. Pap. 33: 105, Material examined – VENEZUELA, La- 1951. ra, Sanare, Sabana Redonda Arriba, on leaves  Pseudocercospora solani-asperi (R.E. of Solanum hirtum Vahl (Solanaceae), Jun. D. Baker & W.T. Dale) Deighton, Mycol. Pap. 2010, R. Urtiaga 377 (HAL 2498 F, holotype). 140: 113, 1976. Notes – Deighton (1976) examined type material and numerous other collections of Pseudocercosporella capsellae (Ellis & Pseudocercospora on Solanum spp. and Everh.) Deighton provided a detailed description and some Material examined – VENEZUELA, La- drawings of the morphologically variable ra, Barquisimeto, market, on leaves of Brassica Pseudocercospora trichophila (F. Stevens) rapa subsp. pekinensis Kitam. (Brassicaceae), Deighton, reduced several Cercospora spp. to Jul. 2011, R. Urtiaga 446 (HAL 2476 F). synonymy with the latter species and discussed Notes – New to Venezuela (Braun 1995, it in detail. In the type material of P. IMI Descriptions of Fungi and Bacteria 161, trichophila and its synonyms, colonies are Sheet 1605, 2004). chiefly hypophyllous, stromata are lacking and the conidiophores are formed singly, arising Zasmidium asclepiadis U. Braun & Urtiaga, from superficial hyphae. In the present sp. nov. Fig. 17 collection from Venezuela, the hypophyllous MycoBank, MB 800015. fructification agrees perfectly with typical P. Etymology – epithet derived from the trichophila, but additional epiphyllous caes- host genus, Asclepias. pituli are formed which are punctiform, com- Differt ab omnibus speciebus Zasmidii posed of immersed stromata and fasciculate (et Stenellae) ad Asclepiadaceas (Z. cerope- conidiophores. The conidia agree with those in giae, Z. cynanchi, Z. gongronematis, Z. penta- typical collections of P. trichophila. We prefer tropidis et Z. telosmae) conidiophoris valde to assign such collections with additional fasci- brevioribus, modo 5–40 µm longis et conidio- culate conidiophores arising from stromata to a phoris simplicibus, 0–2-septatis. special morphological variety of the latter On living and faded leaves, lesions species. Var. punctata undoubtedly occurs on subcircular or with diffuse, irregular discolo- other Solanum spp. as well. Hsieh & Goh rations, brownish to reddish. Colonies hypo- (1990: 321, Fig. 243) illustrated, for instance, phyllous, effuse, thin, grey to dingy greyish an agreeing collection from Taiwan on Sola- brown, in older colonies hyphae, conidiophores num aculeatissimum Jacq. Conidia in var. and conidia sometimes with a reddish tinge. trichophila are (3–)4–5(–7) µm wide. Collec- Mycelium internal and external; superficial tions on Solanum asperum Rich. (Brazil and hyphae emerging through stomata, sparingly Trinidad), described as Cercospora solani- branched, septate, thin-walled, vegetative asperi (Trinidad, Arima Forest Reserve, 25 hyphae 1–3 µm wide, hyaline to very pale Oct. 1947, W.T. Dale, Fungi of Trinidad 1729, olivaceous, rough-walled, fertile hyphae or IMI 24507, type, examined), are barely distinct hyphal cells with conidiophores somewhat from P. trichophila, but due to somewhat wider, darker and mostly more or less smooth; narrower conidia, Deighton (1976) introduced stromata lacking. Conidiophores solitary, ari- the combination Pseudocercospora solani- sing from superficial hyphae, lateral, rarely

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Fig. 16 – Pseudocercospora trichophila var. punctata. Based on type material. a Superficial hyphae with solitary conidiophores. b Conidiophore fascicles. c Conidiophores. d Conidia. – Bar = 10 µm. terminal, with 1–2(–3) conidiophores per cell, hyaline or subhyaline, thin-walled, smooth, occasionally with a few fasciculate conidio- apex subacute to subobtuse, base short obco- phores emerging through stomata, erect, nically truncate, 1–1.5 µm wide, hila slightly straight, subcylindrical to moderately geni- thickened and darkened. culate-sinuous, simple or sometimes once to Material examined – VENEZUELA, olivaceous, olivaceous-brown, brown or without locality, on leaves of Asclepias curas- reddish brown, thin-walled, smooth, later savica L. (Asclediadaceae), 23 Jan. 1970, R. several times branched, 5–40 × 2.5–6 µm, Urtiaga 915 (IMI 146225 = K(M) 173065, unbranched conidiophores 0–2-septate, longer holotype). branched conidiophores often with additional Notes – Several species of Zasmidium septa, pigmentation variable, subhyaline, pale have been described on hosts belonging to the sometimes somewhat rough-walled; conidio- Asclediadaceae, but all of them have much phores integrated, terminal, conidiophores longer, pluriseptate conidiophores. Further- aseptate, i.e. reduced to conidiogenous cells, or more, Zasmidium gongronematis (J.M. Yen & conidiogenous cells integrated in superficial Gilles) U. Braun comb. nov. (Bas.: Cercospora hyphae, with lateral peg-like protuberances, 5– gongronematis J.M. Yen & Gilles, Bull. 20 µm long, with a single to several conspi- Trimest. Soc. Mycol. France 137: 69, 1974; cuous conidiogenous loci, circular in outline, MycoBank, MB 800043), a typical Zasmidium thickened and darkened, 1–1.5 µm diam. with planate conidiogenous loci and conidial Conidia formed singly, rarely in short chains, hila, differs from Z. asclepiadis in having narrowly cylindrical-obclavate, short conidia broader (4–5 µm), cylindrical conidia (Yen sometimes narrowly ellipsoid-fusoid, (6–)10– 1975). Z. pentatropidis (K. Srivast., A.K. 60(–100) × 2–3.5 µm, (0–)1–6(–8)-septate, Srivast. & Kamal) Kamal (Kamal 2010) and Z.

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Fig. 17 – Zasmidium asclepiadis. Based on type material. a Superficial hyphae with solitary conidiophores. b Conidiophores and hyphae emerging through a stoma. c Conidiophores. d Conidia. – Bar = 10 µm. telosmae (K. Srivast., A.K. Srivast. & Kamal) southern Brazil with particular reference Kamal (Kamal 2010) are characterized by to fungal pathogens for biological having verruculose conidia, which are also control. Mycological Research 99, 343– wider (3–6 µm) in the latter species. In Stenella 352. ceropegiae M.S. Patil & Sawant (Patil & Braun U. 1993 – New genera of phytopatho- Sawanat 1991) on Ceropegia bulbosa Roxb. in genic deuteromycetes. Cryptogamic Bo- India, the conidiophores are formed in tany 4, 107–114. fascicles, but superficial hyphae with solitary Braun U. 1995 – A monograph of Cerco- conidiophores are lacking. The Indian S. sporella, Ramularia and allied genera cynanchi J.M. Yen, A.K. Kar & B.K. Das (Yen (phytopathogenic hyphomycetes). Vol. 1. et al. 1982b) is characterized by its small, 7–23 IHW-Verlag Eching. × 2.5–3.5 µm, 0–1-septate conidia. The latter Braun U. 1999 – Taxonomic notes on some two species have not yet been re-examined. species of the Cercospora complex (VI). Type material was not available. Therefore, the Cryptogamie, Mycologie 20, 155–177. generic affinity of these species is still unclear. Braun U, Crous PW. 2007 – The diversity of cercosporoid hyphomycetes – new spe- Acknowledgements cies, combinations, names and nomen- We are much obliged to the mycology clatural clarifications. Fungal Diversity curator of the Kew Herbarium (K) for the 26, 55–72. possibility to examine collections previously Braun U, Freire FCO. 2002 – Some cerco- deposited by R. Urtiaga at IMI. sporoid hyphomycetes from Brazil – II. Cryptogamie, Mycologie 23, 295–328. References Braun U, Freire FCO. 2004 – Some cerco- sporoid hyphomycetes from Brazil – III. Barreto RW, Evans HC. 1995 – The mycobiota Cryptogamie, Mycologie 25, 221–244. of the weed Mikania micrantha in Braun U, Urtiaga R. 2008 – New species and

326 Mycosphere Doi 10.5943/mycosphere/3/3/5 new records of cercosporoid Hypho- Deighton FC. 1973 – Studies on Cercospora mycetes from Venezuela. Feddes Reper- and allied genera. IV. Cercosporella torum 119, 484–506. Sacc., Pseudocercosporella gen. nov. and Braun U, Mouchacca J, McKenzie EHC. 1999 Pseudocercosporidium gen. nov. Myco- – Cercosporoid hyphomycetes from New logical Papers 133, 1–62. Caledonia and some other South Pacific Deighton FC. 1974 – Studies on Cercospora islands. New Zealand Journal of Botany and allied genera. V. Mycovellosiella 37, 297–327. Rangel, and a new species of Ramu- Braun U, Crous PW, Kamal. 2003 – New lariopsis. Mycological Papers 137, 1–75. species of Pseudocercospora, Pseudo- Deighton FC. 1976 – Studies on Cercospora cercosporella, Ramularia and Stenella and allied genera. VI. Pseudocercospora (cercosporoid hyphomycetes). Myco- Speg., Pantospora Cif. and Cerco- logical Progress 2, 197–208. septoria Petr. Mycological Papers 140, Braun U, Hill F, Schubert K. 2006 – New 1–168. species and new records of biotrophic Deighton FC. 1979 – Studies on Cercospora micromycetes from Australia, Fiji, New and allied genera. VII. New species and Zealand and Thailand. Fungal Diversity redispositions. Mycological Papers 144, 22, 13–35. 1–56. Braun U, Freire FCO, Urtiaga R. 2010 – New species and new records of cercosporoid Deighton FC. 1983 – Studies on Cercospora hyphomycetes from Brazil, New Zealand and allied genera. VIII. Further notes on and Venezuela. Polish Botanical Journal Cercoseptoria and some new species and 55, 281–291. redispositions. Mycological Papers 151, Chupp C. 1934 – Cercosporae. In: Mycological 113. explorations of Venezuela (eds Chardon Dianese JC, Furlanetto C, Santos LTP 1999 – LE, Toro RA). Monographs of the Pseudocercospora zeyheriae, a new University of Puerto Rico, Series B, 2, combination for Cercospora zeyrae. 241–255. Mycological Research 103, 40–42. Chupp C. 1954 – A monograph of the fungus Doyle JJ, Chappill JA, Bailey CD, Kajita T genus Cercospora. Ithaca. 2000 – Towards a comprehensive phylo- Crous PW, Braun U. 1996 – Cercosporoid geny of legumes: evidence from RBCL fungi from South Africa. Mycotaxon 57, sequences and non-molecular data. In: 233–321. Advances in Legume Systematics (eds. Crous PW, Braun U. 2003 – Mycosphaerella Herendeen PS, Bruneau A). Royal and its anamorphs: 1. Names published in Botanic Gardens, Kew 1–20. Cercospora and Passalora. CBS Bio- Ellis MB. 1976 – More Dematiaceous Hypho- diversity Series 1: 1–569. mycetes. CAB, Kew. Crous PW, Câmara MPS. 1998 – Cercosporoid García CE, Pons N, de Rojas CB. 1996 – fungi from Brazil. 2. Mycotaxon 68, Cercospora y hongos similares sobre 299–310. especies de Ipomoea. Fitopatología Crous PW, Alfenas AC, Barreto RW. 1997 – Venezolana 9, 22–36. Cercosporoid fungi from Brazil. 1. Guo YL, Hsieh WH. 1995 – The genus Mycotaxon 64, 405–430. Pseudocercospora in China. Mycosyste- Crous PW, Benchimol RL, Albuquerque FC, ma Monographicum Series 2, 1–388. Alfenas AC. 2000 – Foliicolous ana- Gupta BK, Kamal. 1987 – Some new species morphs of Mycosphaerella from South of Pseudocercospora from India. In: America. Sydowia 52, 78–91. Perspectives in Mycological Research – Deighton FC. 1967 – Studies on Cercospora I, Prof. G.P. Agarwal Festschrift Volume and allied genera. II. Passalora, Cerco- (eds. Hasija BK, Rajak RC, Singh SM). sporidium and some species of Fusi- New Delhi 19–33. cladium on Euphorbia. Mycological Hsieh WH, Goh TK. 1990 – Cercospora and Papers 112, 1–80. Similar Fungi from Taiwan. Taipei.

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