MYCOTAXON. A new species of Gloeocantharellus (Fungi-Basidiomycetes) from Colombian Amazonia

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1 MYCOTAXON Volume x, pp. Month-Month 200x A new species of Gloeocantharellus (Fungi-Basidiomycetes) from Colombian Amazonia AÍDA MARCELA VASCO-P. avascop@yahoo.com Laboratorio de Taxonomía de Hongos Instituto de Biología, Universidad de Antioquia A.A.1226, Medellín, Colombia ANA ESPERANZA FRANCO-MOLANO afranco@quimbaya.udea.edu.co Laboratorio de Taxonomía de Hongos Instituto de Biología, Universidad de Antioquia A.A.1226, Medellín, Colombia Abstract - A new taxon of Gloeocantharellus is proposed as G. uitotanus. The species was collected in a terra firme forest in the northern part of the Middle Caquetá in the Amazonian region of Colombia. This species is characterized by having a combination of characters not found in other species of the genus: dry pileus surface which is smooth, areolate and bright peach red to reddish brown; and basidiospores (8-) x (4-) (-6.4) µm, that are ellipsoidal to amygdaliform with rugulose walls. The genus Gloeocantharellus has five species with mainly neotropical and pantropical distribution. Gloeocantharellus uitotanus constitutes the sixth species of the genus, and it is known only from Colombia. Key words - Amazonas, Basidiomycetes, Gomphaceae Resumen - Una nueva especie de Gloeocantharellus es propuesta como G. uitotanus. El espécimen que representa este taxón se encontró sobre suelo, en bosque de terra firme, en la zona norte de la región del Medio Caquetá, Amazonía Colombiana. Esta especie se caracteriza por tener una combinación de caracteres diferentes a las encontradas en otras especies del género, como son la superficie del píleo seca, glabra, areolada, de color rojo brillante a rojo-marrón y basidiosporas de (8-) x (4-) (-6.4) µm, elipsoides a amigdaliformes y con paredes rugulosas. El género Gloeocantharellus tiene cinco especies de distribución tropical y pantropical principalmente. G. uitotanus se constituye en la sexta especie del género y hasta el momento solo se ha encontrado en Colombia.

2 Introduction Colombia is expected to have a highly diverse mycota due to the great variation in climatic zones, but our knowledge of this diversity is still very limited and only a small fraction of the expected taxa have been reported. Most of these reports have been based on collections from mountainous regions as most of the mycological studies have been limited to these areas (Franco-M. & Uribe-C. 2000; Henao-M. 1989, 1990, 1997; Tobón 1991; Uribe-M. 1995). Data on the diversity, biology and ecology of the mycota from Colombian lowland forests are lacking. Although the Amazonian region and the Pacific Chocó area are characterized by high biodiversity of flora and fauna (Etter 1997; Rangel-Ch. et al. 1995a, 1995b; Duivenvoorden & Lips 1993), very few mycological investigations have occurred in these areas. New fungal taxa have been identified in a study of the uses and handling of fungi by the Uitoto ethnic group, carried out in the region of Middle Caquetá, along the banks of the Caquetá river, situated between the departments of Caquetá and Amazonas. In this paper Gloeocantharellus uitotanus is described as a terrestrial macrofungus collected in a terra firme tropical rain forest. The indigenous people are not reported to use this fungus, but they recognize this species as the land mushroom or fungus of earth (hongo de tierra) (Vasco-P. 2002) in reference to the habitat of this fungus. Methods Color designations given within parentheses are from Kornerup & Wanscher (1983). Line drawings were made with the aid of a drawing tube. All micromorphological structures were measured in 3% KOH. Letter abbreviations describing basidiospore size are: n = number of spores measured, x = mean length x mean width, Q = mean length/width ratio. Taxonomy Gloeocantharellus Singer is a poorly known genus that belongs to the Gomphaceae Donk. The genus was proposed by Singer in 1945 to include species characterized by the presence of warty spores and cyanophilous gloeocystidia (Singer 1945; Singer 1986; Kirk et al. 2001). Singer considered Gloeocantharellus as an intermediate genus between Gomphus and Cantharellus because it has characteristics shared by these two genera (Singer 1945; Corner 1966). The cantharelloid basidiomes and the bifurcate lamellae are similar to Cantharellus, but the hymenophore is not interveined. The basidiospores of Gloeocantharellus are verrucose or warty and the cystidia, also called gloeocystidia, are immersed in the hymenophoral trama (as in Gomphus), have oily contents, and are cyanophilous (Singer 1945; Corner 1966). Corner (1966, 1969) proposed that Gloeocantharellus is more closely related to Paxillus than to Cantharellus, since they share an agaricoid form, the presence of caulocystidia and subhymenial cystidia. However, the presence of cheilocystidia and the ornamented spores separates these genera considerably (Corner 1966, 1969;

3 Petersen 1971, Watling & Meijer 1997). Phylogenetic studies of the family Gomphaceae show that Gloeocantharellus is very close and basal to Gomphus (Pine et al. 1999; Villegas et al. 1999). Five species of Gloeocantharellus have been described to date. Gloeocantharellus purpurascens (A. H. Smith & Hesler) Singer was described from North America and has also been reported from French Guyana (Corner 1966; Petersen 1971). Gloeocantharellus corneri (Singer) Corner was described from Sri Lanka and has also been reported from Brazil and French Guyana (Corner 1969; Petersen 1971; Pegler 1986; Watling & Meijer 1997). Three other species of Gloeocantharellus are only known from SE Asia and Melanesia: G. echinosporus Corner from Malaysia, Indonesia, Solomon Islands and Melanesia (Corner 1969; Pegler 1986), G. lateritius (Petch) Corner from Ceylon (Petersen 1971), and G. okapaensis (Corner) Corner from New Guinea and Solomon Islands (Corner 1969). Gloeocantharellus uitotanus from Colombia is the sixth species. A species of Gomphus described from Guadaloupe that has recently been found in Belize, Gomphus viride (Pat.) Singer, may also belong in the genus Gloeocantharellus (D. J. Lodge, pers. com., 19 July 2003). Gloeocantharellus uitotanus Vasco-Palacios & Franco-Molano sp. nov. (Fig. 1. A- D) Gloeocantharellus uitotanus ae G. okapaensis maxime simile est; ex quod autem in egestatibus pileocystidiarum et in mensura basidiarum disimile est. Basidiarum in Gloeocantharellus uitotanus x 7-10 µm mensurae sunt, aliter in G. okapaensis x µm basidiarum sunt mensurae. Etymology: Named after the Uitoto indigenous ethnic group that lives in the region of the Middle Caquetá in the Amazonas Department of Colombia. COLOMBIA. DPTO AMAZONAS: Los Monos, at the south side of the Caquetá river, m. altitude, 24 September 2001, gregarious, growing on ground in argillaceous soil, A. Vasco-P. 376 (HOLOTYPE, HUA). Description Pileus (-6) cm diam., plano-convex, slightly depressed at the center; surface humid and bright when young, dry and bright in mature specimens, smooth, areolate, bright peach red to reddish brown (near 8C8-9C8), paler towards the margin; margin entire, inrolled when young. Context up to 1 cm thick, white, unchanging when bruised or exposed, continuous with the stipe. Odor and taste not distinctive. Lamellae up to 2 mm broad, subdistant, subdecurrent, some anastomosing towards the stipe, pale yellow (4A2); margin entire or fimbriate. Stipe 5-7 cm long., cm diam. near the apex, and 0.8 cm diam. at the base, tapered downward; surface dry, with scanty hairs at the apex; hairs reddish brown on pale yellow (3A2) background; interior white, unchanging, fleshy, and solid. No reaction was observed with KOH. Spore print not obtained. Basidospores (Fig. 1B) (8-) x (4-) (-6.4) µm (x = 9.5 x 5 µm, n = 40, Q = 1.89), ellipsoid to amygdaliform, hyaline in KOH, cyanophilous in Cotton blue,

4 inamyloid in Melzer s reagent; spore wall somewhat thick, rugulose. Basidia (Fig. 1C) x 7-10 µm, clavate, with 4-sterigmata, thin-walled, hyaline, inamyloid, with clamp connections at the base; sterigmata up to 7 µm long. Pileipellis an ixocutis of a compact layer of repent and interwoven hyphae; hyphae up to 4 µm diam., with thin, smooth, inamyloid walls, reddish in H 2 O, and reddish but soon hyaline in KOH. Trama of pileus monomitic, composed of interwowen, generative, thin-walled, hyaline, inamyloid hyphae, µm diam. Hymenophoral trama irregular, composed of thin-walled, inamyloid hyphae, up to 4 µm diam; oleiferous hyphae clavate, flexuose to cylindrical, up to 5 µm diam, with granular and yellow contents in KOH, cyanophilous. Gloeocystidia (Fig. 1D) x µm, inmersed 55 µm in the trama and projecting about 20 µm beyond the basidia, elongated, fusiform to sublanceolate at apex, with yellow content in H 2 O and KOH, cyanophilous, thin-walled. Stipitipellis formed of a compact layer of vertically arranged interwoven hyphae giving rise to bundles of caulocystidia. Caulocystidia 30-42(-50) x 5-9 µm narrowly clavate, clavate to cilindric, with yellow content in H 2 O and KOH. Clamp connections abundantly present. Gloeocantharellus uitotanus is very similar to G. okapaensis from which it differs in the morphology of pileipellis, chemical reaction of the pileipellis and stipitipellis and size of the basidia. While in Gloeocantharellus uitotanus the pileipellis is an ixocutis, the hyphae of the pileipellis and stipitipellis do not react with KOH and basidia are x 7-10 µm. In G. okapaensis the pileipellis is a dermis formed by hyphae more or less divergent over the center of the pileus, the pileipellis and stipitipellis turn reddish in KOH and the basidia are larger (55-70 x µm). Gloeocantharellus okapaensis has been reported from New Guinea and Solomon Islands (Corner 1969). Gloeocantharellus corneri, reported from French Guiana and Brazil, differs from G. uitotanus by the dull colour of the pileus, and because the pileipellis, stipitipellis and gloeocystidia turn reddish brown in KOH. Gloeocantharellus corneri also has bigger spores and basidia. Gloeocantharellus purpurascens reported from French Guiana, differs from G. uitotanus in having pileocystidia of two types, and bigger basidia (55-70 x µm). Gloeocantharellus echinosporus reported from Malayasia, Indonesia, Solomon Islands and Melanesia differs from G. uitotanus by having echinulate basidiospores and because the gloeocystidia, the pileipellis and the stipitipellis turn reddish in KOH. Gloeocantharellus lateritius, known only from Ceylon, differs from G. uitotanus in having a finely felty pileus surface, and the gloeocystidia, pileipellis and stipitipellis turn reddish in KOH as in G. corneri, G. okapaensis and G. echinosporus.

5 Fig. 1. Gloeocantharellus uitotanus A Habit; B Basidospores; C Basidia; D Gloeocystidia Acknowledgements This work was possible due to the support of the Instituto de Ciencias Naturales de la Universidad Nacional de Colombia, Instituto de Biología de la Universidad de Antioquia, and NWO and WOTRO grant WB to Dr. Teun Boekhout (CBS, Utrecht, The Netherlands). The authors wish to acknowledge Vicente Makuritofe, Emilia Faribia`go, Isolina Guerrero, Magdalena Farirama, Reinaldo Ruiz and the other indigenous people who actively collaborated

6 in the field. Thanks are extended to Carlos Alberto López, Aída Palacios and Luis Guillermo Vasco for the unconditional support and encouragement throughout the work and to Drs Jaime Uribe, Teun Boekhout, D. Jean Lodge, Gregory Mueller and Roy E. Halling for the critical review of the manuscript. Drawings were made by Gloria Mora from the Herbarium of the University of Antioquia. Literature cited Corner E JH A monograph of cantharelloid fungi. Ann. Bot. Mem. 2: Corner E JH Notes on cantharelloid fungi. Nova Hedwigia. 18: Duivenvoorden J, Lips JM Ecología del paisaje del Medio Caquetá. Memoria explicativa de los mapas. Serie estudios en la Amazonía Colombiana III A. Tropenbos Colombia. 301 pp. Etter E Bosque húmedo tropical. En: Informe nacional sobre el estado de la biodiversidad. Colombia. Tomo I. Diversidad Biológica. Editor. Instituto de Investigación de Recursos Biológicos A. von Humboldt. Bogotá, Colombia. Franco-M. AE, Uribe-C. E Hongos Agaricales y Boletales de Colombia. Biota Colombiana 1 (1): Henao-M LG Notas sobre afiloforales Colombianos (Basidiomicetes: Aphyllophorales). Caldasia 16: 1-9. Henao-M LG Notas sobre afiloforales Colombianos (Basidiomicetes: Aphyllophorales) II. Caldasia 16: Henao-M LG Afiloforales de Colombia III. Amauroderma (Basidiomicetes: Ganodermataceae) en el Herbario Nacional Colombiano. Caldasia 19: Kirk PM, Canon PF, David JC, Stalpers JA Ainsworth & Bisby s Dictionary of fungi. CAB International. 9ª Ed. Surrey UK. 981 pp. Kornerup A, Wanscher J Methuen handbook of colour. 3 rd ed. Eyre Methuen. London. 252 p. 30 pl. Pegler D Agaric flora from Sri Lanka. Kew Bull., Addit. Ser. 12: pp. Petersen R The genera Gomphus and Gloeocantharellus in North America. Nova Hedwigia 21: Pine EM, Hibbett DS, Donoghue MJ Phylogenetic relationships of cantharelloid and clavarioid Homobasidiomycetes based on mitochondrial and nuclear rdna sequences. Mycologia 91: Rangel-Ch. O., Aguilar-P M, Sánchez-C H, Lowy-C P, Garzón-C A, Sánchez LA. 1995a. Región de la Amazonía pp En: Colombia, diversidad biótica I. O. Rangel-Ch., Editor. Editorial Guadalupe, Santafé de Bogotá. Rangel-Ch. O., Aguilar-P M, Sánchez-C H, Lowy-C P. 1995b. Región Costa Pacífica. pp En: Colombia, diversidad biótica I. O. Rangel-Ch., Editor. Editorial Guadalupe, Santafé de Bogotá. Singer R New genera of fungi-ii. Lloydia, 8: Singer R The Agaricales in modern taxonomy. Königstein, Koeltz Scientific Books. Germany. 981pp. Tobón LE Ascomicetes de Colombia: Discomicetes del departamento de Antioquia. Caldasia. 16: Uribe-M J Catálogo de Myxomycetes registrados para Colombia. Caldasia 18: Vasco-P AM Estudio etnobiológico de los hongos macromicetes entre los Uitoto de la región de Araracuara (Amazonia colombiana). Tesis de Pregrado. Universidad Nacional de Colombia, Bogotá. 199 pp. Villegas M, De Luna E, Cifuentes J, Estrada-Torres A Phylogenetic studies in Gomphaceae sensu lato (Fungi: Basidiomycetes). Mycotaxon 70 (2): Watling R, De Meijer AR Macromycetes from the State of Paraná, Brasil. 5. Poroid and lamellate boletes. Edinburgh J. Bot. 54 (2):