Bioluminescent Mycena Species from Sa˜O Paulo, Brazil
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Mycologia, 99(2), 2007, pp. 317–331. # 2007 by The Mycological Society of America, Lawrence, KS 66044-8897 Bioluminescent Mycena species from Sa˜o Paulo, Brazil Dennis E. Desjardin1 made to Parque Estadual Turı´stico do Alto Ribeira Department of Biology, San Francisco State University, (PETAR), Sa˜o Paulo State, Brazil, in search of San Francisco, California 94132, USA bioluminescent basidiomycetes. Our collecting sites Marina Capelari were over calcareous soil in evergreen hygrophyllous Instituto de Botaˆnica, Caixa Postal 4005, 01061-970 forest classified as Dense Ombrophylous Forest by Joly Sa˜o Paulo, SP, Brazil et al (1999). Dominant angiosperm families in this habitat include Myrtaceae, Leguminosae. Rubiaceae, Cassius Stevani Melastomataceae, Lauraceae, Sapotaceae, Euphorbia- Instituto de Quı´mica da Universidade de Sa˜o Paulo, ceae, Moraceae and Annonaceae (Oliveira Filho and Caixa Postal 26077, 05599-970, Sa˜o Paulo, SP, Brazil Fontes 2000, Aidar et al 2001). This forest type constitutes the most ancient and diverse ecosystem for Orchidaceae in South America (Brieger 1969) with Abstract: Six species of bioluminescent agarics are high diversity and endemism indices (Mori et al described and illustrated from a single site in primary 1981). To date we have documented the presence of Atlantic Forest habitat in the Parque Estadual eight bioluminescent species of agarics in PETAR. We Turı´stico do Alto Ribeira, Sa˜o Paulo State, Brazil. are unaware of any other site in the world with equal These include two new taxa of Mycena, viz. M. asterina or greater diversity of bioluminescent basidiomycetes. and M. lucentipes. Luminescence in Mycena fera, M. Herein we describe two new species of Mycena, report singeri and M. discobasis is reported for the first time. for the first time luminescence in Mycena fera Maas In addition an undeterminable luminescent Mycena Geest. & de Meijer, M. singeri Lodge and M. discobasis species is described and additional specimens of Me´trod, and provide data on two other luminescent Gerronema viridilucens are documented. An account- agarics, viz. an undeterminable Mycena species and ing of known bioluminescent species of Mycena and Gerronema viridilucens Desjardin, Capelari & Stevani. a discussion of why they luminesce are presented. Key words: agarics, antioxidant defense, diver- sity, fungi, taxonomy MATERIALS AND METHODS All basidiomes were collected at night in near total darkness INTRODUCTION and were detected in the field as sources of green luminescence. Specimens were placed in plastic collecting Numerous species of Mycena form luminescent boxes in the field for transport back to the field laboratory. mycelium and/or basidiomes. By our count no fewer All fresh specimens then were photographed in lighted than 26 species of Mycena have been reported as conditions with aid of a flash, and in dark conditions with bioluminescent (TABLE I), belonging to a least 13 20–50 min exposures. Photos were taken with a Nikon N80 taxonomic sections within Mycena sensu lato. To this camera equipped with a Nikkor AF Zoom 28–105 mm lens, accounting we add another six species of luminescent with Fuji Superia ASA 400 or ASA 800 color print film at f5.5 and f11. Fresh specimens were maintained in cool condi- Mycena that belong to an additional three sections of tions until the next morning when macromorphological the genus. The new reports represent material data were documented in daylight. Color terms and collected recently in old growth Atlantic Forest notations in parentheses are those of Kornerup and habitat in Sa˜o Paulo State, Brazil. Wanscher (1978). The terms used to describe lamellae At one time more than one million square kilo- spacing refer to the number of lamellae that reach from the meters of Brazil were covered in Atlantic Forest stipe to the pileus margin (primaries) and do not include the habitat, considered one of the three most threatened lamellulae, whose spacing is indicated by the number of ecosystems in the world. Until relatively recently more series present. All measurements and colors reported for than 82% of Sa˜o Paulo State in southeastern Brazil microscopic features were made from dried material was covered by forest, but today only about 7% of the rehydrated in 100% ethanol followed by distilled water, 3% KOH, 3% KOH + Congo red, or Melzer’s reagent. Spore region remains forested, primarily in mountainous statistics include: x, the arithmetic mean of the spore length regions near the coast (SOS Mata Atlantica 1998). In by spore width (6 standard deviation) for n spores measured Dec–Apr 2001–2006 numerous collecting trips were in a single specimen; xmr, the range of spore means and xmm, the mean of spore means (6SD) where more than one Accepted for publication 30 Nov 2006. specimen is available; Q, the quotient of spore length and 1 Corresponding author. E-mail: [email protected] spore width in any one spore, indicated as a range of 317 318 MYCOLOGIA TABLE I. Species of Mycena reported as bioluminescent in the literature Taxon1 Mycelium Basidiomes Citation Reporting Luminescence Sect. Basipedes M. illuminans Henn. ? + Corner 1954, 1994 M. stylobates (Pers. : Fr.) Kummer + 2 Bothe 1931 Sect. Calodontes M. pura (Pers. : Fr.) Kummer + 2 Treu and Agerer 1990 M. rosea (Bull.) Gramberg + 2 Treu and Agerer 1990 Sect. Citricolores M. citricolor (Berk. & M.A. Curtis) Sacc. + 2 Buller 1934 (as Omphalia flavida) Sect. Diversae M. lucentipes ++ this paper Sect. Exornatae M. chlorophos (Berk. & M.A. Curtis) Sacc. ++ Corner 1954 M. discobasis Me´trod ? + this paper Sect. Fragilipedes M. polygramma (Bull. : Fr.) S.F. Gray + 2 Bothe 1931, Treu and Agerer 1990 M. zephirus (Fr. : Fr.) Kummer + 2 Bothe 1931, Treu and Agerer 1990 Sect. Galactopoda M. haematopus (Pers. : Fr.) Kummer ++ Treu and Agerer 1990, Bermudes et al 1992 Sect. Hygrocyboideae M. epipterygia (Scop. : Fr.) S.F. Gray + 2 Bothe 1931 Sect. Lactipedes M. galopus (Pers. : Fr.) Kummer + 2 Bothe 1931, Treu and Agerer 1990 Sect. Mycena M. inclinata (Fr.) Que´l. + 2 Wassink 1948 (as M. galericulata var. calopus) M. maculata P. Karst. + Treu and Agerer 1990 M. tintinnabulum (Fr.) Que´l. + 2 Bothe 1930 Sect. Roridae M. irritans E. Horak 2 + Horak 1978 M. lamprospora (Corner) E. Horak 2 + (spores only) Corner 1950, 1994, Horak 1978 M. pruinoso-viscida Corner ? + Corner 1954, 1994 M. pruinoso-viscida var. rabaulensis Corner ? + (spores only) Corner 1954, 1994 M. rorida (Fr.) Que´l. + 2 Josserand 1953 M. sublucens Corner 2 + Corner 1954 Sect. Rubromarginatae M. lux-coeli Corner ? + Corner 1954 M. noctilucens Corner ? + Corner 1954, 1994 M. noctilucens var. magnispora Corner ? + Corner 1994 M. olivaceomarginata (Massee apud Cooke) Massee + 2 Wassink 1978 (as M. avenacea) M. singeri Lodge ? + this paper M. species ? + this paper Sect. Sacchariferae M. asterina ? + this paper Sect. Sanguinolentae M. sanguinolenta (Alb. & Schwein. : Fr.) Kummer + 2 Bothe 1931 Sect. Supinae M. fera Maas Geest. & de Meijer ? + this paper Manipularis-group M. manipularis (Berk.) Me´trod nom. inval. ++ Corner 1954 non M. manipularis (Berk.) Sacc. 5Poromycena manipularis (Berk.) Heim 5Filoboletus manipularis (Berk.) Singer M. manipularis var. microporus Kawamura ex Corner ? + Corner 1954 nom. inval. DESJARDIN ET AL:BIOLUMINESCENT MYCENA SPECIES 319 TABLE I. Continued Taxon1 Taxon1 Mycelium Basidiomes Citation Reporting Luminescence Incertae Sedis2 M. daisyogunensis Kobayasi ? + Kobayasi 1951 M. pseudostylobates Kobayasi + ? Kobayasi 1951 1 Synonyms of the taxa listed here are not included. Ex: Mycena dilitata (Fr. : Fr.) Gillet was cited by Bothe (1931) as having luminescent mycelium, but that epithet is now considered a synonym of M. stylobates. 2 Haneda (1955) reported a number of luminescent Mycena species from Japan, referred to Kominami (M. photogena)or Kawamura (M. citrinella var. illumina, M. microillumina, M. phosphora, M. yapensis). None of these epithets were published validly and their taxonomic affinities remain uncertain. variation in n spores measured; Qm, the mean of Q-values in lose overall, lacking universal veil spines at maturity, a single specimen; Qmr, the range of Qm values and Qmm, the dull, dry; disk pure white or pale yellowish white mean of Qm values where more than one specimen is (,4A2), pure white elsewhere, sometimes developing available. Specimens are deposited in SP and SFSU. pale yellowish white tones overall when dried; strongly greenish-white luminescent. Context extremely thin, TAXONOMY white. Lamellae ascending, subfree, close to subdistant with 1–2 series of lamellulae, narrow (0.5–0.75 mm), Mycena asterina Desjardin, Capelari et Stevani, sp. white, nonmarginate; strongly greenish-white lumi- nov. FIGS. 1a–b, 7a–g nescent. Stipe 10–30 3 0.2–0.5 mm, central, filiform, Primordium 0.5–1.5 mm latus 3 0.2–2.0 mm altus, hemisphaericum, album pallide flavo-tinctum, elementis terete, cylindrical for most of the length, base veli universalis cylindraceis dense obtectis. Pileus 1.0– subclavate to subbulbous, sometimes arising from 3.5 mm latus, obtusus conicus raro convexus, striatus vel a small circular pad of mycelium, fragile, translucent, sulcatus, siccus, minute pulverulentus, glabrescens, albus, dull, dry; apex glabrous, base with scattered, white centro pallide flavo-tinctis, viridis luminosus. Caro perte- strigose hairs; pure white overall or seldom with the nuis, alba. Lamellae adscendentes, subliberae, confertae vel base faintly pale grayish white under the white hairs; subdistantes, angustatae (0.5–0.75 mm), albae, margine not luminescent. concolores, viridae luminosae. Stipes 10–30 3 0.2– Basidiospores (FIG. 7b) 8.2–11.5 3 3.5–5 mm(xmr 5 0.5 mm, fragilis, cylindraceus, albus; apice aequali, glaber; 8.8–10 3 4.0–4.3 mm, x 5 9.4 6 0.8 3 4.1 6 basi pubescens, subbulbosus, haud luminosus. Basidios- mm 0.2 mm, Q 5 2.0–3.0, Q 5 2.22–2.34, Q 5 2.28 6 porae 8.2–11.5 3 3.5–5 mm, ellipsoideae vel subfusoideae, mr mm 5 leves, hyalinae, amyloideae. Basidia 12–15 3 9–11 mm, 0.08, n 20 spores per 2 specimens), a few spores subglobosa vel late clavata, 2- vel 4-sporigera, fibulata.