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A New Poroid Species of Resupinatus from Puerto Rico, with a Reassessment of the Cyphelloid Genus Stigmatolemma

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A New Poroid Species of Resupinatus from Puerto Rico, with a Reassessment of the Cyphelloid Genus Stigmatolemma Mycologia, 97(5), 2005, pp. 000–000. # 2005 by The Mycological Society of America, Lawrence, KS 66044-8897 A new poroid species of Resupinatus from Puerto Rico, with a reassessment of the cyphelloid genus Stigmatolemma R. Greg Thorn1 their place in the cyphellaceous genus Stigmatolemma…’’ Department of Biology, University of Western Ontario, (Donk 1966) London, Ontario, N6A 5B7 Canada Jean-Marc Moncalvo INTRODUCTION Centre for Biodiversity and Conservation Biology, Royal Ontario Museum and Department of Botany, University Resupinatus S.F. Gray is a small genus of euagarics of Toronto, Toronto, Ontario, M5S 2C6 Canada (Hibbett and Thorn 2001) with 49 specific and Scott A. Redhead varietal epithets as of Apr 2005, excluding autonyms Systematic Mycology and Botany Section, Eastern Cereal and invalid names (www.indexfungorum.org). Fruit- and Oilseed Research, Agriculture and Agri-Food ing bodies of Resupinatus are small—a few mm to Canada, Ottawa, Ontario, K1A 0C6 Canada 2 cm in breadth—and generally pendent or resupi- D. Jean Lodge nate on the undersides of rotting logs and other Center for Forest Mycology Research, USDA Forest woody materials or herbaceous debris. Historically, Service-FPL, P.O. Box 1377, Luquillo, Puerto Rico, members of Resupinatus were treated within the USA 00773-1377 broad concept of Pleurotus (Fr.) P. Kumm. (e.g. Pila´t 1935, Coker, 1944). In modern times, the genus has Marı´a P. Martı´n been characterized by a gelatinous zone in the pileus, Real Jardı´n Bota´nico, CSIC, Plaza de Murillo 2, 28014 Madrid, Spain hyaline inamyloid spores and the absence of metuloid cystidia. The genus Hohenbuehelia Schulzer shares the gelatinized layer and inamyloid spores, but has Abstract: A fungus with gelatinous poroid fruiting metuloid cystidia (Singer 1986, Thorn and Barron bodies was found in Puerto Rico and determined by 1986). Singer (1986) treated the two genera as macro- and micromorphology to be most similar to separate members of the tribe Resupinateae, family members of the lamellate agaric genus Resupinatus. Tricholomataceae. In the Resupinateae, Singer This species is described as a new species, Resupinatus (1986) also included three cyphelloid genera with porosus. Phylogenetic analyses of ribosomal DNA cup-shaped fruiting bodies lacking gills, namely sequences support the inclusion of this fungus in Stigmatolemma Kalchbr., Stromatocyphella W.B. Cooke, the clade containing Resupinatus, and indicate that and Aphyllotus Singer. In contrast, Ku¨hner (1980) this monophyletic group also includes members of treated Hohenbuehelia as a synonym of Resupinatus,in Asterotus and the cyphelloid genus Stigmatolemma. the tribe Resupinateae, family Pleurotaceae. Thorn Resupinatus porosus is another example of tropical and Barron (1986) showed that Hohenbuehelia and poroid representatives of lamellate agaric genera. Resupinatus differ in a fundamental way-members of Resupinatus is emended to include species with Hohenbuehelia posses a nematophagous anamorph poroid (R. porosus) or merulioid (R. merulioides) referrable to Nematoctonus Drechsler, whereas Resu- hymenophore as well as those with laterally stipitate pinatus species are non-nematophagous and lack (Asterotus) or cyphelloid (Stigmatolemma) fruiting a conidial anamorph. Despite this difference, Thorn bodies. Seven new combinations in Resupinatus are and Barron (1986) treated Hohenbuehelia and Resu- proposed to accommodate well-known species of pinatus as members of the same tribe, Resupinateae, Stigmatolemma. and suggested that both genera were closely allied to Key words: Agarics, Henningsomyces, hymeno- Pleurotus. phore evolution, phylogeny, Porotheleum, rDNA, More recent analyses of ribosomal DNA sequences tropical fungi (Hibbett and Vilgalys 1993, Hibbett et al 1997, Moncalvo et al 2000, Thorn et al 2000, Hibbett and ‘‘Resupinatus is an agaric genus with small to minute Thorn 2001, Moncalvo et al 2002, Bodensteiner et al cupulate fruitbodies. If one takes away the gills, which 2004, Binder et al 2005) have greatly clarified and may be few, one has constructed species that would find changed our concepts of relationships among the pleurotoid fungi and other agarics. Thorn et al Accepted for publication 30 Aug 2005. (2000) showed using sequences from nuclear rDNA 1 Corresponding author. E-mail: [email protected] that Pleurotus and Hohenbuehelia form a monophyletic 0 Mycologia myco-97-05-16.3d 11/1/06 20:29:24 1 0MYCOLOGIA group referred to as the family Pleurotaceae. Both (1987) described Resupinatus merulioides, a species genera are nematophagous, Pleurotus by toxic drop- from Japan with a merulioid instead of strictly lets and Hohenbuehelia by adhesive knobs; a single lamellate hymenophore. Other microscopic charac- known Hohenbuehelia has both adhesive knobs and ters of this species greatly resemble the type species, toxic droplets (Thorn et al 2000). Sequences from Resupinatus applicatus (Batsch : Fr.) S.F. Gray. Red- other members of the Resupinateae, including head and Nagasawa (1987) remarked that the almost Asterotus Singer and Resupinatus, formed a mono- poroid hymenophore of R. merulioides is a unique phyletic group within the Tricholomataceae sensu feature within Resupinatus, but used the combination lato, well separated from the Pleurotaceae (Thorn et of gelatinized, fuscous tissues, globose, inamyloid al 2000). This clade also includes species of Stigma- spores, absence of metuloid cystidia and the presence tolemma (Moncalvo et al 2002, Bodensteiner et al of coralloid pileipellis elements with scattered tibii- 2004). As yet, no sequences from the other cyphelloid form cystidioid ends as evidence for placing this members of the Resupinateae, Aphyllotus and Stroma- species in Resupinatus as redefined by Thorn (1986). tocyphella, have been available for phylogenetic Here, we describe a truly poroid representative of analysis. Resupinatus from Puerto Rico. The first collection of The smallest species of Resupinatus, such as R. this species was found growing on wet wood by kavinii (Pila´t) M.M. Moser, form gregarious, small a mycology class of A. Perez (Univ. Interamericana at cups, 1–2 mm in diam, with a few reduced lamellae. Arecibo) in the Toro Negro Commonwealth Forest at Fruiting bodies of Stigmatolemma are minute cups, 1000 m above sea level (asl) in the Cordillera Central 0.1–1.5 mm in diam, gregarious or crowded on the of Puerto Rico. Since then several other collections substratum, and often surrounded by or seated in have been made in the Luquillo Mountains of Puerto a tomentose mat of hyphae called a subiculum. Rico at 250–380 m asl at El Verde and the Bisley Basidia line the inner surface of the cups and there Watersheds. As a consequence of including a poroid are no wrinkles or lamellae. As in Resupinatus,the species within Resupinatus, based upon phylogenetic trama is fuscous and gelatinous, and the basidio- analyses, we provide an emended description of the spores are hyaline and inamlyloid. Romagnesi (1950, genus, and transfer well known species of Stigmato- 1953, both cited in Donk 1962a) noted the similarities lemma to Resupinatus. between R. kavinii (as Scytinotopsis kavinii)and Stigmatolemma poriaeforme (Pers.: Fr.) Singer (as MATERIALS AND METHODS Solenia poriaeformis). The generic name Stigmato- lemma was poorly known and disused until Talbot Micromorphology.—Small portions of specimens of Resu- (1956) redescribed the type species as Porotheleum pinatus and Stigmatolemma were rehydrated in distilled incanum (Kalchbr.) Sacc. Up to that time, species of water and then sectioned to observe micromorpholog- ical features of the hymenium, trama and pileipellis. Stigmatolemma had been classified primarily in Sections were mounted in Melzer’s reagent, 0.05% (w/v) Cyphella (Burt 1915, Bourdot and Galzin 1928), Cotton Blue in lactophenol and 2% (w/v) KOH for Porotheleum (Cunningham 1953, Cooke 1957) or microscopy (Kirk et al 2001). Basidiospores were Solenia (Burt 1924). Singer (1962) resurrected measured at 10003 magnification in Melzer’s reagent. Stigmatolemma, provided a modern description for Size ranges presented are the median 80%,with the genus, and treated it as a ‘‘reduced’’ (cyphelloid) extremes in parentheses. Capitalized color terms in member in the tribe Resupinateae, together with descriptions are from Ridgway (1912). Herbarium Asterotus, Hohenbuehelia and Resupinatus.Donk acronyms follow Holmgren et al (1990). (1962a) emended the genus and provided discussion DNA extraction and sequencing.—Genomic DNA was of four species, S. incanum Kalchbr., S. conspersum isolated from dried specimens of R. porosus using the (Pers. : Fr.) Donk, S. taxi (Le´v.) Donk, and S. E.Z.N.A. Fungal MiniPrep Kit (Omega-Biotech, Dora- urceolatum (Wallr. ex Fr.) Donk. ville, Georgia) as described in Martı´n and Winka In both Resupinatus and Stigmatolemma, the pilei- (2000). The internal transcribed spacer regions of the pellis is a loose trichodermium of coralloid-diverticu- nuclear ribosomal DNA (ITS) including the 5.8S rDNA late hyphae that secrete hyaline to brownish crystal- gene were amplified using primers ITS1F and ITS4 9 line material from fine cylindric or tapering pegs. In (White et al 1990, Gardes and Bruns 1993), and the 5 1000-base region of the nuclear large ribosomal RNA addition, the diverticulate elements at the cup gene (nLSU-rDNA) was amplified using primers LROR margins in Stigmatolemma (see Redhead 1973) greatly and LR5 (Vilgalys and Hester 1990). Amplifications resemble cheilocystidia from gill edges in Resupinatus were done using Ready-to-GoH
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