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Megacollybia (Agaricales) Rep. Tottori Mycol. Inst. 45 : 1–57, 2007. Megacollybia (Agaricales) KAREN W. HUGHES1, RONALD H. PETERSEN1, JUAN LUIS MATA2, NADEZHDA V. PSURTSEVA3, ALEXANDER E. KOVALENKO3, OLGA V. MOROZOVA3, EDGAR B. LICKEY 4, JOAQUIN CIFUENTES BLANCO5, DAVID P. LEWIS6, EIJI NAGASAWA7, ROY E. HALLING8, SEIJI TAKEHASHI9, M. CATHERINE AIME10, TOLGOR BAU11, TERRY HENKEL12 Abstract The genus Megacollybia, originally proposed for M. (Collybia) platyphylla, has traditional- ly been treated as monotaxic. A phylogenetic reconstruction based on ITS rDNA sequences indicates that several species are involved, with strong phylogeographic signal. Although morphological characters are largely qualitative, examination of basidiomata suggests that specimens included in discrete clades can be distinguished at the species level. On these bases (phylogenetic, morphological), several new taxa are proposed: M. clitocyboidea, M. texensis, M. fusca, M. subfurfuracea, M. rodmani (with f. murina) and M. marginata. Tricholomopsis fallax is transferred to Megacollybia; M. platyphylla remains the type species of the genus but appears to be restricted to Europe, Scandinavia and western and central Russia. Key words: Taxonomy, systematics, biogeography, phylogeny, phylogeography 1 Ecology and Evolutionary Biology, University of Tennessee, Knoxville, TN 37996-1100 USA. 2 Department of Biology, University of South Alabama, Mobile, AL 36688 USA 3 Komarov Botanical Institute, 2 Prof. Popov Street, St Petersburg, 197376 Russia 4 Department of Biology, Bridgewater College, Bridgewater, VA 22812 USA 5 Herbario FCME, Facultad de Ciencias UNAM, AP 86-95, C.P. 14391, Tlalpan, Mexico D.F., Mexico 6 262 CR 3062, Newton, TX 75966-7003 USA 7 Tottori Mycological Institute, 211 Kokoge, Tottori 689-1125, Japan 8 New York Botanical Garden, Bronx, NY 10458-5126, USA 9 1-3-10-3 Kanayama, Teine-ku, Sapporo, Hokkaido 006-0041, Japan 10 Department of Plant Pathology and Crop Physiology, Louisiana State University Agricultural Center, Baton Rouge, LA 70803 USA 11 Institute of Mycology, Jilin Agricultural University, 2888 Xincheng St., Changhun City, Peoples’ Republic of China 12 Department of Biological Sciences, Humboldt State University, Arcata, CA 95521 USA 1 K.H. HUGHES et al. Introduction Moncalvo, et al. (2002), in which Tricholomop- sis rutilans is found in a poorly supported clade By the time Kotlaba and Pouzar (1972) (24, /tricholomopsis) with Clavaria fusiformis, segregated Megacollybia, Collybia (Tricho- Marasmius rhyssophyllus and Collybia aurea, of lomopsis, Oudemansiella, Megacollybia) which all of the latter three can find more com- platyphylla had become a popular name in use fortable morphogeneric placements. Mega- throughout Europe and eastern North collybia platyphylla, conversely, is in a clade America. As long as workers from these (27, /hydropoid) with three Gerronema taxa, regions did not seriously collect in each oth- Clitocybula oculus, two Hydropus taxa and a ers’ territory this single name seemed applica- Porotheleum and these findings have recently ble for what was considered to be a been confirmed by the six-locus analyses of transAtlantic organism. The name was also Matheny et al. (2006). Tricholomopsis fallax applied to far east-Asian basidiomata which ITS sequences are most closely related to seemed to resemble those from Europe. Megacollybia platyphylla by blast search, then Comparing photos by Phillips (1981: 44, as to Clitocybula and Gerronema but not to Tricholomopsis platyphylla) of the mushroom Tricholomopsis sensu stricto. from Europe, by Phillips (1991: 46 under the Petersen and Gordon (1994) reported that same name) for the mushroom in North eastern North American Megacollybia had not America, and Halling and Mueller (2005: 151, yielded to monokaryon culture, while spores as M. platyphylla) for the mushroom in Costa from EuroScandinavian collections not only Rica, it is a wonder that the same epithet has underwent in vitro spore germination, but been applied for such disparate morphological crossed in culture and exhibited a tetrapolar forms. The photo from Huffman et al. (1989: mating system. Combining these data with Fig. 93) may show yet another taxon from the morphology, it was suggested that a more thor- midsection of North America. ough investigation of Megacollybia sensu lato Singer (1986) and Smith (1960) considered could be profitable. Any study of Megacolly- that Tricholomopsis sheltered a complex of bia, whether morphological or molecular, must about 30 species [T. rutilans (typus generis), T. start with knowledge of western European decora, etc.] of brightly colored, centrally stipi- Megacollybia platyphylla since the epithet was tate agarics fruiting on wood, but included, adopted from Persoon. Material from west-cen- sometimes as a separate section Platyphyllae tral Germany should serve as the foundation for (Singer, 1986: 253), T. platyphylla (typus sec- comparison of other collections. In the litera- tionis). Singer (1986) listed several names ture, good descriptions of the western European under sect. Platyphyllae, most of which he con- organism have been furnished by Boekhout (in sidered to belong in other genera or were Bas et al., 1999), Breitenbach and Kränzlin unpublished. Oblique reference to South (1991) and Gulden (in Hansen and Knudsen, American species of Platyphyllae, “...confirm- 1992). ing the position of the whole group within Our project had the following objectives: 1) Tricholomopsis” (Singer 1986: 252, ftnt) fur- to investigate whether a single species epithet nished no data. could accommodate many collections of The conjecture that Tricholomopsis and Megacollybia platyphylla from across its pur- Megacollybia are separate entities has been sup- ported range; 2) to determine infrageneric ported by molecular data, most convincingly by clades and their relationships within 2 Megacollybia (Agaricales) Megacollybia; 3) to explore whether phylogeo- Other herbarium acronyms are from Index graphic signal was present in a phylogenetic Herbariorum (http://sciweb.nybg.org/sci- reconstruction of the genus; 4) to propose and ence2/IndexHerbariorum.asp). TFB = number describe whatever additional taxa might be from Tennessee fieldbooks assigned to speci- necessary as a result of the study; and 5) to mens for documentation (i.e. notes, photos, explore the relationship of Megacollybia with etc.), supplanted by TENN numbers when Tricholomopsis fallax , a closely related accessioned into TENN herbarium. GSMNP = species reported from the North American Great Smoky Mountains National Park, strad- temperate western coast (British Columbia, dling the border between Tennessee and North Washington, Idaho, Oregon) and the dry Carolina, USA. In species descriptions, char- southwest (Arizona) . acters in Italics are considered diagnostic for Macro- and micromorphology were com- the taxon. bined with an ITS-based phylogenetic recon- Abbreviations of basidiospore statistics are struction to investigate collections from eastern as follows: x = number of spores measured; Q and western North America, Scandinavia, west- = spore length divided by spore width; Qm = ern and eastern Europe, western, central and far median Q value over the spore population; Lm = eastern Russia, east Asia, Central America and median spore length over the spore population. northern South America. Using the western Collections from which DNA sequences European organism as prototype, results were obtained are listed in Table 1. Specimens include: 1) the eastern North American organ- examined morphologically are listed under ism requires a new species epithet (M. each taxon in “Specimens examined.” rodmani); 2) temperate east Asia shelters a Basidiomata used for both purposes appear in species (M. marginata) closely related to but both places. distinct from the northern and western European organism; 3) northern South America Molecular protocols produces yet another species (M. fusca); and 4) Procedures for DNA extractions were there are at least three lineages represented described in Mata et al. (2007). PCR amplifica- across North America. Tricholomopsis fallax tion of the ribosomal ITS1-5.8S-ITS2 region falls within Megacollybia and transfer to was carried out with primers ITS1F and ITS4. Megacollybia is proposed. Thus, reflected in DNA sequencing followed manufacturer’s the ITS-based phylogeny, Megacollybia must directions for Big Dye Terminator mix (ABI) include at least seven species, with more to with forward primer ITS5 and reverse primer come as additional geographic areas are ITS4. Intermediate sequencing primers ITS2 explored. and ITS3 were used as necessary. Primers ITS2, ITS3, ITS4 and ITS5 are described in White et Materials and Methods al. (1990); primer ITS1F is described in Gardes and Bruns (1993). Cloning was accomplished Color notations in quotation marks are from with a Promega pGem-T vector and JM109 Ridgway (1912); those cited alphanumerically competent cells (Promega Corp., Madison, WI) (i.e. 7B3) are from Kornerup and Wanscher following manufacturer’s instructions when (1967). BF = bright field microscopy; PhC = direct sequencing did not resolve a sequence. phase contrast microscopy. TENN = herbarium Five clones per collection were sequenced. of the University of Tennessee, Knoxville. Restriction sites were determined by the GCG 3 K.H. HUGHES et al. Table 1 — GenBank Numbers for collections used in this study 4 Megacollybia (Agaricales) 5 K.H. HUGHES et al. 6 Megacollybia (Agaricales) 7 K.H. H UGHES 8 et al. crefers to clone number; ss refers to single
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