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Lepiota in California

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Lepiotain California: species with a hymeniform pileus coveringElse C. Vellinga1111 Koshland Hall 3102, Department of Plant andMicrobial Biology, University of California atBerkeley, Berkeley, California 94720-3102Abstract: EightLepiotaspecies with a hymeniformpileus covering that are known in California arepresented and discussed.Lepiota phaeodermaisdescribed as new; this species is characterized by adark pileus surface, which splits open into smallpatches around the umbo, and the absence of anannulus.Lepiota neophana, a species with an annulusand a closed brown pileus surface, andL. lilacea, witha conspicuous dark annulus, are both recorded forthe first time west of the Rocky Mountains. Theidentity ofL. neophanais reviewed and clarified basedon morphological and molecular data. The typecollection ofL. rufipeswas studied, andL. rufipeswas placed in synonymy withCystolepiota seminuda.Akey to the species is given.Key words:Agaricaceae, biodiversity,Callitropsismacrocarpa, nrITS and EF1-a sequences, taxonomy,type studiesINTRODUCTIONLepiota(Pers.) Gray is represented in the highlydiverse state of California by at least 25 species, 30%of which are still undescribed. This genus is lessspecies-rich thanLeucoagaricus(Locq.) ex Singer(Vellinga 2004a), which also contains many unde-scribed species in California. SeveralLepiotaspecieshave been described from the state in the past (e.g.Murrill 1912; Sundberg 1971, 1989; Vellinga 2007),but an overview of the genus is still lacking.The group of species with a hymeniform pileuscovering was the focus of this article. Basidiocarps ofthese species are small, with a pileus covering thateither covers the pileus homogeneously or splits openinto small patches or flat scales, depending on thepresence or absence of an intercellular matrix thatbinds the individual cells. An annulus can be presentor absent; the spores are ellipsoid or bullet-shaped,and small (less than 8mm long), and the spore wallsdo not react strongly with Melzer’s reagent andCongo red. One species has only one nucleus perspore (Ku¨hner 1945, Vellinga and Huijser 1999),whereas the other species have two, as is usual inLepiota. Cheilocystidia are present or absent. Themorphological differences among the species areoften subtle (Vellinga and Huijser 1999), but nrITSsequence data support the morphologically recog-nized species. However species with a hymeniformpileipellis do not form a monophyletic group(Vellinga 2003).This group has been accommodated in 1–3 sectionsbased on morphological characters, according tovarious authors. Singer (1986) put all species insectionCristatae(Ku¨ hner ex Wasser) Bon, regardlessof spore shape, but Bon (1993) placed them in threesections (in two subgenera) by distinguishing specieswith bullet-shaped spores (in sect.Cristatae) fromspecies with ellipsoid spores in subgenusParalepiotulaBon. He then divided this subgenus in two: sect.Integrellae(Ku¨ hner ex Bon) Bon for species with apileus covering that does not split open and only onenucleus per spore and sect.LilaceaeBon for specieswhose pileus covering breaks open into scales andhave binucleate spores.Phylogenetic analyses based on molecular datafailed to resolve many of these differences (Vellinga2003). However they clearly showed that species withbullet-shaped spores and a hymeniform pileus cover-ing, such asL. cristata, belong with ellipsoid-sporedspecies and a similar pileus covering and not withspecies with spurred spores and a trichodermal pileuscovering. The bullet-shaped spores arose once in thisclade of ellipsoid-spored species; the more pro-nouncedly spurred spores in the group of specieswith trichodermal or cutis-like pileus covering (L.boudieriBres.,L. castaneaQue´l.) evolved indepen-dently.The molecular data (Vellinga 2003) also did notsupport a separate section forL. rufipes, which differsfrom the other species in the uninucleate spores. Theisolated and unresolved positions ofLepiota lilaceaBres.,L. ochraceofulvaP.D. Orton andL. pyrochroaMalenc¸on, outside the core group of species with ahymeniform pileus covering, came as a surprise(Vellinga 2003) because morphologically they arenot distinct in essential characters from the rest of thehymeniform species.Despite several articles on species in this groupfrom California (Sundberg 1971, 1989; Vellinga2001a, b) naming problems still persist. The identityofL. neophanaMorgan, described from Preston,Ohio, (Morgan 1906) is at the center of the problemSubmitted 16 Jul 2009; accepted for publication 24 Sep 2009.1Corresponding author. E-mail: [email protected],102(3), 2010, pp. 664–674. DOI: 10.3852/09-180#2010 by The Mycological Society of America, Lawrence, KS 66044-8897664because at least three different interpretations of thisspecies have been published (Smith 1954, Sundberg1989, Bizio et al. 1993). This species again has beenstudied and is here redescribed from modernCalifornian collections, according to the originaldescription by Morgan (1906). As a result the speciesdescribed asL. neophana sensuSundberg (1989) is inneed of a new name and is described asL.phaeoderma.Seven of the eight species presented here co-occurin one small area, containing several Montereycypress (Callitropsis macrocarpa(Hartw.) D.P. Little;syn.Cupressus macrocarpaHartw.;Hesperocyparismacrocarpa(Hartw.) Bartel) plantations on an east-facing slope south of San Francisco, and one of theseseven is not known from any other location inCalifornia. These Monterey cypress groves are notablefor being rich in species ofLepiotaandLeucoagaricus(Vellinga 2004a).The species are treated in checklist format inalphabetical order, with a nomenclator and referenc-es to descriptions and illustrations in the literature.Species new to science and the newly recorded speciesare fully described. Additional morphological infor-mation is provided for several species, including thenumber of nuclei per spore, an important diagnosticcharacter. An identification key is provided to aidrecognition of the species.MATERIALS AND METHODSStandard methods for describing basidiocarps were appliedwith the terminology of Vellinga and Noordeloos (2001).Color annotations in macroscopic descriptions are fromMunsell2 soil color charts (1975). The notation [115,7,6]means that measurements were made on 115 spores inseven samples in six collections. These abbreviations areused: L for number of lamellae, l for number of lamellulaebetween two lamellae, avl for average length, avw foraverage width, Q


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