IMA FUNGUS · 5(2): 415–423 (2014) doi:10.5598/imafungus.2014.05.02.06 Phylogenetic and morphological comparison of Pluteus variabilicolor and ARTICLE P. castri (Basidiomycota, Agaricales) Tomaso Lezzi1, Alfredo Vizzini2, Enrico Ercole2, Vincenzo Migliozzi3, and Alfredo Justo4 1CSM AMINT. Via Q. Filopanti 2, I-00152 Roma, Italy 2Dipartimento di Scienze della Vita e Biologia dei Sistemi, Università di Torino, Viale P.A. Mattioli 25, I-10125 Torino, Italy; corresponding author e-mail: [email protected] 3Viale G. Marconi 196, I-00146 Roma, Italy 4Biology Department, Clark University, Lasry Biological Science Center, 950 Main St, Worcester, MA 01610, USA Abstract: Based on several collections from Italy, a detailed description (including also macrophotographs, Key words: microphotographs, and drawings) of the morphological characters of the poorly known Pluteus variabilicolor, Agaricomycetes originally described from Hungary, is provided. The analysis of the ITS sequences placed this species within the Italy P. leoninus clade of sect. Hispidoderma, in spite of the presence of clavate elements in the pileipellis. According ITS sequences to the molecular comparison of the type collections, Pluteus castri, a species recently described on the basis of pluteoid clade material collected in Japan and Central Russia, is reduced to a synonym of P. variabilicolor. Pluteus sect. Celluloderma sect. Hispidoderma subsect. Mixtini taxonomy Article info: Submitted: 17 October 2014; Accepted: 24 November 2014; Published: 28 November 2014. INTRODUCTION like elements in the pileipellis. Finally, Vellinga & Schreurs (1985) proposed an infrageneric classification for Pluteus differing from The genus Pluteus, typified by P. cervinus, includes saprobic Singer’s classificatory scheme only with respect to the placement agaricoid fungi characterized by free lamellae, no universal veil of species with non-metuloid cystidia and a filamentous (a partial veil may be present in a few species), pinkish spore-print, pileipellis: they reduced sect. Hispidoderma to a subsection of inverse hymenophoral trama, and inamyloid, cyanophilic spores Celluloderma, including only species with a hymenodermal or (Vellinga & Schreurs 1985, Orton 1986, Singer 1986, Vellinga trichodermal pileipellis, and established sect. Villosi for species 1990, Heilmann-Clausen 2012). It belongs to the pluteoid clade with a pileipellis which was a differentiated cutis. This system as circumscribed by several authors on the basis of molecular has not been widely accepted by later authors (e.g. Banerjee data (Moncalvo et al. 2000, 2002, Bodensteiner et al. 2004, 1992, Banerjee & Sundberg 1993, Citérin & Eyssartier 1998, Binder et al. 2006, Matheny et al. 2006, Justo et al. 2011a, b, Rodríguez & Guzmán-Dávalos 2001). Vizzini et al. 2011). The infrageneric taxonomy is based primarily Justo et al. (2011a, b) showed a general agreement between on the characteristics of the hymenial cystidia and pileipellis. the morphological subdivision of Pluteus and the molecular Historically, three sections based on morphological features phylogeny they obtained, even though: (1) some species are widely accepted (Homola 1975, Horak & Heinemann 1978, with non-metuloid pleurocystidia and a pileipellis which was a Orton 1986, Singer 1986, Citérin & Eyssartier 1998, Malysheva differentiated cutis (formerly sect. Villosi) were placed either et al. 2009, Minnis & Sundberg 2009, 2010): namely sect. in sect. Celluloderma, together with the species characterized Pluteus, with thick-walled (metuloid) pleurocystidia, usually by a hymenidermal pileipellis, or in sect. Pluteus with metuloid with apical hooks, and pileipellis as a cutis; sect. Hispidoderma bearing species; and (2) the division of sect. Celluloderma into with non-metuloid pleurocystidia, and a pileipellis consisting of subsections Mixtini and Eucellulodermini was not supported elongated, filamentous elements organized either as a cutis, by the phylogenetic analyses of the molecular data; species a hymeniderm or a trichoderm; and sect. Celluloderma with with a mixture of vesiculose-clavate and elongate elements in non-metuloid pleurocystidia and a pileipellis composed of the pileipellis were not monophyletic and were distributed over swollen clavate or spheropedunculate elements organized in sections Celluloderma and Hispidoderma. a hymeniderm, with transitions to an epithelium. Singer (1956, Here we fully describe, based on recent Italian collections, 1958, 1986) sometimes also recognized two subsections in the rarely reported Pluteus variabilicolor, which was originally section Celluloderma: Mixtini and Eucellulodermini, differing assigned to subsect. Mixtini of sect. Celluloderma by Babos respectively in the presence or absence of elongated cystidium- (1978). A phylogenetic analysis of ITS sequences was performed © 2014 International Mycological Association You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). Non-commercial: You may not use this work for commercial purposes. No derivative works: You may not alter, transform, or build upon this work. For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. VOLUME 5 · NO. 2 415 Lezzi et al. ARTICLE 416 1/100 Pluteus roseipes Estonia UDB011423 0.87/- Pluteus roseipes Estonia UDB011721 0.76/- Pluteus leoninus Estonia UDB011811 Pluteus leoninus Estonia UDB019490 0.8/78 Pluteus leoninus France HM562077 Pluteus leoninus Spain HM562045 0.85/- KC147679 /leoninus 0.94/85 Pluteus roseipes California 0.98/74 Pluteus roseipes California KC147681 Pluteus leoninus complex 0.82/- Pluteus leoninus Idaho HM562215 0.72/- 0.95/88 Pluteus flavofuligineus Michigan HM562187 Pluteus flavofuligineus New York HM562071 0.7/57 leoninus Pluteus aff. leoninus I Illinois HM562188 1/100 Pluteus aff. leoninus I Missouri HM562190 clade Pluteus aff. leoninus II Japan HM562139 Pluteus variabilicolor TL20130521-­‐01 Italy KP192911 1/100 0.98/84 Pluteus variabilicolor VM20130504-­‐01 Italy KP192914 Pluteus variabilicolor BP-­‐FN 56936 Isotype Hungary KP192912 Pluteus variabilicolor VM20111105-­‐01 Italy KP192913 /variabilicolor 1/100 Pluteus castri Holotype Japan HM562092 0.86/54 Pluteus castri Japan HM562099 1/100 Pluteus castri Russia FJ774077 1/100 Pluteus chrysaegis K13587 India JN603206 /chrysaegis Pluteus conizatus var. africanus Thoen5250 Dem Rep Congo HM562142 Pluteus pantherinus Japan HM562089 P. pantherinus Pluteus longistriatus Brazil HM562149 1/96 Pluteus longistriatus Brazil HM562158 1/100 Pluteus longistriatus Brazil HM562172 Pluteus longistriatus Missouri HM562082 1/95 Pluteus aff. semibulbosus Japan HM562090 Pluteus semibulbosus Russia FJ774080 0.98/75 Pluteus sp. RSPF Brazil 0330 JQ065028 Pluteus sp. UC 1861231 California KC147677 1/94 Pluteus heteromarginatus Florida HM562058 1/100 Uncultured fungus clone TSBF1 52 Brazil HQ257439 Pluteus velutinus K12851 India JN603205 1/100 1/100 Pluteus velutinus TNSF12365 Japan HM562114 Pluteus velutinus TNSF12372 Japan HM562127 /100 1 1/100 Pluteus plautus LE213024 Russia FJ774076 Pluteus plautus Spain HM562055 plautus/longistriatus 1/100 Pluteus granulatus Russia FJ774086 clade Pluteus granulatus Spain HM562048 1/100 Pluteus granularis Illinois HM562069 1/100 Pluteus granularis Illinois HM562189 umbrosus clade Pluteus umbrosus Spain HM562140 Pluteus seticeps HM562199 Pluteus diettrichii HM562143 Outgroups ( sect. Celluloderma) 0.07 expected changes per site Fig. 1. Bayesian tree analysis of Pluteus sect. Hispidoderma. BPP values (in bold) ≥ 0.7 and MLB values ≥ 50 % are shown on the branches. Clade names follow Justo et al. (2011a) and Pradeep et al. (2012). Newly sequenced collections are in bold. IMA FUNGUS Pluteus variabilicolor, phylogenetic and morphological comparison to: (1) provide molecular insights on its status and taxonomic shown). Alignments were generated using MAFFT (Katoh ARTICLE placement within Pluteus; and (2) assess whether this species, et al. 2002) with default conditions for gap openings and with a mixture of elongated and short elements in the pileipellis, gap extension penalties. The sequence alignments were was phylogenetically closer to species in sect. Hispidoderma or then imported into MEGA v. 5.10 (Tamura et al. 2011) for in sect. Celluloderma. manual adjustment. Pluteus diettrichii (HM562143) and The ITS sequence of the isotype collection was also obtained Pluteus seticeps (HM562199) were used as outgroup taxa to confirm our identifications of the recent collections. following Justo et al. (2011a) and Pradeep et al. (2012). Best-fit models were estimated by both the Akaike information criterion (AIC) and the Bayesian information criterion MATERIAL AND METHODS (BIC) with jModelTest v. 0.1.1 (Posada 2008) to provide a substitution model for the alignment. Phylogenetic analyses Morphology were performed using the Bayesian Inference (BI) and Microscopic descriptions are based on fresh and dried Maximum likelihood (ML) approaches. The BI was performed specimens. The optical microscope used was a Nikon with MrBayes v. 3.1.2 (Huelsenbeck & Ronquist 2001) with Eclipse