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Five New Species of Inocybe (Agaricales) from Tropical India

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Five New Species of Inocybe (Agaricales) from Tropical India Mycologia ISSN: 0027-5514 (Print) 1557-2536 (Online) Journal homepage: http://www.tandfonline.com/loi/umyc20 Five new species of Inocybe (Agaricales) from tropical India K.P. Deepna Latha & Patinjareveettil Manimohan To cite this article: K.P. Deepna Latha & Patinjareveettil Manimohan (2016) Five new species of Inocybe (Agaricales) from tropical India, Mycologia, 108:1, 110-122, DOI: 10.3852/14-358 To link to this article: https://doi.org/10.3852/14-358 View supplementary material Published online: 20 Jan 2017. Submit your article to this journal Article views: 43 View related articles View Crossmark data Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=umyc20 Mycologia, 108(1), 2016, pp. 110–122. DOI: 10.3852/14-358 # 2016 by The Mycological Society of America, Lawrence, KS 66044-8897 Five new species of Inocybe (Agaricales) from tropical India K.P. Deepna Latha Species of Inocybe are characterized by brownish basi- Patinjareveettil Manimohan1 diomata, at times with a lilac or purplish tint, the coar- Department of Botany, University of Calicut, Kerala, 673 sely fibrillose or squamulose texture of the pileus and 635, India the stipe, brownish lamellae, a brown spore-print and occurrence on soil. Several species have thick-walled cystidia with crystalline, apical encrustations and some Abstract: Five new species of Inocybe, I. iringolkavensis, species have a distinctive odor. While several species I. keralensis, I. kuruvensis, I. muthangensis and I. wayana- have smooth basidiospores, many species are character- densis, are described from Kerala state, India, based on ized by gibbous or nodulose basidiospores. Although it morphological and molecular data. All are associated originally was placed in the Cortinariaceae (Kuyper with trees belonging to Dipterocarpaceae. Inocybe irin- 1986, Singer 1986), recent phylogenetic analyses sup- golkavensis is characterized by nodulose to somewhat port Jülich’s (1982) placement of Inocybe in its own – stellate basidiospores, 1 4-spored basidia, and caulo- family, the Inocybaceae (Matheny 2005, 2009). Accord- cystidia restricted to the stipe apex. Inocybe keralensis ing to Kirk et al. (2008), about 500 species of the genus has a yellowish brown pileus, lamellae with whitish, ser- are described worldwide and new species are discov- rate edges, smooth, ellipsoidal basidiospores and a ered on a regular basis. Although most species are duplex pileipellis with the superficial hyphae devoid known from temperate and arctic-alpine regions, of encrustations and encrusted hyphae beneath. The some are known from the tropics (Matheny et al. 2009). diagnostic features of I. kuruvensis include a dark Although India encompasses some of the biologically brown pileus, stipe with a whitish base and grayish richest and most endangered terrestrial ecoregions brown, floccose-fibrillose surface, nodulose basidios- (Mittermeier et al. 2005), several areas remain unex- pores with saddle-shaped projections and faintly plored or underexplored for many groups of fungi. encrusted paracystidia with refractive contents. Violet Available literature on Indian agarics consists mostly basidiomata with a rimose, hygrophanous pileus, den- of isolated and sporadic records. The situation in Ker- sely pruinose stipe with a marginate-bulbous base, ala state, a narrow strip of land on the southwestern cor- and nodulose basidiospores are the major features of ner of India with a tropical, maritime and monsoonal I. muthangensis. Inocybe wayanadensis is characterized climate, is somewhat different. The diversity of agarics by small, whitish basidiomata, a viscid pileus with a of this region was explored intensively in the past three rimulose surface, a densely pruinose and fibrillose decades resulting in a recent checklist included 616 stipe with a marginate-bulbous base, nodulose basidios- species in 112 genera (Farook et al. 2013). Mono- pores, thick-walled pleuro- cheilo- and caulocystidia graphic accounts of several agaric genera were pub- and an ixotrichoderm-type pileipellis. The phyloge- lished, for instance Hymenagaricus (Heinemann and netic relationships of these new species are inferred Little Flower 1984), Micropsalliota (Heinemann and from an analysis of nuc rDNA sequences of the inter- Leelavathy 1991), Entoloma (Manimohan et al. 1995, nal transcribed spacers (ITS1-5.8S-ITS2) and the 28S 2006), Psilocybe (Thomas et al. 2002), Hygrocybe (Leelav- gene. Except I. keralensis, which belongs to the Pseu- athy et al. 2006), Lepiota (Kumar and Manimohan dosperma clade, all other species belong to the Ino- 2009a), Leucocoprinus and Leucoagaricus (Kumar and cybe clade. This study represents the second report Manimohan 2009b). Remarkably most of the forested of an Inocybe species (I. muthangensis) that combines areas of Kerala include parts of the western Ghats, violet basidiomata with nodulose basidiospores. one of the biodiversity hotspots of the world. To date Key words: Basidiomycota, Inocybaceae, phylogeny, 19 species of Inocybe were reported from Kerala (Farook systematics, tropical agarics et al. 2013). During our studies on the diversity of Ino- cybe in this region, we found five new species that are INTRODUCTION described here based on both morphology and mole- The genus Inocybe (Fr.) Fr. (Inocybaceae, Agaricales) is cular phylogenetic results. one of the most speciose genera of gilled mushrooms and is noteworthy for the ectomycorrhizal ecology MATERIALS AND METHODS and toxicity of most of its species (Matheny 2009). Morphology.—Specimens examined were collected from Submitted 31 Dec 2014; accepted for publication 17 Aug 2015. natural forests and other forested lands. Microscopic exami- 1 Corresponding author. E-mail: [email protected] nations were made on dried material stained with 1% 110 Published online 20 Jan 2017 DEEPNA LATHA ET AL.: NEW INDIAN SPECIES OF INOCYBE 111 aqueous solution of Congo red and mounted in 3% aqueous with RAxML 8.0.26 (Stamatakis 2014) employed in raxml- KOH. Color and pigment topography of cells and hyphae GUI 1.3.1 (Silvestro and Michalak 2012), implementing GTR were observed on tissues mounted in water. For evaluation +I+G model and executing 1000 rapid ML bootstrap repli- of the range of spore dimension, 20 basidiospores from one cates. Bayesian inference (BI) was performed with MrBayes specimen of each collection cited were measured. Basidio‐ 3.2.1 (Ronquist and Huelsenbeck 2003), also implementing spore measurements include both the hilar appendix and the GTR+I+G model. A general time reversible model includ- the nodules. In the description of the basidiospores, Q repre- ing a proportion of invariable sites and gamma distributed sents the range of ratio of basidiospore length/width calcu- rate heterogeneity (GTR+I+G) model of molecular evolution lated for each basidiospore and Qm is the average of the Q was selected with TOPALi 2.5 (Milne et al. 2009). For BI values. Alphanumeric color codes from both Kornerup and four Markov chains were run 3 000 000 generations, sam- Wanscher (1978) (e.g. 5E7) and the Online Auction Color pling every 100th step, with two independent runs per analy- Chart (Anonymous 2004) (e.g. OAC768) accompany color sis. The first 25% trees were burned before calculating names in the descriptions. All examined collections except posterior probability (PP) values. Bootstrap values above one are deposited at Kew (Mycology) Herbarium, and the 70% and Bayesian posterior probability values above 0.95 accession numbers (e.g. K(M) 191734) are indicated. One were considered significant. The aligned sequence data collection is presently maintained in the personal herbarium matrix was deposited in TREEBASE (http://purl.org/phylo/ of the second author. treebase/phylows/study/TB2:S16895). The phylogeny from BI and ML analyses were displayed with FigTree 1.4.2 (Ram- — DNA extraction, PCR, and sequencing. Genomic DNA was baut 2014). extracted from dried holotypes of the five new Inocybe species employing the procedure described by Izumitsu et al. (2012). The nuc rDNA sequences of the internal transcribed spacer RESULTS 5 (ITS ITS1-5.8S-ITS2) and the 28S gene were analyzed. Pri- — mers LROR and LR7 (Vilgalys and Hester 1990) were used Results of BLAST query and phylogenetic analysis. A com- for amplifying the 28S. The ITS was amplified with the pri- parison of ITS and 28S sequences of the five new spe- mer pair ITS1 and ITS4 (White et al. 1990). Amplification cies of Inocybe described here with nucleotide 2 reactions were performed in a GeneAtlas thermal cycler sequences of taxa available in GenBank suggests that (Astec, Fukuoka, Japan). Amplified PCR products were puri- they have distinct ITS and 28S sequences. The closest 2 fied with column purification (GeneJet PCR purification hits for the five new Inocybe species in the MegaBLAST kit, Thermo Fisher Scientific, Mumbai, India) as per manu- queries of the GenBank nucleotide database using the ’ facturer s guidelines and were subjected to automated DNA ITS and 28S sequences are provided (TABLE I). sequencing on an ABI3730xl DNA analyzer (Applied Biosys- Molecular analyses yielded phylogenetic trees that tems, Foster City, California) using the same primers. The showed the relative placement of the five new species. generated sequences were edited manually with BioEdit 7.0.9.0 (Hall 1999). The edited sequences were used for The phylogeny inferred from both maximum likeli- BLAST queries in the GenBank database (www.ncbi.nlm. hood and Bayesian analyses
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