AR TICLE New Species of Auritella (Inocybaceae) From
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·!+! doi:10.5598/imafungus.2017.08.02.06 4 ( 1 1% .#%0 4' # 4$4 Auritella Inocybaceae ARTICLE 6*'64( P. Brandon Matheny1, Terry W. Henkel2, Olivier Séné3, Hailee B. Korotkin1, Bryn T.M. Dentinger4, and M. Catherine Aime5 1Department of Ecology and Evolutionary Biology, University of Tennessee, 569 Dabney Hall, Knoxville, TN 37996-1610, USA; corresponding author e-mail: [email protected] 2Department of Biological Sciences, Humboldt State University, 1 Harpst St., Arcata, CA 95521, USA 3Institute of Agricultural Research for Development, National Herbarium of Cameroon, PO Box 1601, Yaoundé, Cameroon 4Utah Museum of Natural History and Department of Biology, University of Utah, Salt Lake City, UT 84112, USA 5Department of Botany and Plant Pathology, Purdue University, West Lafayette, IN 47907, USA )# Two new species in the genus Auritella (Inocybaceae) are described as new from tropical 3*4 rainforest in Cameroon. Descriptions, photographs, line drawings, and a worldwide taxonomic key to the Africa described species of Auritella are presented. Phylogenetic analysis of 28S rDNA and rpb2 nucleotide Agaricales ?G=GLOGVB@B?=ZZG?@?B@XGB?@[GMHWX<ZGLG@EO?MGOEG?@HB@OBLFGB?ODEFGV@<Auritella occur in the ectomycorrhizas region comprising Cameroon and Gabon and constitute a strongly supported monophyletic subgroup Gilbertiodendron IE@HEL@HGZGL=? HWX<ZGLG@EOBLBXW?E?<[VB@B?=MM<D@?@HGO<L?MGOE[OE@W<[L=>GD<=?O<XXGO@E<L? Guineo-Congolian rainforest attributed to the two new species as well as the monophyly of Australian species of Auritella. This work systematics raises the known number of described species of Auritella to thirteen worldwide, four of which occur in tropics @D<MEOBX&[DEOB<LGEL@D<MEOBXLVEBBLVGEZH@EL@G>MGDB@GBLV@D<MEOBXDGZE<L?<[&=?@DBXEBHE?E?@HG[D?@ ?@=VW@<O<L[D>BLGO@<>WO<DDHEBX?@B@=?<[Auritella using molecular data. $1 Submitted: 20 July 2017; Accepted: 13 October 2017; Published: 18 October 2017. INTRODUCTION rainforest. In contrast, Australian species are dull in colour, ?GGDBXHBGB?=B@HBFE@BDG[GX@W@<>GL@<?G@<[FDEXX<?GEL Eleven species of the mushroom genus Auritella surface texture, the cheilocystidia are thin-walled, and most (Inocybaceae, Agaricales) are currently recognized. The (six of eight known species) are distributed in temperate generic name was originally published in 2006 and now forest regions. encompasses taxa discovered in Africa, Australia, and India During recent mycological expeditions to the Dja (Matheny & Bougher 2006a, b, 2017, Matheny et al. 2012). Biosphere Reserve in southeastern Cameroon, we collected Species of Auritella are united morphologically by their smooth > 250 morphospecies of putatively ECM macrofungi in and typically yellowish brown pigmented basidiospores, long monodominant forests of the ECM tree Gilbertiodendron or elongated cheilocystidia, necropigmented basidia, often dewevrei (Fabaceae subfam. Caesalpinioideae; e.g. Buyck @<=ZH \G?H BF?GLOG <[ MXG=D<OW?@EVEB BLV MBXG<@D<MEOBX et al. 2016, Castellano et al. 2016a, b). Several of the putative and Australian distribution. In addition, all known Auritella morphospecies referable to Inocybaceae were found to be species are putatively ectomycorrhizal (ECM), occurring on noteworthy as subsequent determinations placed these in or in soil in the vicinity of ECM trees of Myrtaceae, Fabaceae, Auritella. Here we describe two of these as new species. Casuarinaceae, and Dipterocarpaceae, and share a distinct This work raises the known number of described species phylogenetic position as one of seven major clades in the of Auritella [D<>&[DEOB @< [<=D F=@ DGO<ZLEG? B@ XGB?@ [G family Inocybaceae (Matheny 2009, Matheny et al. 2009). phylogenetic lineages that correspond to species, all of The genus, as currently known, has a restricted, partial which are known only from the Guineo-Congolian rainforests Gondwanan geographical distribution, occurring in tropical of Cameroon and Gabon in Central Africa. Phylogenetic regions of Africa and India and temperate and tropical analyses of 28S-rRNA, rpb2 and the ITS gene region areas of Australia. Tropical African and Indian species are presented, along with morphological descriptions, are distinguished by their typically long stipe, scaly pileus illustrations, and discussion of the new species relative and often scaly stipe surfaces, bright basidiome colours, to other species of Auritella recognized from Africa, India, trichoderm pileipellis, thick-walled cheilocystidia composed and Australia. Historical biogeographic scenarios are also of chains of hyphal cells, and occurrence in lowland tropical discussed. © 2017 International Mycological Association You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: 7<=>=?@B@@DEF=@G@HGI<DJEL@HG>BLLGD?MGOE[GVFW@HGB=@H<D<DXEOGL?<DF=@L<@ELBLWIBW@HB@?=ZZG?@?@HB@@HGWGLV<D?GW<=<DW<=D=?G<[@HGI<DJ Non-commercial: 7<=>BWL<@=?G@HE?I<DJ[<DO<>>GDOEBXM=DM<?G? No derivative works: 7<=>BWL<@BX@GD@DBL?[<D><DF=EXV=M<L@HE?I<DJ 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 8 · NO. 2 287 #'*et al. ;<=<;59= phylogenetic analysis. After homogenizing the taxon lists in F<@HVB@B?G@?IGO<LGD@GV@HG[XG?@<MHWXEM[<D>B@=?ELZ $$$ SeaView v3.2 (Galtier et al. 1996, Gouy et al. 2010). Basidiomata were collected in Cameroon during the Aug.– ITS data from 32 samples were aligned using Clustal X Sept. early rainy season of 2014 and the Nov.–Dec. late rainy 2.0.9 (Larkin et al. 2007) and manually adjusted in MacClade. ARTICLE season of 2016 from the Dja Biosphere Reserve, Northwest Sites deemed too ambiguous to align were removed before Sector, near the village of Somalomo, Upper Dja River Basin, phylogenetic analysis. Auritella foveata was used to root the within a 2 km radius of a base camp located at 3°21'29.8'' ITS tree based on the combined 28S+rpb2 ML tree result. N; 12°43'46.9'' W, 650 m a.s.l., in forests dominated by All ITS sequences used in this study are included in Table 1. Gilbertiodendron dewevrei (Peh et al. 2014). Descriptions of Models of molecular evolution for each data set used macromorphological features were made from fresh material a GTR model of a DNA substitution matrix and a GAMMA EL@HG[GXV(<X<=D?IGDGO<>MBDGVIE@HMXB@G?EL¨<DLGD=M model of rate heterogeneity under the Maximum Likelihood & Wanscher (1978) and are cited in parentheses (e.g. 7F7). (ML) criterion in RAxML v7.2.8 (Stamatakis 2006) or RAxML DG?H?MGOE>GL?IGDGMH<@<ZDBMHGVEL@HG[GXVXBF<DB@<DW v8.0.0 (Stamatakis 2014) following recommendations in the “L” refers to the number of lamellae that reach the stipe. After RAxML user manual. 1000 rapid bootstraps were performed. documentation basidiomata were dried with silica gel. Bootstrap proportions >70 % were considered as evidence of Dried basidiomata were sectioned and rehydrated in strong support for phylogenetic groupings. After determination 3 % KOH in order to examine and measure anatomical <[ L< ?@D<LZXW ?=MM<D@GV EL@GDZGLG O<L\EO@ F<@H BLV features following Matheny & Bougher (2006a, 2017) on a rpb2 gene regions were concatenated in MacClade and Nikon Eclipse 80i compound microscope using NIS Elements saved as a phylip formatted data set in SeaView. A RAxML (D) imaging software. In the taxonomic descriptions, mean analysis with 1000 bootstraps was performed again following basidiospore dimensions are italicized, and Q refers to the the procedures above after partitioning the data by the 28S quotient of the length of basidiospores divided by their width. region and each codon position. Due to high divergence of Holotypes and duplicate collections of African materials the ITS between A. aureoplumosa and A. foveata, the ITS are deposited in the Cameroon National Herbarium (YA); alignment was analyzed separately and not concatenated isotypes and other duplicates are deposited in HSC, TENN, with the 28S+rpb2 BXEZL>GL@ G=? [XG? BLV FEMBD@E@E<L? PUL and/or UT. @DGG [XG? BDG BBEXBFXG B@ http://mathenylab.utk.edu/Site/ Alignments_%26_Data_Sets.html or from the lead author =8#02.#%($#0# upon request. New DNA sequences have been submitted to @" GenBank. Procedures for DNA extractions, PCR, and direct sequencing are detailed in Judge et al. (2010) and Baroni & Matheny (2011). As in the latter work, we sequenced both ITS1 and <; ITS2 and the intervening 5.8S ribosomal RNA gene (ITS), the 5’ portion (initial 700–1350 bp) of the nuclear encoded Thirty-seven new sequences were submitted to GenBank large subunit ribosomal RNA (28S), and the most variable (accession nos. KT378200—KT378216, MF374757—MF374768, region between conserved domains six and seven of the MF474176, and MF476239—MF476241). These are shown in RNA polymerase II second largest subunit (rpb2 gene). PCR F<XVIE@HBVVE@E<LBXVG@BEX?ELBFXG HG[LBXO<LOB@GLB@GV and sequencing primers used in this work are the same as data set of 28S and rpb2 sequences included 37 taxa and those used and cited in Judge et al. (2010) and Baroni & 2155 included sites. The 28S partition included sequence Matheny (2011). data for all 37 taxa and 1434 sites. The rpb2 partition included sequence data for 27 taxa and 721 sites. =@#$"%#('*$" The genus Auritella was strongly supported as mono- ##$* phyletic consistent with prior studies with more extensive To infer the phylogenetic placement of Auritella species, we taxon sampling (Matheny et al. 2009, 2012) (Fig. 1). Within assembled separate alignments of 37 28S and 27 rpb2 DNA Auritella, three major subgroupings were recovered that sequences from on-going and much more taxonomically correspond to continental divisions: (1) a deeply diverging inclusive gene datasets in MacClade v4.08a (Maddison & lineage composed of a single branch represented by the Maddison 2005), including two 28S sequences (JQ657775, tropical Indian species, A.