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Muñoz et al. ARTICLE 100 Leucoagaricus flammeotinctoides GU136173 100 78 Leucoagaricus flammeotinctoides GQ258476 Leucoagaricus flammeotinctoides AY243620 98 100 Leucoagaricus pyrrhophaeus GU136199 Leucoagaricus pyrrhophaeus GQ258473 100 75 Leucoagaricus pyrrhulus GQ258474 Leucoagaricus pyrrhulus GU136201 100 Leucoagaricus erythrophaeus GQ258468 97 Leucoagaricus erythrophaeus GQ258470 Lepiota roseifolia GQ203805 Leucoagaricus decipiens GQ203803 100 Leucoagaricus pardalotus GU136202 Leucoagaricus pardalotus GQ258479 100 Lepiota decorata GU136197 88 Lepiota decorata AY243645 68 Leucoagaricus ionidicolor AY176415 Leucoagaricus marriagei GQ329049 100 Leucoagaricus sp. Weber 6019 AY243633 100 Leucoagaricus brunnescens GQ203804 89 100 Leucoagaricus georginae AY176413 Leucoagaricus georginae GU136198 Leucoagaricus jubilaei AY243635 98 Leucoagaricus badhamii GQ329056 78 Leucoagaricus sp. ecv2484 GU136182 100 Leucoagaricus dyscritus GU136181 Leucoagaricus dyscritus GU136180 100 Leucoagaricus hesperius GU139789 Leucoagaricus hesperius GU139788 Lepiota fuliginescens GU136183 98 Lepiota fuliginescens GU136185 52 Lepiota fuliginescens GU136189 Lepiota flammeotincta GU136170 100 Leucoagaricus sp. Vellinga 2746 AY176440 95 Lepiota flammeotincta GU136167 54 Lepiota fuliginescens GU136187 Leucoagaricus sp. Vellinga 2619 AY243637 Lepiota fuliginescens GU136188 Leucoagaricus sp. Huijser s.n. AY243643 Leucoagaricus adelphicus GQ258478 100 68 Leucoagaricus adelphicus AY243622 Piloselli Leucoagaricus adelphicus AY243624 100 Leucoagaricus pilatianus GQ329040 Leucoagaricus pilatianus GQ329057 85 93 Leucoagaricus cupresseus GQ258477 95 Leucoagaricus cupresseus GU136194 87 Leucoagaricus cupresseus AY243629 Leucoagaricus cupresseus AY243627 Leucoagaricus cupresseus AY243632 Leucoagaricus variicolor coll. GM-2485 JX880032 100 Leucoagaricus variicolor coll. AH 40328 (holotype) JX880030 Leucoagaricus variicolor coll. GM-2486 JX880033 Leucoagaricus variicolor coll. GM-2454 JX880031 89 Leucoagaricus rubrotinctus JN944081 58 Leucoagaricus rubrotinctus FJ481050 100 Leucoagaricus rubrotinctus JN944082 98 Leucoagaricus sublittoralis AY176442 Rubrotincti 100 Leucoagaricus littoralis GQ329041 Leucoagaricus wychanskyi AF482874 Cystolepiota seminuda AY176350 0.06 Fig. 1. Maximum Likelihood phylogram obtained from the ITS (ITS1-5.8S-ITS2) sequence alignment of Leucoagaricus spp. Cystolepiota seminuda was used as outgroup taxon. MLB values over 50 % are given above branches. Newly sequenced collections are in bold. excluded) to the basal septum. Microscopic pictures were taken on a Moticam 2500 digital camera connected to a Motic BA300 microscope. Colour notations for the macroscopic descriptions are from Munsell (1994), hereafter shortened as Mu. Herbarium acronyms follow Index Herbariorum, except for GM and AC that refer to the personal herbaria of Guillermo Muñoz and Agustín Caballero. The type collection is housed at AH. The name and description of the new species are deposited in MycoBank (Crous et al. 2004). DNA extraction, PCR amplification, and DNA sequencing Genomic DNA was isolated from 1 mg of a dried herbarium specimen from four collections (AH-40328, GM-2454, GM- 2485, and GM-2486), using the DNeasy Plant Mini Kit (Qiagen, Milan) according to the manufacturer’s instructions. Universal primers ITS1F/ITS4 were used for the ITS region amplification (White et al. 1990, Gardes & Bruns 1993). Amplification reactions were performed in a PE9700 thermal cycler (Perkin-Elmer, Applied Biosystems) following Vizzini et al. (2011). The PCR products were purified with the AMPure XP kit (Beckman) and sequenced by MACROGEN (Seoul, Republic of Korea). The sequences were submitted to GenBank (http://www.ncbi.nlm.nih.gov/genbank/) and their accession numbers are reported in Fig. 1. Sequence alignment and phylogenetic analysis The sequences obtained in this study were checked and assembled using Geneious v. 5.3 (Drummond et al. 2010) and compared to those available in the GenBank database by using the Blastn algorithm. Based on the Blastn results, sequences were selected according to the outcomes of recent phylogenetic studies on Leucoagaricus (Vellinga 2010, Vellinga et al. 2010, 2011). Alignments were generated using MUSCLE (Edgar 2004) with default conditions for gap openings and gap extension penalties. The alignment was then imported into MEGA v. 5.0 (Tamura et al. 2011) for manual adjustment. The phylogenetic analysis was performed using the Maximum Likelihood (ML) approach. Following Vellinga (2010) and Vellinga et al. (2010), a Cystolepiota seminuda sequence (AY176350) was used as outgroup. ML estimation 118 ima fUNGUS
A new Leucoagaricus species from Spain ARTICLE Fig. 2. Leucoagaricus variicolor. Macroscopic characters. A–C. Fresh basidiomata in situ. D. Herbarium specimens. A, D from AH 40328 (holotypus); B from GM-2454; C from GM-2485. Bars = A–B = 50 mm; C–D = 20 mm. was performed through RAxML v. 7.0.4 (Stamatakis 2006) with 1000 bootstrap replicates (Felsenstein 1985) using the GTRGAMMA algorithm to perform a tree inference and search for a good topology. Support values from bootstrapping runs (MLB) were mapped on the globally best tree using the “-f a” option of RAxML and “-x 12345” as a random seed to invoke the novel rapid bootstrapping algorithm. Only MLB over 50 % are reported in the resulting tree (Fig. 1). RESULTS Phylogenetic analysis The amplification of the ITS regions was successful for the four specimens, yielding a PCR product of about 700 bp. The ITS data matrix comprises a total of 59 sequences (including 55 from GenBank). In the obtained ML phylogram (Fig. 1), our four sequences of the new Leucoagaricus clustered together and are distinct and basal to all the existing sequences of previously sequenced species of section Piloselli. Taxonomy Leucoagaricus variicolor G. Muñoz, A. Caball., Contu & Vizzini, sp. nov. MycoBank MB801565 (Figs 2–4) Etymology: The specific epithet “variicolor” refers to the highly variable colours of the pileus surface depending on fresh or dry conditions. Diagnosis: Differs from every other described species of Leucoagaricus sect. Piloselli in having a cream to pink to eggyellow, or ochre pileus in fresh basidiomata that becomes dark rose in the herbarium, the presence of universal veil remnants on the pileus surface and stipe base, subglobose to broadly ellipsoid spores, pyriform to spheropedunculate cheilocystidia, and a unique ITS sequence. Type: Spain: Aragón: Zaragoza, “José Antonio Labordeta” (antea: “Parque Grande”) park, UTM 30TXM7511, N 41.634 W 0.894, alt. 225 m, on argillose-sandy soil, in Pinus halepensis litter, 3 Dec. 2011, G. Muñoz (AH 40328 – holotype; GM- 2453 and AC-4896 – isotypes). Description: Pileus 40–100 mm wide, when young hemispherical-convex to hemispherical, expanding to subplane, finally plane, without an umbo, with an entire, slightly exceeding margin, involute or incurved in young stages; pileus surface dry, almost smooth to finely feltedfibrillose, variable in colour, starting from whitish or pale cream or egg-yellow to ochre (Mu 2.5Y 8/1-2; Mu 5Y 8/1 “White”; Mu 2.5Y 8/2-4 “Pale yellow”) to orange-pink (Mu 5YR 6/6-8 “Yellowish red”; Mu 2.5YR 5/6-8 “Red”); in adult stages volume 3 · no. 2 119
Page 1 and 2: The Global Mycological Journal Volu
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Novel species of Aspergillus sect.
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