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Nova Hedwigia Vol. 96 issue 1–2, 167–180 published online November 8, 2012

Article

© 2012 J. Cramer in Gebr. Borntraeger Verlagsbuchhandlung, Stuttgart, www.borntraeger-cramer.deGermany. DOI: 10.1127/0029-5035/2012/0056 0029-5035/2012/0056 $ 3.50

Two new resupinate Thelephorales (Basidiomycota, Agaricomycetes) from Guinea (West Africa)

Nourou Soulemane Yorou1,2*, Moussa Diabaté3 and Reinhard Agerer1

1 Department for Biology I, Division of Organism Biology: Mycology, Ludwig-Maximilians- Universität München, Menzinger Str. 67, D-80638 Munich, Germany2 Faculty of Agronomy, University of Parakou, BP 123, Parakou, Benin3 Herbier Serg. de Guinée, Centre Régional de Recherche Agronomique pour la Guinée Forestière CRRA Sérédou / IRAG/CRA, BP 1523, Conakry, Guinee

With 5 figures

Yorou, N.S., M. Diabaté & R. Agerer 2012: Two new resupinate Thelephorales (Basidiomycota, Agaricomycetes) from Guinea (West Africa). – Nova Hedwigia 96: 167–180.

Abstract: Tomentella minispora and T. baroensis are described as new species from West Africa based on anatomical evidence, ITS rDNA sequence analyses and molecular-phylogenetic inference. T. minispora is characterized by subglobose to globose small basidiospores with size ranging from 5.5 µm to 7.5 µm, short aculei of up to 0.8 µm and strongly inflated subhymenial hyphae (up to 23 µm). Both, sequence analyses and molecular-phylogenetic inference indicate the proximity of T. minispora to the clampless species T. amyloapiculata and T. fuscocinerea. Tomentella baroensis is characterized by the presence of extremely large young basidia (up to 20 µm wide), ellipsoid to lobed basidiospores of 8–13 µm and short aculei not exceeding 1 µm long. So far such large young basidia are known only from T. terrestris. However, ITS rDNA sequence analyses revealed that T. baroensis deviates from T. terrestris by at least 17 to 18%. Sequence analyses suggest T. ellisii as the most similar species to T. baroensis, whilst molecular-phylogenetic inference places it close to T. pisoniae and T. hjortstamiana, together with T. ellisii originating from deeper branch of the clade. Detailed anatomy and original iconographs are provided for both new species. Anatomical comparisons are made between the newly described species with their anatomically and molecular-phylogenetically close ones.

Key words: Anatomy, ITS rDNA sequence analyses, molecular-phylogeny, Fungi, West Africa.

Introduction

Resupinate Thelephorales (Amaurodon J.Schröt., Tomentellopsis Hjortstam, Pseudo-tomentella Svrček and Tomentella Persoon ex. Pat. (Kõljalg 1996) belong to the Class

*Corresponding author; e-mail: n.s.yorou@bio.lmu.de

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Agaricomycetes (Basidiomycota, Fungi) and have a world-wide distribution (Kõljalg 1996, Larsen 1965, 1967, Malençon 1952, Martini & Hentic 2002, Melo et al. 2006, Stalpers 1993, Suvi et al. 2010, Tedersoo et al. 2008, Yorou 2008). Still, Thelephorales have been overlooked in tropical Africa for long time. The first record/documentation of this fungal group in tropical Africa is probably that one of Martini & Hentic (2002). However, numerous investigations have been undertaken recently to monograph and document tropical African tomentelloid fungi (Gardt et al. 2011, Suvi et al. 2010, Yorou 2008, Yorou & Agerer 2007, 2008, Yorou et al. 2007, 2011a,b,c). Many new species have been described, attesting the high, but still unknown specific diversity of this fungal group in tropical Africa.

In the present paper, we describe two new Tomentella species collected in Guinea, a country we predict to host the highest but still unexplored fungal diversity in West Africa. Three new species have been previously described from the same country (Yorou et al. 2011a,b).

Materials and methods

Specimen Sampling, light microScope StudieS and drawingS: Specimens were collected during the raining season of 2009 under native trees in the forest reserve of Baro (Upper Guinea). Preliminary notes have been recorded using fresh materials and were dried afterwards by mean of a propane gas heated field dryer. Dried specimens were then herbarized under following labels SYN 2273 and SNY 2282. Colour codes of the dried basidiocarps are given according to Kornerup & Wanscher (1978). Type materials are deposited in M (Thiers 2012). We refer to Yorou & Agerer (2011a,b) and Yorou et al. (2011a,b,c) for protocols of light microscopy.

dna extraction, amplification and Sequencing: DNA was extracted using small pieces of dried basidiocarps. Protocol of DNA extraction, amplification and sequencing of the ITS gene is detailed in Yorou et al. (2011a,b,c). ITS rDNA sequence of each investigated specimen is deposited in GenBank NCBI under following accession numbers JQ814474 and JQ814475.

Sequence edition and phylogenetic analySeS: We refer to Yorou et al. (2011a,b,c) for sequence edition and analyses. The most similar sequences of each generated consensus sequence were searched for in UNITE (Abarenkov et al. 2010, Kõljalg et al. 2005) using the "BLASTN" search option. The search was extended to the GenBank EMBL and DDJB by activating the UNITE + INSD. option. The 15 sequences identified to species level were downloaded. The most similar sequences obtained were completed by that one published recently by Yorou et al. (2011a,b,c) from West Africa and those from the Seychelles (Suvi et al. 2010). Sequences were automatically aligned in BioEdit v7.0.5. (Hall 2005). Our final dataset encompasses a total of 84 ITS rDNA thelephoroid sequences with a length of 500 characters. The best scoring tree was obtained by running a fast bootstrap search of 1000 pseudoreplicates using the program RAxML v.7.0.4 (Stamatakis et al. 2008). For details of molecular-phylogenetic inference we refer to Yorou et al. (2011a,b).

Results

itS rdna SequenceS-baSed analySeS of the new SpecieS: The BLASTN search in UNITE + INSD Using the sequence of specimen SYN 2273 displayed a total of 50 best matches. Of all 50 best matches, the most similar (identity rate of 88%, Query coverage 95%) fully identified species is Tomentella amyloapiculata Yorou (EF507284). The following best matches are composed of 49 sequences of unidentified environmental

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samples with identities ranging from 85 to 87%. Sequence analyses, coupled with molecular-phylogenetic placement and anatomical characteristics (see below) suggest that our specimen SYN 2273 is a new species, and consequently is described below as Tomentella minispora. Similarly, T. ellisii (Sacc.) Jülich & Stalpers is from the 50 best matches the only fully identified most similar taxon to specimen SYN 2282. All three sequences HQ406823, AB634268 and UDB0003326 assigned to T. ellisii range among the top 50 with similarities of 88–89% (Query coverage 90–95%). The following most similar sequences are those of uncultured environmental sequences with an identity of 88–95%. We combined these sequence data, the molecular-phylogenetic placement, anatomical studies, and discuss the results with the respective features of similar taxa to describe specimen SYN 2282 as a new species under the name T. baroensis.

molecular phylogenetic placement of the new SpecieS: The final dataset submitted for molecular-phylogenetic analysis has 253 distinct alignment positions. The proportion of gaps and completely undetermined characters is estimated at 0.14%. The best scoring tree found has a maximal likelihood of ML = -2431.386475 (tree length = 3.006210). Substitution rates have been estimated as follow: A↔T: 0.739931, C G: 0.569115, CT: 5.942972, G T: 1.000000 with following bas frequencies: freq (A): 0.223910, freq (C): 0.247349, freq (G): 0.242109, freq (T): 0.286632. The topology of all three generated by the RAxML search is similar. Only the most likely tree is presented here (Fig. 1).

Molecular-phylogenetically, Tomentella minispora is placed in a terminal clade, forming a sister species to T. fuscocinera (Pers.) Donk (sequence accession number AF272942) with a strong bootstrap support of 89. Both species form a sister clade of a still unidentified Tomentella species from Benin (bootstrap support of 70). The recently described new species T. amyloapiculata (sequence accession number EF507262) forms a sister species to all three. The clade composed of the four species (T. minispora, T. fuscocinerea, T. amyloapiculata and Tomentella sp.) is moderately supported by a bootstrap value of 73.

As far as Tomentella baroensis is concerned, it forms a sister species of the group composed of T. pisoniae Suvi & Kõljalg (accession number FM244908) and T. hjortstamiana Suvi & Kõljalg (accession number AM412303) with a strong bootstrap support of 86. A sister of this group (T. pisoniae, T. hjortstamiana and T. baroensis) is Tomentella ellisii (accession AF272913 and UDB003326). T. guinkoii Yorou (accession number JF520430, newly described from Burkina Faso) represents again a sister species of the group comprising the aforementioned species. This internal clade is supported by a very strong bootstrap value of 96.

Description of the new species

Tomentella minispora Yorou sp. nov.

Genbank NCBI, Accession number JQ814474.

Mycobank MB 566334

Basidiocarpa resupinata, tenuis, ad 0.5 mm alta, adherens, crustosa usque ad arachnoidea, continua. Hymenium subtile granulosum, continuum, cinereobrunneum usque ad brunneum. Subiculum tenue,

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Fig. 1. Maximum Likelihood tree showing the placement of the new species within the genus Tomentella. Bootstrap values higher than 50% are shown above the branches. Accession numbers (from UNITE or NCBI) and country of origin of vouchers are indicated after species names. Some clades have been collapsed (black triangles) in order to reduce the expanse of the tree.

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arachnoideum, coloratum ut hymenium, marginibus sterilibus subdeterminatis, palliodore coloratis quam hymenium, farinosum

Rhizomorphae absentes.

Hyphae subiculi fibuligerae, septa simplicia deficientia, 3–5.5(6) µm in diametro, plerumque regulares, hyphae in 2,5% KOH infrequenter sinuosae, in 2.5% KOH infrequenter inflatae, ramificationes cruciformes et anatomosae deficientes, plerumque tenui- ad subcrassi- (0.4–0.5 µm), interdum crassitunictae (0.5–1 µm), in aqua et in 2.5% KOH incoloratae usque ad tenue subluteae, in aqua nunnullae hyphae densis incrustationibus, incrustationes in 2.5% KOH permanentes, nunnullae hyphae in superficie crystallis magnis rufobrunneis (usque ad 1 µm diametro), hyphae non cyanescentes, non congophilae, non amyloideae.

Hyphae subhymenii fibuligerae, 6–13(15) µm in diametro, breves, plerumque usque ad 19(23) µm inflatae, plerumque tenui- usque ad subcrassitunicatae (0.2–0.5 µm), infrequenter crassitunicatae (0.5–1 µm), nunnullae hyphae in aqua et in 2.5% KOH densis incrustationibus, interdum in superficie crytallis fuscobrunneis, hyphae in aqua et in 2.5% KOH incoloratae usque ad tenue subluteae, non cyanescentes, subcongophilae, subcyanophilae, non amyloideae.

Cystidia absentia.

Basidia fibuligera, (15)18–30(35) µm longa, apicibus 7–13(15) µm, basibus 6 – 8(8.5) µm, utriformia, non stipitata, raro sinuosa, septis transversis infrequentibus, basidia in aqua et in 2.5% KOH densis incrustationibus, guttulis numerosis, in aqua et in 2.5% KOH incolorata usque ad tenue sublutea, non cyanescentia, non congophila, non cyanophila, non amyloidea, 4-sterigmatica, sterigmatibus brevibus, 4–5 µm longis, basaliter 1.5–3 µm latis.

Basidiosporae (5.5)6–7(7.5) × 5–6.5(7) µm in aspectu frontali,(5)5.5–7 × 5.5–6.5(7) µm in aspectu laterali, subglobosae usque ad irregulariter globosae in aspectu frontali et laterali, echinulatae; aculeis brevibus (usque ad 0.8 µm), numquam furcatis; basidiosporae in aqua et in 2.5% KOH tenue subluteae, non cyanescentes, non congophilae, non cyanophilae, non amyloideae.

Chlamydosporae absentes.

Basidiocarp resupinate, thin, up to 0.5 mm, adherent to the substrate, crustose to arachnoid, continuous. Hymenophore finely granulose, continuous, hymenium greyish brown (6D3) to brown (6E4), subiculum very thin, arachnoid, concolorous with the hymenium, sterile margin nearly determinate, paler than the hymenophore, farinaceous.

Rhizomorphs absent.

Subicular hyphae clamped, simple septa absent, 3–5.5(6) µm wide, usually regular (Fig. 2), in water and in 2.5% KOH rarely sinuous, in 2.5% KOH rarely inflated, cross-shaped branching and anastomoses absent, commonly thin- to slightly thick-walled (0.2– 0.5 µm), sometimes thick-walled (0.5–1 µm), in water and in 2.5% KOH colourless to very light yellow, some hyphae strongly incrusted in water, incrustation remaining in 2.5% KOH, showing big yellow-red to red-brown crystals (up to 1 µm diameter) on some hyphae, hyphae not cyanescent, not congophilous, not cyanophilous, not amyloid.

Subhymenial hyphae clamped, 6–13(15) µm wide, short, commonly inflated up to 19(23) µm (Fig. 3), usually thin- to slightly thick-walled (0.2–0.5 µm), rarely thick-walled (0.5–1 µm), some in water and in 2.5% KOH strongly incrusted, sometimes with yellow-red to red-brown crystals on the surface, hyphae in water and in 2.5% KOH colourless to light yellow, not cyanescent, slightly congophilous, slightly cyanophilous, not amyloid.

Cystidia absent.

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Basidia clamped at base, (15)18–30(35) µm long, 7–13(15) µm at apex, 6–8(8.5) µm at base, very short and stubbed, utriform, not stalked, rarely sinuous, rarely with transverse septa, some basidia in water and in 2.5% KOH strongly incrusted, basidia commonly with numerous oil droplets, in water and in 2.5% KOH colourless to very light yellow, not cyanescent, congophilous, cyanophilous, not amyloid, 4-sterigmate, sterigmata short, 4–7 µm long and 1.5–3 µm at base.

Fig. 2. Tomentella minispora. a. Basidiospores in lateral view, b. Basidiospores in frontal view, c. Section through the basidiocarp (from SYN 2273). Bar = 10 µm.

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Basidiospores (5.5)6–7(7.5) × 5–6.5(7) µm in frontal face (5)5.5–7 × 5.5–6.5(7) µm in lateral face, subglobose to irregularly globose in frontal and lateral views, echinulate, aculei short (up to 0.8 µm), never forked, basidiospores in water and in 2.5% KOH very light yellow, not cyanescent, not congophilous, not cyanophilous, not amyloid.

Chlamydospores absent.

material Studied and habitat: Holotype: Guinea, upper Guinea, Province of Kakan, Prefecture of Kouroussa, Forest Reserve of Baro, along the road between Kouroussa and Kankan, N10°57' 0.94", W009°64' 60.0", forest dominated by Caesalpiniaceae (Isoberlinia doka Craib & Stapf and Anthonota crassifolia Baill.), Phyllantaceae (Uapaca guineensis Müll.Arg.) and Dipterocarpaceae (Monotes kerstingii Gilg.), on dead logs of Anthonota crassifolia, leg. and det. N.S.Yorou, 18.07.2009, herb. SYN 2273 (M), GenBank NCBI, accession number JQ814474.

ISotype: Mycological Herbarium of the University of Parakou (Benin).

Etymology: In reference to the small size of the basidiospores. Only a few Tomentella species present such small basidiospores (see discussion below).

Tomentella baroensis Yorou sp. nov.

GenBank NCBI, Accession number JQ814475 (to be delivered later)

Mycobank MB 566335

Basidiocarpa resupinata, crassa, ad 2 mm alta, adherens, crustosa, continua. Hymenium laeve usque ad subtile granulosum, continuum, obscure rufum usque ad cinereobrunneum; subiculum percrassum, arachnoideum, obscurius quam hymenium, brunneolum usque at subnigrum, marginibus sterilibus indeterminatis.

Rhizomorphae absentes.

Hyphae subiculi fibuligerae, septa simplicia absentia, 4–6(7) µm in diametro, regulares, numquam inflatae, in 2.5% KOH infrequenter sinuosae, ramificationes cruciformes et anastomosae absentes,

Fig. 3. Tomentella minispora. Optical section through subhymenial hyphae (from SYN 2273). Bar = 10 µm.

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crassitunicatae (0.5–1 µm), in aqua et in 2.5% KOH brunnea usque ad obscure brunnea, laeves, in aqua et in 2.5% KOH sine incrustionibus, non cyanescentes, subcongophilae, non cyanophilae, non amyloideae.

Hyphae subhymenii fibuligerae, 4–7 µm in diametro, nec breves nec inflatae, tenui- usque ad crassitunicatae (0.2–0.8 µm), in aqua et in 2.5% KOH subbrunneae usque ad brunneae, sine incrustationibus in aqua et in 2.5% KOH, interdum cyanescentes, congophilae, cyanophilae, non amyloideae.

Cystidia absentia

Basidia fibuligera, 25–45(50) µm longa, apicibus 7–15 µm, basibus 6–8 µm, clavata, non stipitata, nonnulla sinuosa, septis transversis infreqentibus, basidia iuvenilia usque ad 20 µm lata, basidia in aqua et in 2.5% KOH incolorata usque ad tenue subbrunnea, nunnulla contento ochraceo, non cyanescentia, congophila, cyanophila, non amyloidea, 4-sterigmatica, sterigmatibus 6–8(9) µm longis, basaliter 2–4 µm latis.

Basidiosporae 8–11(13) × 7.5–9(10) µm in aspectu frontali, 8–11 × 7–9 µm in aspectu laterali, ellip-soideae usque ad lobatae in aspectu frontali, ellipsoideae in aspectu laterali, echinulatae, aculeis brevibus, usque ad 1 µm longis, isolatis, basidiosporae in aqua et in 2.5% KOH brunneae, guttulis frequentibus, non cyanescentes, non congophilae, non cyanophilae, non amyloiedae.

Chlamydosporae absentes.

Basidiocarp resupinate, thick, up to 2 mm, adherent to the substrate, crustose, contin-uous. Hymenophore smooth to finely granulose, continuous, hymenium dull red (9C3) to greyish brown (9D3 to 9E4), subiculum very thick, arachnoid, darker than the hymenium, brownish to blackish, sterile margin indeterminate.

Rhizomorphs absent.

Subicular hyphae clamped, simple septa absent, 4–6(7) µm wide, regular, never inflated, in 2.5% KOH rarely sinuous (Fig. 4), cross-shaped branching and anastomoses absent, thick-walled (0.5–1 µm), in water and in 2.5% KOH brown to dark brown, smooth, in water and in 2.5% KOH without incrustations, not cyanescent, slightly congophilous, not cyanophilous, not amyloid.

Subhymenial hyphae clamped, 4–7 µm wide, neither short nor inflated, thin to thick-walled (0.2–0.8 µm), in water and in 2.5% KOH light brown to brown, in water and in 2.5% KOH without incrustations, sometimes cyanescent, congophilous, cyanophilous, not amyloid.

Cystidia absent.

Basidia clamped at base, 25–45(50) µm long, 7–15 µm at apex, 6–8 µm at base, clavate, not stalked, some sinuous, rarely with transverse septa, young basidia very large, up to 20 µm wide (Fig. 5), basidia in water and in 2.5% KOH colourless to very light brown, some commonly with ochraceous content, not cyanescent, congophilous, cyanophilous, not amyloid, 4-sterigmate, sterigmata 6–8(9) µm long × 2–4 µm at base.

Basidiospores 8–11(13) × 7.5–9(10) µm in frontal face, 8–11 × 7–9 µm in lateral face, ellipsoid to lobbed in frontal, ellipsoid in lateral views, echinulate, aculei short, up to 1 µm long, isolated, never forked, basidiospores in water and in 2.5% KOH brown, oil drops common, not cyanescent, not congophilous, not cyanophilous, not amyloid.

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Chlamydospores absent

material Studied and habitat: holotype: Guinea, upper Guinea, Province of Kankan, Prefecture of Kouroussa, forest reserve of Baro, along the road between Kouroussa and Kankan, N10°57'0.94", W009°64'60.0", forest composed of Uapaca guineensis (Phyllantaceae) and Anthonota crassifolia (Caesalpiniaceae), on dead logs, under A. crassifolia, leg. and det. N.S.Yorou, 18.07.2009, herb. SYN 2282 (M); GenBank NCBI, accession number JQ814475.

Fig. 4. Tomentella baroensis. a. Basidiospores in lateral view, b. Basidiospores in frontal view, c. Section through the basidiocarp (from SYN 2282). Bar = 10 µm.

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ISotype: Mcological herbarium of the University of Parakou (Benin).

Etymology: In reference to the origin of the type material. The studied material is collected in the Forest reserve of Baro in Upper Guinea.

Discussion

Important anatomical identification features for Tomentella minispora are the small basidiospores (maximum 7 µm), the short and very inflated subhymenial hyphae (up to 23 µm) and the short basidia (maximum 35 µm). Only a limited number of cystidia-lacking tomentelloid fungi present similar small basidiospores, including T. calcicola (Bourdot & Galzin) M.J.Larsen, T. griseoumbrina Litsch., T. neobourdotii M.J.Larsen, T. cinerascens (P.Karst.) Höhn. & Litsch., and T. botryoides (Schwein.) Bourdot & Galzin (Stalpers 1993, Kõljalg 1996). One important discriminating character between T. minispora and the above mentioned species is the presence of inflated (up to 23 µm) subhymenial hyphae in T. minispora. In addition to this anatomical character, T. cinerascens and T. neobourdotii differ from T. minispora by the presence of monomitic thelephoroid rhizomorphs (according to Agerer 1999) in the subiculum and sometimes at sterile margin (Yorou & Agerer 2011a). Larsen (1974) and Dämmrich (2006) reported on the presence of dimitic rhizomorphs for T. neobourdotii. In addition, T. neobourdotii presents a dark-brown hymenium with a fibrous to arachnoid subiculum and T. cinerascens presents a papillose to granular hymenophore with a whitish subiculum (Yorou & Agerer 2011a, Dämmrich 2006, Kõljalg 1996) in contrast to T. minispora with a greyish brown hymenophore and subiculum. According to Dämmrich (2006) and Kõljalg (1996), T. calcicola forms dimitic rhizomorphs. We were not able to observe rhizomorphs in T. baroensis. An important anatomical difference of T. calcicola with T. minispora is the presence of warted basidiospores in T. calcicola (Dämmrich 2006; Kõljalg 1996) in comparison to the echinulate ones in T. minispora. Tomentella botryoides is not only distinctive

Fig. 5. Tomentella baroensis. Young basidia (from SYN 2282). Bar = 10 µm.

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because of di- to trimitic, slightly differentiated rhizomorphs (Yorou & Agerer 2011a), but the basidiospores are commonly triangular to distinctly lobed, with basidia and sometimes subhymenial hyphae showing a strong cyanescent reaction in KOH (Yorou & Agerer 2011a, Dämmrich 2006) in contrast to T. minispora with subglobose to irregularly globose basidiospores and no cyanescent reaction in subhymenial hyphae and basidia. Anatomically, the combination of characters in T. minispora is strongly reminiscent of those of T. griseoumbrina. In the later species, however, the basidiospores never exceed 5.5 µm. Furthermore, the basidia in T. griseoumbrina present a maximal length of 25 µm (Dämmrich 2006, Kõljalg 1996, Larsen 1974) unlike of up to 35 µm in T. minispora. Additional discriminating features are the presence of strong incrustations and red-brown crystals on the subicular and subhymenial hyphae of T. minispora. Incrustations are lacking in T. griseoumbrina.

According to sequence analyses, T. fuscocinerea and T. amyloapiculata are the most similar identified species to T. minispora. Anatomically, both, T. fuscocinerea and T. amyloapiculata, are characterised by simple septate subicular hyphae and triangular (with widened proximal part) large pale brown to brown basidiospores of 8–12 µm (with long aculei of up to 2 µm). Within this clade, T. minispora deviates anatomically by the consistently present clamps, but also by the small size of basidiospores (maximum of 7 µm) and the small size of the aculei (maximum 0.8 µm). The combination of the clamp forming T. minispora with another collection of T. fuscocinerea and with T. amyloapiculata suggests that lack of clamps is neither a synapomorphic, nor a diagnostic feature for the clade. In general, the genus Tomentella encompasses only a limited number of clampless species, including in addition, notably T. spinosispora Čížek, T. badia (Link) Stalpers, T. fibrosa (Berk. & M.A.Curtis) Kõljalg, and T. furcata Yorou & Agerer (Dämmrich 2006, Stalpers 1993, Yorou & Agerer 2007). Infrequent clamps have been reported on subicular and subhymenial hyphae of T. cinereoumbrina (Bres.) Stalpers (Kõljalg 1996, Dämmrich 2006). In numerous molecular-phylogenetic investigations (Jakucs et al. 2005; Jakucs & Erős-Honti 2008, Yorou et al. 2007, Yorou & Agerer 2007), several clampless species fall within quite different terminal clades, supporting the argument that lack of clamps is not a synapomorphic feature. Within the genus Tomentella at least, the capability to form clamps may have been lost many times independently in the clades to which clamp forming species belong. The overwhelming number of clamp forming species of the genus Tomentella suggests clamps as a plesiomorphic character, which goes in line with other apparently originally clamp bearing relationships (Agerer 1999).

Anatomically, Tomentella baroensis can be easily identified by the presence of large young basidia (up to 25 µm) emerging either from the hymenium or the subhymenium. So far this feature is known only in Tomentella terrestris (Kõljalg 1996, Yorou & Agerer 2011b). Both species present many similarities including the regular and not inflated subicular hyphae of 4–7 µm, the very short aculei (up to 1 µm) of their basidiospores and rarely transverse septa in their basidia (Dämmrich 2006, Yorou & Agerer 2011b). Tomentella baroensis lacks and T. terrestris may lack rhizomorphs (Yorou & Agerer 2011b), though Loci (1997), Stalpers (1993) and Dämmrich (2006) reported on the presence of monomitic rhizomorphs in the subiculum of T. terrestris. However, unlike T. baroensis which presents ellipsoid basidiospores (8–13 µm) in both frontal

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and lateral views, T. terrestris shows rather commonly triangular (in frontal view), smaller basidiospores with size ranging from 7.5 to 9(10) µm. Transverse septa have been reported for basidia of both species, but T. terrestris basidia are much longer, commonly 55–85(90) µm (Yorou & Agerer 2011, Kõljalg 1996), in comparison to those of T. baroensis with 25 to 50 µm maximum. Basidia in T. baroensis are considerably smaller (maximum 25 µm). Sequence analyses reveal high differences between both species. T. baroensis deviates from T. terrestris (AF272901 from USA and AF272911 from Estonia) by at least 17 to 18% regarding ITS rDNA sequences (length of the considered segment = 550 bp). Such high genetic distance between T. baroensis and T. terrestris is supported by the molecular-phylogenetic analyses as both species fall within two distinct clades. Tomentella terrestris is closely related to T. sublilacina (Ellis & Holw.) Wakef. as both species together form a terminal clade (bootstrap support = 66). The molecular-phylogenetic affiliation of T. terrestris and T. sublilacina has been repeatedly demonstrated in molecular-phylogenetic inferences (Yorou et al. 2011a,b,c, Kõljalg et al. 2000, 2001, Jakucs et al. 2005, Peintner & Dämmrich 2011). As far as T. baroensis is concerned, it is molecular-phylogenetically closer to T. hjortstamiana and T. pisoniae than to T. terrestris.

The BLASTN search of the ITS rDNA in public GenBank suggests T. ellisii as the most similar species to T. baroensis. In the molecular-phylogenetic analysis, T. baroensis appears to be closer to T. hjortstamiana and T. pisoniae, whilst both sequences assigned to T. ellisii form the sister clade of the group comprising the aforementioned species (T. baroensis, T. hjortstamiana and T. pisoniae). Morphologically, both, T. baroensis and T. ellisii, present similar hymenophore patterns including a smooth to finely granular, dull red hymenium. Another morphological similarity is the absence of rhizomorphs either in the subiculum or at the sterile margin in both species (Yorou & Agerer 2011b). Subhymenial hyphae of T. ellisii are commonly inflated up to 12 µm whilst they are regular and never exceed 7 µm in T. baroensis. In addition, the basidia are so much longer in T. ellisii (45–80 µm) unlike those of T. baroensis with maximal 50 µm length. Regarding the basidiospores, T. baroensis presents larger basidiospores of 8–13 µm unlike T. ellisii whose basidiospores never exceeds 10 µm.

Though T. baroensis is molecular-phylogenetically close to both, T. pisoniae and T. hjortstamiana, numerous divergent anatomo-morphological features can be observed. Unlike T. hjortstamiana, T. baroensis lacks rhizomorphs. T. pisoniae lacks rhizomorphs, too, but it reveals a granular dark grey hymenium (unlike T. baroensis which present a smooth and finely granular dull red hymenium). Anatomical dissimilarities include the presence of inflated subhymenial hyphae in both T. pisoniae and T. hjortstamiana. T. pisoniae and T. baroensis possess basidiospores with similar size range, but those of T. pisoniae are rather triangular unlike those of T. baroensis that are ellipsoid.

Acknowledgement

We are much indebted to the Deutsche Forschungsgemeinschaft for financial support (DFG project Ag7/19-1).

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Manuscript received March 16, 2012; accepted March 28, 2012.