Five new species of Pseudosperma (Inocybaceae, Agaricales) from Benin and Turkey based on morphological characteristics and phylogenetic evidence

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YOROU, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-3937122/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 03 Apr, 2024 Read the published version in Mycological Progress → Version 1 posted 4 You are reading this latest preprint version Abstract Species of Pseudosperma (Inocybaceae) are widely distributed from temperate to tropical regions. In this study, we describe and illustrate five new species of Pseudosperma : P. beninense , P. cremeo-ochraceum , P. squarrosofulvum , P. stramineum , and P. tiliae , based on comprehensive analyses of morphological and molecular data derived from specimens collected in Benin (West Africa) and Turkey (Western Eurasia). These new species have been found in forests with Isoberlinia spp. and other ectomycorrhizal tree species in Benin and in association with Tilia platyphyllos in Turkey. The phylogenetic relationships of the new species were inferred through analyses of nuclear rDNA sequences, encompassing the internal transcribed spacer (ITS1-5.8S-ITS2) and 28S rDNA regions. Phylogenetic analyses revealed that P. beninense , P. cremeo-ochraceum , P. squarrosofulvum , and P. stramineum from Benin cluster with species from Australia, China, and India within a clade formed exclusively by species known from the palaeotropics and Australia, whereas P. tiliae from Turkey clustered with P. mediterraneum from Italy. Detailed descriptions are provided, supplemented by illustrations and line drawings of key micromorphological features. In addition, a comparative analysis with morphologically similar and phylogenetically closely related species is presented and discussed in detail. Ectomycorrhizal fungi Biodiversity Agarics Molecular systematics Biogeography Taxonomy Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Figure 9 Figure 10 Figure 11 Introduction By a recent molecular investigation based on several independent genetic loci, seven genera were identified within the family Inocybaceae Jülich: Auritella Matheny & Bougher, Inocybe (Fr.) Fr., Inosperma (Kühner) Matheny & Esteve-Rav., Mallocybe (Kuyper) Matheny, Vizzini & Esteve-Rav., Nothocybe Matheny & K.P.D. Latha, Pseudosperma Matheny & Esteve-Rav., and Tubariomyces Esteve-Rav. & Matheny (Matheny et al. 2020 ). The newly established genus Pseudosperma was initially classified as Inocybe section Rimosae sensu stricto (= Pseudosperma clade) (Matheny 2005 ; Larsson et al. 2009 ) in the subgenus Inosperma (Kuyper 1986 ; Bon 1997 ). Species of Pseudosperma are characterized by a rimose pileus, a furfuraceous or pruinose stipe, smooth, elliptical to indistinctly phaseoliform basidiospores, hyaline, non-necropigmented basidia, cylindrical to clavate cheilocystidia with thin walls, and absence of pleurocystidia (Bandini and Oertel 2020 ; Cervini et al. 2020 ; Matheny et al. 2020 ; Saba et al. 2020 ). Pseudosperma species form ectomycorrhizal symbioses with both angiosperm or gymnosperm trees and occur in a variety of habitats, including temperate forests dominated by spp. of Betula , Cedrus , Picea , Pinus , Populus , Quercus , and Salix (Kuyper 1986 ; Stangl 1989 ; Jacobsson 2008 ). Currently 80 species of Pseudosperma are described worldwide (Bandini et al. 2022 , 2023 ), including species recently introduced from Austria (Bandini et al. 2021 ), Australia (Matheny and Bougher 2017 ), China (Yu et al. 2020 ; Mao et al. 2022 ; Yan et al. 2022 ; Zhao et al. 2022 ), Germany (Bandini and Oertel 2020 ; Bandini et al. 2021 , 2022 , 2023 ), India (Latha et al. 2023 ), Italy (Cervini et al. 2020 ; Sanna et al. 2024 ), Pakistan (Jabeen and Khalid 2020 ; Saba et al. 2020 ; Jabeen et al. 2021 ; Naseer et al. 2023 ), Spain (Sanna et al. 2024 ), Sweden (Bandini et al. 2023 ) and Turkey (Kaygusuz et al. 2023 ). Despite this progress, the diversity of Pseudosperma species, particularly in tropical regions such as West Africa, remains poorly explored and documented. In addition to this, there are numerous cryptic and semi-cryptic species waiting for discovery (Ryberg et al. 2008 ; Matheny and Bougher 2017 ; Yan et al. 2022 ). West Africa is a region of species rich ecosystems with forests ranging among the 25 world hotspots that deserve top conservation priorities (Myers et al. 2000 ). The mycological exploration of West Africa, however, shows that the identified fungal species diversity in six countries in this region does not exceed 2% of the existing diversity (Piepenbring et al. 2020 ). For Benin, only 432 fungal species have been documented in (Piepenbring et al. 2020 ). To date, the only species of Pseudosperma reported from Benin is Pseudosperma squamatum (J.E. Lange) Matheny & Esteve-Rav., historically called Inocybe squamata J.E. Lange (Lange 1917 ; Boa 2004 ; Piepenbring et al. 2020 ). No further species of Pseudosperma has ever been cited for any African country according to the literature available to us. This study aims to increase the knowledge of the species diversity of Pseudosperma , with particular emphasis on the description of four new species from Benin and one from Turkey. Thereby, we contribute to the knowledge of morphological diversity, ecology, biogeography, and phylogeny of these hidden fungal treasures of Western Eurasia and West Africa. Materials and methods Sampling and Morphological Studies The specimens from Benin were collected during a mycological survey conducted from June to August 2022. In Turkey, samples were picked up during fieldwork in the Isparta Province in 2022 and 2023. The macroscopic characteristics were obtained from fresh specimens, field notes and photographs taken in situ. Standardized colour values were documented using the Munsell Soil Color Charts (Munsell 1975 ). Microscopic features were observed using 1% Congo red (w/v) or 5% potassium hydroxide (KOH) (w/v). All samples were analysed and photographed with a light microscope. A minimum of thirty basidiospores were measured for each collection. Q values (ratio of length to width of basidiospore) and average values (length and width of basidiospore or cystidia) are presented. SD is the abbreviation for the standard deviation of the length × width. The terminology of Vellinga ( 1988 ) is used for macro- and micro-characters. Index Fungorum ( http://www.IndexFungorum.org ) and the International Index of Plant Names ( https://www.ipni.org ) were used as sources for taxonomic names and nomenclature. The samples from Benin are deposited in the fungarium of the Staatliches Museum für Naturkunde Stuttgart (STU) or the mycological herbarium of the University of Parakou (UNIPAR). The Turkish specimens are stored in the fungarium of the Isparta University of Applied Sciences (ISUF). Molecular Analyses Genomic DNA was isolated from Pseudosperma specimens using the innuPREP Plant DNA Kit (Analytik Jena, Jena, Germany) and the Fungi/Yeast Genomic DNA Isolation Kit (Norgen Biotek Corp, Ontario, Canada). For amplification of nuclear rDNA internal transcribed spacer region ITS1-5.8S-ITS2 (ITS) the primer pair ITS1F/ITS4 (White et al. 1990 ; Gardes and Bruns 1993 ), and for the nuclear 28S rDNA (LSU) the primer pair LR0R/LR5 (Vilgalys and Hester 1990 ; Rehner and Samuels 1994 ) were used. Polymerase chain reaction (PCR) procedures were performed according to the methods described by Kaygusuz et al. ( 2020 ). The PCR products were sequenced at Microsynth Seqlab (Göttingen, Germany) and the Sanger DNA sequencing service of Source Bioscience (Berlin, Germany), using the same primers. The obtained DNA sequences were aligned and analysed using ClustalX (Thompson et al. 1997 ) and MEGA X v.10.0.5 (Kumar et al. 2018 ) and subsequently submitted to GenBank. A total of 18 DNA sequences (nine from the nrITS and nine from the nrLSU) from nine collections were newly generated. BLASTn searches were conducted in the NCBI GenBank. For phylogenetic analyses, sequences with high similarity (with maximum identities larger than 82%) to the new sequences were retrieved from GenBank ( www.ncbi.nlm.nih.gov ) and UNITE ( https://unite.ut.ee/ , Kõljalg et al. 2005 ) databases, along with sequences from recent publications (Ryberg et al. 2008 ; Larsson et al. 2009 ; Matheny 2009 ; Matheny et al. 2009 ; Vauras and Larsson 2011 ; Sjökvist et al. 2012 ; Kropp et al. 2013 ; Osmundson et al. 2013 ; Pradeep et al. 2016 ; Matheny and Bougher 2017 ; Tibpromma et al. 2017 ; Bau and Fan 2018 ; Li et al. 2018; Ullah et al. 2018 ; Matheny and Kudzma 2019 ; Matheny et al. 2020 ; Bandini and Oertel 2020 ; Cervini et al. 2020 ; Jabeen and Khalid 2020 ; Saba et al. 2020 ; Yu et al. 2020 ; Bandini et al. 2021 , 2022 , 2023 ; Mao et al. 2022 ; Zhao et al. 2022 ; Kaygusuz et al. 2023 ; Latha et al. 2023 ; Naseer et al. 2023 ). The nrITS and nrLSU rDNA sequences were separately aligned using MAFFT 7.11 (Katoh et al. 2019 ) applying the E-INS-i iterative method, followed by manual corrections in AliView V.1.28 (Larsson 2014 ). Mallocybe agardhii (N. Lund) Matheny & Esteve-Rav. (AB980912) and M . picea L. Fan & N. Mao (BJTC FM555) were designated as outgroups. Multiple sequence alignments were inspected using MEGA X 10.0.5 prior to subsequent analyses. A single combined dataset of nrITS-nrLSU rDNA sequences was assembled for phylogenetic analysis. The optimal evolutionary model for each segment was determined using MrModeltest 2.3 (Nylander 2004 ). Phylogenetic assessments employed both Maximum Likelihood (ML) and Bayesian Inference (BI) approaches on the concatenated genes. The ML analysis was conducted with IQ-TREE v.1.6.12 (Minh et al. 2020 ), using the Ultrafast Bootstrap (UFBoot) (Hoang et al. 2018 ) approach with five thousand bootstrap iterations. The BI analysis using the Markov Chain Monte Carlo (MCMC) method was conducted in MrBayes 3.2.5 (Ronquist et al. 2012 ) over 1 × 10 7 generations, sampling trees every thousand generations. Phylogenetic trees were visualized using FigTree v1.4.4 (Rambaut 2018 ), with only Maximum Likelihood Bootstrap (MLB) values above 75% and Bayesian Posterior Probabilities (BPP) exceeding 0.90 being indicated. Results Phylogeny The combined nrITS and nrLSU rDNA sequence dataset for Pseudosperma species, including 18 new sequences of the specimens from Benin and Turkey, consisted of 107 taxa with 2480 characters, of which 631 were parsimony-informative, 274 singleton sites, and 1575 constant sites. The best models in ML were TVM+F+I+G4 for nrITS and nrLSU rDNA, and GTR+I+G in the BI analysis. This matrix exhibited 1222 unique alignment patterns. The estimated nucleotide substitution rates were as follows: A-C = 1.06255, A-G = 5.05462, A-T = 1.63234, C-G = 0.42882, C-T = 5.05462, G-T = 1.00000. Base frequencies were determined as A = 0.263, C = 0.190, G = 0.247, T = 0.300. The gamma distribution shape parameter α was calculated to be 0.617. Phylogenetic trees derived from ML and BI analyses showed largely congruent topologies. The topology resulting from the ML analysis was selected for presentation, with statistical support values indicated by Maximum Likelihood Bootstrap (MLB) and Bayesian Posterior Probabilities (BPP) values (Fig. 1). Molecular analyses based on the combined dataset revealed that Pseudosperma specimens from Benin and Turkey are genetically distinct from other species within the genus represented by molecular sequences data. Their sequences form part of five independent lineages, as shown in Fig. 1. The first lineage with high statistical support (MLB = 100%, BPP = 1.0) consists of three specimens of the new species called Pseudosperma tiliae together with P. mediterraneum (Kuyper) Bandini, B. Oertel & U. Eberh. from Italy in the same subclade. The second lineage (MLB = 99%, BPP = 1.0) consists of Pseudosperma stramineum andfive undescribed and unpublished sequences (HLA0707, HLA0386, 486ad3d1, 79c39691 and ASV_39) from Benin. The third lineage is formed by a single sequence labelled as Pseudosperma squarrosofulvum (AR-22-024) from Benin. The fourth lineage (MLB = 100%, BPP = 1.0) includes the new species Pseudosperma beninense (AR-22-037) from Benin and an undescribed and unpublished sequence from an uncultured fungus (ASV_330). Pseudosperma beninense , P. squarrosofulvum ,and P. stramineum form a distinct, statistically highly supported clade of exclusively Beninese species (MLB = 100%, BPP = 1.0). The last lineage (MLB = 100%, BPP = 1.0) comprises the new species Pseudosperma cremeo-ochraceum (AR-22-088) and three undescribed and unpublished sequences (HLA0454, ASV_313, 881fdb9c) from Benin. The new sequences from Benin are located in a clade formed by a total of ten known species that all originate from the palaeotropics (and subtropics).The sequences of the recently collected Pseudosperma specimens consistently form distinct lineages in all phylogenetic analyses and can not be assigned to any existing Pseudosperma species concept by morphological characteristics. Therefore, we propose them as species new to science and provide detailed descriptions of these species in the following. Figure 1 [near here] Taxonomy Pseudosperma beninense Kaygusuz, Bandini, Rühl, Sarawi, Yorou & M. Piepenbr., sp. nov. (Figs. 2 and 3) MycoBank: MB 851970 Etymology: The specific epithet refers to the locality where the type specimen was collected. Holotype: Benin, Borgou Department, Fôret Classée de l'Ouémé Supérieur, on soil in savannah forest dominated by Isoberlinia doka Craib & Stapf, I . tomentosa (Harms)Craib & Stapf, Monotes kerstingii Gilg and Uapaca togoensis Pax, at 09°15'37.9"N, 002°11'03.9"E, 340 m asl., 18 June 2022, leg. A. Rühl, C. Manz, D. Dongnima, F. Hampe & S. Sarawi (AR-22-037, STU). GenBank accession numbers PP060393 (nrITS) and PP060408 (nrLSU). Diagnosis: Differs from P. squamatum by smaller basidiomata with brown to straw-brown pileus, whitish to light yellow stipe, longer basidiospores (on average 12.5 × 7.0 μm) mostly with acute apex, longer (on av. 51 × 14 μm) and oblong to cylindrical or narrowly clavate cheilocystidia, and by distinct nrITS and nrLSU rDNA sequences. Description: Pileus 7–14 mm diam., when young paraboloid, later hemispherical to convex, with or without low and broad umbo, margin inflexed when young, later long deflexed, surface dry, tomentose-lanose to subtomentose, radially fibrillose to rimulose outwards, colour light brown (2.5Y 8–7/6, 8/8) to straw brown (2.5Y 6/4–6), slightly darker at the centre (2.5Y 6/8). Lamellae moderately crowded, adnexed, subventricose, whitish to yellowish-white or light yellow, edge slightly eroded, whitish. Stipe 17–25 × 1.2–1.8 mm, central, cylindrical with subbulbous base up to 2.5 mm diam., solid, surface whitish to light yellow, pruinose only near the apex. Colour of exsiccate: pileus very light yellow to light straw yellow (5Y 8/2–4, 7/4), lamellae and stipe whitish. Smell unrecorded. Basidiospores 10.7–14.7 µm (av. 12.5 µm, SD ± 0.4 µm) × 6.2–8.3 µm (av. 7.0 µm, SD ± 0.5 µm); Q = 1.4–2.1 (av. 1.8, SD ± 0.1) (n = 90 of 1 coll.), mainly (sub)amgydaliform with acute apex, also subcylindrical and subellipsoid, with guttules, smooth, thick-walled, dark yellowish brown in 5% KOH. Basidia 35–42 × 10–13 µm, clavate, 4-spored, thin-walled, hyaline. Cheilocystidia 37–65 µm (av. 51 µm, SD ± 7.0 µm) × 9–20 µm (av. 14 µm, SD ± 2.3 µm); Q = 2.4–5.8 (av. 4.2, SD ± 0.6) (n = 45 of 1 coll.), mostly oblong to cylindrical or narrowly clavate, sometimes with subcapitate apex, sometimes in chains of 1–3 cells, thin-walled, hyaline or very pale yellowish-brown in 5% KOH. Paracystidia 20–35 × 9–14 µm, cylindrical to broadly clavate, in chains of 2–4 cells, thin-walled, hyaline or very pale yellowish-brown in 5% KOH. Pileipellis cutis, consisting of long cylindrical or narrowly fusiform terminal cells, with sharply pointed apex, 60–170(220) × 11–25 µm, smooth, thin-walled, pale yellowish-brown in 5% KOH. Caulocystidia a cutis with transitions to a trichoderm composed predominantly of multiseptate cylindrical to inflated hyphae, 10–100 × 6.5–12 µm, sometimes with subcapitate apex, often in bundles, smooth, thin-walled, hyaline in 5% KOH. Stipitipellis a cutis of parallel hyphae, 5–18 µm wide, thin-walled, hyaline in 5% KOH. Clamp connections present in all parts examined. Figure 2 [near here] Habitat and distribution: Basidiomata solitary, terrestrial, on wet and sandy soils, growing in a forest dominated by species of Caesalpiniaceae ( Isoberlinia spp.), Dipterocarpaceae ( Monotes kerstingii ), and Phyllanthaceae ( Uapaca togoensis ). Currently known only from Benin. Additional specimen examined: Based on the phylogenetic analysis, one further specimen from West Africa (ASV_330) belongs to Pseudosperma beninense . Figure 3 [near here] Discussion: Phylogenetic analyses inferred from nrITS and nrLSU rDNA sequences show that Pseudosperma beninense forms a monophyletic subclade within Pseudosperma and is closely related to P. squarrosofulvum and P. stramineum , two other new species presented in this study. When the nrITS DNA sequences generated from Pseudosperma beninense are compared with the sequences of P. squarrosofulvum and P. stramineum , 56 nucleotide differences (83% similarity) were observed in the nrITS DNA sequences of P. squarrosofulvum and 52 differences (85% similarity) in the sequences of P. stramineum . Morphologically, Pseudosperma squarrosofulvum differs from P. beninense by its pileus surface, which is covered by yellow ochre to brownish-yellow fibrillose scales, longer basidiospores (on av. 12.9 × 6.2 µm), somewhat shorter and narrower cheilocystidia(on av. 44 × 11 µm), mostly utriformparacystidia, pileipellis hyphae often with encrusted walls, and mostly cylindrical or narrowly clavate caulocystidia. Pseudosperma stramineum has a predominantly straw-yellow to buff pileus with a distinct squamulose-squarrose surface, pale brown lamellae when old, mostly oblong basidiospores, smaller cheilocystidia (on av. 40 × 14 μm), a pileipellis with strongly incrusted walls, and somewhat shorter caulocystidia (up to 90 μm in length). Morphologically, the species closest to the new species Pseudosperma beninense is P. squamatum , which differs mainly by larger basidiomata, with a pileus measuring 30–70 mm in diameter, yellowish to yellow-ochraceous pileus, often with orange tinged, a longer stipe (up to 70 mm), shorter basidiospores (on av. 9.9 × 6.1 µm), shorter cheilocystidia (on av. 44 × 14 µm) that are subclavate to subglobose, and a habitat with a clay soil as well as an associated with Populus sp. (Lange 1917; pers. observation of D. Bandini). In addition, P . beninense is distant from P . squamatum following phylogenetic analyses (Fig. 1). Other tropical or subtropical species that are morphologically somewhat similar to Pseudosperma beninense are P. fissuratum (Matheny & Bougher) Matheny & Esteve-Rav., P. gracilissimum (Matheny & Bougher) Matheny & Esteve-Rav., P. palaeotropicum (E. Turnbull & Watling) Matheny & Esteve-Rav. and P. renisporum (E. Horak) Matheny & Esteve-Rav. Pseudosperma fissuratum , originally described from Australia, differs from P. beninense by a typically bicoloured pileus, the presence of a velipellis, shorter basidiospores (on av. 11.4 × 6.2 µm), longer cheilocystidia (up to 72 µm in length), and ecologically by an association with Eucalyptus sp.(Matheny and Bougher 2017). Another Australian species, Pseudosperma gracilissimum , has a markedly conical pileus, slightly shorter basidiospores (on av. 9.9 × 5.9 µm), and is associated with Acacia , Allocasuarina , Corymbia , Eucalyptus, Lophostemon ,and Melaleuca (Matheny and Bougher 2017). Pseudosperma palaeotropicum , initially discovered from Singapore and later reported from Australia and Malaysia, has larger basidiomata with a pileus measuring 20–40 mm in diameter, a longer stipe (40–70 × 4.0–6.0 µm), considerably shorter basidiospores (7.0–8.3 × 4.8–6.5 µm), and is typically associated with species of Dipterocarpaceae (Turnbull 1995). Pseudosperma renisporum , originally described from New Zealand, has a squamulose pileus centre, bean-shaped and shorter basidiospores (9.0–12.0 × 4.5–6.5 μm), and is associated with species of Leptospermum and Nothofagus (Horak 1978). Pseudosperma cremeo-ochraceum Kaygusuz, Bandini, Rühl, Sarawi, Yorou & M. Piepenbr., sp. nov. (Figs. 4 and 5) MycoBank: MB 851972 Etymology: The specific epithet refers to the cream to ochraceous colour of the surface of the pileus. Holotype: Benin, Borgou Department, Fôret l'Ouémé Supérieur, on the soil in savannah forest dominated by Isoberlinia doka , I . tomentosa , Monotes kerstingii and Uapaca togoensis , at 08°36'04.7"N, 002°36'00.7"E, 340 m asl., 28 June 2022, leg. A. Rühl, C. Manz, D. Dongnima, F. Hampe & S. Sarawi (AR-22-088, STU). GenBank accession numbers PP060394 (nrITS) and PP060409 (nrLSU). Diagnosis: Most similar to the tropical Australian species Pseudosperma gracilissimum , but differing by a silky fibrillose pileus, longer basidiospores (on av. 12.9 × 7.8 µm), mostly narrowly utriform to utriform cheilocystidia with subcapitate apex, pileipellis elements without encrusted walls, the presence of caulocystidia, a different habitat dominated by Isoberlinia spp., M . kerstingii and U . togoensis , and by distinct nrITS and nrLSU rDNA sequences. Figure 4 [near here] Description: Pileus 10–15 mm diam., broadly conical to hemispherical, expanded plano-convex, usually with a low umbo, with a straight and translucently striate margin reaching up to 1/4 or 2/4 of the radius, colour cream (2.5Y 8–7/2) to yellowish brown (2.5Y 7/6–10) or ochraceous (2.5Y 6/6–8), becoming darker at the centre with age, always creamy white (2.5Y 8/2–4) to ivory white (2.5Y 7/2) at the edge, surface dry, radially silky-fibrillose. Lamellae moderately crowded to subdistant, adnexed, subventricose, pale ivory white to pale yellow grey, becoming yellowish white, edge somewhat eroded, whitish. Stipe 12–20 × 0.6–1.2 mm, central, cylindrical, sometimes subbulbous at the base, straight or curved towards the base of the stipe, surface sordid white or cream to light brown when old, slightly pruinose only near the apex. Colour of exsiccate: pileus sordid white coloured, lamellae and stipe whitish. Smell unrecorded. Basidiospores 11.0–16.0 µm (av. 12.9 µm, SD ± 1.2 µm) × 6.5–9.3 µm (av. 7.8 µm, SD ± 0.6 µm); Q = 1.4–1.9 (av. 1.7, SD ± 0.1) (n = 100 of 2 coll.), mostly oblong, with central germ pore, with guttules, smooth, thick-walled, yellowish brown in 5% KOH. Basidia 35–45 × 11–13 µm, clavate, 4-spored, thin-walled, hyaline. Cheilocystidia 40–80 µm (av. 51 µm, SD ± 6.0 µm) × 10–17 µm (av. 13.0 µm, SD ± 2.0 µm); Q = 3.1–5.3 (av. 4.1, SD ± 0.6) (n = 40 of 1 coll.), scattered, narrowly utriform to utriform mostly with subcapitate apex, hyaline or very pale yellowish-brown in 5% KOH. Paracystidia 25–40 × 10–18 µm, utriform with obtuse or subcapitate apex or fusiform, thin-walled, hyaline or very pale yellowish-brown in 5% KOH. Pileipellis a hymeniderm to epithelium formed by broadly fusiform to cylindrical terminal elements, with obtuse to mucronate apex, 47–95 × 20–40 µm, smooth, thin-walled, pale yellowish-brown in 5% KOH. Caulocystidia 15–35 × 10–15 µm, narrowly clavate, on clusters of erect hyphae, smooth, thin-walled, hyaline in 5% KOH. Stipitipellis a cutis of subparallel hyphae, 6–12 µm wide, thin-walled, hyaline in 5% KOH. Clamp connections present in all parts examined. Figure 5 [near here] Habitat and distribution: Basidiocarpsgregarious, usually terrestrial, on wet and sandy soils, in woodlands dominated by Caesalpiniaceae ( Isoberlinia spp.), Dipterocarpaceae ( Monotes kerstingii ), and Phyllanthaceae ( Uapaca togoensis ). Currently only known from Benin. Additional specimen examined: Benin, Borgou Department, Fôret Classée de l'Ouémé Supérieur, on the soil in savannah forest dominated by Isoberlinia spp., 08°36'04.7"N, 002°36'00.7"E, 340 m asl., 28 June 2022, leg. A. Rühl, C. Manz, D. Dongnima, F. Hampe & S. Sarawi (AR-22-089, UNIPAR). According to the phylogenetic analysis, one further specimen from Benin (HLA0454) collected by H.L. Aignon and two sequences obtained from soil (ASV_313 and 881fdb9c) in Benin also belong to Pseudosperma cremeo-ochraceum . Discussion In the concatenated nrITS-nrLSU rDNA phylogeny (Fig. 1), Pseudosperma tiliae is sister to P. mediterraneum and nested with P. holoxanthum (Grund & D.E. Stuntz) Matheny & Esteve-Rav., P. melliolens (Kühner) Matheny & Esteve-Rav., P. rimosum (Bull.) Matheny & Esteve-Rav., and P. sororium (Kauffman) Matheny & Esteve-Rav. However, morphologically P. mediterraneum , originally described from Italy, differs from P. tiliae by a pale buff to ochraceous pileus, slightly longer basidiospores (on av. 13.2 × 6.6 µm) with a higher Q-value (Q = 2), shorter cheilocystidia (36–57 × 13–26 µm), and a habitat on dune sand associated with Pinus pinea (Kuyper 1986). The genetic distance between Pseudosperma tiliae and P. mediterraneum is 3%, corresponding to 18 base divergences in 600 nucleotides, indicating that these are different species. Pseudosperma holoxantha , originally described from the USA, differs from P . tiliae by a longer (up to 100 mm in length) and pale yellow to straw yellow stipe, shorter basidiospores (9.0–13.0 × 6.0–8.0 µm), longer cheilocystidia (up to 110 µm in length), and growth with conifers (Grund and Stuntz 1981). Pseudosperma melliolens differs by a brown to brownish coloured pileus, greyish or reddish stipe, shorter basidiospores (9.0–13.5 × 6.0–8.0 µm) and shorter cheilocystidia (up to 55 µm in length) (Kühner 1988). Pseudosperma rimosum differs by usually less stout habitus, often dull fallow pileus colour, only faint and fugacious greyish velipellis (Bulliard 1789). Pseudosperma sororium , originally described from the USA, has larger basidiomata (up to 70 mm in diam.) with a subconical to conical-campanulate pileus, and distinctly longer basidiospores (9.0–16.0 × 5.0–8.0 µm) (Murrill et al. 1924). Other European species that are morphologically similar to Pseudosperma tiliae are P. arenicola (R. Heim) Matheny & Esteve-Rav., P . mimicum (Massee) Matheny & Esteve-Rav., P. musilii Bandini, B. Oertel & Schmidt-Stohn, P. pamukkalense Kaygusuz, Bandini & Knudsen, P. pseudoorbatum (Esteve-Rav. & García Blanco) Matheny & Esteve-Rav, and P . spectrale Bandini & B. Oertel. Pseudosperma arenicola differs by larger pileus (up to 68 mm in diam.) and stipe (up to 75 mm long), distinctly longer basidiospores (on av. 13–15.4 × 6.3–8.0 µm), longer (up to 105 µm) and mostly cylindrical cheilocystidia, cylindrical caulocystidia, and a habitat associated with Salix repens L, Populus canadensis Moench, or sometimes Pinus maritima Mill. (Kuyper 1986). Pseudosperma mimicum differs by larger pileus (up to 80 mm in diam.), yellow-brown lamellae, much longer basidiospores (14.0–16.0 × 6.0–8.0 µm), and the absence of cystidia (Massee 1904). Pseudosperma musilii differs by its dingy straw to dark brown pileus with a strongly rimose surface, shorter basidiospores (on av. 11.3 × 6.7 µm), and shorter cheilocystidia (on av. 48 × 12 µm) (Bandini et al. 2023). Pseudosperma pamukkalense differs by subphaseoliform basidiospores, a predominant association with Pinus nigra subsp. pallasiana (Lamb.) Holmboe, and genetic differences (nrITS locus 81.5% identity) (Kaygusuz et al. 2023). Pseudosperma pseudoorbatum has a white to ivory-white pileus, notably longer basidiospores (on av. 14.6 × 7.1 µm), and a habitat with Pinus forests ( Pinus pinaster Aiton and P . pinea L.) (Esteve-Raventós et al. 2003). Pseudosperma spectrale has a whitish to straw-coloured pileus, slightly shorter basidiospores (on av. 12.1 × 7.0 µm), and a habitat with conifers (Bandini et al. 2022). Conclusions It was previously suggested that species of Inocybaceae began to spread over large areas of northern and southern South America, Australia, and New Zealand during the Palaeogene or later periods (Matheny 2009; Matheny et al. 2009). Recent multi-gene phylogenetic analyses confirmed close affinities between Pseudosperma taxa from tropical Asia, tropical Australia, and tropical China (Zhao et al. 2022). Similarly, in the present study, phylogenetic analyses generated from nrITS and nrLSU rDNA sequences revealed rather close phylogenetic relationships among Pseudosperma species indigenous to the tropical regions of Benin, Australia, India, and China. The new species Pseudosperma beninense , P. cremeo-ochraceum , P. squarrosofulvum , and P. stramineum were found in ectomycorrhizal forests with tropical climate of Benin and are phylogenetically close to each other in a clade that in addition to the species from Benin includes six further species from tropical regions of the palaeotropics and Australia. These six species are P. araneosum from Australia (Matheny and Bougher 2017), Pseudosperma brunneosquamulosum , P. luteobrunneum , and P. rubrobrunneum from India (Tibpromma et al. 2017), and Pseudosperma fulvidiscum and P. singulare from China (Zhao et al. 2022). The ongoing discovery and characterization of tropical to subtropical taxa within the genus will improve our understanding of their biogeographical distribution and evolutionary history. Declarations Acknowledgements We would like to thank Frank Lappe and Anika Rüb for their technical assistance, and Cathrin Manz, Felix Hampe, Boris Olou, and Sylvestre Badou for their support in the field. We are grateful to Daouda Dognima for careful driving through Benin´s ecosystems. This study results from the project FunTrAf that is generously supported by the German Federal Ministry of Education and Research (BMBF, 01DG20015FunTrAf). The first author also acknowledges the financial support of the Scientific and Technical Research Council of Turkey (TUBITAK) for the 2219 International Postdoctoral Research Fellowship Programme (Grant No. 1059B192202880). Author contribution All authors contributed to the conception and design of the study. Oğuzhan Kaygusuz, Adrian Rühl, Sepas Sarawi, Nourou S. Yorou, and Meike Piepenbring contributed to material preparation and data collection. Morphological characteristics were examined by Oğuzhan Kaygusuz and Ditte Bandini. Molecular lab work and phylogenetic analyses were conducted by Oğuzhan Kaygusuz, Adrian Rühl, and Sepas Sarawi. The first draft of the manuscript was written by Oğuzhan Kaygusuz and Meike Piepenbring, which was then improved by changes, edits, suggestions, and comments from Ditte Bandini, Adrian Rühl, Sepas Sarawi and Nourou S. Yorou. All authors read and approved the final manuscript. Funding FunTrAf by the German Federal Ministry of Education and Research (BMBF, 01DG20015FunTrAf) and 2219 International Postdoctoral Research Fellowship Programme (Grant No. 1059B192202880) by the Scientific and Technical Research Council of Turkey (TUBITAK). Data availability The DNA sequences produced in this study are available on NCBI GenBank (https://www.ncbi.nlm.nih.gov). Code availability Not applicable. Ethics approval and consent to participate Not applicable. Consent for publication Informed consent was obtained from all individual participants included in the study. Competing interests The authors declare no competing interests. References Bandini D, Oertel B (2020) Three new species of the genus Pseudosperma (Inocybaceae). Czech Mycol 72(2):221–250. https://doi.org/10.33585/cmy.72205 Bandini D, Oertel B, Eberhardt U (2021) Even more fibre-caps (2): Thirteen new species of the family Inocybaceae. Mycologia Bavarica 21:27–98 Bandini D, Oertel B, Eberhardt U (2022) Noch mehr Risspilze (3): Einundzwanzig neue Arten der Familie Inocybaceae. Mycologia Bavarica 22:31–138 Bandini D, Oertel B, Eberhardt U (2023) Even more fibre-caps (4): Fourteen new species of the family Inocybaceae. Mycologia Bavarica 23:1–50 Bau T, Fan Y-G (2018) Three new species of Inocybe sect. Rimosae from China. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-3937122","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":273315044,"identity":"42215407-b2a2-4412-b9f0-3771e23b5c46","order_by":0,"name":"Oğuzhan Kaygusuz","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA7klEQVRIiWNgGAWjYDACHhiDmf3gAxCfj3gt7D3JBiA+G/FaeA6YSYBoglr4eQ4/k/zaVpc4f0ZCWuXXHDsZNgbmh49u4NEi2dtmJi3bdjixcUbisduy25KBDmMzNs7Bo8XgPIOZtGTbgcRmiYS025LbmIFaeNik8Wth/wbUUpfYJpFgViy5rZ4ILWd7zCQ/tjEn9gC9z/hx22HCWiR7zhRbM5w7bDwDGMjSjNuO87AxE/ALP0/6xps/yupk5zezH/z4c1u1PT9788PH+LQAAYs0LG6YwQxm/MrBSj7+gLIYf+BVOApGwSgYBSMVAACkGUVdgLACjwAAAABJRU5ErkJggg==","orcid":"https://orcid.org/0000-0003-3455-2965","institution":"Goethe-Universität Frankfurt am Main: Goethe-Universitat Frankfurt am Main","correspondingAuthor":true,"prefix":"","firstName":"Oğuzhan","middleName":"","lastName":"Kaygusuz","suffix":""},{"id":273315045,"identity":"83fc439d-5868-4776-9eed-aeb6eac54f85","order_by":1,"name":"Ditte BANDINI","email":"","orcid":"","institution":"Panorama str 47, Wiesenbach","correspondingAuthor":false,"prefix":"","firstName":"Ditte","middleName":"","lastName":"BANDINI","suffix":""},{"id":273315046,"identity":"c12e15fd-f589-47b1-a508-13031a86eee6","order_by":2,"name":"Adrian RÜHL","email":"","orcid":"","institution":"Goethe-Universität Frankfurt am Main: Goethe-Universitat Frankfurt am Main","correspondingAuthor":false,"prefix":"","firstName":"Adrian","middleName":"","lastName":"RÜHL","suffix":""},{"id":273315047,"identity":"0bff235f-0c51-43b5-8cdc-530e3f8485cc","order_by":3,"name":"Sepas SARAWI","email":"","orcid":"","institution":"Goethe-Universität Frankfurt am Main: Goethe-Universitat Frankfurt am Main","correspondingAuthor":false,"prefix":"","firstName":"Sepas","middleName":"","lastName":"SARAWI","suffix":""},{"id":273315048,"identity":"bf99a944-34c0-4771-a8a1-ea38de7a3c41","order_by":4,"name":"Nourou S. YOROU","email":"","orcid":"","institution":"Université de Parakou: Universite de Parakou","correspondingAuthor":false,"prefix":"","firstName":"Nourou","middleName":"S.","lastName":"YOROU","suffix":""},{"id":273315049,"identity":"6df2fdc2-e05f-4632-93e0-705a7cdbbf48","order_by":5,"name":"Meike PIEPENBRING","email":"","orcid":"","institution":"Goethe-Universität Frankfurt am Main: Goethe-Universitat Frankfurt am Main","correspondingAuthor":false,"prefix":"","firstName":"Meike","middleName":"","lastName":"PIEPENBRING","suffix":""}],"badges":[],"createdAt":"2024-02-07 14:25:14","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-3937122/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-3937122/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s11557-024-01964-2","type":"published","date":"2024-04-03T15:00:50+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":51362308,"identity":"561457b8-3ef8-45dd-851f-3d1451e013f5","added_by":"auto","created_at":"2024-02-20 09:04:36","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1539182,"visible":true,"origin":"","legend":"\u003cp\u003ePhylogenetic position of the Beninese and Turkish collections of \u003cem\u003ePseudosperma\u003c/em\u003ebased on nrITS and nrLSU rDNA sequences using Maximum Likelihood (ML) and Bayesian analyses.\u003cstrong\u003e \u003c/strong\u003eMLB ≥ 75% and BPP ≥ 0.90 values are indicated on the branches. \u003cem\u003eMallocybe agardhii\u003c/em\u003e (AB980912) and \u003cem\u003eM\u003c/em\u003e.\u003cem\u003epicea\u003c/em\u003e (BJTC FM555) were selected as outgroups. The newly generated sequences are marked in red bold. A clade formed by species from the palaeotropics and Australia is marked by a grey background.\u003c/p\u003e","description":"","filename":"FigPage01.jpg","url":"https://assets-eu.researchsquare.com/files/rs-3937122/v1/9351af81b66fb137894edc32.jpg"},{"id":51362311,"identity":"13f52ecc-89e8-451c-9302-93f710deca59","added_by":"auto","created_at":"2024-02-20 09:04:36","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":2706967,"visible":true,"origin":"","legend":"\u003cp\u003eBasidiomata of \u003cem\u003ePseudosperma beninense \u003c/em\u003e(AR-22-037, holotype). Bar = 5 mm.\u003c/p\u003e","description":"","filename":"FigPage02.jpg","url":"https://assets-eu.researchsquare.com/files/rs-3937122/v1/41919fdff6ac8f40544bd106.jpg"},{"id":51362309,"identity":"76e8dea1-775f-4d28-a8d7-a1a3a75e5d03","added_by":"auto","created_at":"2024-02-20 09:04:36","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":805607,"visible":true,"origin":"","legend":"\u003cp\u003eMicroscopic features of \u003cem\u003ePseudosperma beninense \u003c/em\u003e(AR-22-037, holotype)\u003cem\u003e.\u003c/em\u003e \u003cstrong\u003ea-\u003c/strong\u003e Basidiospores. \u003cstrong\u003eb-\u003c/strong\u003e Basidia. \u003cstrong\u003ec-\u003c/strong\u003eParacystidia. \u003cstrong\u003ed-\u003c/strong\u003e Cheilocystidia. \u003cstrong\u003ee-\u003c/strong\u003e Pileipellis. \u003cstrong\u003ef-\u003c/strong\u003eCaulocystidia. Bars = 10 μm.\u003c/p\u003e","description":"","filename":"FigPage03.jpg","url":"https://assets-eu.researchsquare.com/files/rs-3937122/v1/5a82b26cffb2b942b255a53d.jpg"},{"id":51362784,"identity":"a8cb69b2-950c-47bf-85cb-d44a78f21aea","added_by":"auto","created_at":"2024-02-20 09:12:36","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":9529395,"visible":true,"origin":"","legend":"\u003cp\u003eBasidiomata of \u003cem\u003ePseudosperma stramineum\u003c/em\u003e. \u003cstrong\u003ea-\u003c/strong\u003e Collection AR-22-008 (holotype). \u003cstrong\u003eb-\u003c/strong\u003e Collection AR-22-009. \u003cstrong\u003ec-\u003c/strong\u003e Collection AR-22-015. Bars = 10 mm.\u003c/p\u003e","description":"","filename":"FigPage04.jpg","url":"https://assets-eu.researchsquare.com/files/rs-3937122/v1/099bc316864a8d2ce0600a62.jpg"},{"id":51362307,"identity":"49c07fac-669e-4749-b769-a3cebffbc3d5","added_by":"auto","created_at":"2024-02-20 09:04:36","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":742774,"visible":true,"origin":"","legend":"\u003cp\u003eMicroscopic features of \u003cem\u003ePseudosperma stramineum\u003c/em\u003e(AR-22-008, holotype)\u003cem\u003e.\u003c/em\u003e \u003cstrong\u003ea-\u003c/strong\u003e Basidiospores. \u003cstrong\u003eb-\u003c/strong\u003eBasidia. \u003cstrong\u003ec-\u003c/strong\u003e Caulocystidia. \u003cstrong\u003ed-\u003c/strong\u003e Cheilocystidia. \u003cstrong\u003ee-\u003c/strong\u003eParacystidia. \u003cstrong\u003ef-\u003c/strong\u003e Pileipellis. Bars = 10 μm.\u003c/p\u003e","description":"","filename":"FigPage05.jpg","url":"https://assets-eu.researchsquare.com/files/rs-3937122/v1/3b006f172590bd388f528070.jpg"},{"id":51362312,"identity":"853e5f45-99cd-4711-ba9f-b4e2411bbca8","added_by":"auto","created_at":"2024-02-20 09:04:36","extension":"jpg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":3729158,"visible":true,"origin":"","legend":"\u003cp\u003eBasidiomata of \u003cem\u003ePseudosperma squarrosofulvum \u003c/em\u003e(AR-22-024, holotype). Bar = 10 mm.\u003c/p\u003e","description":"","filename":"FigPage06.jpg","url":"https://assets-eu.researchsquare.com/files/rs-3937122/v1/2adef0f713924470fdca3b82.jpg"},{"id":51362313,"identity":"26ece872-10e3-42cd-9363-f9c9bab99cc6","added_by":"auto","created_at":"2024-02-20 09:04:36","extension":"jpg","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":848753,"visible":true,"origin":"","legend":"\u003cp\u003eMicroscopic features of \u003cem\u003ePseudosperma squarrosofulvum \u003c/em\u003e(AR-22-024, holotype)\u003cem\u003e.\u003c/em\u003e \u003cstrong\u003ea-\u003c/strong\u003e Basidiospores. \u003cstrong\u003eb-\u003c/strong\u003e Basidia. \u003cstrong\u003ec-\u003c/strong\u003eParacystidia. \u003cstrong\u003ed-\u003c/strong\u003e Cheilocystidia. \u003cstrong\u003ee-\u003c/strong\u003e Caulocystidia. \u003cstrong\u003ef-\u003c/strong\u003ePileipellis. Bars = 10 μm.\u003c/p\u003e","description":"","filename":"FigPage07.jpg","url":"https://assets-eu.researchsquare.com/files/rs-3937122/v1/964bf90e9b37b37f203fe701.jpg"},{"id":51362316,"identity":"2fa4eb78-b091-4976-bb19-bc5189c01250","added_by":"auto","created_at":"2024-02-20 09:04:37","extension":"jpg","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":4987645,"visible":true,"origin":"","legend":"\u003cp\u003eBasidiomata of \u003cem\u003ePseudosperma cremeo-ochraceum\u003c/em\u003e.\u003cem\u003e \u003c/em\u003e\u003cstrong\u003ea, b-\u003c/strong\u003e Collection AR-22-088 (holotype). \u003cstrong\u003ec, d-\u003c/strong\u003e Collection AR-22-089. Bars = 5 mm.\u003c/p\u003e","description":"","filename":"FigPage08.jpg","url":"https://assets-eu.researchsquare.com/files/rs-3937122/v1/a967e44fbf2440c5ebd205a8.jpg"},{"id":51362785,"identity":"3cd834e2-6e7f-4b8c-9be8-948dacc0b540","added_by":"auto","created_at":"2024-02-20 09:12:36","extension":"jpg","order_by":9,"title":"Figure 9","display":"","copyAsset":false,"role":"figure","size":845685,"visible":true,"origin":"","legend":"\u003cp\u003eMicroscopic features of \u003cem\u003ePseudosperma cremeo-ochraceum\u003c/em\u003e(AR-22-088, holotype)\u003cem\u003e.\u003c/em\u003e \u003cstrong\u003ea-\u003c/strong\u003e Basidiospores. \u003cstrong\u003eb-\u003c/strong\u003eBasidia. \u003cstrong\u003ec-\u003c/strong\u003e Cheilocystidia. \u003cstrong\u003ed-\u003c/strong\u003e Paracystidia. \u003cstrong\u003ee-\u003c/strong\u003ePileipellis. Bars = 10 μm.\u003c/p\u003e","description":"","filename":"FigPage09.jpg","url":"https://assets-eu.researchsquare.com/files/rs-3937122/v1/ee46789b39ace78457fc22c2.jpg"},{"id":51362786,"identity":"7238c59e-86b4-4a5d-b04c-ab4a1847cf78","added_by":"auto","created_at":"2024-02-20 09:12:37","extension":"jpg","order_by":10,"title":"Figure 10","display":"","copyAsset":false,"role":"figure","size":7868597,"visible":true,"origin":"","legend":"\u003cp\u003eBasidiomata of \u003cem\u003ePseudosperma tiliae\u003c/em\u003e in the field. \u003cstrong\u003ea-\u003c/strong\u003eCollection OKA-TR3501 (holotype). \u003cstrong\u003eb-\u003c/strong\u003e Collection OKA-TR3502. \u003cstrong\u003ec-\u003c/strong\u003e Collection OKA-TR3503. Bars = 10 mm.\u003c/p\u003e","description":"","filename":"FigPage10.jpg","url":"https://assets-eu.researchsquare.com/files/rs-3937122/v1/f90ed5714b21a1162af7283f.jpg"},{"id":51362317,"identity":"ee6f359d-d707-47e2-b006-6a0504725132","added_by":"auto","created_at":"2024-02-20 09:04:37","extension":"jpg","order_by":11,"title":"Figure 11","display":"","copyAsset":false,"role":"figure","size":882192,"visible":true,"origin":"","legend":"\u003cp\u003eMicroscopic features of \u003cem\u003ePseudosperma tiliae\u003c/em\u003e(OKA-TR3501, holotype)\u003cem\u003e.\u003c/em\u003e \u003cstrong\u003ea-\u003c/strong\u003e Basidiospores.\u003cstrong\u003e b-\u003c/strong\u003eBasidia. \u003cstrong\u003ec-\u003c/strong\u003e Cheilocystidia. \u003cstrong\u003ed-\u003c/strong\u003e Paracystidia. \u003cstrong\u003ee-\u003c/strong\u003ePileipellis. \u003cstrong\u003ef-\u003c/strong\u003e Caulocystidia. Bars = 10 μm.\u003c/p\u003e","description":"","filename":"FigPage11.jpg","url":"https://assets-eu.researchsquare.com/files/rs-3937122/v1/c10ce31637cd198fc6a82249.jpg"},{"id":54303668,"identity":"0a933603-dbe5-4db5-86aa-9a3b028140fa","added_by":"auto","created_at":"2024-04-08 15:09:06","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2592576,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3937122/v1/e2ef9d75-9382-4a39-ae0d-d0896f2945ea.pdf"}],"financialInterests":"","formattedTitle":"Five new species of Pseudosperma (Inocybaceae, Agaricales) from Benin and Turkey based on morphological characteristics and phylogenetic evidence","fulltext":[{"header":"Introduction","content":"\u003cp\u003eBy a recent molecular investigation based on several independent genetic loci, seven genera were identified within the family Inocybaceae J\u0026uuml;lich: \u003cem\u003eAuritella\u003c/em\u003e Matheny \u0026amp; Bougher, \u003cem\u003eInocybe\u003c/em\u003e (Fr.) Fr., \u003cem\u003eInosperma\u003c/em\u003e (K\u0026uuml;hner) Matheny \u0026amp; Esteve-Rav., \u003cem\u003eMallocybe\u003c/em\u003e (Kuyper) Matheny, Vizzini \u0026amp; Esteve-Rav., \u003cem\u003eNothocybe\u003c/em\u003e Matheny \u0026amp; K.P.D. Latha, \u003cem\u003ePseudosperma\u003c/em\u003e Matheny \u0026amp; Esteve-Rav., and \u003cem\u003eTubariomyces\u003c/em\u003e Esteve-Rav. \u0026amp; Matheny (Matheny et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The newly established genus \u003cem\u003ePseudosperma\u003c/em\u003e was initially classified as \u003cem\u003eInocybe\u003c/em\u003e section \u003cem\u003eRimosae\u003c/em\u003e sensu stricto (=\u0026thinsp;\u003cem\u003ePseudosperma\u003c/em\u003e clade) (Matheny \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Larsson et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) in the subgenus \u003cem\u003eInosperma\u003c/em\u003e (Kuyper \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1986\u003c/span\u003e; Bon \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1997\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSpecies of \u003cem\u003ePseudosperma\u003c/em\u003e are characterized by a rimose pileus, a furfuraceous or pruinose stipe, smooth, elliptical to indistinctly phaseoliform basidiospores, hyaline, non-necropigmented basidia, cylindrical to clavate cheilocystidia with thin walls, and absence of pleurocystidia (Bandini and Oertel \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Cervini et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Matheny et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Saba et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). \u003cem\u003ePseudosperma\u003c/em\u003e species form ectomycorrhizal symbioses with both angiosperm or gymnosperm trees and occur in a variety of habitats, including temperate forests dominated by spp. of \u003cem\u003eBetula\u003c/em\u003e, \u003cem\u003eCedrus\u003c/em\u003e, \u003cem\u003ePicea\u003c/em\u003e, \u003cem\u003ePinus\u003c/em\u003e, \u003cem\u003ePopulus\u003c/em\u003e, \u003cem\u003eQuercus\u003c/em\u003e, and \u003cem\u003eSalix\u003c/em\u003e (Kuyper \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1986\u003c/span\u003e; Stangl \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e1989\u003c/span\u003e; Jacobsson \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2008\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eCurrently 80 species of \u003cem\u003ePseudosperma\u003c/em\u003e are described worldwide (Bandini et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2022\u003c/span\u003e, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), including species recently introduced from Austria (Bandini et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), Australia (Matheny and Bougher \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), China (Yu et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Mao et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Yan et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Zhao et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), Germany (Bandini and Oertel \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Bandini et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2021\u003c/span\u003e, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2022\u003c/span\u003e, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), India (Latha et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), Italy (Cervini et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Sanna et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), Pakistan (Jabeen and Khalid \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Saba et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Jabeen et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Naseer et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), Spain (Sanna et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), Sweden (Bandini et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) and Turkey (Kaygusuz et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Despite this progress, the diversity of \u003cem\u003ePseudosperma\u003c/em\u003e species, particularly in tropical regions such as West Africa, remains poorly explored and documented. In addition to this, there are numerous cryptic and semi-cryptic species waiting for discovery (Ryberg et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Matheny and Bougher \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Yan et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWest Africa is a region of species rich ecosystems with forests ranging among the 25 world hotspots that deserve top conservation priorities (Myers et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). The mycological exploration of West Africa, however, shows that the identified fungal species diversity in six countries in this region does not exceed 2% of the existing diversity (Piepenbring et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). For Benin, only 432 fungal species have been documented in (Piepenbring et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). To date, the only species of \u003cem\u003ePseudosperma\u003c/em\u003e reported from Benin is \u003cem\u003ePseudosperma squamatum\u003c/em\u003e (J.E. Lange) Matheny \u0026amp; Esteve-Rav., historically called \u003cem\u003eInocybe squamata\u003c/em\u003e J.E. Lange (Lange \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1917\u003c/span\u003e; Boa \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Piepenbring et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). No further species of \u003cem\u003ePseudosperma\u003c/em\u003e has ever been cited for any African country according to the literature available to us.\u003c/p\u003e \u003cp\u003eThis study aims to increase the knowledge of the species diversity of \u003cem\u003ePseudosperma\u003c/em\u003e, with particular emphasis on the description of four new species from Benin and one from Turkey. Thereby, we contribute to the knowledge of morphological diversity, ecology, biogeography, and phylogeny of these hidden fungal treasures of Western Eurasia and West Africa.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eSampling and Morphological Studies\u003c/h2\u003e \u003cp\u003eThe specimens from Benin were collected during a mycological survey conducted from June to August 2022. In Turkey, samples were picked up during fieldwork in the Isparta Province in 2022 and 2023. The macroscopic characteristics were obtained from fresh specimens, field notes and photographs taken in situ. Standardized colour values were documented using the Munsell Soil Color Charts (Munsell \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1975\u003c/span\u003e). Microscopic features were observed using 1% Congo red (w/v) or 5% potassium hydroxide (KOH) (w/v). All samples were analysed and photographed with a light microscope. A minimum of thirty basidiospores were measured for each collection. Q values (ratio of length to width of basidiospore) and average values (length and width of basidiospore or cystidia) are presented. SD is the abbreviation for the standard deviation of the length \u0026times; width. The terminology of Vellinga (\u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e1988\u003c/span\u003e) is used for macro- and micro-characters. Index Fungorum (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.IndexFungorum.org\u003c/span\u003e\u003cspan address=\"http://www.IndexFungorum.org\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) and the International Index of Plant Names (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.ipni.org\u003c/span\u003e\u003cspan address=\"https://www.ipni.org\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) were used as sources for taxonomic names and nomenclature. The samples from Benin are deposited in the fungarium of the Staatliches Museum f\u0026uuml;r Naturkunde Stuttgart (STU) or the mycological herbarium of the University of Parakou (UNIPAR). The Turkish specimens are stored in the fungarium of the Isparta University of Applied Sciences (ISUF).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eMolecular Analyses\u003c/h2\u003e \u003cp\u003eGenomic DNA was isolated from \u003cem\u003ePseudosperma\u003c/em\u003e specimens using the innuPREP Plant DNA Kit (Analytik Jena, Jena, Germany) and the Fungi/Yeast Genomic DNA Isolation Kit (Norgen Biotek Corp, Ontario, Canada). For amplification of nuclear rDNA internal transcribed spacer region ITS1-5.8S-ITS2 (ITS) the primer pair ITS1F/ITS4 (White et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Gardes and Bruns \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1993\u003c/span\u003e), and for the nuclear 28S rDNA (LSU) the primer pair LR0R/LR5 (Vilgalys and Hester \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Rehner and Samuels \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e1994\u003c/span\u003e) were used. Polymerase chain reaction (PCR) procedures were performed according to the methods described by Kaygusuz et al. (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The PCR products were sequenced at Microsynth Seqlab (G\u0026ouml;ttingen, Germany) and the Sanger DNA sequencing service of Source Bioscience (Berlin, Germany), using the same primers. The obtained DNA sequences were aligned and analysed using ClustalX (Thompson et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1997\u003c/span\u003e) and MEGA X v.10.0.5 (Kumar et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) and subsequently submitted to GenBank.\u003c/p\u003e \u003cp\u003eA total of 18 DNA sequences (nine from the nrITS and nine from the nrLSU) from nine collections were newly generated. BLASTn searches were conducted in the NCBI GenBank. For phylogenetic analyses, sequences with high similarity (with maximum identities larger than 82%) to the new sequences were retrieved from GenBank (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e\u003ca href=\"http://www.IndexFungorum.org\" target=\"_blank\"\u003ewww.ncbi.nlm.nih.gov\u003c/a\u003e\u003c/span\u003e\u003cspan address=\"http://www.ncbi.nlm.nih.gov\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) and UNITE (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://unite.ut.ee/\u003c/span\u003e\u003cspan address=\"https://unite.ut.ee/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e, K\u0026otilde;ljalg et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2005\u003c/span\u003e) databases, along with sequences from recent publications (Ryberg et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Larsson et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Matheny \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Matheny et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Vauras and Larsson \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Sj\u0026ouml;kvist et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Kropp et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Osmundson et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Pradeep et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Matheny and Bougher \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Tibpromma et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Bau and Fan \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Li et al. 2018; Ullah et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Matheny and Kudzma \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Matheny et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Bandini and Oertel \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Cervini et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Jabeen and Khalid \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Saba et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Yu et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Bandini et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2021\u003c/span\u003e, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2022\u003c/span\u003e, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Mao et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Zhao et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Kaygusuz et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Latha et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Naseer et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The nrITS and nrLSU rDNA sequences were separately aligned using MAFFT 7.11 (Katoh et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) applying the E-INS-i iterative method, followed by manual corrections in AliView V.1.28 (Larsson \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). \u003cem\u003eMallocybe agardhii\u003c/em\u003e (N. Lund) Matheny \u0026amp; Esteve-Rav. (AB980912) and \u003cem\u003eM\u003c/em\u003e. \u003cem\u003epicea\u003c/em\u003e L. Fan \u0026amp; N. Mao (BJTC FM555) were designated as outgroups. Multiple sequence alignments were inspected using MEGA X 10.0.5 prior to subsequent analyses. A single combined dataset of nrITS-nrLSU rDNA sequences was assembled for phylogenetic analysis. The optimal evolutionary model for each segment was determined using MrModeltest 2.3 (Nylander \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Phylogenetic assessments employed both Maximum Likelihood (ML) and Bayesian Inference (BI) approaches on the concatenated genes. The ML analysis was conducted with IQ-TREE v.1.6.12 (Minh et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), using the Ultrafast Bootstrap (UFBoot) (Hoang et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) approach with five thousand bootstrap iterations. The BI analysis using the Markov Chain Monte Carlo (MCMC) method was conducted in MrBayes 3.2.5 (Ronquist et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) over 1 \u0026times; 10\u003csup\u003e7\u003c/sup\u003e generations, sampling trees every thousand generations. Phylogenetic trees were visualized using FigTree v1.4.4 (Rambaut \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), with only Maximum Likelihood Bootstrap (MLB) values above 75% and Bayesian Posterior Probabilities (BPP) exceeding 0.90 being indicated.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cstrong\u003ePhylogeny\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe combined nrITS and nrLSU rDNA sequence dataset for \u003cem\u003ePseudosperma\u003c/em\u003e species, including 18 new sequences of the specimens from Benin and Turkey, consisted of 107 taxa with 2480 characters, of which 631 were parsimony-informative, 274 singleton sites, and 1575 constant sites. The best models in ML were TVM+F+I+G4 for nrITS and nrLSU rDNA, and GTR+I+G in the BI analysis. This matrix exhibited 1222 unique alignment patterns. The estimated nucleotide substitution rates were as follows: A-C = 1.06255, A-G = 5.05462, A-T = 1.63234, C-G = 0.42882, C-T = 5.05462, G-T = 1.00000. Base frequencies were determined as A = 0.263, C = 0.190, G = 0.247, T = 0.300. The gamma distribution shape parameter α was calculated to be 0.617. Phylogenetic trees derived from ML and BI analyses showed largely congruent topologies. The topology resulting from the ML analysis was selected for presentation, with statistical support values indicated by Maximum Likelihood Bootstrap (MLB) and Bayesian Posterior Probabilities (BPP) values (Fig. 1).\u003c/p\u003e\n\u003cp\u003eMolecular analyses based on the combined dataset revealed that \u003cem\u003ePseudosperma\u003c/em\u003e specimens from Benin and Turkey are genetically distinct from other species within the genus represented by molecular sequences data. Their sequences form part of five independent lineages, as shown in Fig. 1. The first lineage with high statistical support (MLB = 100%, BPP = 1.0) consists of three specimens of the new species called \u003cem\u003ePseudosperma tiliae\u0026nbsp;\u003c/em\u003etogether with \u003cem\u003eP. mediterraneum\u003c/em\u003e (Kuyper) Bandini, B. Oertel \u0026amp; U. Eberh. from Italy in the same subclade. The second lineage (MLB = 99%, BPP = 1.0) consists of \u003cem\u003ePseudosperma\u003c/em\u003e \u003cem\u003estramineum\u003c/em\u003e andfive\u0026nbsp;undescribed and unpublished sequences (HLA0707, HLA0386, 486ad3d1, 79c39691 and ASV_39) from Benin. The third lineage is formed by a single sequence labelled as \u003cem\u003ePseudosperma squarrosofulvum\u003c/em\u003e (AR-22-024) from Benin. The fourth lineage (MLB = 100%, BPP = 1.0) includes the new species \u003cem\u003ePseudosperma beninense\u0026nbsp;\u003c/em\u003e(AR-22-037) from Benin and an undescribed and unpublished sequence from an uncultured fungus (ASV_330). \u003cem\u003ePseudosperma beninense\u003c/em\u003e, \u003cem\u003eP. squarrosofulvum\u003c/em\u003e,and \u003cem\u003eP. stramineum\u003c/em\u003e form a distinct, statistically highly supported clade of exclusively Beninese species (MLB = 100%, BPP = 1.0). The last lineage (MLB = 100%, BPP = 1.0) comprises the new species \u003cem\u003ePseudosperma\u0026nbsp;cremeo-ochraceum\u0026nbsp;\u003c/em\u003e(AR-22-088) and three undescribed and unpublished sequences (HLA0454, ASV_313, 881fdb9c) from Benin. The new sequences from Benin are located in a clade formed by a total of ten known species that all originate from the palaeotropics (and subtropics).The sequences of the recently collected \u003cem\u003ePseudosperma\u0026nbsp;\u003c/em\u003especimens consistently form distinct lineages in all phylogenetic analyses and can not be assigned to any existing \u003cem\u003ePseudosperma\u003c/em\u003e species concept by morphological characteristics. Therefore, we propose them as species new to science and provide detailed descriptions of these species in the following.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFigure 1 [near here]\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTaxonomy\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003ePseudosperma beninense\u0026nbsp;\u003c/em\u003e\u003c/strong\u003eKaygusuz, Bandini, Rühl, Sarawi, Yorou\u0026nbsp;\u0026amp;\u0026nbsp;M. Piepenbr.,\u0026nbsp;sp. nov.\u0026nbsp;(Figs. 2\u0026nbsp;and\u0026nbsp;3)\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eMycoBank:\u003c/em\u003e\u003c/strong\u003eMB 851970\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eEtymology:\u003c/em\u003e\u003c/strong\u003e The specific epithet refers to the locality where the type specimen was collected.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eHolotype:\u003c/em\u003e\u003c/strong\u003eBenin, Borgou Department, Fôret Classée de l'Ouémé Supérieur, on soil in savannah forest dominated by \u003cem\u003eIsoberlinia doka\u0026nbsp;\u003c/em\u003eCraib \u0026amp; Stapf,\u003cem\u003e\u0026nbsp;I\u003c/em\u003e.\u003cem\u003e\u0026nbsp;tomentosa\u0026nbsp;\u003c/em\u003e(Harms)Craib \u0026amp; Stapf,\u003cem\u003e\u0026nbsp;Monotes kerstingii\u003c/em\u003e Gilg and\u0026nbsp;\u003cem\u003eUapaca togoensis\u0026nbsp;\u003c/em\u003ePax, at\u0026nbsp;09°15'37.9\"N, 002°11'03.9\"E, 340 m asl., 18 June 2022, leg. A. Rühl, C. Manz, D. Dongnima, F. Hampe \u0026amp; S. Sarawi (AR-22-037, STU). GenBank accession numbers PP060393 (nrITS) and PP060408 (nrLSU).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eDiagnosis:\u003c/em\u003e\u003c/strong\u003eDiffers from\u0026nbsp;\u003cem\u003eP. squamatum\u003c/em\u003e by\u0026nbsp;smaller basidiomata with brown to straw-brown pileus, whitish to light yellow stipe,\u0026nbsp;longer\u0026nbsp;basidiospores\u0026nbsp;(on average 12.5 × 7.0 μm) mostly with acute apex,\u0026nbsp;longer\u0026nbsp;(on av. 51 × 14 μm) and\u0026nbsp;oblong to cylindrical or narrowly clavate\u0026nbsp;cheilocystidia, and by distinct nrITS and nrLSU rDNA sequences.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eDescription:\u003c/em\u003e\u003c/strong\u003ePileus 7–14 mm diam.,\u0026nbsp;when young paraboloid, later hemispherical to convex, with or without low and broad umbo, margin inflexed when young, later long deflexed, surface dry, tomentose-lanose to subtomentose, radially fibrillose to rimulose outwards, colour light brown\u0026nbsp;(2.5Y 8–7/6, 8/8)\u0026nbsp;to straw brown\u0026nbsp;(2.5Y 6/4–6), slightly darker at the centre\u0026nbsp;(2.5Y 6/8). Lamellae moderately crowded, adnexed, subventricose, whitish to yellowish-white\u0026nbsp;or light yellow,\u0026nbsp;edge slightly\u0026nbsp;eroded,\u0026nbsp;whitish. Stipe 17–25 × 1.2–1.8 mm, central, cylindrical with subbulbous base up to 2.5 mm diam., solid,\u0026nbsp;surface whitish to light yellow, pruinose only near the apex.\u0026nbsp;Colour of exsiccate: pileus very light yellow to light straw yellow\u0026nbsp;(5Y 8/2–4, 7/4), lamellae and stipe\u0026nbsp;whitish.\u0026nbsp;Smell\u0026nbsp;unrecorded.\u003c/p\u003e\n\u003cp\u003eBasidiospores 10.7–14.7 µm (av. 12.5 µm, SD\u0026nbsp;\u003cem\u003e±\u0026nbsp;\u003c/em\u003e0.4 µm) × 6.2–8.3 µm (av. 7.0 µm, SD\u0026nbsp;\u003cem\u003e±\u0026nbsp;\u003c/em\u003e0.5 µm); Q = 1.4–2.1 (av. 1.8, SD\u0026nbsp;\u003cem\u003e±\u0026nbsp;\u003c/em\u003e0.1) (n = 90 of 1 coll.),\u0026nbsp;mainly (sub)amgydaliform\u0026nbsp;with acute apex,\u0026nbsp;also subcylindrical and subellipsoid, with guttules, smooth, thick-walled, dark yellowish brown in 5% KOH. Basidia 35–42 × 10–13 µm, clavate, 4-spored, thin-walled, hyaline. Cheilocystidia 37–65 µm (av. 51 µm, SD\u0026nbsp;\u003cem\u003e±\u0026nbsp;\u003c/em\u003e7.0 µm) × 9–20 µm (av. 14 µm, SD\u0026nbsp;\u003cem\u003e±\u0026nbsp;\u003c/em\u003e2.3 µm); Q = 2.4–5.8 (av. 4.2, SD\u0026nbsp;\u003cem\u003e±\u0026nbsp;\u003c/em\u003e0.6) (n = 45 of 1 coll.), mostly oblong to cylindrical or narrowly clavate, sometimes with\u0026nbsp;subcapitate apex, sometimes in chains of 1–3 cells,\u0026nbsp;thin-walled, hyaline or\u0026nbsp;very pale yellowish-brown in\u0026nbsp;5% KOH.\u0026nbsp;Paracystidia 20–35 × 9–14 µm, cylindrical to broadly clavate, in chains of 2–4 cells,\u0026nbsp;thin-walled, hyaline or\u0026nbsp;very pale yellowish-brown in\u0026nbsp;5% KOH. Pileipellis cutis, consisting of long cylindrical or narrowly fusiform terminal cells, with\u0026nbsp;sharply pointed apex, 60–170(220) × 11–25 µm,\u0026nbsp;smooth,\u0026nbsp;thin-walled,\u0026nbsp;pale yellowish-brown in\u0026nbsp;5% KOH. Caulocystidia\u0026nbsp;a cutis with transitions to a trichoderm\u0026nbsp;composed predominantly of\u0026nbsp;multiseptate cylindrical\u0026nbsp;to inflated hyphae, 10–100 × 6.5–12 µm, sometimes with\u0026nbsp;subcapitate apex, often in bundles, smooth, thin-walled, hyaline\u0026nbsp;in\u0026nbsp;5% KOH.\u0026nbsp;Stipitipellis a cutis of parallel hyphae, 5–18 µm wide, thin-walled, hyaline\u0026nbsp;in\u0026nbsp;5% KOH. Clamp connections present in all parts examined.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFigure 2 [near here]\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eHabitat\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u003cem\u003eand\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u003cem\u003edistribution:\u003c/em\u003e\u003c/strong\u003eBasidiomata solitary, terrestrial, on wet and sandy soils, growing in a forest dominated by species of Caesalpiniaceae (\u003cem\u003eIsoberlinia\u003c/em\u003e spp.), Dipterocarpaceae (\u003cem\u003eMonotes kerstingii\u003c/em\u003e), and Phyllanthaceae (\u003cem\u003eUapaca togoensis\u003c/em\u003e). Currently known only from Benin.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eAdditional specimen examined:\u003c/em\u003e\u003c/strong\u003e Based on the phylogenetic analysis, one further specimen from West Africa (ASV_330) belongs to \u003cem\u003ePseudosperma\u0026nbsp;\u003c/em\u003e\u003cem\u003ebeninense\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFigure 3 [near here]\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eDiscussion:\u003c/em\u003e\u003c/strong\u003ePhylogenetic analyses inferred from nrITS and nrLSU rDNA sequences\u0026nbsp;show that \u003cem\u003ePseudosperma beninense\u0026nbsp;\u003c/em\u003eforms a monophyletic subclade within \u003cem\u003ePseudosperma\u003c/em\u003e and is closely related to \u003cem\u003eP.\u0026nbsp;\u003c/em\u003e\u003cem\u003esquarrosofulvum\u0026nbsp;\u003c/em\u003eand \u003cem\u003eP.\u0026nbsp;\u003c/em\u003e\u003cem\u003estramineum\u003c/em\u003e, two other new species presented in this study. When the nrITS DNA sequences generated from \u003cem\u003ePseudosperma beninense\u0026nbsp;\u003c/em\u003eare compared with the sequences of \u003cem\u003eP.\u0026nbsp;\u003c/em\u003e\u003cem\u003esquarrosofulvum\u0026nbsp;\u003c/em\u003eand \u003cem\u003eP.\u0026nbsp;\u003c/em\u003e\u003cem\u003estramineum\u003c/em\u003e, 56 nucleotide differences (83% similarity) were observed in the nrITS DNA sequences of \u003cem\u003eP.\u0026nbsp;\u003c/em\u003e\u003cem\u003esquarrosofulvum\u0026nbsp;\u003c/em\u003eand 52 differences (85% similarity) in the sequences of \u003cem\u003eP.\u0026nbsp;\u003c/em\u003e\u003cem\u003estramineum\u003c/em\u003e.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eMorphologically,\u003cem\u003e\u0026nbsp;Pseudosperma\u0026nbsp;\u003c/em\u003e\u003cem\u003esquarrosofulvum\u0026nbsp;\u003c/em\u003ediffers from \u003cem\u003eP. beninense\u0026nbsp;\u003c/em\u003eby its pileus\u0026nbsp;surface, which is covered by yellow ochre to brownish-yellow fibrillose scales,\u0026nbsp;longer\u0026nbsp;basidiospores\u0026nbsp;(on av. 12.9\u0026nbsp;×\u0026nbsp;6.2\u0026nbsp;µm), somewhat shorter and narrower\u0026nbsp;cheilocystidia(on av. 44\u0026nbsp;×\u0026nbsp;11\u0026nbsp;µm), mostly\u0026nbsp;utriformparacystidia, pileipellis hyphae often with encrusted walls, and mostly\u0026nbsp;cylindrical or narrowly clavate\u0026nbsp;caulocystidia.\u003cem\u003e\u0026nbsp;Pseudosperma\u0026nbsp;\u003c/em\u003e\u003cem\u003estramineum\u003c/em\u003e has a predominantly straw-yellow to buff pileus with a distinct\u0026nbsp;squamulose-squarrose surface,\u0026nbsp;pale brown lamellae when old, mostly oblong basidiospores, smaller cheilocystidia\u0026nbsp;(on av. 40 × 14 μm), a pileipellis with\u0026nbsp;strongly incrusted walls, and somewhat shorter caulocystidia\u0026nbsp;(up to 90\u0026nbsp;μm in\u0026nbsp;length).\u003c/p\u003e\n\u003cp\u003eMorphologically, the species closest to the new species\u003cem\u003ePseudosperma\u003c/em\u003e\u003cem\u003e\u0026nbsp;beninense\u0026nbsp;\u003c/em\u003eis\u0026nbsp;\u003cem\u003eP. squamatum\u003c/em\u003e, which differs mainly by larger basidiomata, with a pileus measuring 30–70 mm in diameter, yellowish to yellow-ochraceous pileus, often with orange tinged, a longer stipe (up to 70 mm),\u0026nbsp;shorter\u0026nbsp;basidiospores (on av. 9.9\u0026nbsp;×\u0026nbsp;6.1\u0026nbsp;µm),\u0026nbsp;shorter cheilocystidia\u0026nbsp;(on av. 44\u0026nbsp;×\u0026nbsp;14\u0026nbsp;µm) that are subclavate to subglobose, and\u0026nbsp;a habitat with a\u0026nbsp;clay soil\u0026nbsp;as well as an\u0026nbsp;associated with\u0026nbsp;\u003cem\u003ePopulus\u003c/em\u003e sp. (Lange 1917; pers. observation of D. Bandini). In addition, \u003cem\u003eP\u003c/em\u003e.\u003cem\u003e\u0026nbsp;beninense\u0026nbsp;\u003c/em\u003eis distant from \u003cem\u003eP\u003c/em\u003e. \u003cem\u003esquamatum\u003c/em\u003e following phylogenetic analyses (Fig. 1).\u003c/p\u003e\n\u003cp\u003eOther tropical or subtropical species that are morphologically somewhat similar to\u0026nbsp;\u003cem\u003ePseudosperma\u003c/em\u003e \u003cem\u003ebeninense\u003c/em\u003e are\u0026nbsp;\u003cem\u003eP. fissuratum\u003c/em\u003e (Matheny \u0026amp; Bougher) Matheny \u0026amp; Esteve-Rav., \u003cem\u003eP. gracilissimum\u003c/em\u003e (Matheny \u0026amp; Bougher) Matheny \u0026amp; Esteve-Rav.,\u0026nbsp;\u003cem\u003eP. palaeotropicum\u003c/em\u003e (E. Turnbull \u0026amp; Watling) Matheny \u0026amp; Esteve-Rav. and\u0026nbsp;\u003cem\u003eP. renisporum\u003c/em\u003e (E. Horak) Matheny \u0026amp; Esteve-Rav.\u0026nbsp;\u003cem\u003ePseudosperma fissuratum\u003c/em\u003e, originally described from Australia,\u0026nbsp;differs from \u003cem\u003eP.\u003c/em\u003e \u003cem\u003ebeninense\u003c/em\u003e by a typically bicoloured pileus, the presence of a velipellis, shorter basidiospores (on av. 11.4\u0026nbsp;×\u0026nbsp;6.2\u0026nbsp;µm), longer\u0026nbsp;cheilocystidia (up to 72 µm in length),\u0026nbsp;and ecologically by an association with\u0026nbsp;\u003cem\u003eEucalyptus\u003c/em\u003e sp.(Matheny and Bougher 2017). Another Australian species, \u003cem\u003ePseudosperma gracilissimum\u003c/em\u003e, has a markedly conical pileus, slightly shorter basidiospores (on av. 9.9\u0026nbsp;×\u0026nbsp;5.9\u0026nbsp;µm), and is associated with\u0026nbsp;\u003cem\u003eAcacia\u003c/em\u003e, \u003cem\u003eAllocasuarina\u003c/em\u003e\u003cem\u003e, Corymbia\u003c/em\u003e, \u003cem\u003eEucalyptus, Lophostemon\u003c/em\u003e,and \u003cem\u003eMelaleuca\u003c/em\u003e (Matheny and Bougher 2017). \u003cem\u003ePseudosperma palaeotropicum\u003c/em\u003e, initially discovered from Singapore and later reported from Australia and Malaysia,\u0026nbsp;has\u0026nbsp;larger basidiomata with a pileus measuring 20–40 mm in diameter, a longer stipe (40–70\u0026nbsp;×\u0026nbsp;4.0–6.0\u0026nbsp;µm),\u0026nbsp;considerably shorter\u0026nbsp;basidiospores (7.0–8.3\u0026nbsp;×\u0026nbsp;4.8–6.5\u0026nbsp;µm), and is typically associated with species of Dipterocarpaceae (Turnbull 1995).\u0026nbsp;\u003cem\u003ePseudosperma renisporum\u003c/em\u003e, originally described from\u0026nbsp;New Zealand,\u0026nbsp;has a squamulose pileus centre, bean-shaped and shorter\u0026nbsp;basidiospores (9.0–12.0 × 4.5–6.5 μm),\u0026nbsp;and is associated with species of\u0026nbsp;\u003cem\u003eLeptospermum\u003c/em\u003e and \u003cem\u003eNothofagus\u003c/em\u003e (Horak 1978).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003ePseudosperma cremeo-ochraceum\u0026nbsp;\u003c/em\u003e\u003c/strong\u003eKaygusuz, Bandini, Rühl, Sarawi, Yorou\u0026nbsp;\u0026amp;\u0026nbsp;M. Piepenbr.,\u0026nbsp;sp. nov.\u0026nbsp;(Figs. 4\u0026nbsp;and\u0026nbsp;5)\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eMycoBank:\u003c/em\u003e\u003c/strong\u003e MB 851972\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eEtymology:\u003c/em\u003e\u003c/strong\u003e The specific epithet refers to the cream to ochraceous colour of the surface of the pileus.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eHolotype:\u003c/em\u003e\u003c/strong\u003eBenin, Borgou Department, Fôret l'Ouémé Supérieur, on the soil in savannah forest dominated by \u003cem\u003eIsoberlinia doka\u003c/em\u003e,\u003cem\u003e\u0026nbsp;I\u003c/em\u003e.\u003cem\u003e\u0026nbsp;tomentosa\u003c/em\u003e,\u003cem\u003e\u0026nbsp;Monotes kerstingii\u0026nbsp;\u003c/em\u003eand\u0026nbsp;\u003cem\u003eUapaca togoensis\u003c/em\u003e, at 08°36'04.7\"N, 002°36'00.7\"E, 340 m asl., 28 June 2022, leg. A. Rühl, C. Manz, D. Dongnima, F. Hampe \u0026amp; S. Sarawi (AR-22-088,\u0026nbsp;STU). GenBank accession numbers PP060394 (nrITS) and PP060409 (nrLSU).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eDiagnosis:\u003c/em\u003e\u003c/strong\u003eMost similar to the tropical Australian species \u003cem\u003ePseudosperma gracilissimum\u003c/em\u003e, but differing by a silky fibrillose pileus, longer\u0026nbsp;basidiospores (on av. 12.9\u0026nbsp;×\u0026nbsp;7.8\u0026nbsp;µm), mostly\u0026nbsp;narrowly utriform to utriform cheilocystidia\u0026nbsp;with subcapitate apex,\u0026nbsp;pileipellis elements without encrusted walls, the\u0026nbsp;presence of\u0026nbsp;caulocystidia, a different habitat dominated by\u0026nbsp;\u003cem\u003eIsoberlinia\u0026nbsp;\u003c/em\u003espp.,\u003cem\u003e\u0026nbsp;M\u003c/em\u003e.\u003cem\u003e\u0026nbsp;kerstingii\u0026nbsp;\u003c/em\u003eand\u0026nbsp;\u003cem\u003eU\u003c/em\u003e.\u003cem\u003e\u0026nbsp;togoensis\u003c/em\u003e,\u0026nbsp;and by distinct nrITS and nrLSU rDNA sequences.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFigure 4 [near here]\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eDescription:\u003c/em\u003e\u003c/strong\u003e Pileus 10–15 mm diam., broadly conical to\u0026nbsp;hemispherical, expanded plano-convex, usually with a low umbo, with a\u0026nbsp;straight and translucently striate margin reaching up to 1/4 or 2/4 of the radius, colour\u0026nbsp;cream (2.5Y 8–7/2) to yellowish brown (2.5Y 7/6–10) or ochraceous (2.5Y 6/6–8), becoming darker at the centre with age, always creamy white (2.5Y 8/2–4) to ivory white (2.5Y 7/2) at the edge, surface dry, radially silky-fibrillose. Lamellae moderately crowded to subdistant, adnexed, subventricose,\u0026nbsp;pale ivory white to pale yellow grey, becoming yellowish white, edge somewhat\u0026nbsp;eroded, whitish. Stipe 12–20 × 0.6–1.2 mm, central, cylindrical, sometimes subbulbous at the base, straight or curved towards the base of the stipe,\u0026nbsp;surface sordid white or cream to light brown when old, slightly pruinose only near the apex.\u0026nbsp;Colour of exsiccate: pileus\u0026nbsp;sordid white\u0026nbsp;coloured, lamellae and stipe\u0026nbsp;whitish.\u0026nbsp;Smell\u0026nbsp;unrecorded.\u003c/p\u003e\n\u003cp\u003eBasidiospores 11.0–16.0 µm (av. 12.9 µm, SD\u0026nbsp;\u003cem\u003e±\u0026nbsp;\u003c/em\u003e1.2 µm) × 6.5–9.3 µm (av. 7.8 µm, SD\u0026nbsp;\u003cem\u003e±\u0026nbsp;\u003c/em\u003e0.6 µm); Q = 1.4–1.9 (av. 1.7, SD\u0026nbsp;\u003cem\u003e±\u0026nbsp;\u003c/em\u003e0.1) (n = 100 of 2 coll.), mostly oblong,\u0026nbsp;with central germ pore,\u0026nbsp;with guttules, smooth, thick-walled, yellowish brown in 5% KOH. Basidia 35–45 × 11–13 µm, clavate, 4-spored, thin-walled, hyaline. Cheilocystidia 40–80 µm (av. 51 µm, SD\u0026nbsp;\u003cem\u003e±\u0026nbsp;\u003c/em\u003e6.0 µm) × 10–17 µm (av. 13.0 µm, SD\u0026nbsp;\u003cem\u003e±\u0026nbsp;\u003c/em\u003e2.0 µm); Q = 3.1–5.3 (av. 4.1, SD\u0026nbsp;\u003cem\u003e±\u0026nbsp;\u003c/em\u003e0.6) (n = 40 of 1 coll.), scattered, narrowly utriform to utriform mostly\u0026nbsp;with subcapitate apex,\u0026nbsp;hyaline or\u0026nbsp;very pale yellowish-brown in\u0026nbsp;5% KOH.\u0026nbsp;Paracystidia 25–40 × 10–18 µm, utriform with obtuse or subcapitate apex or fusiform, thin-walled, hyaline or\u0026nbsp;very pale yellowish-brown in\u0026nbsp;5% KOH. Pileipellis\u0026nbsp;a hymeniderm to epithelium formed by broadly fusiform to cylindrical terminal elements, with obtuse to mucronate apex, 47–95 × 20–40 µm,\u0026nbsp;smooth,\u0026nbsp;thin-walled,\u0026nbsp;pale yellowish-brown in\u0026nbsp;5% KOH. Caulocystidia 15–35 × 10–15 µm,\u0026nbsp;narrowly clavate,\u0026nbsp;on clusters of erect hyphae, smooth, thin-walled, hyaline\u0026nbsp;in\u0026nbsp;5% KOH.\u0026nbsp;Stipitipellis a cutis of subparallel hyphae, 6–12 µm wide, thin-walled, hyaline\u0026nbsp;in\u0026nbsp;5% KOH. Clamp connections present in all parts examined.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFigure 5 [near here]\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eHabitat\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u003cem\u003eand\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u003cem\u003edistribution:\u003c/em\u003e\u003c/strong\u003eBasidiocarpsgregarious, usually terrestrial, on wet and sandy soils, in woodlands dominated by Caesalpiniaceae (\u003cem\u003eIsoberlinia\u003c/em\u003e spp.), Dipterocarpaceae (\u003cem\u003eMonotes kerstingii\u003c/em\u003e), and Phyllanthaceae (\u003cem\u003eUapaca togoensis\u003c/em\u003e).\u0026nbsp;Currently only known from Benin.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eAdditional specimen examined:\u003c/em\u003e\u003c/strong\u003e Benin, Borgou Department, Fôret Classée de l'Ouémé Supérieur, on the soil in savannah forest dominated by \u003cem\u003eIsoberlinia\u0026nbsp;\u003c/em\u003espp., 08°36'04.7\"N, 002°36'00.7\"E, 340 m asl., 28 June 2022, leg. A. Rühl, C. Manz, D. Dongnima, F. Hampe \u0026amp; S. Sarawi (AR-22-089, UNIPAR). According to the phylogenetic analysis, one further specimen from Benin (HLA0454) collected by H.L. Aignon and two sequences obtained from soil (ASV_313\u0026nbsp;and 881fdb9c) in Benin also belong to \u003cem\u003ePseudosperma cremeo-ochraceum\u003c/em\u003e.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eIn the concatenated nrITS-nrLSU rDNA phylogeny (Fig. 1), \u003cem\u003ePseudosperma tiliae\u003c/em\u003e is sister to \u003cem\u003eP. mediterraneum\u003c/em\u003e and nested with \u003cem\u003eP. holoxanthum\u003c/em\u003e (Grund \u0026amp; D.E. Stuntz) Matheny \u0026amp; Esteve-Rav., \u003cem\u003eP. melliolens\u003c/em\u003e (K\u0026uuml;hner) Matheny \u0026amp; Esteve-Rav.,\u003cem\u003e\u0026nbsp;P. rimosum\u0026nbsp;\u003c/em\u003e(Bull.) Matheny \u0026amp; Esteve-Rav.,\u003cem\u003e\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;P. sororium\u003c/em\u003e (Kauffman) Matheny \u0026amp; Esteve-Rav. However, morphologically\u003cem\u003e\u0026nbsp;P. mediterraneum\u003c/em\u003e, originally described from Italy, differs from \u003cem\u003eP.\u003c/em\u003e \u003cem\u003etiliae\u003c/em\u003e by a pale buff to ochraceous pileus,\u0026nbsp;slightly\u0026nbsp;longer\u0026nbsp;basidiospores (on av. 13.2\u0026nbsp;\u0026times;\u0026nbsp;6.6\u0026nbsp;\u0026micro;m) with a higher Q-value (Q = 2),\u0026nbsp;shorter\u0026nbsp;cheilocystidia\u0026nbsp;(36\u0026ndash;57 \u0026times; 13\u0026ndash;26 \u0026micro;m), and a habitat on dune sand associated with \u003cem\u003ePinus pinea\u0026nbsp;\u003c/em\u003e(Kuyper 1986). The genetic distance between\u0026nbsp;\u003cem\u003ePseudosperma\u003c/em\u003e \u003cem\u003etiliae\u003c/em\u003e and\u003cem\u003e\u0026nbsp;P. mediterraneum\u003c/em\u003e is 3%, corresponding to 18 base divergences in 600 nucleotides, indicating that these are different species.\u0026nbsp;\u003cem\u003ePseudosperma\u0026nbsp;holoxantha\u003c/em\u003e, originally described from the USA, differs from \u003cem\u003eP\u003c/em\u003e. \u003cem\u003etiliae\u003c/em\u003e by a\u0026nbsp;longer (up to 100\u0026nbsp;mm in length) and pale yellow to straw yellow stipe,\u0026nbsp;shorter\u0026nbsp;basidiospores (9.0\u0026ndash;13.0\u0026nbsp;\u0026times;\u0026nbsp;6.0\u0026ndash;8.0\u0026nbsp;\u0026micro;m), longer cheilocystidia (up to 110\u0026nbsp;\u0026micro;m in length), and growth with conifers (Grund and Stuntz 1981).\u0026nbsp;\u003cem\u003ePseudosperma\u0026nbsp;melliolens\u003c/em\u003e differs by\u0026nbsp;a\u0026nbsp;brown to brownish coloured pileus, greyish or reddish stipe,\u0026nbsp;shorter\u0026nbsp;basidiospores (9.0\u0026ndash;13.5\u0026nbsp;\u0026times;\u0026nbsp;6.0\u0026ndash;8.0\u0026nbsp;\u0026micro;m) and\u0026nbsp;shorter\u0026nbsp;cheilocystidia (up to 55\u0026nbsp;\u0026micro;m in length) (K\u0026uuml;hner 1988).\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003cem\u003ePseudosperma\u0026nbsp;rimosum\u003c/em\u003e differs by\u0026nbsp;usually less stout habitus, often dull fallow pileus colour, only faint and fugacious greyish velipellis (Bulliard\u0026nbsp;1789). \u003cem\u003ePseudosperma\u0026nbsp;sororium\u003c/em\u003e, originally described from the USA, has\u0026nbsp;larger basidiomata (up to 70 mm in diam.) with a subconical to conical-campanulate pileus,\u0026nbsp;and\u0026nbsp;distinctly\u0026nbsp;longer\u0026nbsp;basidiospores (9.0\u0026ndash;16.0\u0026nbsp;\u0026times;\u0026nbsp;5.0\u0026ndash;8.0\u0026nbsp;\u0026micro;m) (Murrill et al. 1924).\u003c/p\u003e\n\u003cp\u003eOther European species that are morphologically similar to\u0026nbsp;\u003cem\u003ePseudosperma\u003c/em\u003e \u003cem\u003etiliae\u003c/em\u003e are \u003cem\u003eP. arenicola\u003c/em\u003e (R. Heim) Matheny \u0026amp; Esteve-Rav., \u003cem\u003eP\u003c/em\u003e.\u003cem\u003e\u0026nbsp;mimicum\u003c/em\u003e (Massee) Matheny \u0026amp; Esteve-Rav.,\u003cem\u003e\u0026nbsp;P. musilii\u0026nbsp;\u003c/em\u003eBandini, B. Oertel \u0026amp; Schmidt-Stohn, \u003cem\u003eP. pamukkalense\u003c/em\u003e Kaygusuz, Bandini \u0026amp; Knudsen,\u0026nbsp;\u003cem\u003eP. pseudoorbatum\u0026nbsp;\u003c/em\u003e(Esteve-Rav. \u0026amp; Garc\u0026iacute;a Blanco) Matheny \u0026amp; Esteve-Rav,\u003cem\u003e\u0026nbsp;\u003c/em\u003eand\u0026nbsp;\u003cem\u003eP\u003c/em\u003e.\u0026nbsp;\u003cem\u003espectrale\u003c/em\u003e Bandini \u0026amp; B. Oertel.\u0026nbsp;\u003cem\u003ePseudosperma arenicola\u0026nbsp;\u003c/em\u003ediffers by larger\u0026nbsp;pileus (up to 68\u0026nbsp;mm in diam.) and stipe (up to 75\u0026nbsp;mm long), distinctly\u0026nbsp;longer\u0026nbsp;basidiospores (on av. 13\u0026ndash;15.4\u0026nbsp;\u0026times;\u0026nbsp;6.3\u0026ndash;8.0\u0026nbsp;\u0026micro;m), longer (up to 105\u0026nbsp;\u0026micro;m) and mostly cylindrical cheilocystidia, cylindrical caulocystidia, and a habitat associated with \u003cem\u003eSalix repens\u0026nbsp;\u003c/em\u003eL, \u003cem\u003ePopulus canadensis\u003c/em\u003e Moench, or sometimes \u003cem\u003ePinus maritima\u0026nbsp;\u003c/em\u003eMill.\u003cem\u003e\u0026nbsp;\u003c/em\u003e(Kuyper 1986).\u003cem\u003e\u0026nbsp;Pseudosperma\u0026nbsp;\u003c/em\u003e\u003cem\u003emimicum\u003c/em\u003e differs by larger\u0026nbsp;pileus (up to 80\u0026nbsp;mm in diam.), yellow-brown lamellae, much\u0026nbsp;longer\u0026nbsp;basidiospores (14.0\u0026ndash;16.0\u0026nbsp;\u0026times;\u0026nbsp;6.0\u0026ndash;8.0\u0026nbsp;\u0026micro;m), and the absence of cystidia (Massee 1904). \u003cem\u003ePseudosperma musilii\u003c/em\u003e differs by its\u0026nbsp;dingy straw to dark brown pileus with a strongly rimose surface, shorter basidiospores (on av. 11.3\u0026nbsp;\u0026times;\u0026nbsp;6.7\u0026nbsp;\u0026micro;m), and shorter cheilocystidia\u0026nbsp;(on av. 48\u0026nbsp;\u0026times;\u0026nbsp;12\u0026nbsp;\u0026micro;m) (Bandini et al. 2023).\u0026nbsp;\u003cem\u003ePseudosperma pamukkalense\u003c/em\u003e differs by\u0026nbsp;subphaseoliform basidiospores, a predominant association with \u003cem\u003ePinus nigra\u003c/em\u003e subsp.\u003cem\u003e\u0026nbsp;pallasiana\u003c/em\u003e (Lamb.) Holmboe, and genetic differences (nrITS locus\u0026nbsp;81.5% identity) (Kaygusuz et al. 2023).\u003cem\u003e\u0026nbsp;Pseudosperma pseudoorbatum\u003c/em\u003e has a white to ivory-white pileus, notably longer basidiospores (on av.\u0026nbsp;14.6\u0026nbsp;\u0026times;\u0026nbsp;7.1\u0026nbsp;\u0026micro;m), and\u0026nbsp;a habitat with \u003cem\u003ePinus\u0026nbsp;\u003c/em\u003eforests (\u003cem\u003ePinus pinaster\u003c/em\u003e Aiton and \u003cem\u003eP\u003c/em\u003e. \u003cem\u003epinea\u003c/em\u003e L.) (Esteve-Ravent\u0026oacute;s et al. 2003).\u0026nbsp;\u003cem\u003ePseudosperma\u0026nbsp;\u003c/em\u003e\u003cem\u003espectrale\u003c/em\u003e has a whitish to straw-coloured pileus, slightly shorter basidiospores (on av. 12.1 \u0026times; 7.0 \u0026micro;m), and a habitat with conifers (Bandini et al. 2022).\u003c/p\u003e"},{"header":"Conclusions","content":"\u003cp\u003eIt was previously suggested that species of Inocybaceae began to spread over large areas of northern and southern South America, Australia, and New Zealand during the Palaeogene or later periods (Matheny 2009; Matheny et al. 2009). Recent multi-gene phylogenetic analyses confirmed close affinities between \u003cem\u003ePseudosperma\u003c/em\u003e taxa from tropical Asia, tropical Australia, and tropical China (Zhao et al. 2022). Similarly, in the present study, phylogenetic analyses generated from nrITS and nrLSU rDNA sequences revealed rather close phylogenetic relationships among \u003cem\u003ePseudosperma\u003c/em\u003e species indigenous to the tropical regions of Benin, Australia, India, and China. The new species \u003cem\u003ePseudosperma beninense\u003c/em\u003e, \u003cem\u003eP. cremeo-ochraceum\u003c/em\u003e, \u003cem\u003eP. squarrosofulvum\u003c/em\u003e, and \u003cem\u003eP. stramineum\u003c/em\u003e were found in ectomycorrhizal forests with tropical climate of Benin and are phylogenetically close to each other in a clade that in addition to the species from Benin includes six further species from tropical regions\u0026nbsp;of the palaeotropics and Australia. These six species are \u003cem\u003eP. araneosum\u003c/em\u003e from Australia (Matheny and Bougher 2017), \u003cem\u003ePseudosperma brunneosquamulosum\u003c/em\u003e, \u003cem\u003eP. luteobrunneum\u003c/em\u003e, and \u003cem\u003eP. rubrobrunneum\u003c/em\u003e from India (Tibpromma et al. 2017), and \u003cem\u003ePseudosperma fulvidiscum\u003c/em\u003e and \u003cem\u003eP. singulare\u003c/em\u003e from China (Zhao et al. 2022). The ongoing discovery and characterization of tropical to subtropical taxa within the genus will improve our understanding of their biogeographical distribution and evolutionary history.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe would like to thank Frank Lappe and Anika R\u0026uuml;b for their technical assistance, and Cathrin Manz, Felix Hampe, Boris Olou, and Sylvestre Badou for their support in the field. We are grateful to Daouda Dognima for careful driving through Benin\u0026acute;s ecosystems. This study results from the project FunTrAf that is generously supported by the German Federal Ministry of Education and Research (BMBF, 01DG20015FunTrAf). The first author also acknowledges the financial support of the Scientific and Technical Research Council of Turkey (TUBITAK) for the 2219 International Postdoctoral Research Fellowship Programme (Grant No. 1059B192202880).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contribution\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll authors contributed to the conception and design of the study. Oğuzhan Kaygusuz, Adrian R\u0026uuml;hl, Sepas Sarawi, Nourou S. Yorou, and Meike Piepenbring contributed to material preparation and data collection. Morphological characteristics were examined by Oğuzhan Kaygusuz and Ditte Bandini. Molecular lab work and phylogenetic analyses were conducted by Oğuzhan Kaygusuz, Adrian R\u0026uuml;hl, and Sepas Sarawi. The first draft of the manuscript was written by Oğuzhan Kaygusuz and Meike Piepenbring, which was then improved by changes, edits, suggestions, and comments from Ditte Bandini, Adrian R\u0026uuml;hl, Sepas Sarawi and Nourou S. Yorou. All authors read and approved the final manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFunTrAf by the German Federal Ministry of Education and Research (BMBF, 01DG20015FunTrAf) and 2219 International Postdoctoral Research Fellowship Programme (Grant No. 1059B192202880) by the Scientific and Technical Research Council of Turkey (TUBITAK).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe DNA sequences produced in this study are available on NCBI GenBank (https://www.ncbi.nlm.nih.gov).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCode availability\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics approval and consent to participate\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eInformed consent was obtained from all individual participants included in the study.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eBandini D, Oertel B (2020) Three new species of the genus \u003cem\u003ePseudosperma\u003c/em\u003e (Inocybaceae). 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PCR Protocols: A Guide to Methods and Applications, Academic Press, New York, pp 315\u0026ndash;322. https://doi.org/10.1016/B978-0-12-372180-8.50042-1\u003c/li\u003e\n\u003cli\u003eYan Y-Y, Zhang Y-Z, Vauras J, Zhao L-N, Fan Y-G, Li H-J, Xu F (2022) \u003cem\u003ePseudosperma arenarium \u003c/em\u003e(Inocybaceae), a new poisonous species from Eurasia, based on morphological, ecological, molecular and biochemical evidence. MycoKeys 92:79\u0026ndash;93. https://doi.org/10.3897/mycokeys.92.86277\u003c/li\u003e\n\u003cli\u003eYu WJ, Chang C, Qin LW, Zeng NK, Wang SX, Fan YG (2020) \u003cem\u003ePseudosperma citrinostipes \u003c/em\u003e(Inocybaceae), a new species associated with Keteleeria from southwestern China. Phytotaxa 450(1):8\u0026ndash;16. https://doi.org/10.11646/phytotaxa.450.1.2\u003c/li\u003e\n\u003cli\u003eZhao LN, Yu WJ, Deng LS, Hu JH, Ge YP, Zeng NK, Fan YG (2022) Phylogenetic analyses, morphological studies, and muscarine detection reveal two new toxic \u003cem\u003ePseudosperma\u003c/em\u003e (Inocybaceae, Agaricales) species from tropical China. Mycological Progress 21(75). https://doi.org/10.1007/s11557-022-01822-z\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"mycological-progress","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mypr","sideBox":"Learn more about [Mycological Progress](https://www.springer.com/journal/11557)","snPcode":"","submissionUrl":"https://www.editorialmanager.com/mypr/default.aspx","title":"Mycological Progress","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Ectomycorrhizal fungi, Biodiversity, Agarics, Molecular systematics, Biogeography, Taxonomy","lastPublishedDoi":"10.21203/rs.3.rs-3937122/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-3937122/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eSpecies of \u003cem\u003ePseudosperma\u003c/em\u003e (Inocybaceae) are widely distributed from temperate to tropical regions. In this study, we describe and illustrate five new species of \u003cem\u003ePseudosperma\u003c/em\u003e: \u003cem\u003eP. beninense\u003c/em\u003e, \u003cem\u003eP. cremeo-ochraceum\u003c/em\u003e, \u003cem\u003eP. squarrosofulvum\u003c/em\u003e, \u003cem\u003eP. stramineum\u003c/em\u003e, and \u003cem\u003eP. tiliae\u003c/em\u003e, based on comprehensive analyses of morphological and molecular data derived from specimens collected in Benin (West Africa) and Turkey (Western Eurasia). These new species have been found in forests with \u003cem\u003eIsoberlinia\u003c/em\u003e spp. and other ectomycorrhizal tree species in Benin and in association with \u003cem\u003eTilia platyphyllos\u003c/em\u003e in Turkey. The phylogenetic relationships of the new species were inferred through analyses of nuclear rDNA sequences, encompassing the internal transcribed spacer (ITS1-5.8S-ITS2) and 28S rDNA regions. Phylogenetic analyses revealed that \u003cem\u003eP. beninense\u003c/em\u003e, \u003cem\u003eP. cremeo-ochraceum\u003c/em\u003e, \u003cem\u003eP. squarrosofulvum\u003c/em\u003e, and \u003cem\u003eP. stramineum\u003c/em\u003e from Benin cluster with species from Australia, China, and India within a clade formed exclusively by species known from the palaeotropics and Australia, whereas \u003cem\u003eP. tiliae\u003c/em\u003e from Turkey clustered with \u003cem\u003eP. mediterraneum\u003c/em\u003e from Italy. Detailed descriptions are provided, supplemented by illustrations and line drawings of key micromorphological features. In addition, a comparative analysis with morphologically similar and phylogenetically closely related species is presented and discussed in detail.\u003c/p\u003e","manuscriptTitle":"Five new species of Pseudosperma (Inocybaceae, Agaricales) from Benin and Turkey based on morphological characteristics and phylogenetic evidence","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-02-20 09:04:31","doi":"10.21203/rs.3.rs-3937122/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewersInvited","content":"","date":"2024-02-16T02:05:51+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Mycological Progress","date":"2024-02-08T09:17:40+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-02-07T01:11:54+00:00","index":"","fulltext":""},{"type":"submitted","content":"Mycological Progress","date":"2024-02-06T06:21:54+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"mycological-progress","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mypr","sideBox":"Learn more about [Mycological Progress](https://www.springer.com/journal/11557)","snPcode":"","submissionUrl":"https://www.editorialmanager.com/mypr/default.aspx","title":"Mycological Progress","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"9ed31819-9064-4de5-83d4-e12bb37dff19","owner":[],"postedDate":"February 20th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2024-04-08T15:02:40+00:00","versionOfRecord":{"articleIdentity":"rs-3937122","link":"https://doi.org/10.1007/s11557-024-01964-2","journal":{"identity":"mycological-progress","isVorOnly":false,"title":"Mycological Progress"},"publishedOn":"2024-04-03 15:00:50","publishedOnDateReadable":"April 3rd, 2024"},"versionCreatedAt":"2024-02-20 09:04:31","video":"","vorDoi":"10.1007/s11557-024-01964-2","vorDoiUrl":"https://doi.org/10.1007/s11557-024-01964-2","workflowStages":[]},"version":"v1","identity":"rs-3937122","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-3937122","identity":"rs-3937122","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}