<hi rend="italic" style="typo_italique" >Leucocoprinus beninensis</hi > Sarawi, sp. nov., a new species of Agaricaceae from Benin<hi rend="italic" style="typo_italique" >Leucocoprinus beninensis</hi > Sarawi, sp. nov., a new species from BeninSepas SARAWIsarawi@bio.uni-frankfurt.de0000-0003-1574-6393Nourou Soulemane YOROU0000-0001-6997-811XMeike PIEPENBRING0000-0002-7043-57692026-02-12T14:17:00Service des publications scientifiques23/09/202407/01/20251-92/02/2025472/02/2025https://doi.org/10.5252/cryptogamie-mycologie2026v47a1http://cryptogamie.com/mycologie/47/1urn:lsid:zoobank.org:pub:DF691E15-4977-0C00-FF0A-BACB48C0F87F18680964dcf26aab-644e-4f32-ab75-5a0e24c41bc0314370

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border:0.75pt solid #000000;border:0.75pt solid #000000;Agaricaleslepiotaceous fungimolecular phylogenytropicsnew species.Agaricaleschampignons lépiotesphylogénie moléculairetropiquesespèce nouvelle.RévisionLeucocoprinus beninensis Sarawi, sp. nov., a new species of Agaricaceae from BeninSepas SARAWIMycology Working Group, Institute of Ecology, Evolution and Diversity, Goethe University Frankfurt am Main, Max-von-Laue-Straße 13, 60438 Frankfurt am Main (Germany)Nourou Soulemane YOROUResearch Unit Tropical Mycology and Plant-Soil Fungi Interactions (MyTIPS), Faculty of Agronomy, University of Parakou, BP 123, Parakou (Benin)Meike PIEPENBRINGMycology Working Group, Institute of Ecology, Evolution and Diversity, Goethe University Frankfurt am Main, Max-von-Laue-Straße 13, 60438 Frankfurt am Main (Germany)

Abstract. Leucocoprinus beninensis Sarawi, sp. nov., is described here as a new species from Benin, West Africa, based on morphological and phylogenetic investigation. Leucocoprinus beninensis Sarawi, sp. nov., differs from known species of Leucocoprinus Pat. and closely related species of Leucoagaricus (Locq. Ex) Singer by the combination of basidiomata with reddish brown, light brown to brown squamules on a white background, a change of colour of the basidiomata to blue-green or grey-blue when dried, the absence of cheilocystidia, and small amygdaliform to ellipsoid basidiospores. Molecular phylogenetic analyses of the nrDNA ITS region reveal that sequences of Lc. beninensis Sarawi, sp. nov., cluster with sequences of Lc. cf. caerulescens (Peck) J.F. Liang, Zhu L. Yang & J. Xu, Lc. cinnamomeodiscus Asif, Saba & M. Raza, Lc. irinellus (Chalange) Kun L. Yang, Jia Y. Lin & Zhu L. Yang, Lc. sericifer (Locq.) Asif, Saba & Vellinga, and Lc. viriditinctus (Berk. & Broome) Asif, Saba & Vellinga.

KEYWORDS: Agaricales, lepiotaceous fungi, molecular phylogeny, tropics, new species.

RÉSUMÉ. Leucocoprinus beninensis Sarawi, sp. nov., est décrit ici comme une nouvelle espèce du Bénin, en Afrique de l’Ouest, sur la base de données morphologiques et phylogénétiques. Leucocoprinus beninensis Sarawi, sp. nov., diffère des espèces connues de Leucocoprinus Pat. et d’espèces voisines de Leucoagaricus (Locq. Ex) Singer par la combinaison de basidiomes avec des squamules brun rougeâtre, brun clair à brun sur un fond blanc, un changement de couleur des basidiomes en bleu-vert ou gris-bleu lorsqu’ils sont séchés, l’absence de cheilocystides, et de petites basidiospores amygdaliformes à ellipsoïdes. Les analyses phylogénétiques moléculaires de la région ITS de nrADN révèlent que les séquences de Lc. beninensis Sarawi, sp. nov., se regroupent avec celles de Lc. cf. caerulescens (Peck) J.F. Liang, Zhu L. Yang & J. Xu, Lc. cinnamomeodiscus Asif, Saba & M. Raza, Lc. irinellus (Chalange) Kun L. Yang, Jia Y. Lin & Zhu L. Yang, Lc. sericifer (Locq.) Asif, Saba & Vellinga et Lc. viriditinctus (Berk. & Broome) Asif, Saba & Vellinga.

MOTS CLÉS: Agaricales, champignons lépiotes, phylogénie moléculaire, tropiques, espèce nouvelle.

Introduction

The genus Leucocoprinus ‌Pat. (Agaricaceae) includes 50 known species with a worldwide distribution (He et al. 2019). Morphologically, species of the genus Leucocoprinus are characterized by basidiomata with coprinoid shape, with sulcate pileus margin, free lamellae, white spore print, dextrinoid basidiospores, and absence of clamp connections (Singer 1986; Vellinga 2001).

Due to similar morphological characteristics, species of the genera Leucocoprinus and Leucoagaricus (Locq. ex) Singer were grouped in the older genus Leucocoprinus in the past, but Leucocoprinus species were separated from Leucoagaricus species based on fragile, coprinoid basidiomata with striate pileus margin and pseudoparaphysis in the hymenium (Singer 1986). However, phylogenetic studies have demonstrated that the genera Leucocoprinus and Leucoagaricus are polyphyletic (Johnson & Vilgalys 1998; Johnson 1999; Vellinga 2004; Vellinga et al. 2011; Ge et al. 2015; Ma et al. 2022). Subsequently, Redhead (2023) and Asif et al. (2024) recombined the majority of Leucoagaricus species to Leucocoprinus. However, Yang et al. (2024) recognize both genera and described several new genera based on multilocus phylogenetic analyses, and we follow this approach.

In temperate regions, the genus Leucocoprinus has been extensively studied (Candusso & Lanzoni 1990). Recently, several new species of Leucocoprinus, named as species of Leucoagaricus, have been described from the subtropics and tropics, suggesting the presence of a great diversity of species in these regions (Dennis 1952; Heinemann 1973, 1977; Pegler 1977, 1983; Vellinga et al. 2011; Jabeen et al. 2020; Ma et al. 2022; Sysouphanthong & Thongklang 2022; Ashraf et al. 2023; Rehman et al. 2023). For Benin, West Africa, only Leucoagaricus ‌americanus ‌(Peck) Vellinga and Leucocoprinuscretaceus ‌(Bull.) Locq. have been reported, but no Leucoagaricus or Leucocoprinus species was yet described from Benin (Piepenbring et al. 2020).

Material and methods
Morphological examination

Specimens of Leucocoprinus sp. were collected during a mycological survey in June to July 2022 at different sites in Benin, West Africa. The macroscopic characteristics of the fresh basidiomata were recorded in the field and colour codes were assigned using Kornerup & Wanscher (1978). The basidiomata were air-dried on an electric food dehydrator at 40 °C. Microscopic features were examined using a Nikon Eclipse 80i light microscope. Microscopic characteristics of the dried material were studied in water or 5-10 % KOH. Some sections were stained with ammoniacal 10 % Congo Red. Melzer’s reagent was used to study the dextrinoidity and Cresyl blue (0.1 % in H2O) for the metachromasy of the basidiospores. Measurements are given with extreme values in brackets. The specimens are deposited in the Herbarium FR, Frankfurt am Main, Germany.

DNA extraction, PCR amplification and sequencing

For DNA extraction, small pieces of dried basidiomata were frozen with liquid nitrogen and ground to a fine powder using a MM301 Mixer Mill (Retsch GmbH, Germany). The extraction was performed using the innuprep plant DNA kit (IST Innuscreen, Germany) following the manufacturers’ instructions. The PCR was performed using the VWR Taq DNA polymerase (VWR, Germany) in a peqStar gradient thermal cycler (PEQLAB, Germany). For the amplification of the nuclear ribosomal internal transcribed spacer (nrITS) region, the primer pairs ITS5/ITS4B were used (White et al. 1990; Gardes & Bruns 1993). PCR products were verified by gel electrophoresis using a 1 % (w/v) agarose gel. The purified PCR products were sequenced by Sanger sequencing at Microsinth SeqLab (Germany).

Phylogenetic analysis

The obtained DNA sequences were edited, assembled, and quality checked using Geneious Prime V2021.2.2 (Biomatters Ltd., New Zealand). Additional sequences of Leucoagaricus, Leucocoprinus, Micropsalliota, and Cystolepiota ‌Singer species were obtained from GenBank (Table 1). The final data sets were then aligned in MAFFT v7 (Katoh & Standley 2013) using the E-INS-i algorithm for the nrITS dataset (Katoh & Toh 2008). The alignments were checked and pruned in Mega7 (Kumar et al. 2016). A Maximum-Likelihood analysis was conducted for each dataset using RAxML 8.2.12 (Stamatakis 2014) applying the GTRCAT model with 25 per site rate categories and 1000 bootstrap iterations (Felsenstein 1985). A Bayesian Markov Chain Monte Carlo (MCMC) analysis was performed using MrBayes 3.2.7a on XSEDE (Huelsenbeck & Ronquist 2001; Ronquist et al. 2012), with two runs of 10 million generations with four chains each, a sampling frequency every 1000 generations, and a stoprule for topological convergence diagnostics falling below 0.01. The GTR model and a gamma distribution was used. The first 25 % generations were discarded as burn-in. The Maximum-Likelihood and Bayesian Markov Chain Monte Carlo analyses were performed via Cipres Science Gateway (Miller et al. 2010). Phylogenetic trees were visualized using TreeGraph2 (Stöver & Müller 2010).

Results
Phylogenetic results

The Maximum-Likelihood phylogeny (Fig. 1) based on ITS sequences of species of Leucoagaricus, Leucocoprinus, and further closely relates genera comprises 63 sequences, including 61 from GenBank. The sequences of the specimens from Benin cluster together with Lc. cinnamomeodiscus ‌Asif, Saba & M. Raza, Lc. irinellus ‌(Chalange) Kun L. Yang, Jia Y. Lin & Zhu L. Yang, Lc. sericifer ‌(Locq.) Asif, Saba & Vellinga, Lc. viriditinctus ‌(Berk. & Broome) Asif, Saba & Vellinga, and two sequences deposited as Lc. caerulescens ‌(Peck) J.F. Liang, Zhu L. Yang & J. Xu (basionym: La.coerulescens) in GenBank. The phylogenetic position of the specimens from Benin within the cluster is well supported by the classical bootstrap values as well as by the bayesian PP.

Taxonomy‌ Agaricaceae Genus Pat. Leucocoprinus ‌Chevall Pat.
Leucocoprinus ‌beninensis ‌Sarawi ‌sp. nov., (Fig. 2)DF691E15-4977-0C00-FF0A-BACB48C0F87F
Holotype

Benin•Borgou Department, Okpara, Forêt de Okpara; on sandy soil, in woodland dominated by Isoberlinia doka Craib & Stapf and Uapaca togoensis Pax; 9°14’13.2”N, 2°43’26.4”E; 200 m alt.; 12.VI.2022; S. Sarawi, S. Badou, D. Dongnima, F. Hampe, C. Manz, B. Olou, A. Rühl SeSa113; FR-0246101, isotype: UNIPAR; GenBank PX634060 (ITS).

Etymology

The epithet beninensis refers to Benin, the country in which the type specimens of this species were collected.

Additional material

Benin•Collines Department, near Kilibo; on sandy soil, in woodland dominated by Isoberlinia doka and Uapaca togoensis; 8°36’57.6”N, 2°39’54”E; 320 m alt.; 21.VI.2022; S. Sarawi, D. Dongnima, F. Hampe, C. Manz, A. Rühl SeSa186; FR-0246102; GenBank PX634061 (ITS).

MycoBank

MB#861479.

Ecology, habitat, and distribution

Basidiomata solitary or gregarious, pressumably saprotrophic, on soil, in woodland. Currently only known from Benin.

Description

Basidiomata small to medium sized. Pileus 5-25 mm diam., initially campanulate or hemispherical, then convex and eventually plano-convex, umbonate, pileus covered by almost erect, tomentose squamules, light reddish brown to light brown (7C2-4, D3-5, 8C2-4, D3-6), sometimes greyish-brown (7E2-3, F2-3) with a darker reddish brown to brown pileus centre (7E5-6, 8E4-6, F4-6), sometimes purplish-brown or grey (9E3-5, F1-5), on a slightly striate, white background. Lamellae free, moderately crowded, ventricose, 1-3 mm deep, white. Stipe 10-30 × 1-3 mm, cylindrical, equal, not bulbous at the base, smooth, white, towards the base yellowish to light brown (6A1-3; B1-2). Annulus persistent, ascending, white. Odour and taste not recorded. Basidiospores (80/2/2) (5.3–)5.8–6.4-6.9(–7.6) × (3.8–)4.1–4.3-4.5(–4.8) µm, Q = 1.33-1.69, Qav = 1.46-1.51, amygdaliform, ellipsoid to ovoid, hyaline, smooth, without germ pore, dextrinoid, non-metachromatic. Basidia (16–)17-20(–23) × (6.7–)7.6-9.0(–10.0) µm, clavate, hyaline, with oily content, mostly 4-spored, sometimes 1- or 2-spored, with up to 5 µm long sterigmata, pseudoparaphyses absent. Lamella edge fertile, consisting of a homogeneous mixture of basidia and basidioles. Cheilocystidia absent. Pleurocystidia absent. Pileus covering a hymeniderm made up of chains of cylindrical to subglobose elements, sometimes disorderly arranged, terminal elements narrowly clavate to clavate, sometimes with acute apex, (23–)29-43(–47) × (6.5–)8.5-16.0(–20.0) µm, with pale brown to brown intracellular pigment and sometimes with encrusted brown pigments. Clamp connections absent in all parts of the basidiomata.

Discussion

In this study, a species of Leucocoprinus from Benin is described as new to science based on morphological and phylogenetic studies for the first time. Basidiomata of Leucocoprinus ‌beninensis ‌Sarawi ‌sp. nov.,, are characterized by reddish brown, light brown to brown squamules on a white background on the pileus. Notably, these basidiomata, predominantly the lamellae, turn greenish-blue to greyish-blue after drying. Further, they exhibit an ascending annulus, dextrinoid, non-metachromatic, amygdaliform to ellipsoid basidiospores, and lack cheilocystidia.

Phylogenetically, Lc. beninensis ‌Sarawi ‌sp. nov.,, forms a distinct clade together with Lc. cinnamomeodiscus, Lc. irinellus, Lc. sericifer, Lc. viriditinctus, and two sequences deposited in GenBank as Lc. caerulescens (Fig. 1). The latter two sequences are labelled as “Lc. cf. caerulescens” probably due to their change of colour, basidiospore size, and cheilocystidia. No sequence data of the holotype of La. caerulescens is available, however, Liang et al. (2010) investigated the holotype and provided a detailed morphological description. Morphologically, Lc. beninensis ‌Sarawi ‌sp. nov.,, differs from most known species by the colour of its basidiomata, absence of cheilocystidia, and basidiospore size (Table 2). Leucocoprinus ‌beninensis ‌Sarawi ‌sp. nov.,, has significant smaller basidiospores than Lc. irinellus, Lc. caerulenscens, Lc. sericifer, and Lc. viriditinctus and has larger basidiospores than Lc. cinnamomeodiscus. The absence of cheilocystidia is a characteristic only shared by the closely related species Lc. irinellus, which differs by the colour of the basidiomata and larger basidiospores. Additionally, basidiomata of Lc. beninensis ‌Sarawi ‌sp. nov.,, have a less intense change in colour after drying than the closely related species.

Several further species of Leucoagaricus, Leucocoprinus, and Lepiota as described in older studies in East and Central Africa and other (sub-)tropical regions are morphologically similar to Lc. beninensis ‌Sarawi ‌sp. nov.,, but lack molecular sequence data. According to information available, basidiomata of these species do not change in colour by bruising or drying. Leucoagaricus ‌striatulus ‌Heinem., known from Central Africa, is macromorphologically similar to Lc. beninensis ‌Sarawi ‌sp. nov.,, but differs by pinkish brown colours, when dried cream with dark brown squamules, and more distinctive amygdaliform, metachromatic basidiospores (Heinemann 1973). La. fuligineus ‌Pegler and La. rhodocephalus ‌(Berk.) Pegler known from East Africa are also morphologically similar. Leucoagaricus ‌fuligineus differs from Lc. beninensis ‌Sarawi ‌sp. nov.,, by more robust basidiomata, a non-striate pileus margin, and metachromatic basidiospores (Pegler 1977). Leucoagaricus ‌rhodocephalus differs from Lc. beninensis ‌Sarawi ‌sp. nov.,, by pinkish to reddish colours, more robust basidiomta, and strongly amygdaliform basidiospores (Pegler 1977). Another morphologically similar species is Lc. venezuelanus ‌Dennis, known from Martinique, Venezuela, and Zaire (Central Africa). Leucocoprinus ‌venezuelanus differs from Lc. beninensis ‌Sarawi ‌sp. nov.,, by its greyish brown to orange-brown colours, minute brown squamules on the annulus, and the presence of a germ pore (Dennis 1952). Beeli (1932) recorded Lepiota ‌montagnei ‌(Kalchbr.) Sacc. for Central Africa based on two collections. Later, Beeli separated them due to morphological differences by introducing L. ‌montagnei var. congolensis ‌Beeli (Beeli 1936). Later Heinemann (1977) renamed one of these two collections as Lc. submontagnei ‌Heinem. based on studies of the holotype and a watercolour drawing. Leucocoprinus ‌submontagnei (former L. ‌montagnei var. congolensis) resembles Lepiota ‌micropholis ‌(Berk. & Broome) Sacc., known from Trinidad and Venezuela, but differ by larger basidiospores (Dennis 1952). Lepiota ‌micropholis differs from Lc. beninensis ‌Sarawi ‌sp. nov.,, by its dark purplish brown to black colours and smaller basidiospores (Dennis 1952). Leucocoprinus ‌submontagnei differs from Lc. beninensis ‌Sarawi ‌sp. nov.,, by basidiomata with dark brown-violet colours when fresh and ochre-brown colours when dried and slightly larger, metachromatic basidiospores (Heinemann 1977).

These results highlight the need for further exploration of unstudied fungal species in the forests of Benin. Additionally, they emphasise the importance of re-investigating and re-collecting previously documented African species to enhance our understanding through modern molecular techniques.

Fig. 1. — Maximum-Likelihood phylogram of Leucocoprinus ‌Pat. species and the outgroup based on ITS sequences. Maximum-Likelihood bootstrap values (> 70) are shown above branches and posterior probabilities obtained from Bayesian analysis (> 90 %) are shown below. Scale bar: estimated changes per nucleotide. 10.5281/zenodo.18680966
Table 1. — Specimens used for phylogenetic analyses and the corresponding GenBank accession 355 numbers.SpeciesVoucherOriginITSReferencesCystolepiota ‌bucknallii ‌(Berk. & Broome) Singer & ClémençonSeSa12GermanyOL527684Sarawi et al. 2022Candelolepiota sinica Kun L. Yang, Jia Y. Lin & Zhu L. YangHKAS 52738ChinaKY039574Yang & Ge 2017Leucoagaricus ‌albosquamosus ‌Y.R. Ma, Z.W. Ge & T.Z. LiuCFSZ20662ChinaOM976879Ma et al. 2022Leucoagaricus ‌aurantioruber ‌Y.R. Ma, Z.W. Ge & T.Z. LiuCFSZ:19756, holotypeChinaNR182724Ma et al. 2022Leucoagaricus ‌badius ‌S. Hussain, Pfister, Afshan & KhalidLAH:SH148, holotypePakistanNR160453Hussain et al. 2018Leucoagaricus ‌brunneocanus ‌Fei Yu, J.F. Liang & Z.W. GeZ. L. Yang 3972ChinaKP096238Ge et al. 2015Leucoagaricus ‌bulbiger ‌Justo, Bizzi & AngeliniJBSD:130924, holotypeDominican RepublicNR173881Justo et al. 2021Leucoagaricus ‌caeruleovertens ‌Justo, Bizzi & AngeliniANGE 734, holotypeDominican RepublicMN483032Justo et al. 2021Leucoagaricus ‌candidus ‌Y.R. Ma, Z.W. Ge & T.Z. LiuCFSZ20964, holotypeChinaOM976877Ma et al. 2022Leucoagaricus cf. caerulescens ‌(Peck) J.F. Liang, Zhu L. Yang & J. XuMushroom Observer # 116307MexicoMH298921Rockefeller 12.V.2018 GenBank direct submissionLeucoagaricus cf. coerulescens ‌(Peck) J.F. Liang, Zhu L. Yang & J. XuMushroom Observer # 115977MexicoMF629828Rockefeller 16.VIII.2017 GenBank direct submissionLeucoagaricus ‌croceus ‌S.M. Tang & K.D. HydeMFLU:19-1331, holotypeThailandNR182408Manawasinghe et al. 2022Leucoagaricus ‌crystallifer ‌VellingaSFC 1010003-02SpainKY350216Caballero et al. 2009 24.XII.2016 GenBank direct submissionLeucoagaricus ‌griseodiscus ‌(Bon) Bon & Migl.MCVE:13719ItalyGQ329059Garbelotto et al. 2009 24.II.2010 GenBank direct submissionLeucoagaricus ‌lahorensiformis ‌S. Hussain, H. Ahmad, Afshan & KhalidSHL2, holotypePakistanKU647729Hussain et al. 2018Leucoagaricus ‌lahorensis ‌Qasim, Amir & NawazLAH 2481336, paratypePakistanKJ701795Qasim et al. 2015Leucoagaricus ‌menieri ‌(Sacc.) Singer6275NetherlandsKP300879Vellinga 20.V.2015 GenBank direct submissionLeucoagaricus ‌orientiflavus ‌Z.W. GeZ.W. Ge 2063, holotypeChinaNR119764Schoch et al. 2014Leucoagaricus ‌pabbiensis ‌Usman & KhalidLAH35302, holotypePakistanMG973423Usman & Khalid 2018Leucoagaricus ‌purpureolilacinus ‌HuijsmanE.C. Vellinga 2291 (L)NetherlandsAF482869Vellinga et al. 2003Leucoagaricus ‌rubrobrunneus ‌E.F. Malysheva, Svet. & E.M. BulakhLE262863, holotypeRussiaNR120166Schoch et al. 2014Leucoagaricus ‌rubroconfusus ‌Migl. & Coccia82002ItalyMN483035Justo et al. 2021Leucoagaricus ‌rubrotinctus ‌(Peck) Singer420526MF0068ChinaMG719611Wang et al. 2017 31.XII.2018 GenBank direct submissionLeucoagaricus ‌sardous ‌(Zecchin & Migl.) Cons. & ContuMCVE 20105, holotypeItalyKU041693Muñoz et al. 2012Leucoagaricus ‌serenus ‌(Fr.) Bon & BoiffardE.C. Vellinga 1930 (L)BelgiumAF482871Vellinga et al. 2003Leucoagaricus ‌subcrystallifer ‌Z. W. Ge & Zhu L. YangHKAS:49373, holotypeChinaNR155320Ge et al. 2015Leucoagaricus ‌subpurpureolilacinus ‌Z.W. Ge & Zhu L. YangHKAS:48285, holotypeChinaNR155318Ge et al. 2015Leucoagaricus ‌subvolvatus ‌(Malençon & Bertault) BonL1TürkiyeMN134030Dizkirici et al. 2019Leucoagaricus ‌sultanii ‌S. Hussain, H. Ahmad & KhalidLAH-SH115, holotypePakistanNR160452Hussain et al. 2018Leucoagaricus ‌umbonatus ‌S. Hussain, H. Ahmad & AfshanLAH:SHL1, holotypePakistanNR160454Hussain et al. 2018Leucoagaricus ‌viridiflavus ‌(Petch) T.K.A. Kumar & Manim.AK80aIndiaGU574745Liang et al. 2010Leucoagaricus ‌wychanskyi ‌(Pilát) Bon & BoiffardIX/X-1987NetherlandsAF482874Vellinga et al. 2003Leucocoprinus ‌adelphicus ‌(Vellinga) RedheadE.C. Vellinga 3153United StatesGQ258478Vellinga et al. 2010Leucocoprinus ‌atroazureus ‌(J.F. Liang, Zhu L. Yang & J. Xu) Asif, Saba & VellingaHKAS:42670, holotypeChinaNR182317Liang et al. 2010Leucocoprinus ‌beninensis ‌Sarawi ‌sp. nov.,SeSa113, holotypeBeninPX634060This studyLeucocoprinus ‌beninensis ‌Sarawi ‌sp. nov.,SeSa186BeninPX634061This studyLeucocoprinus ‌birnbaumii ‌(Corda) SingerPDD:117582New ZealandMW191767Johnston et al. 2020 04.XI.2020 GenBank direct submissionLeucocoprinus ‌cepistipes ‌(Sowerby) Pat.A18ChinaKC414250Shang et al. 2012 25.IX.2013 GenBank direct submissionLeucocoprinus ‌cinnamomeodiscus ‌Asif, Saba & M. RazaAsif-SP14AS11, holotypePakistanOQ947831Asif et al. 2024Leucocoprinus ‌cretaceus ‌(Bull.) Locq.JLF10042United StatesOM106913Frank 2022 09.I.2022 GenBank direct submissionLeucocoprinus ‌croceovelutinus ‌Bon & BoiffardE.C. Vellinga 3131 (UC)United StatesEU166351Vellinga & Sundberg 2008Leucocoprinus ‌erythrophaeus ‌(Vellinga) RedheadE.C. Vellinga 3248 (UC)United StatesGQ258470Vellinga et al. 2010Leucocoprinus ‌flammeotinctoides ‌(Vellinga) RedheadE.C. Vellinga 3247 (UC)United StatesGU136174Vellinga 2010 04.VI.2012Leucocoprinus ‌flavovirens ‌(J.F. Liang, Zhu L. Yang & J. Xu) Kun L. Yang, Jia Y. Lin & Zhu L. YangHKAS 50024, paratypeChinaEU416295Liang et al. 2010Leucocoprinus ‌gaillardii ‌(Bon & Boiffard) Asif, Saba & VellingaMCVE:736ItalyGQ329042Garbelotto et al. 2009 24.02.2010 GenBank direct submissionLeucocoprinus ‌gongylophorus ‌(Möller) HeimCylo005PanamaKF572000Kooij et al. 2015Leucocoprinus ‌irinellus ‌(Chalange) Kun L. Yang, Jia Y. Lin & Zhu L. YangChalange 97082101, isotypeFranceAY243648Vellinga 04.VI.2012 GenBank direct submissionLeucocoprinus ‌jubilaei ‌(Joss.) WasserJ. & A. Guinberteau 99101101FranceAY243635Vellinga 04.VI.2012 GenBank direct submissionLeucocoprinus ‌leucothites ‌(Vittad.) RedheadE.C. Vellinga 2050 (L)NetherlandsAF482865Vellinga et al. 2003Leucocoprinus ‌marriagei ‌(D.A. Reid) BonE.C. Vellinga 2005 (L)NetherlandsAF482866Vellinga et al. 2003Leucocoprinus ‌melanotrichus ‌(Malençon & Bertault) Migl. & DonatoE.C. Vellinga 2262 (L)NetherlandsAY176417Vellinga 2004Leucocoprinus ‌sabinae ‌(Angelini, Justo & Vizzini) Kun L. Yang, Jia Y. Lin & Zhu L. YangANGE305, paratypeDominican RepublicKM983666Justo et al. 2015Leucocoprinus ‌sericifer ‌(Locq.) Asif, Saba & VellingaE.C. Vellinga 2116 (L)NetherlandsAY176426Vellinga 2004Leucocoprinus ‌virens ‌(Y.R. Ma, Z.W. Ge & T.Z. Liu) Asif, Saba & VellingaCFSZ19794_2, paratypeChinaOM976880Ma et al. 2022Leucocoprinus ‌viriditinctus ‌(Berk. & Broome) Asif, Saba & VellingaHKAS 50033, paratypeChinaEU419375Liang et al. 2010Macropsalliota ‌americanus ‌(Peck) Kun L. Yang, Jia Y. Lin & Zhu L. YangE.C. Vellinga 2454 (UCB)United StatesAY176407Vellinga 2004Macropsalliota ‌meleagris ‌(Gray) Kun L. Yang, Jia Y. Lin & Zhu L. YangE.C. Vellinga 1990 (L)NetherlandsAY176419Vellinga 2004Micropsalliota ‌delicatula ‌R.L. Zhao, J. Xin Li & M.Q. HeHMAS:290752, holotypeChinaNR177529Li et al. 2021Micropsalliota ‌digitatocystis ‌R.L. Zhao, J. Xin Li & M.Q. HeHMAS:290750, holotypeChinaNR177530Li et al. 2021Micropsalliota ‌furfuracea ‌R.L. Zhao, Desjardin, K. Soytong & K.D. HydeHFJAU1570ChinaOM650275Yan et al. 2022Micropsalliota ‌globocystis ‌Heinem.HFJAU1518ChinaOM650277Yan et al. 2022Micropsalliota ‌megaspora ‌R.L. Zhao, Desjardin, K. Soytong & K.D. HydeHFJAU0712ChinaMN508425Yan et al. 2022Micropsalliota ‌wuyishanensis ‌J.Q. YanHFJAU3048, holotypeChinaOM650298Yan et al. 2022
Table 2. — Morphological differences matrix of Leucocoprinusbeninensis ‌Sarawi ‌sp. nov., and closely related species. — indicates missing information.‌Lc. beninensis Sarawi ‌sp. nov., (this study)Lc. cinnamomeodiscus ‌Asif, Saba & M. Raza (Asif et al. 2024)Lc. irinellus ‌(Chalange) Kun L. Yang, Jia Y. Lin & Zhu L. Yang (Chalange 1999)Lc. sericifer ‌(Locq.) Asif, Saba & Vellinga (Vellinga 2001)Lc. virens ‌(Y.R. Ma, Z.W. Ge & T.Z. Liu) Asif, Saba & Vellinga (Ma et al. 2022)Lc. viriditinctus ‌(Berk. & Broome) Asif, Saba & Vellinga (Liang et al. 2010)Lc. caerulescens ‌(Peck) J.F. Liang, Zhu L. Yang & J. Xu (Liang et al. 2010)La. cf. caerulescens ‌(Peck) J.F. Liang, Zhu L. Yang & J. Xu (MH298921)La. cf. coerulenscens ‌(Peck) J.F. Liang, Zhu L. Yang & J. Xu (MF629828)Pileus centre colourReddish brown, light brown, brownCinnamon-like or dull orangeOchraceous brown to brownPale ochraceous cream to yellowishBrown to grey-brownPale brown, reddish brown to dark brownBrownishReddish brown to brown–Basidiospore5.8-6.9 × 4.1-4.54.7-5.5 × 3.4-3.88.0-9.0 × 4.5-5.06.0-9.5 × 3.0-4.58.0-10.0 × 4.5-5.07.0-8.5 × 4.0-5.07.5-9.5 × 4.0-5.0–7.0-8.0 × 4.0-4.5Basidiospore propertiesAmygdaliform to ellipsoid, without germ pore, dextrinoid, non- metachromaticEllipsoid to oblong, without germ pore, dextrinoidEllipsoide to subovoid, metachromaticOblong amygdaliform to cylindrical- amygdaliform, without germ pore, dextrinoid, metachromaticAmygdaliform to ellipsoid, without germ pore, dextrinoid, metachromaticAmygdaliform to ovoid, without germ pore, dextrinoid, metachromaticAmygdaliform to ovoid, without germ pore, dextrinoid, metachromatic–Amygdaliform to ellipsoid, without germ pore, dextrinoid, metachromaticCheilocystidiaAbsentPresentAbsentPresentPresentPresentPresentPresentPresentColouring reactionsGreenish-blue to greyish-blue when dried–With ammonia first turning purple and than getting bluish-grey–Dark green to light green when bruised or driedDark blue when bruised or driedBluish green when dried–Vinaceous red then greenish when bruised and dark olive-grey when dried
Fig. 2. — Basidiomata and microstructures of Leucocoprinus ‌beninensis ‌Sarawi ‌sp. nov., (SeSa113, holotype): A, basidiomata (SeSa113, holotype); B, basidioma (SeSa186), seen from above (left) and from below the pileus (right); C, basidiospores; D, basidia; E, pileus covering. Scale bars: A, 1.5 cm; B, 0.5 cm; C, 3 µm; D, 10 µm; E, 20 µm. Photo credits: Cathrin Manz.10.5281/zenodo.18863073
Acknowledgements

This study was supported by the Federal Ministry of Education and Research (BMBF, Germany) as part of the projects “Diversity and Uses of Tropical African Fungi: Edible Mushrooms of Benin–FunTrAf” (01DG20015) and FunTrAf_2 (01DG24003). We are grateful to all members of the MyTIPS laboratory in Parakou, Benin, and the local people who generously assisted us during fieldwork. Additionally, we would like to thank Adrian Rühl, Cathrin Manz, and Felix Hampe for their great help during fieldwork, especially Cathrin Manz for taking pictures of the species described here. We also thank the referees and the editor-in-chief for their valuable work.

ReferencesAshraf S., Naseer A., Usman M. & Khalid A. N. 2023. — Two new species of genus Leucoagaricus (Agaricaceae, Agaricales) from Pakistan. MycoKeys 96: 159-171. https://doi.org/10.3897/mycokeys.96.101745Ashraf ‌S., Naseer ‌A., Usman ‌M. & Khalid ‌A. N.2023Two new species of genus Leucoagaricus (Agaricaceae, Agaricales) from PakistanMycoKeys96159171https://doi.org/10.3897/mycokeys.96.101745Asif M., Saba M., Raza M. & Vellinga E. C. 2024. — Molecular insights into fungal diversity reveal three novel species of Leucocoprinus from southern Punjab, Pakistan. Mycologia 116: 601-620. https://doi.org/10.1080/00275514.2024.2351769Asif ‌M., Saba ‌M., Raza ‌M. & Vellinga ‌E. C.2024Molecular insights into fungal diversity reveal three novel species of Leucocoprinus from southern Punjab, PakistanMycologia116601620https://doi.org/10.1080/00275514.2024.2351769Beeli M. 1932. — Fungi Goossensiani: IX genre Lepiota. Bulletin de la Société Royale de Botanique de Belgique 64: 206-222.Beeli ‌M.1932Fungi Goossensiani: IX genre LepiotaBulletin de la Société Royale de Botanique de Belgique64206222Beeli M. 1936. — Flore iconographique des Champignons du Congo: Fascicule 2, Lepiota & Annularia par M. Beeli (août 1936). Jardin botanique de l’État, Bruxelles: 29-48.Beeli ‌M.1936Flore iconographique des Champignons du Congo: Fascicule 2, Lepiota & Annularia par M. Beeli (août 1936)Jardin botanique de l’État, Bruxelles2948Candusso M. & Lanzoni G. 1990. — Lepiota s.l. Libreria editrice Giovanna Biella, Saronno, 743 p.Candusso ‌M. & Lanzoni ‌G.1990Lepiota sLibreria editrice Giovanna Biella, SaronnoChalange R. 1999. — Un nouveau Leucoagaricus thermophile à odeur d’iris, Leucoagaricus irinellus. Bulletin de la Société Mycologique de France 115: 385-392.Chalange ‌R.1999Un nouveau Leucoagaricus thermophile à odeur d’iris, Leucoagaricus irinellusBulletin de la Société Mycologique de France115385392Dennis R. W. G. 1952. — Lepiota and allied genera in Trinidad, British West Indies. Kew Bulletin 7: 459-499.Dennis ‌R. W. G.1952Lepiota and allied genera in Trinidad, British West IndiesKew Bulletin7459499Dizkirici A., Kalmer A. & Acar I. 2019. — Morphologic and molecular diagnosis of some Leucoagaricus species and revealing a new record from Turkey. Mantar dergisi 10: 143-150.Dizkirici ‌A., Kalmer ‌A. & Acar ‌I.2019Morphologic and molecular diagnosis of some Leucoagaricus species and revealing a new record from TurkeyMantar dergisi10143150Felsenstein J. 1985. — Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39: 783-791. https://doi.org/10.1111/j.1558-5646.1985.tb00420.xFelsenstein ‌J.1985Confidence limits on phylogenies: an approach using the bootstrapEvolution39783791https://doi.org/10.1111/j.1558-5646.1985.tb00420.xGardes M. & Bruns T. D. 1993. — ITS primers with enhanced specificity for basidiomycetes-application to the identification of mycorrhizae and rusts. Molecular ecology 2: 113-118. https://doi.org/10.1111/j.1365-294X.1993.tb00005.xGardes ‌M. & Bruns ‌T. D.1993ITS primers with enhanced specificity for basidiomycetes-application to the identification of mycorrhizae and rustsMolecular ecology2113118https://doi.org/10.1111/j.1365-294X.1993.tb00005.xGe Z.-W., Yang Z.-L., Qasim T., Nawaz R., Khalid A. N. & Vellinga E. C. 2015. — Four new species in Leucoagaricus (Agaricaceae, Basidiomycota) from Asia. Mycologia 107: 1033-1044. https://doi.org/10.3852/14-351Ge ‌Z.-W., Yang ‌Z.-L., Qasim ‌T., Nawaz ‌R., Khalid ‌A. N. & Vellinga ‌E. C.2015Four new species in Leucoagaricus (Agaricaceae, Basidiomycota) from AsiaMycologia10710331044https://doi.org/10.3852/14-351He M.-Q., Zhao R.-L., Hyde K. D., Begerow D., Kemler M., Yurkov A., McKenzie E. H. C., Raspé O., Kakishima M., Sánchez-Ramírez S., [...] & Kirk P. M. 2019. — Notes, outline and divergence times of Basidiomycota. Fungal Diversity 99: 105-367. https://doi.org/10.1007/s13225-019-00435-4He ‌M.-Q., Zhao ‌R.-L., Hyde ‌K. D., Begerow ‌D., Kemler ‌M., Yurkov ‌A., McKenzie ‌E. H. C., Raspé ‌O., Kakishima ‌M., Sánchez-Ramírez ‌S., [...] & Kirk ‌P. M.2019Notes, outline and divergence times of BasidiomycotaFungal Diversity99105367https://doi.org/10.1007/s13225-019-00435-4Heinemann P. 1973. — Flore illustrée des Champignons d’Afrique centrale: Fascicule 2,Leucocoprineae p.p. (Agaricaceae) par P. Heinemann (août 1973),Cystoderma(Tricholomataceae) par P. Heinemann & D. Thoen (août 1973). Jardin botanique de l’État, Bruxelles: 29-48.Heinemann ‌P.1973Flore illustrée des Champignons d’Afrique centrale: Fascicule 2,Leucocoprineae p.p. (Agaricaceae) par P. Heinemann (août 1973),Cystoderma(Tricholomataceae) par P. Heinemann & D. Thoen (août 1973)Jardin botanique de l’État, Bruxelles2948Heinemann P. 1977. — Flore illustrée des Champignons d’Afrique centrale: Fascicule 5, Leucocoprinus(Agaricaceae) par P. Heinemann (septembre 1977), Asproinocybe(Tricholomataceae) par P. Heinemann & D. Thoen (septembre 1977). Jardin botanique de l’État, Bruxelles: 87-101.Heinemann ‌P.1977Flore illustrée des Champignons d’Afrique centrale: Fascicule 5, Leucocoprinus(Agaricaceae) par P. Heinemann (septembre 1977), Asproinocybe(Tricholomataceae) par P. Heinemann & D. Thoen (septembre 1977)Jardin botanique de l’État, Bruxelles87101Huelsenbeck J. P. & Ronquist F. 2001. — MrBayes: Bayesian inference of phylogenetic trees. Bioinformatics 17: 754-755. https://doi.org/10.1093/bioinformatics/17.8.754Huelsenbeck ‌J. P. & Ronquist ‌F.2001MrBayes: Bayesian inference of phylogenetic treesBioinformatics17754755https://doi.org/10.1093/bioinformatics/17.8.754Hussain S., Jabeen S., Khalid A. N., Ahmad H., Afshan N. U. S., Sher H. & Pfister D. H. 2018. — Underexplored regions of Pakistan yield five new species of Leucoagaricus. Mycologia 110: 387-400. https://doi.org/10.1080/00275514.2018.1439651Hussain ‌S., Jabeen ‌S., Khalid ‌A. N., Ahmad ‌H., Afshan ‌N. U. S., Sher ‌H. & Pfister ‌D. H.2018Underexplored regions of Pakistan yield five new species of LeucoagaricusMycologia110387400https://doi.org/10.1080/00275514.2018.1439651Jabeen S., Waseem B., Hamid M., Yasmeen A. & Yasmeen A. 2020. — First record of Leucoagaricusnivalis from Pakistan. Bangladesh Journal of Plant Taxonomy 27: 453-459. https://doi.org/10.3329/bjpt.v27i2.50684Jabeen ‌S., Waseem ‌B., Hamid ‌M., Yasmeen ‌A. & Yasmeen ‌A.2020First record of Leucoagaricusnivalis from PakistanBangladesh Journal of Plant Taxonomy27453459https://doi.org/10.3329/bjpt.v27i2.50684Johnson J. 1999. — Phylogenetic relationships within Lepiotasensu lato based on morphological and molecular data. Mycologia 91: 443-458. https://doi.org/10.1080/00275514.1999.12061038Johnson ‌J.1999Phylogenetic relationships within Lepiotasensu latobased on morphological and molecular dataMycologia91443458https://doi.org/10.1080/00275514.1999.12061038Johnson J. & Vilgalys R. 1998. — Phylogenetic systematics of Lepiotasensu lato based on nuclear large subunit rDNA evidence. Mycologia 90: 971-979. https://doi.org/10.1080/00275514.1998.12026994Johnson ‌J. & Vilgalys ‌R.1998Phylogenetic systematics of Lepiotasensu latobased on nuclear large subunit rDNA evidenceMycologia90971979https://doi.org/10.1080/00275514.1998.12026994Justo A., Angelini C., Bizzi A. & Vizzini A. 2015. — Leucoagaricussabinae (Agaricaceae), a new species from the Dominican Republic. North American Fungi 10: 1-15. https://doi:10.2509/naf2015.010.005Justo ‌A., Angelini ‌C., Bizzi ‌A. & Vizzini ‌A.2015Leucoagaricussabinae (Agaricaceae), a new species from the Dominican RepublicNorth American Fungi10115https://doi:10.2509/naf2015.010.005Justo A., Angelini C. & Bizzi A. 2021. — The genera Leucoagaricus and Leucocoprinus in the Dominican Republic. Mycologia 113: 348-389. https://doi.org/10.1080/00275514.2020.1819142Justo ‌A., Angelini ‌C. & Bizzi ‌A.2021The genera Leucoagaricus and Leucocoprinus in the Dominican RepublicMycologia113348389https://doi.org/10.1080/00275514.2020.1819142Katoh K. & Standley D. M. 2013. — MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular biology and evolution 30: 772-780. https://doi.org/10.1093/molbev/mst010Katoh ‌K. & Standley ‌D. M.2013MAFFT multiple sequence alignment software version 7: improvements in performance and usabilityMolecular biology and evolution30772780https://doi.org/10.1093/molbev/mst010Katoh K. & Toh H. 2008. — Recent developments in the MAFFT multiple sequence alignment program. Briefings in bioinformatics 9: 286-298. https://doi.org/10.1093/bib/bbn013Katoh ‌K. & Toh ‌H.2008Recent developments in the MAFFT multiple sequence alignment programBriefings in bioinformatics9286298https://doi.org/10.1093/bib/bbn013Kooij P. W., Aanen D. K., Schiøtt M. & Boomsma J. J. 2015. — Evolutionarily advanced ant farmers rear polyploid fungal crops. Journal of Evolutionary Biology 28:1911-1924. https://doi.org/10.1111/jeb.12718Kooij ‌P. W., Aanen ‌D. K., Schiøtt ‌M. & Boomsma ‌J. J.2015Evolutionarily advanced ant farmers rear polyploid fungal cropsJournal of Evolutionary Biology2819111924https://doi.org/10.1111/jeb.12718Kornerup A. & Wanscher J. H. 1978. — Methuen handbook of colour. 3rd edition. Eyre Methuen, London, 254 p.Kornerup ‌A. & Wanscher ‌J. H.1978Methuen handbook of colour. 3rd editionEyre Methuen, LondonKumar S., Stecher G. & Tamura K. 2016. — MEGA7: Molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular biology and evolution 33: 1870-1874. https://doi.org/10.1093/molbev/msw054Kumar ‌S., Stecher ‌G. & Tamura ‌K.2016MEGA7: Molecular evolutionary genetics analysis version 7.0 for bigger datasetsMolecular biology and evolution3318701874https://doi.org/10.1093/molbev/msw054Li J.-X., He M.-Q. & Zhao R.-L. 2021. — Three new species of Micropsalliota (Agaricaceae, Agaricales) from China. Phytotaxa 491: 167-176. https://doi.org/10.11646/phytotaxa.491.2.6Li ‌J.-X., He ‌M.-Q. & Zhao ‌R.-L.2021Three new species of Micropsalliota (Agaricaceae, Agaricales) from ChinaPhytotaxa491167176https://doi.org/10.11646/phytotaxa.491.2.6Liang J. F., Yang Z.-L., Xu J. & Ge Z.-W. 2010. — Two new unusual Leucoagaricus species (Agaricaceae) from tropical China with blue-green staining reactions. Mycologia 102: 1141-1152. https://doi.org/10.3852/09-021Liang ‌J. F., Yang ‌Z.-L., Xu ‌J. & Ge ‌Z.-W.2010Two new unusual Leucoagaricus species (Agaricaceae) from tropical China with blue-green staining reactionsMycologia10211411152https://doi.org/10.3852/09-021Ma Y., Liu T., Yu X., Wei T. & Ge Z. W. 2022. — Six new species of Leucoagaricus (Agaricaceae) from Northeastern China. Diversity 14: 314. https://doi.org/10.3390/d14050314Ma ‌Y., Liu ‌T., Yu ‌X., Wei ‌T. & Ge ‌Z. W.2022Six new species of Leucoagaricus (Agaricaceae) from Northeastern ChinaDiversity14314https://doi.org/10.3390/d14050314Manawasinghe I. S., Calabon M. S., Jones E. B. G., Zhang Y. X., Liao C. F., Xiong Y. R., Chaiwan N., Kularathnage N. D., Liu N. G., Tang S. M., [...] & Hyde K. D. 2022. — Mycosphere notes 345-386. Mycosphere 13: 454-557. https://doi.org/10.5943/mycosphere/13/1/3Manawasinghe ‌I. S., Calabon ‌M. S., Jones ‌E. B. G., Zhang ‌Y. X., Liao ‌C. F., Xiong ‌Y. R., Chaiwan ‌N., Kularathnage ‌N. D., Liu ‌N. G., Tang ‌S. M., [...] & Hyde ‌K. D.2022Mycosphere notes 345Mycosphere13454557https://doi.org/10.5943/mycosphere/13/1/3Miller M. A., Pfeiffer W. & Schwartz T. 2010. — Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Gateway Computing Environments Workshop (GCE) 14.11.2010: 1-8. https://doi.org/10.1109/GCE.2010.5676129Miller ‌M. A., Pfeiffer ‌W. & Schwartz ‌T.2010Creating the CIPRES Science Gateway for inference of large phylogenetic treesGateway Computing Environments Workshop (GCE) 14.11.201018https://doi.org/10.1109/GCE.2010.5676129Muñoz G., Caballero A., Contu M. & Vizzini A. 2012. — A new Leucoagaricus species of section Piloselli (Agaricales, Agaricaceae) from Spain. IMA Fungus 3: 117-123. https://doi.org/10.5598/imafungus.2012.03.02.03Muñoz ‌G., Caballero ‌A., Contu ‌M. & Vizzini ‌A.2012A new Leucoagaricus species of section Piloselli (Agaricales, Agaricaceae) from SpainIMA Fungus3117123https://doi.org/10.5598/imafungus.2012.03.02.03Pegler D. N. 1977. — A preliminary agaric flora of East Africa, in Pegler D. N. (ed.), Kew Bulletin Additional Series VI. Her Majesty’s Stationery Office, London, 615 p.Pegler ‌D. N.1977A preliminary agaric flora of East AfricainPegler ‌D. N.Kew Bulletin AdditionalHer Majesty’s Stationery Office, LondonPegler D. N. 1983. — Agaric flora of the lesser Antilles, in Pegler D. N. (ed.), Kew Bulletin Additional Series IX. Her Majesty’s Stationery Office, London, 235 p.Pegler ‌D. N.1983Agaric flora of the lesser AntillesinPegler ‌D. N.Kew Bulletin AdditionalHer Majesty’s Stationery Office, LondonPiepenbring M., Maciá-Vicente J. G., Codjia J. E. I., Glatthorn C., Kirk P., Meswaet Y., Minter D., Olou B. A., Reschke K., Schmidt M. & Yorou N. S. 2020. — Mapping mycological ignorance-checklists and diversity patterns of fungi known for West Africa. IMA fungus 11: 13. https://doi.org/10.1186/s43008-020-00034-yPiepenbring ‌M., Maciá-Vicente ‌J. G., Codjia ‌J. E. I., Glatthorn ‌C., Kirk ‌P., Meswaet ‌Y., Minter ‌D., Olou ‌B. A., Reschke ‌K., Schmidt ‌M. & Yorou ‌N. S.2020Mapping mycological ignorance-checklists and diversity patterns of fungi known for West AfricaIMA fungus1113https://doi.org/10.1186/s43008-020-00034-yQasim T., Amir T., Nawaz R., Niazi A. R. & Khalid A. N. 2015. — Leucoagaricuslahorensis, a new species of L. sect. Rubrotincti. Mycotaxon 130: 533-541. https://doi.org/10.5248/130.533Qasim ‌T., Amir ‌T., Nawaz ‌R., Niazi ‌A. R. & Khalid ‌A. N.2015Leucoagaricuslahorensis, a new species of L. sect. RubrotinctiMycotaxon130533541https://doi.org/10.5248/130.533Redhead S. A. 2023. — Nomenclatural novelties. Index Fungorum 551: 1.Redhead ‌S. A.2023Nomenclatural noveltiesIndex Fungorum5511Rehman A., Usman M., Afshan N. U. S. & Khalid A. N. 2023. — Leucoagaricusgujratensis sp. nov. (Agaricaceae, Agaricales) from Pakistan. Phytotaxa 589: 39-50. https://doi.org/10.11646/phytotaxa.589.1.4Rehman ‌A., Usman ‌M., Afshan ‌N. U. S. & Khalid ‌A. N.2023Leucoagaricusgujratensis sp. nov. (Agaricaceae, Agaricales) from PakistanPhytotaxa5893950https://doi.org/10.11646/phytotaxa.589.1.4Ronquist F., Teslenko M, van der Mark P., Ayres D. L., Darling A., Höhna S., Larget B., Liu L., Suchard M. A. & Huelsenbeck J. P. 2012. — MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic biology 61: 539-542. https://doi.org/10.1093/sysbio/sys029Ronquist ‌F., Teslenko ‌M, van der Mark ‌P., Ayres ‌D. L., Darling ‌A., Höhna ‌S., Larget ‌B., Liu ‌L., Suchard ‌M. A. & Huelsenbeck ‌J. P.2012MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model spaceSystematic biology61539542https://doi.org/10.1093/sysbio/sys029Sarawi S., Shi Y.-N., Lotz-Winter H., Reschke K., Bode H. B. & Piepenbring M. 2022. — Occurrence and chemotaxonomical analysis of amatoxins in Lepiota spp. (Agaricales). Phytochemistry 195: 113069. https://doi.org/10.1016/j.phytochem.2021.113069Sarawi ‌S., Shi ‌Y.-N., Lotz-Winter ‌H., Reschke ‌K., Bode ‌H. B. & Piepenbring ‌M.2022Occurrence and chemotaxonomical analysis of amatoxins in Lepiota spp. (Agaricales)Phytochemistry195113069https://doi.org/10.1016/j.phytochem.2021.113069Schoch C. L., Robbertse B., Robert V., Vu D., Cardinali G., Irinyi L., Meyer W., Nilsson R. H., Hughes K., Miller A. N., [...] & Federhen S. 2014. — Finding needles in haystacks: linking scientific names, reference specimens and molecular data for fungi. Database 2014: bau061. https://doi.org/10.1093/database/bau061Schoch ‌C. L., Robbertse ‌B., Robert ‌V., Vu ‌D., Cardinali ‌G., Irinyi ‌L., Meyer ‌W., Nilsson ‌R. H., Hughes ‌K., Miller ‌A. N., [...] & Federhen ‌S.2014Finding needles in haystacks: linking scientific names, reference specimens and molecular data for fungiDatabase2014https://doi.org/10.1093/database/bau061Singer R. 1986. — The Agaricales in modern taxonomy. 4th ed. Koeltz Scientific Books, Oberreifenberg, 833 p.Singer ‌R.1986The Agaricales in modern taxonomy. 4th edKoeltz Scientific Books, OberreifenbergStamatakis A. 2014. — RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30: 1312-1313. https://doi.org/10.1093/bioinformatics/btu033Stamatakis ‌A.2014RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogeniesBioinformatics3013121313https://doi.org/10.1093/bioinformatics/btu033Stöver B. C. & Müller K. F. 2010. — TreeGraph 2: combining and visualizing evidence from different phylogenetic analyses. BMC bioinformatics 11: 7. https://doi.org/10.1186/1471-2105-11-7Stöver ‌B. C. & Müller ‌K. F.2010TreeGraph 2: combining and visualizing evidence from different phylogenetic analysesBMC bioinformatics117https://doi.org/10.1186/1471-2105-11-7Sysouphanthong P. & Thongklang N. 2022. — Two new species of Leucoagaricus (Agaricaceae) from the Lao People’s Democratic Republic. Current Research in Environmental & Applied Mycology 12: 65-74. https://doi.org/10.5943/cream/12/1/6Sysouphanthong ‌P. & Thongklang ‌N.2022Two new species of Leucoagaricus (Agaricaceae) from the Lao People’s Democratic RepublicCurrent Research in Environmental & Applied Mycology126574https://doi.org/10.5943/cream/12/1/6Usman M. & Khalid A. N. 2018. — Leucoagaricuspabbiensis sp. nov. from Punjab, Pakistan. Pakistan. Mycotaxon 133: 354- 363. https://doi.org/10.5248/133.355Usman ‌M. & Khalid ‌A. N.2018Leucoagaricuspabbiensis sp. nov. from PunjabPakistan. Pakistan. Mycotaxon133354363https://doi.org/10.5248/133.355Vellinga E. C. 2001. — Leucoagaricus; Leucocoprinus, in Noordeloos M. E., Kuyper T. W. & Vellinga E. C. (eds), Flora Agaricina Neerlandica 5. Balkema Publishers, Rotterdam: 76-108.Vellinga ‌E. C.2001Leucoagaricus; LeucocoprinusinNoordeloos ‌M. E., Kuyper ‌T. W. & Vellinga ‌E. C.Flora Agaricina Neerlandica 5Balkema Publishers, Rotterdam76108Vellinga E. C. 2004. — Genera in the family Agaricaceae: evidence from nrITS and nrLSU sequences. Mycological Research 108: 354-377. https://doi.org/10.1017/S0953756204009700Vellinga ‌E. C.2004Genera in the family Agaricaceae: evidence from nrITS and nrLSU sequencesMycological Research108354377https://doi.org/10.1017/S0953756204009700Vellinga E. C. & Sundberg W. 2008. — Lepiotaceous fungi in California, U.S.A. 6. Lepiota castanescens. Mycotaxon 103: 97-108.Vellinga ‌E. C. & Sundberg ‌W.2008Lepiotaceous fungi in California, U.S.A. 6Lepiota castanescens. Mycotaxon 10397108Vellinga E. C., de Kok R. P. J. & Bruns T. D. 2003. — Phylogeny and taxonomy of Macrolepiota (Agaricaceae). Mycologia 95: 442-456. https://doi.org/10.1080/15572536.2004.11833089eVellinga ‌E. C., de Kok ‌R. P. J. & Bruns ‌T. D.2003Phylogeny and taxonomy of Macrolepiota (Agaricaceae)Mycologia95442456https://doi.org/10.1080/15572536.2004.11833089eVellinga E. C., Contu M. & Vizzini A. 2010. — Leucoagaricus decipiens and La. erythrophaeus, a new species pair in sect. Piloselli. Mycologia 102: 447-454. https://doi.org/10.3852/09-164Vellinga ‌E. C., Contu ‌M. & Vizzini ‌A.2010Leucoagaricus decipiens and La. erythrophaeus, a new species pair in sect. PiloselliMycologia102447454https://doi.org/10.3852/09-164Vellinga E. C., Sysouphanthong P. & Hyde K. D. 2011. — The family Agaricaceae: phylogeneis and two new white-spored genera. Mycologia 103: 494-509. https://doi.org/10.3852/10-204Vellinga ‌E. C., Sysouphanthong ‌P. & Hyde ‌K. D.2011The family Agaricaceae: phylogeneis and two new white-spored generaMycologia103494509https://doi.org/10.3852/10-204White T. J., Bruns T. D. & Taylor L. J. 1990. — Amplification direct sequencing of fungal ribosomal RNA genes for phylogenetics, in Innis M. A., Gelfand D. H., Sninsky J. J. & White T. J. (eds), PCR protocols: a guide to methods applications. Academic Press, New York: 315-322. https://doi.org/10.1016/b978-0-12-372180-8.50042-1White ‌T. J., Bruns ‌T. D. & Taylor ‌L. J.1990Amplification direct sequencing of fungal ribosomal RNA genes for phylogeneticsinInnis ‌M. A., Gelfand ‌D. H., Sninsky ‌J. J. & White ‌T. J.PCR protocols: a guide to methods applicationsAcademic Press, New York315322https://doi.org/10.1016/b978-0-12-372180-8.50042-1Yan J.-Q., Zeng Z.-H., Hu Y.-P., Ke B.-R., Zeng H. & Wang S.-N. 2022. — Taxonomy and multi-gene phylogeny of Micropsalliota (Agaricales, Agaricaceae) with description of six new species from China. Frontiers in Microbiology 13: 1011794. https://doi.org/10.3389/fmicb.2022.1011794Yan ‌J.-Q., Zeng ‌Z.-H., Hu ‌Y.-P., Ke ‌B.-R., Zeng ‌H. & Wang ‌S.-N.2022Taxonomy and multi-gene phylogeny of Micropsalliota (Agaricales, Agaricaceae) with description of six new species from ChinaFrontiers in Microbiology131011794https://doi.org/10.3389/fmicb.2022.1011794Yang Z.-L. & Ge Z.-W. 2017. — Six new combinations of lepiotaceous fungi from China. Mycosystema 36: 542-551. https://doi.org/10.13346/j.mycosystema.160221Yang ‌Z.-L. & Ge ‌Z.-W.2017Six new combinations of lepiotaceous fungi from ChinaMycosystema36542551https://doi.org/10.13346/j.mycosystema.160221Yang K. L., Lin J. Y., Li G.-M., Li T. & Yang Z.-L. 2024. — Rediscovering Leucoagaricus sinicus, with the recognition of Leucoagaricus and Leucocoprinus as separate genera, and two new genera in Agaricaceae (Basidiomycota). Phytotaxa 676: 199-255. https://doi.org/10.11646/phytotaxa.676.3.1Yang ‌K. L., Lin ‌J. Y., Li ‌G.-M., Li ‌T. & Yang ‌Z.-L.2024Rediscovering Leucoagaricus sinicus, with the recognition of Leucoagaricus and Leucocoprinus as separate genera, and two new genera in Agaricaceae (Basidiomycota)Phytotaxa676199255https://doi.org/10.11646/phytotaxa.676.3.1