Recognition of Tylopilus alboater once relied heavily on the morphological features of the basidiomata, which resulted in considerable confusion due to morphological stasis and plasticity. In this study, we examined specimens morphologically identified as T. alboater from Asia, North America, and Australia using phylogenomics and multi-locus sequence data. To clarify its phylogenetic placement within Boletaceae, we conducted a phylogenomic analysis based on 45 genomes (including three newly sequenced T. alboater genomes) comprising representatives from the eight subfamilies of Boletaceae. Additionally, we constructed a concatenated dataset (nrLSU + tef1-α + rpb1 + rpb2), incorporating representative species from all genera within the subfamily to which T. alboater belongs, to infer its phylogeny. Our phylogenomic analysis revealed that specimens morphologically identified as T. alboater exhibit polyphyly, clustering entirely within the subfamily Boletoideae. Our multi-locus phylogenetic analyses further indicated that these specimens represent eight distinct species distributed across five generic lineages within the subfamily Boletoideae, including one new genus, Neoporphyrellus, proposed in this study, as well as one new species, N. sinoalboater, and two new combinations, N. alboater and N. atronicotianus. The remaining five species are nested in four known genera, including two new species of Abtylopilus, Ab. indonesiensis and Ab. australiensis, and three known species belonging to Anthracoporus, Indoporus, and Tylopilus, respectively. These findings highlight the taxonomic complexity of the T. alboater complex and emphasize the importance of integrating phylogenomic and multi-locus approaches for accurate fungal systematics. Color photos of fresh basidiomata, line drawings of microscopic features, and detailed descriptions of the new taxa are presented.
A multigene phylogenetic assessment of North American species of Mallocybe is presented based on analyses of rpb1 , rpb2 , ITS, and 28S rDNA nucleotide data. This framework enables a systematic revision of the genus for 16 eastern North American species and captures taxonomic and phylogenetic diversity in a global context. A grade of two unusual and poorly known North American species stems from the most recent common ancestor of the genus that gives rise to three core subgroups named here as clades Unicolores, Nothosperma, and Mallocybe. The grade of taxa includes the poorly known Lepista praevillosa from Florida and a new species from the southern Appalachians, M. montana , both of which appear to be narrow-range endemics. Clade Nothosperma is characterized by Australian and New Zealand species, whereas clade Unicolores is composed of six species from eastern North America and East Asia. Clade Mallocybe is dominated by numerous north temperate taxa and constitutes the sister group to clade Nothosperma. These major clades are distinguished by a combination of phylogeny, morphology, geographic distribution, and ecology. In addition, four North American species are described as new: M. leucothrix, M. luteobasis, M. montana , and M. tomentella . Several names originating in North America, long ignored or misunderstood in the literature, are revitalized and established by type comparisons and modern reference material collected from or near type localities. In addition, 11 species were subjected to mass spectrometry muscarine assays, none of which contained detectable amounts of muscarine except for two: M. sabulosa and M. praevillosa . This confirms a diffuse phylogenetic distribution of muscarine within the genus. Taxonomic descriptions are presented for 16 species, several synonymies proposed, and four new combinations made. A key to species of eastern North American Mallocybe is presented, along with illustrations of important diagnostic features.
Molecular characterization of type specimens is a powerful tool used in clarifying species identity/circumscription, as well as establishing the taxonomic and phylogenetic status of organisms in question. However, DNA sequencing of aged herbarium collections can be a challenge due to the quantity and quality of DNA still present in the specimens. Herein, we report a custom DNA isolation protocol suitable for processing minute quantities of old specimen tissue and its utilization via high-throughput sequencing technologies to obtain, for the first time, the genome assembly of the 134-year-old holotype of Boletus subvelutipes Peck, a North American fleshy pored mushroom of taxonomic and historical significance. A side-by-side evaluation of our DNA isolation method with that of a commercial "kit" by Qiagen is also presented. By relying on the type material, we have established the genetic identity of B. subvelutipes, as well as providing preliminary phylogenetic evidence for its generic affinities in Neoboletus within Boletaceae. The reference genome of the B. subvelutipes holotype provides a resource for future comparative genomic studies, taxonomic revisions in Boletaceae, and other evolutionary studies of fungi.
The Global Consortium for the Classification of Fungi and fungus-like taxa is an international initiative of more than 550 mycologists to develop an electronic structure for the classification of these organisms. The members of the Consortium originate from 55 countries/regions worldwide, from a wide range of disciplines, and include senior, mid-career and early-career mycologists and plant pathologists. The Consortium will publish a biannual update of the Outline of Fungi and fungus-like taxa, to act as an international scheme for other scientists. Notes on all newly published taxa at or above the level of species will be prepared and published online on the Outline of Fungi website (https://www.outlineoffungi.org/), and these will be finally published in the biannual edition of the Outline of Fungi and fungus-like taxa. Comments on recent important taxonomic opinions on controversial topics will be included in the biannual outline. For example, 'to promote a more stable taxonomy in Fusarium given the divergences over its generic delimitation', or 'are there too many genera in the Boletales?' and even more importantly, 'what should be done with the tremendously diverse 'dark fungal taxa?' There are undeniable differences in mycologists' perceptions and opinions regarding species classification as well as the establishment of new species. Given the pluralistic nature of fungal taxonomy and its implications for species concepts and the nature of species, this consortium aims to provide a platform to better refine and stabilise fungal classification, taking into consideration views from different parties. In the future, a confidential voting system will be set up to gauge the opinions of all mycologists in the Consortium on important topics. The results of such surveys will be presented to the International Commission on the Taxonomy of Fungi (ICTF) and the Nomenclature Committee for Fungi (NCF) with opinions and percentages of votes for and against. Criticisms based on scientific evidence with regards to nomenclature, classifications, and taxonomic concepts will be welcomed, and any recommendations on specific taxonomic issues will also be encouraged; however, we will encourage professionally and ethically responsible criticisms of others' work. This biannual ongoing project will provide an outlet for advances in various topics of fungal classification, nomenclature, and taxonomic concepts and lead to a community-agreed classification scheme for the fungi and fungus-like taxa. Interested parties should contact the lead author if they would like to be involved in future outlines.
SUMMARY Investigating the mechanisms that underpin the diversity and distribution patterns of species is fundamental in ecology and evolution. However, the study of fungi, particularly the ectomycorrhizal group, has been relatively constrained in this field. We conducted a high-resolution phylogenomic analysis of Boletales, an ecologically and economically significant group of fungi, covering 83 genera across 15 families. We subsequently investigated its evolutionary history using sequences at four genes obtained from 984 species across 121 genera within 15 families. The findings unveiled that Boletales likely originated in Early Jurassic and underwent two remarkable episodes of rapid diversification, commencing in early Eocene (∼54 Mya) and early Miocene (∼17 Mya) epochs. The notable surges were predominantly driven by ectomycorrhizal clades, with a specific emphasis on East Asia and North America. These expansions were strongly correlated with the warm-humid paleoclimates during the Early Eocene Climatic Optimum and Mid-Miocene Climatic Optimum, as well as the rapid expansion of Fagales and Pinaceae hosts. This study provides novel insights into the spatiotemporal evolution of fungi, highlighting the synergistic impact of abiotic factors, such as warm and humid paleoclimates, and the biotic factor of rapid diversification of host plants on the fungal diversification.
Through a long-term sampling of the ectomycorrhizal fungi in remnant fragments of both lowland oak forest and montane cloud forest, in central Veracruz, Mexico, a new species, Tylopilus tropicalis of the Tylopilus balloui group, was discovered, along with a new record of T. oradivensis originally described from Costa Rica. Macro- and micromorphological features, a phylogenetic analysis of a concatenated ITS, 28S and RPB1 dataset, as well as ecological distribution, support the hypotheses. The new species proposed here grows under Quercus sapotifolia, and T. oradivensis appears near Q. xalapensis in the tropical oak forest and in the montane cloud forest, respectively. Both species treated here are described, illustrated, and discussed. An artificial key is provided to distinguish both species from other closely related species.
A systematic revision of North American species with morphological similarities to Inocybe grammata was conducted based on materials collected in eastern and western North America and Central America. Eight taxa are confirmed in the group proposed here as I. sect. Albodiscae sect. nov., species of which are often characterized by a bicolorous pileus with a distinct pallid disc, the margin covered with superficial silky fibrils, entirely pruinose stipe with a slight or obvious marginate bulbous base, angular to angular-nodulose basidiospores with comparatively few nodules, and thick-walled cystidia. Species in the section are mainly distinguished by basidiome size, pileus color and texture, variation in basidiospore length and number of nodules, to some extent geographic location, and plant association. The section includes the widely distributed I. grammata (= I. albodisca), which associates with conifers and birch in eastern North America; I. albodiscoides sp. nov. from the Pacific Northwest, previously subsumed under I. grammata and I. albodisca; I. floridana from northern Florida, reported for the first time in more than 75 years; I. acriolens, an apparent associate of hemlock and pine in southeast Canada and the northeast U.S.; I. grammatoides, an aspen associate in northern regions of the U.S. and southeast Canada; I. velicopia sp. nov., a widely distributed associate of oak and chestnut in the eastern U.S. and Costa Rica; I. panamica sp. nov. from oak and oak-Oreomunnea (Juglandaceae) forests in Costa Rica and Panama; and I. vestalis, a European species sister to the rest of sect. Albodiscae but lacking the distinct bicolored pileus. Independent phylogenies of ITS+28S, rpb2, and rpb1 are inferred. Taxonomic descriptions, illustrations, and/or notes of the North American taxa are provided.
Sutorius is a poroid genus in Boletaceae that typically has chocolate brown to reddish brown or purplish brown basidiomata with a finely scaly stipe and produces a reddish brown spore deposit. During the survey on diversity of boletes in Northern and Northeastern Thailand, several Sutorius collections were obtained. Combined evidence from morphology and phylogenetic analyses of a combined three-gene data set (atp6, tef1 and rpb2) of the Sutorius collections along with selected Boletaceae in the Pulveroboletus group indicated that Thai collections represent seven new Sutorius species. The analyses also indicated that Tylopilus maculatoides belongs in Sutorius. Therefore, the transfer of T. maculatoides to Sutorius is proposed. Full descriptions and illustrations of the seven new species and S. maculatoides are presented in this study. With the seven new species and the new combination, eight of the eleven described Sutorius species are known to occur in Northern and Northeastern Thailand, whereas only one species is known from each of two continents, the Americas and Australia.
Species of Tricholoma sections Genuina and Megatricholoma are characterized by having pilei that are some shade of brown, and by white or yellow lamellae that also discolor some shade of brown. From Costa Rica in sect. Genuina, we describe as new T. luteopallidum, T. cacumense, and T. talamancense, and confirm the occurrence of Tricholoma stans, and from the United States describe as new Tricholoma brunneoluteum. Tricholoma roseoacerbum from sect. Megatricholoma is con-firmed for Costa Rica. Morphology as well as ITS sequences are employed to confirm the species’ identifications or circumscription of the new species.
The systematic position of the enigmatically mycoparasitic genus Squamanita ( Agaricales , Basidiomycota ) together with Cystoderma , Phaeolepiota , Floccularia , and Leucopholiota is largely unknown. Recently they were recognized as Squamanitaceae , but previous studies used few DNA markers from a restricted sample of taxa from the family and lacked a formal taxonomic treatment. In this study, with newly generated sequences of the type of the genus Squamanita , S. schreieri , and several additional species of the family, the phylogeny is reinvestigated with a concatenated (18S-5.8S-nrLSU-RPB2-TEF1 - α) dataset. This study reveals that Cystoderma , Phaeolepiota , Squamanita , Floccularia , and Leucopholiota are a monophyletic clade with strong statistical support in Bayesian analysis and form Squamanitaceae . Phaeolepiota nested within Cystoderma ; Squamanita , Leucopholiota , and Floccularia clustered together as two monophyletic subclades; and Squamanita was present as a monophyletic clade with strong statistical support in both Maximum Likelihood and Bayesian analyses. The family name Squamanitaceae is formally emended and a detailed taxonomic treatment is presented to accommodate the five genera. Meanwhile, another concatenated (18S-ITS-nrLSU-RPB2-TEF1 - α) dataset is used to investigate phylogenetic relationships and species delimitation in Squamanita . Our data indicates that “ S. umbonata ” from the Northern hemisphere forms two species complexes, one complex includes six specimens from North America, Europe, and East Asia, the other includes two specimens from Central America and North America respectively. Futhermore, species of Squamanita can parasitize species of Amanita , besides other fungal species. Squamanita mira parasitizes A . kitamagotake ( A. sect. Caesareae ), while S. orientalis and S. sororcula are parasites of species belonging to the A . sepiacea complex ( A. sect. Validae ). “ Squamanita umbonata ” from Italy occurs on A. excelsa ( A. sect. Validae ). Three new species of Squamanita from East Asia, viz. S. mira , S. orientalis and S. sororcula are documented with morphological, multi-gene phylogenetic, and ecological data, along with line drawings and photographs, and compared with similar species. A key for identification of the global Squamanita species is provided.
Three new species of Gyroporus, one from the southern hemisphere (Gyroporus madagascariensis sp. nov.) and two from the northern hemisphere (Gyroporus borealis sp. nov. and Gyroporus smithii sp. nov.), are described. G. madagascariensis is a brownish bolete currently known from Madagascar; it has a mottled pileus somewhat reminiscent of G. mcnabbii, another species known from the southern hemisphere. G. borealis is known from the northeastern United States and is also recorded from the northwestern United States under planted landscape trees. G. smithii corresponds to an orange-colored species that has often been encountered in the United States west and south of the Appalachian Mountains (east of the Great Plains) and is likely often mistaken for G. borealis, which can display orange coloration. Additionally, this study presents evidence that G. roseialbus Murrill and G. subalbellus Murrill, previously synonymized by Rolf Singer, are in fact distinct species.
Lactifluusrussulisporus Dierickx & De Crop and Lactifluuscaliendrifer Froyen & De Crop are described from eucalypt forests in Queensland, Australia and different forest types in Thailand, respectively. Both species have recently been published on Index Fungorum and fit morphologically and molecularly in L.sect.Luteoli, a section within L.subg.Gymnocarpi that encompasses species with alboochraceous basidiomes, white latex that stains brown and typical capitate elements in the pileipellis and/or marginal cells.
Erythrophylloporus is a lamellate genus in the family Boletaceae that has been recently described from China based on E.cinnabarinus , the only known species. Typical characters of Erythrophylloporus are reddish-orange to yellowish-red basidiomata, including lamellae, bright yellow basal mycelium and smooth, broadly ellipsoid, ellipsoid to nearly ovoid basidiospores. During our survey on diversity of Boletaceae in Thailand, several yellowish-orange to reddish- or brownish-orange lamellate boletes were collected. Based on both morphological evidence and molecular analyses of a four-gene dataset ( atp 6, tef 1, rpb 2 and cox 3), they were recognised as belonging in Erythrophylloporus and different from the already known species. Two new species, E.paucicarpus and E.suthepensis are therefore introduced from Thailand with detailed descriptions and illustrations. Moreover, two previously described Phylloporus species, P.aurantiacus and P.fagicola , were also revised and recombined in Erythrophylloporus . A key to all known Erythrophylloporus species is provided.
The Basidiomycota constitutes a major phylum of the kingdom Fungi and is second in species numbers to the Ascomycota. The present work provides an overview of all validly published, currently used basidiomycete genera to date in a single document. An outline of all genera of Basidiomycota is provided, which includes 1928 currently used genera names, with 1263 synonyms, which are distributed in 241 families, 68 orders, 18 classes and four subphyla. We provide brief notes for each accepted genus including information on classification, number of accepted species, type species, life mode, habitat, distribution, and sequence information. Furthermore, three phylogenetic analyses with combined LSU, SSU, 5.8s, rpb1, rpb2, and ef1 datasets for the subphyla Agaricomycotina, Pucciniomycotina and Ustilaginomycotina are conducted, respectively. Divergence time estimates are provided to the family level with 632 species from 62 orders, 168 families and 605 genera. Our study indicates that the divergence times of the subphyla in Basidiomycota are 406–430 Mya, classes are 211–383 Mya, and orders are 99–323 Mya, which are largely consistent with previous studies. In this study, all phylogenetically supported families were dated, with the families of Agaricomycotina diverging from 27–178 Mya, Pucciniomycotina from 85–222 Mya, and Ustilaginomycotina from 79–177 Mya. Divergence times as additional criterion in ranking provide additional evidence to resolve taxonomic problems in the Basidiomycota taxonomic system, and also provide a better understanding of their phylogeny and evolution.
The ectomycorrhizal genus Lactifluus is known to contain many species complexes, consisting of morphologically very similar species, which can be considered cryptic or pseudocryptic. In this paper, a thorough molecular study is performed of the clade around Lactifluus deceptivus (originally described by Peck from North America) or the deceptive milkcaps. Even though most collections were identified as L. deceptivus , the clade is shown to contain at least 15 species, distributed across Asia and America, indicating that the L. deceptivus clade represents a species complex. These species are morphologically very similar and are characterized by a tomentose pileus with thin-walled hyphae and a velvety stipe with thick-walled hyphae. An ITS1 sequence was obtained through Illumina sequencing for the lectotype of L. deceptivus , dating from 1885, revealing which clade represents the true L. deceptivus . In addition, it is shown that three other described species also belong to the L. deceptivus clade: L. arcuatus , L. caeruleitinctus and L. mordax, and molecularly confirmed that L. tomentoso-marginatus represents a synonym of L. deceptivus . Furthermore, two new Neotropical species are described: Lactifluus hallingii and L. domingensis .
Boletus longipes was described in 1909 by George Massee from a specimen collected in Singapore. The species was not mentioned again until E.J.H. Corner's book in 1972 on Malaysian boletes. Corner had collected it many times during his tenure in Singapore, and he synonymized Boletus tristis with B. longipes described nineyears after B. longipes by Patouillard and Baker from the same site. Among the distinguishing characters of B. longipes were deep vinous red spore deposit, red oxidation reaction of the hymenophore when bruised, and spores that displayed a strong violet color reaction in contact with KOH. C.B. Wolfe ultimately moved both species to a new genus Austroboletus. During recent efforts to circumscribe Austroboletus in Australasia using morphological and molecular phylogenetic inferences, it became clear that B. longipes was neither in harmony with Boletus, Porphyrellus, nor Austroboletus. Rather, it is a new genus, which we describe here as Ionosporus, allied to Borofutus, Spongiforma, and Rhodactina of subfamily Leccinoideae. In addition, recent collections from Queensland and New South Wales, Australia, that are morphologically similar to I. longipes, are inferred to be a separate new species, I. australis.
Three new species of Tricholoma are described from Costa Rican montane forests with additional collections cited from the United States. The new species are Tricholoma felschii sp. nov., Tricholoma costaricense sp. nov. and Tricholoma atratum sp. nov. A discussion of Tricholoma luteomaculosum is also included. These taxa share morphological features of strong farinaceous odor and taste, pseudoparenchymatous pileal subcutis, and rugulose pileus at some stage of development. In addition, cheilocystidia occur in two of the taxa but not the third. Phylogenies are presented based on ITS sequences.