High rate of gene family evolution in proximity to the origin of ectomycorrhizal symbiosis in Inocybaceae.

The genomes of ectomycorrhizal (ECM) fungi have a reduced number of genes encoding Carbohydrate-Active EnZymes (CAZymes), expansions in transposable elements (TEs) and small secreted proteins (SSPs) compared with saprotrophs. Fewer genes for specific peptidases and lipases in ECM fungi are also reported. It is unclear whether these changes occur at the shift to the ECM habit or are more gradual throughout the evolution of ECM lineages. We generated a genomic dataset of 20 species in the ECM lineage Inocybaceae and compared them with six saprotrophic species. Inocybaceae genomes have fewer CAZymes, peptidases, lipases, secondary metabolite clusters and SSPs and higher TE content than their saprotrophic relatives. There was an increase in the rate of gene family evolution along the branch with the transition to the ECM lifestyle. This branch had very high rate of evolution in CAZymes and had the largest number of contractions. Other significant changes along this branch included expansions in transporters, transposons-related genes and communication genes such as fungal kinases. There is a high concentration of changes in proximity to the transition to the ECM lifestyle, which correspond to the identified key changes for the gain of this lifestyle.

More smooth-spored species of Inocybe (Agaricales, Ba sidiomycota): type studies and 12 new species from Europe.

Twelve new species of Inocybe (I. adorabilis, I. comis, I. demetris, I. filiana, I. galactica, I. morganae, I. othini, I. ovilla, I. proteica, I. somae, I. suryana and I. venerabilis) are described from Europe on the basis of detailed morphological and molecular investigation. A portrait of the recently described I. ianthinopes is given. All species are smooth-spored and some pruinose only in the apical part of the stipe, and some on entire length. The new species are compared to 24 type specimens (17 characterized by at least partial ITS sequence data), all of which are described and revised here. Epitypes were selected for two species, I. hirtella and I. sindonia. Based on our studies, we confirm that I. kuehneri and I. sindonia on one hand, and I. subalbidodisca and I. ochroalba on the other, are synonyms and furthermore suggest that I. abietis is synonymous with I. catalaunica, I. exilis with I. rufobrunnea, I. hirtellarum with I. mycenoides, I. lapidicola with I. deianae, I. ochraceolutea with I. sindonia, I. stangliana with I. pelargonium, I. subrubens with I. subhirtella and I. sulfovirescens with I. langei. All of the new species are supported by phylogenetic analyses. Among the 16 previously described species accepted here, 10 are represented by types in the phylogenetic analyses and five by own collections corresponding to the type. Two species, I. eutheloides (remaining doubtful) and I. pallidolutea are only treated morphologically. In summary, we describe as new or verify the taxonomic status and provide or corroborate morphological concepts for 37 smooth-spored species of Inocybe. Citation: Bandini D, Oertel B, Eberhardt U. 2022. More smooth-spored species of Inocybe (Agaricales, Basidiomycota): type studies and 12 new species from Europe. Persoonia 48: 91-149. https://doi.org/10.3767/persoonia.2022.48.03.

Mallocybe heimii ectomycorrhizae with Cistus creticus and Pinus halepensis in Mediterranean littoral sand dunes - assessment of phylogenetic relationships to M. arenaria and M. agardhii.

Ectomycorrhizal symbiosis appears extensively in the Northern Hemisphere, where Mediterranean ecosystems constitute an important ecological area of considerable biodiversity value. Littoral sand dunes are among high-risk habitats, and ectomycorrhizal lifestyle contributes significantly to supporting life in such regions. Mallocybe heimii (Bon) Matheny & Esteve-Rav. (Inocybaceae, Basidiomycota) and the very similar M. arenaria (Bon) Matheny & Esteve-Rav. grow in poor, usually sandy soils, in association with angiosperms or gymnosperms. Basidiomata originally identified under these names were collected from littoral sand dunes of Greece, and their morpho-anatomical characteristics were examined in conjunction with material derived from other European regions. Sequences from basidiomata and root tips corresponding to the nuclear rDNA internal transcribed spacer region (ITS) and large subunit (LSU) were obtained and analyzed. Phylogenetic results demonstrated that material identified as M. heimii or M. arenaria form a single well-supported group, while M. agardhii (N. Lund) Matheny & Esteve-Rav. is confirmed to be distinct from M. arenaria (the latter was initially described as a variety of the former, i.e., I. agardhii var. arenaria Bon). A detailed tree of the genus Mallocybe was generated on the basis of concatenated ITS and LSU sequences, and relationships of selected taxa are discussed in the light of morphological and sequence data. In addition, the first morphotype descriptions of M. heimii ectomycorrhizae with Cistus creticus L. and Pinus halepensis Miller are hereby provided. Both morphotypes exhibited the typical characteristics of Inocybe/Mallocybe ectomycorrhizae; however, differences were noted, the most significant being the presence of clamps on mantle hyphae and the type of anastomoses.

Five new species of Inocybe (Agaricales) from tropical India.

Five new species of Inocybe, I. iringolkavensis, I. keralensis, I. kuruvensis, I. muthangensis and I. wayanadensis, are described from Kerala state, India, based on morphological and molecular data. All are associated with trees belonging to Dipterocarpaceae. Inocybe iringolkavensis is characterized by nodulose to somewhat stellate basidiospores, 1-4-spored basidia, and caulocystidia restricted to the stipe apex. Inocybe keralensis has a yellowish brown pileus, lamellae with whitish, serrate edges, smooth, ellipsoidal basidiospores and a duplex pileipellis with the superficial hyphae devoid of encrustations and encrusted hyphae beneath. The diagnostic features of I. kuruvensis include a dark brown pileus, stipe with a whitish base and grayish brown, floccose-fibrillose surface, nodulose basidiospores with saddle-shaped projections and faintly encrusted paracystidia with refractive contents. Violet basidiomata with a rimose, hygrophanous pileus, densely pruinose stipe with a marginate-bulbous base, and nodulose basidiospores are the major features of I. muthangensis. Inocybe wayanadensis is characterized by small, whitish basidiomata, a viscid pileus with a rimulose surface, a densely pruinose and fibrillose stipe with a marginate-bulbous base, nodulose basidiospores, thick-walled pleuro- cheilo- and caulocystidia and an ixotrichoderm-type pileipellis. The phylogenetic relationships of these new species are inferred from an analysis of nuc rDNA sequences of the internal transcribed spacers (ITS1-5.8S-ITS2) and the 28S gene. Except I. keralensis, which belongs to the Pseudosperma clade, all other species belong to the Inocybe clade. This study represents the second report of an Inocybe species (I. muthangensis) that combines violet basidiomata with nodulose basidiospores.

Unraveling the Inocybe praetervisa group through type studies and ITS data: Inocybe praetervisoides sp. nov. from the Mediterranean region.

Species in the Inocybe praetervisa group are characterized by producing nodulose to angular basidiospores and a bulbous, marginate, white stipe devoid of any pinkish to reddish tinge. Species delimitation problems and common misinterpretations in the I. praetervisa group have not yet been resolved through type studies and analysis of molecular data. This study seeks to clarify the taxonomy and nomenclature of species around I. praetervisa. Analyses of the nuc rDNA internal transcribed regions (ITS) recovered two major groups within the I. praetervisa group that can be separated on the basis of cystidial morphology. The study of three authentic and topotypic specimens in the Bresadola herbarium revealed that the name I. praetervisa has been misapplied often. The ITS region of one of the specimens was obtained, and this specimen is designated as epitype in support of a lectotype. Inocybe rivularis is demonstrated to be a later synonym of I. praetervisa, while Inocybe phaeocystidiosa is the correct name for the species most often misdetermined as I. praetervisa. Inocybe salicis-herbaceae and I. praetervisa var. flavofulvida are shown to be synonyms of I. phaeocystidiosa based on ITS sequence data from type collections. A new species sister to I. phaeocystidiosa with a Mediterranean distribution is described as I. praetervisoides. Cystidial morphology, distribution of caulocystidia, basidiospore morphology and ecology are shown to be the main diagnostic characters for separating the species. Inocybe praetervisa and I. phaeocystidiosa have a transoceanic distribution in Europe and North America, whereas I. praetervisoides so far is known only from the Mediterranean region.

Two new species of Inocybe from Mediterranean Cistaceae heathlands.

This study explored a heathland region in Portugal, and through morphology, biogeography, and multilocus phylogeny, two new species of Inocybaceae are described. The first species, Inocybe iberilepora, belongs to "I. flocculosa group," whereas the second species, Inocybe phaeosquamosa, belongs to a relatively isolated and understudied clade, distantly related to I. furfurea and allies. Both species are tied to a west Mediterranean distribution and ecology, associating with the local Cistaceae ecosystems. By characterizing these new species, our research contributes to the understanding of European Funga and enriches the knowledge of the genus Inocybe on a global scale.

Inocybe section Rimosae in Utah: phylogenetic affinities and new species.

Results of a study on species of Inocybe section Rimosae sensu lato in Utah are presented. Eight species, seven from the Pseudosperma clade (section Rimosae sensu stricto) and one from the Inosperma clade (section Rimosae pro parte), are documented morphologically and phylogenetically. Five of the eight species, I. aestiva, I. breviterincarnata, I. cercocarpi, I. niveivelata and I. occidentalis-all members of the Pseudosperma clade-are described as new from Utah and other western states. Two European species, I. spuria and I. obsoleta, are confirmed from Utah. Inocybe aurora, originally described from Nova Scotia, is synonymized with I. obsoleta. The only member of the Inosperma clade recorded from Utah is I. lanatodisca, a widely distributed species for which three geographical clusters were detected. The phylogenetic analyses indicate that the Pseudosperma clade includes 53 clusters or species worldwide and that the Inosperma clade includes 47 such clusters. Many of these probably correspond to undescribed species. A key to species of section Rimosae sensu lato from Utah is provided together with illustrations of the eight species found in the state.

Re-Valuation of the Taxonomic Status of Species within the Inocybe similis Complex.

The taxonomy of Inocybe similis and closely allied species is addressed using morphological and molecular data (nrITS and nrLSU DNA). The holotypes of I. chondrospora and I. vulpinella and the isotype of I. immigrans were studied and sequenced. Our results suggest the synonymy between I. similis and I. vulpinella as well as that between I. chondrospora and I. immigrans.

New Species of Mallocybe and Pseudosperma from North China.

Within the family Inocybaceae, many species of Mallocybe and Pseudosperma have been reported, but there are only a few reports on these two genera from north China. In this study, six collections of Mallocybe and 11 collections of Pseudosperma were studied by morphological and phylogenetic methods. Phylogenetic analyses based on sequence data from three or two different loci (ITS, LSU, and rpb2 for Mallocybe; ITS and LSU for Pseudosperma) are performed to infer species relationships within genera Mallocybe and Pseudosperma, respectively. Results indicate that eight species of Mallocybe and Pseudosperma are found in Shanxi province, north China; two new species of Mallocybe, M. depressa and M. picea, are described. Overall, six species belong to Pseudosperma, of which three are new: P. gilvum, P. laricis and P. pseudoniveivelatum.

Five new species of Inosperma from China: Morphological characteristics, phylogenetic analyses, and toxin detection.

Many species of Inosperma cause neurotoxic poisoning in humans after consumption around the world. However, the toxic species of Inosperma and its toxin content remain unclear. In the present study, we proposed five new Inosperma species from China, namely, I. longisporum, I. nivalellum, I. sphaerobulbosum, I. squamulosobrunneum, and I. squamulosohinnuleum. Morphological and molecular phylogenetic analyses based on three genes (ITS, nrLSU, rpb2) revealed that these taxa are independent species. A key to 17 species of Inosperma in China is provided. In addition, targeted screening for the most notorious mushroom neurotoxins, muscarine, psilocybin, ibotenic acid, and muscimol, in these five new species was performed by using ultrahigh-performance liquid chromatography-tandem mass spectrometry (UPLC-MS/MS). Our results show that the neurotoxin contents in these five species varied: I. sphaerobulbosum contains none of the tested neurotoxins; I. nivalellum is muscarine positive; I. longisporum and I. squamulosohinnuleum contain both ibotenic acid and muscimol, and I. squamulosobrunneum only contains muscimol; psilocybin was not detected in these five new species.

Four new nodulose-spored species of Inocybe (Agaricales) from West Africa.

We describe four new nodulose-spored species of Inocybe from tropical regions of Africa: I. beninensis, I. flavipes, I. fuscobrunnea and I. pallidiangulata. The new species are recognised based on morphological data and phylogenetic analyses of ITS, 28S and RPB2 sequences. Phylogenetic analyses indicated that I. flavipes and I. beninensis are part of a subclade leading to the I. calida group. Inocybe fuscobrunnea appears sister to the I. asterospora group. Inocybe pallidiangulata is nested within a clade of mainly tropical species from South Asia, Africa, and South America, close to the subclade of I. lilacinosquamosa and I. ayangannae from Guyana. Complete descriptions and illustrations, including photographs and line drawings, and a key to nodulose-spored taxa of tropical African species of Inocybe are provided. Citation: Aïgnon HL, Jabeen S, Verbeken A, Matheny PB, Yorou NS, Ryberg M (2022). Four new nodulose-spored species of Inocybe (Agaricales) from West Africa. Fungal Systematics and Evolution 10: 1-18. doi: 10.3114/fuse.2022.10.01.

Two new toxic yellow Inocybe species from China: morphological characteristics, phylogenetic analyses and toxin detection.

Some species of Inocybe s. str. caused neurotoxic poisoning after consumption around the world. However, there are a large number of species in this genus that have not been studied for their toxicity or toxin content. In this study, we report two new toxic yellow Inocybe s. str. species from China based on morphological characteristics, phylogenetic analyses and toxin detection. Among the two species, Inocybesquarrosolutea is reported as a newly recorded species of China. We also describe a new species, I.squarrosofulva, which is morphologically similar to I.squarrosolutea. The new species is characterized by its ochraceous squarrose pileus, distinctly annulate cortina on the stipe, nodulose basidiospores and thick-walled pleurocystidia. Muscarine in the fruitbodies was detected by UPLC-MS/MS, the content in I.squarrosolutea and I.squarrosofulva were 136.4 ± 25.4 to 1683.0 ± 313 mg/kg dry weight and 31.2 ± 5.8 to 101.8 ± 18.9 mg/kg dry weight, respectively.

Four new species of sequestrate Inocybe from Chilean Nothofagaceae forests.

Sequestrate fungi have enclosed hypogeous, subhypogeous, or epigeous basidiomes and have lost the ability to actively discharge their spores. They can be distinguished as gasteroid (basidiome fully enclosed with a loculated hymenophore) or secotioid (basidiome with some agaricoid or pileate-stipitate features, but the lamellae are misshapen and unexposed or mostly unexposed at maturity). There are only four reports of sequestrate taxa within the ectomycorrhizal family Inocybaceae, three from Australia and one from western North America. Recent field work in Nothofagaceae forests in the Chilean coastal range revealed novel sequestrate forms of Inocybe. We examined specimens using a combination of morphological and molecular data from nuc rDNA internal transcribed spacer region ITS1-5.8S-ITS2 (ITS) and portions of nuc 28S rDNA (28S) and the gene encoding the second largest subunit of RNA polymerase II (rpb2). Here, we describe four new sequestrate Inocybe species, I. ranunculiformis, I. anfractuosa, I. illariae, and I. nahuelbutensis. Results of our phylogenetic analysis resolved the four new species as distinct species-level clades with strong support, suggesting that these fungi have convergently evolved sequestrate forms independently. The species described here were all placed along with members of the "smooth-spored temperate austral clade," which includes almost exclusively Australasian and South American species of Inocybe.

Inocybe brijunica sp. nov., a New Ectomycorrhizal Fungus from Mediterranean Croatia Revealed by Morphology and Multilocus Phylogenetic Analysis.

A new ectomycorrhizal species was discovered during the first survey of fungal diversity at Brijuni National Park (Croatia), which consists of 14 islands and islets. The National Park is located in the Mediterranean Biogeographical Region, a prominent climate change hot-spot. Inocybe brijunica sp. nov., from sect. Hysterices (Agaricales, Inocybaceae), is described based on morphology and multilocus phylogenetic data. The holotype collection was found at the edge between grassland and Quercus ilex forest with a few planted Pinus pinea trees, on Veli Brijun Island, the largest island of the archipelago. It is easily recognized by a conspicuous orange to orange-red-brown membranaceous surface layer located at or just above the basal part of the stipe. Other distinctive features of I. brijunica are the medium brown, radially fibrillose to rimose pileus; pale to medium brown stipe with fugacious cortina; relatively small, amygdaliform to phaseoliform, and smooth basidiospores, measuring ca. 6.5-9 × 4-5.5 µm; thick-walled, utriform, lageniform or fusiform pleurocystidia (lamprocystidia) with crystals and mostly not yellowing in alkaline solutions; cheilocystidia of two types (lamprocystidia and leptocystidia); and the presence of abundant caulocystidia only in the upper 2-3 mm of the stipe. Phylogenetic reconstruction of a concatenated dataset of the internal transcribed spacer region (ITS), the nuclear 28S rRNA gene (nrLSU), and the second largest subunit of RNA polymerase II (rpb2) resolved I. brijunica and I. glabripes as sister species.

Nodulose-spored Inocybe from the Rocky Mountain alpine zone molecularly linked to European and type specimens.

Inocybe (Inocybaceae) is one of the most diverse ectomycorrhizal genera in arctic and alpine habitats where the primary hosts are Salix, Betula, and Dryas. Subgenus Inocybe is common in these habitats and typically characterized by the presence of thick-walled pleurocystidia. Here, we focus on species that have angular or nodulose spores. Historically, over 30 taxa from this group have been reported from arctic and alpine habitats. Many names have been synonymized, whereas molecular analysis has revealed new species. Nuc rDNA internal transcribed spacer ITS1-5.8S-ITS2 (ITS) sequence data of 26 type specimens in this group now allow for further taxonomic clarification and comparison across continents of disjunct populations. Here, we compare ITS sequence data and the D1-D2 portion of nuc 28S rDNA (28S) from Rocky Mountain specimens with those of types and European reference material. We report 10 species from the Rocky Mountain alpine zone, all of which are conspecific with known European boreal, montane, or alpine species, and four are described as new; all have intercontinental distributions. Nodulose-spored Inocybe taxa that occur in the Rocky Mountain alpine zone include I. alpinomarginata, sp. nov., I. arctica, I. giacomi, I. leonina, I. murina, sp. nov., I. occulta, I. paragiacomi, sp. nov., I. phaeocystidiosa, I. purpureobadia, and I. subgiacomi, sp. nov. Remarkably, these species occur at elevations up to 4000 m and at latitudes as low as 36°N, hundreds of miles from the Arctic, the European alpine, and original type localities. Distributions are explained in part by host distributions and historical glaciation patterns. A key and full descriptions for Rocky mountain species are provided to promote species recognition.

The Inocybe geophylla group in North America: a revision of the lilac species surrounding I. lilacina.

The Inocybe geophylla group is circumscribed based on phylogenetic analysis of DNA sequences largely sampled from North America and Europe. Twenty-nine phylogenetic species are uncovered after analysis of combined nuc 28S rDNA (28S) and RNA polymerase II second largest subunit (rpb2) DNA sequence data. Species in the I. geophylla group share the presence of a cortina, silky-fibrillose pileus and stipe, pruinose stipe apex, spermatic odor, thick-walled hymenial cystidia, and smooth amygdaliform or elliptical basidiospores. Within the group, as many as five phylogenetic species attributable to I. lilacina and allies form a strongly supported clade based on analysis of nuc ITS1-5.8S-ITS2 rDNA (ITS [internal transcribed spacer]), 28S, and rpb2 data. However, all lilac-colored species do not form a monophyletic group. Sufficient morphological and ecological data are present to document four of the I. lilacina subgroup species, two of which are described from North America as new: I. ionocephala and I. sublilacina. Inocybe lilacina is recircumscribed based on sequencing the holotype and is distributed in the eastern United States under pines and/or hardwoods. Inocybe pallidicremea is a widespread and common conifer associate in mostly northern parts of North America, to which the name I. lilacina was previously applied. Descriptions, photographs, line drawings, and a taxonomic key to lilac species in the I. lilacina subgroup from North America are provided. Well-documented collections, especially notes on gross morphology and ecology, are needed to continue to assess and describe the high taxonomic variation in the I. lilacina subgroup and its allies worldwide.