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Revision of the taxonomically complicated European Melanoleuca taxa with well-developed macrocystidia (subgenus Melanoleuca) is presented. The species are delimited by macro- and micromorphological characters and by multilocus molecular data based on internal transcribed spacer (ITS), second largest subunit of RNA polymerase I (rpb2), and translation elongation factor 1-alpha (tef1) genes. The study is based on both type material and recent collections. Phylogenetically, these taxa form 10 clades recognized as 11 separate species: M. albomarginata, M. ammophila, M. bataillei, M. cavipes, M. friesii, M. granadensis, M. melaleuca, M. pallidicutis, M. polioleuca, M. strictipes, and M. variabilis. Three species, M. albomarginata, M. ammophila, and M. variabilis, are described as new. Melanoleuca melaleuca is discussed here as the type species of the genus. Discussion of other older names and their taxonomic status is also provided.

Melanoleuca is one of the taxonomically most complicated genera of Agaricomycetes. The aim of this study is taxonomic revision of European specimens confused with M. angelesiana A.H. Sm. and delimitation of four species, M. brachyspora Harmaja, M. graminicola (Velen.) Kühner & Maire, M. rufipes Bon and M. stridula (Fr.) Singer. The research is based on macro- and micromorphological characters and the multigene phylogenetic analyses of a combined (ITS, rpb2, and tef1) dataset. According to the result, the two new species are proposed – M. monticola sp. nov. which accomodates European specimens previously confused with American species M. angelesiana and M. romanensis sp. nov. from Italy. Moreover, American specimens identified as M. angelesiana belongs to three different phylogenetic species (M. angelesiana sensu typi, M. acystidiata aff. and one more separate lineage). M. graminicola is emended and the respective epitype designated. An identification key of the European species of subg. Urticocystis is provided.

We present the first phylogenetic evaluation of the genus Dermoloma, which is resolved as monophyletic and closely related to Pseudotricholoma, a poorly known Dermoloma-like lineage within the family Tricholomataceae. The position of Dermoloma is confirmed by the placement of the type species, Dermoloma cuneifolium, represented by multiple samples including the neotype. Based on our phylogenetic analyses, we recognised 25 European operational taxonomic units (OTUs) but could only assign species names to ten of them based on ex-type sequences. Furthermore, only five additional published Dermoloma names of uncertain status are available for the remaining 16 potential European species, thus demonstrating an unexpected amount of taxonomic diversity. Samples from Europe and North America seem to be endemic on a continental scale. North American samples formed six unique OTUs, but only one could be reliably named, Dermoloma hymenocephalum. Dermoloma is morphologically defined by basidiomata with brown, grey and white colours with a farinaceous odour and a pluristratous hymeniderm type of pileipellis. Our phylogenetic analyses support the subdivision of the genus into two subgenera and four sections, and species with inamyloid basidiospores are placed in subg. Dermoloma and those with amyloid basidiospores in subg. Amylospora. Both subgenera are further divided into two sections. The analysis of spore morphology shows that sect. Conica of subg. Dermoloma and sect. Nigrescentia of subg. Amylospora have a very distinctive spore shape. Sect. Atrobrunnea of subg. Amylospora showed relatively high variability of spores among species, but spores of sect. Dermoloma were similar and not useful for species discrimination.

This study evaluates the morphological concept of Crepidotus variabilis and similar taxa based on basidiospore and cheilocystidia characters. A phylogenetic analysis of the ITS region clearly distinguished C. variabilis var. variabilis from the two other distinct taxa of the same lineage, C. variabilis var. trichocystis and C. neotrichocystis. We also recognised one additional morphologically similar and undescribed species that belongs to the C. variabilis lineage. All four taxa can also be recognised morphologically using basidiospore and cheilocystidia characters. Our detailed statistical comparison of these characters improves the knowledge of their variability and specifies more precisely the morphological circumscription of the taxa within this group.

Supplementary material for Phylogeny of Crepidotus applanatus look-alikes reveals a con-vergent morphology evolution and a new species C. pini. Table S1: Details on the specimens (arranged alphabetically and within Slovakia from west to east) examined in this study. Specimens marked with asterisk symbol have been sequenced; Table S2: The list of ITS and LSU nrDNA sequences used in the phylogenetic analysis. Taxon names (in alphabetical order) are presented in their original form (C. = Crepidotus). Sequences prepared in this study are in bold; Table S3: Estimated ranges of spore dimensions based on 30 measurements represented as average ± standard deviation, in parenthesis are minimum and maximum values, the average is underlined, Q = the length/width ratio. Specimens marked with asterisk have been sequenced; Table S4: Estimated ranges of cheilocystidia dimensions based on 30 measurements represented as average ± standard deviation, the average is underlined. Morphological groups of cheilocystidia: cla – clavate, cyl – cylindrical, utr – utriform, ven – ventricose, for – forked, lob – lobate, pyr – pyriform, glo – globose (undifferentiated here whether it is a broadly or narrowly type of the cheilocystidia). Specimens marked with asterisk have been sequenced.

Revision of the taxonomically complicated European

The study deals with the last unexplored morphological group of the genus Hodophilus defined by absence of distinct odours, absence of yellow colours and absence of darker dots on the stipe. The phylogenetic reconstruction of the whole genus based on nrITS, nrLSU and RPB2 sequences placed all European members having these morphological characters in a monophyletic group defined here as a new section H. sect. Phaeophylli. The remaining European members of the genus are placed in two additional groups classified as section H. sect. Hodophilus and the new section H. sect. Micacei. Five species are recognised within section Phaeophylli which is typified by H. phaeophyllus that is lecto- and epitypified. Three new species belonging to this section are described: H. carpathicus, H. decurrentior and H. stramineus. Hodophilus decurrentior is the only species showing distinct morphological differences under the microscope. The identification of other species of the section depends mainly on the colour of basidiomata. An updated key to all European members of the genus is provided.

Since 2007, the quality of Russula descriptions has improved and the use of molecular support for species delimitation and the number of published new species has increased. However, the description style is not consistent and has regional or author-specific patterns. Most recent publications still favour descriptions of spores compared to hymenium and pileipellis elements, and usually only the spore size is provided with statistical support. This study proposes standards for descriptions of the microscopic structure of Russula species (Russulaceae, Agaricomycetes). We present the description template, the template measurements table, the specific terminology and the essential chemical reagents. The proposed standards were tested by mycologists from 11 countries who met at the Russula Microscopy Workshop in Slovakia. Descriptions of 26 species from 9 countries and four continents were prepared, among them R. amarissima, R. castanopsidis, R. seperina and R. subtilis are re-described and 15 species are introduced as new: R. abietiphila, R. amerorecondita, R. aurantioflava, R. echidna, R. flavobrunnescens, R. fluvialis, R. fortunae, R. garyensis, R. gemmata, R. laevis, R. madrensis, R. olivaceohimalayensis, R. purpureogracilis, R. sancti-pauli and R. wielangtae. Seven descriptions for candidate new species are provided without a formal name assignment. Pairwise comparison of species described in this study with available similar descriptions of related species suggests that microscopic characters from all parts of the basidiomata can be equally important for species recognition and they deserve the same treatment including number of measurements and statistics. The majority of recent studies does not recognise differences between the pileus margin and centre, but more than one-third of the species described in this study show distinct differences between the pileus areas, emphasizing the importance to specify the origin of pileipellis observations. This study proved that there is frequently insufficient difference in the ITS barcode between closely related species and that it is necessary to use more genetic markers combined with ecological and geographical data. © 2019, School of Science.

We present the first phylogenetic evaluation of the genus Dermoloma , which is resolved as monophyletic and closely related to Pseudotricholoma , a poorly-known Dermoloma -like lineage within the family Tricholomataceae. The position of Dermoloma is confirmed by the placement of the type species, D. cuneifolium , represented by multiple samples including the neotype. Based on our phylogenetic analyses, we recognised 25 European operational taxonomic units (OTUs), but could only assign species names to ten of them based on ex-type sequences. Furthermore, only five additional published Dermoloma names of uncertain status are available for the remaining 16 potential European species, thus demonstrating an unexpected amount of taxonomic diversity. Samples from Europe and North America seem to be endemic on a continental scale. North American samples formed six unique OTUs, but only one could be reliably named, D. hymenocephalum . Dermoloma is morphologically defined by basidiomata with brown, grey and white colours with a farinaceous odour and a pluristratous hymeniderm type of pileipellis. Our phylogenetic analyses support the subdivision of the genus into two subgenera and four sections, species with inamyloid basidiospores are placed in subg. Dermoloma and those with amyloid basidiospores in subg. Amylospora . Both subgenera are further divided in two sections. The analysis of spore morphology shows that sect. Conica of subg. Dermoloma and sect. Nigrescentia of subg. Amylospora have a very distinctive spore shape. Sect. Atrobrunnea of subg. Amylospora showed relatively high variability of spores among species, but spores of sect. Dermoloma were similar and not useful for species discrimination.

Phylogenetic reconstruction of Hodophilus species with a yellow colour on the stipe based on nrITS, nrLSU and rpb2 sequences revealed six European species. All these species correspond to the widely accepted European concept of a single species Hodophilus micaceus. Four of these species are described and illustrated. H. micaceus and H. phaeoxanthus are recognised as two separate species and H. albofloccipes as a synonym of the latter. Two species, H. anatinus and H. cambriensis, are described as new. Possible endemism of H. micaceus and H. cambriensis to the British Isles is discussed. All analysed North American samples represent different species to those found in Europe. The North American species Hygrophorus rugulosus is combined in the genus Hodophilus. The preliminary key uses position and development of the yellow colour during maturation as the most important distinguishing character. The presence of the yellow colour is discussed as a possible synapomorphic character.

Among 45 Russula species and infraspecific taxa described by J. Blum, only few are currently accepted. Here we present a case study on R. formosa nom. illeg. (homotypic synonym of R. blumiana), R. decipiens var. ochrospora nom. inval. and R. roseobrunnea. The study is based on sequences and morphological observations of authentic herbarium material determined by J. Blum. The sequence data demonstrate that R. decipiens var. ochrospora is probably identical with the type variety and R. roseobrunnea is probably conspecific with R. rutila. Russula blumiana is recognised and described in detail as a good species related to R. badia.

In this paper, the authors present modern microscopical studies for 28 type specimens of Russula species described by C.H. Peck, thereby completing the revision of all 44 Russula taxa described by this author and for which type specimens were available. Our results suggest that with eventual exception of R. granulata var. lepiotoides (a likely synonym of the type variety) and R. magnifica (a likely synonym of R. polyphylla), all other Russulas described by Peck correspond to well-defined North American taxa. Having been described more than one century ago, Peck's names represent the majority of the earliest described North American Russula species and our type revisions are therefore important both for nomenclatural aspects, as well as typification and identification purposes. All studied taxa are illustrated in detail and accompanied by notes including a short recapitulation of previous type revisions, as well as our own re-evalutation of their systematic position as a result of this study.

Russula is one of the most speciose genera of mushroom-forming fungi, but phylogenetic relationships among species and subgeneric groupings are poorly understood. Our multi-locus phylogenetic reconstruction places R. firmula, R. rubra, R. rutila and R. veternosa in a well-supported Rubrinae clade, belonging to the Integrae clade of the Crown clade of the genus Russula. Traditional morphology-based classifications placed these four species in two different subsections based on the presence or absence of incrustations on pileocystidia. The Integrae clade also contains R. integra and related species that are traditionally placed in other groups based on their mild taste. Ancestral state reconstruction suggests that the common ancestor of the Crown clade and the Integrae clade probably did not have any incrustations in the pileipellis, had a mild taste, yellow spore print and were associated with angiosperms. All four species of the Rubrinae clade are defined by a darker yellow or ochre spore print, acrid taste and incrustations on pileocystidia. This last character contradicts the former splitting of the group because incrustations were apparently overlooked in R. firmula and R. veternosa. Incrustation type is now highlighted as being important for the delimitation of species and groups within the Crown clade. Pink or red staining of the incrustations in sulphovanillin is present in all species of the Rubrinae clade and a majority of the analysed species of the Integrae clade. The delimitation of the Rubrinae clade and its species circumscriptions are summarised here in a new diagnostic key.

Hodophilus atropunctus is traditionally defined as the only species of this genus with dark brown or black dots on the stipe. Multi-locus phylogenetic reconstruction recognised two distinct clades morphologically corresponding to this species concept. The limited morphological description in the protologue of H. atropunctus and absence of a type specimen were limitations in an assignment of this name to one of the recognised phylogenetic species. The emended species concept and the selection of a neotype are based on careful analyses of the colour of the basidiomata and how this changes during maturation and drying. The name H. atropunctus is assigned to the paler of the two species which also shows colour change across the pileus and along the length of the stipe when dry. The second darker species is described here as new, H. variabilipes, but only seven out of 14 collections examined belonging to this taxon had distinct dark coloured dots on the stipe surface.

Seven Melanoleuca taxa related to M. exscissa were studied based on macromorphological and micromorphological characters, and DNA sequences of three genes of recent collections and herbarium specimens. One species, M. griseobrunnea from the Korea is described as new to science. The type specimens of M. cinerascens, M. diverticulata, M. kuehneri, M. meridionalis, M. subexcentrica, M. stepposa and M. tristis are revised by micromorphological description and DNA sequencing of ITS region. The type analysis confirmed M. cinerascens, M. kuehneri and M. meridionalis are conspecific with M. exscissa and M. subexcentrica is conspecific with M. diverticulata. The lectotype and epitype of M. exscissa and neotype of M. rasilis are proposed here.

The Russula globispora lineage is a morphologically and phylogenetically well-defined group of ectomycorrhizal fungi occurring in various climatic areas. In this study we performed a multi-locus phylogenetic study based on collections from boreal, alpine and arctic habitats of Europe and Western North America, subalpine collections from the southeast Himalayas and collections from subtropical coniferous forests of Pakistan. European and North American collections are nearly identical and probably represent a single species named R. dryadicola distributed from the Alps to the Rocky Mountains. Collections from the southeast Himalayas belong to two distinct species: R. abbottabadensis sp. nov. from subtropical monodominant forests of Pinus roxburghii and R. tengii sp. nov. from subalpine mixed forests of Abies and Betula. The results suggest that speciation in this group is driven by a climate disjunction and adaptation rather than a host switch and geographical distance.

The Russula globispora lineage is a morphologically and phylogenetically well-defined group of ectomycorrhizal fungi occurring in various climatic areas. In this study we performed a multi-locus phylogenetic study based on collections from boreal, alpine and arctic habitats of Europe and Western North America, subalpine collections from the southeast Himalayas and collections from subtropical coniferous forests of Pakistan. European and North American collections are nearly identical and probably represent a single species named R. dryadicola distributed from the Alps to the Rocky Mountains. Collections from the southeast Himalayas belong to two distinct species: R. abbottabadensis sp. nov. from subtropical monodominant forests of Pinus roxburghii and R. tengii sp. nov. from subalpine mixed forests of Abies and Betula. The results suggest that speciation in this group is driven by a climate disjunction and adaptation rather than a host switch and geographical distance. Electronic supplementary material The online version of this article (10.1186/s43008-019-0003-9) contains supplementary material, which is available to authorized users.

Four European Hodophilus species with an odour similar to naphthalene, a strong unpleasant odour similar to that of mothballs, are recognized based on sequence and/or morphological data. The traditional concept defines Ho. foetens as the only Hodophilus species with a naphthalene odour in Europe. This name is now assigned to one of the studied species based on morphological examination of the holotype specimen. A recently collected specimen is proposed as the epitype. The other three species with a naphthalene odour are described here as new: Ho. pallidus, Ho. subfoetens and Ho. tenuicystidiatus. They are distinguishable in the field based on a combination of lamellae number and colour of basidiomata. All four species are grouped in the Ho. foetens superclade, one of two superclades, together with the Ho. micaceus superclade, in the genus Hodophilus. All are different species from North American taxa with a naphthalene-like odour recognised in a previous study. The Ho. foetens superclade also includes one species identified as Ho. atropunctus that does not have a distinctive odour. The type collection of Ho. albofloccipes, a recently described European species with a naphthalene odour, is placed together with some collections without a distinctive odour in the Ho. micaceus superclade.

Five North American Hodophilus species with naphthalene-like odours are now recognized based on sequence and (or) morphological data and molecular annotation of type collections. Two well-supported eastern North American species do not match any of the studied types and are described here as new: Hodophilus hesleri and Hodophilus smithii. The previously described Hodophilus paupertinus is found to represent an autonomous species and appears restricted to western North America. Hodophilus subfuscescens is found to be an independent lineage in eastern North America. A morphological type study of Hodophilus peckianus shows that it is a distinct species and not represented among recent collections. Multilocus phylogenetic analyses of European and North American material of species with naphthalene odours reveal no species with transatlantic distributions. Overall, Hodophilus comprises two superclades (the Hodophilus foetens superclade and the Hodophilus micaceus superclade) and 16 terminal clades that correspond to phylogenetic species. This study introduces a new approach for morphological delimitation of agaricoid Clavariaceae combining shape and dimensions of particular elements in the pileipellis and caulocystidia. All previously described taxa included in this study, which were previously treated in the genera Hygrophorus, Camarophyllopsis, or Hygrotrama, are formally transferred to Hodophilus.

In September 2011, an international team of cryptogam experts visited seven national nature reserves in five mountain areas of Slovakia: Havesova and Stužica in the Poloniny Mts., Vihorlat in the Vihorlatske vrchyMts., Oblik in the Slanske vrchyMts., Dobrocský prales and Klenovský Vepor in the Veporske vrchy Mts. and Badinsky prales in the Kremnicke vrchy Mts. The reserves were selected to represent examples of the best protected old-growth beech forests in the country. The aim was to study the diversity of wood-inhabiting fungi on fallen beech logs and epiphytic lichens on standing beech trees. In total, 215 fungal species and 128 lichens were recorded on beech wood and bark, and 27 fungi and 26 lichens on additional substrates. The site of the highest conservation value is Stužica with 126 fungi and 79 lichens recorded on beech, of which 12 fungi and 19 lichens are indicators of high nature conservation value. Combined with historical records, a total of 19 non-lichenised fungal indicators are now reported from the site, making it the highest ranked natural beech forest in Europe. The second most important reserve for fungal diversity is Havesova with 121 species, including 14 indicator species recorded on beech wood. For lichens, the second most important reserve is Klenovský Vepor with 69 species including 18 lichen indicators recorded on beech. Nine fungus species are here reported as new to Slovakia: Asterostroma medium, Entoloma hispidulum, E. pseudoparasiticum, Gloeohypochnicium analogum, Hohenbuehelia valesiaca, Hymenochaete ulmicola, Hypocrea parmastoi, Melanomma spiniferum and Scutellinia colensoi. Lichen species Alyxoria ochrocheila is reported as new to Slovakia and Lecanographa amylacea, which was considered extinct in the Slovak Red list, was also recorded. This is the first list of wood-inhabiting fungi and epiphytic lichens of old-growth beech forests in Slovakia, and hence an important contribution to the exploration of biodiversity in Slovakia.

A short history to the study of Russula in the Western United States is provided and for the first time an exhaustive list of all 49 taxa described from this part of the country has been compiled. As a start to a revision of these species, a first series of four holotypes (one from California and three from the Pacific Northwest) have here been studied microscopically, resulting in some very substantial changes concerning species concept and consequent systematic placement. Russula flava var. pacifica appears unrelated to the type variety. It is therefore excluded as member of Integroidinae, subgen. Polychromidia, and deserves probably to be upgraded to species level within subgen. Russula. The Californian R. paxilloides is lectotypified and is a good member of subgen. Russula, but is closer to sect. Persicinae than to the white-spored Vinaceae as previously suggested. Russula maxima, widely accepted as a species of subsect. Subcompactinae (subgen. Heterophyllidia), certainly does not belong ther...

The types or authentic material of four members of Russula subsect. Maculatinae recognized by Romagnesi were studied: R. decipiens, R. decipiens var. vermiculata nom. inval., R. deceptiva and R. romagnesii. All studied specimens showed several similar micromorphological characters, particularly important one-celled pileocystidia, suggesting their close relationship. Based on the differences in spore ornamentation and terminal cells of hyphae in pileipellis near the cap margin, two morphotypes were recognized. The first one, represented by the neotypus of R. decipiens and authentic material of R. decipiens var. vermiculata nom. inval., has lower spore ornamentation with more numerous warts and apically constricted, terminal cells of hyphae in pileipellis as compared to the second morphotype represented by the holotypus of R. deceptiva and paratypus of R. romagnesii.

A taxonomic and nomenclatural revision of some representatives of Clavariaceae is presented based on extensive collecting in central and western Europe. Five species originally described from Europe are identified, redescribed and delimited: Clavaria fragilis, Ramariopsis crocea, R. corniculata, R. helvola and R. pulchella. Lectotypes, epitypes or neotypes are designated for all these species. Descriptions are based on macro- and micromorphological characters and supplemented with DNA analyses of the nrLSU regions from 20 specimens. The molecular phylogenetic analyses reconstructed a phylogram showing relation- ships among the discussed species as well as some closely related taxa. The taxonomic value of the ratio of length and width of spores (Q-value) is discussed.

Melanoleuca brevipes (basionym Agaricus brevipes) represents one of the most frequently identified Melanoleuca species. However, there are several concepts of this taxon, and thus the identity of this species is unclear. This species was studied, based on the original table by Bulliard, macromorphological and micromorphological characters, and DNA sequences of recent collections and herbarium specimens. DNA sequences of ITS, RPB2 and tef1 markers revealed that specimens identified as M. brevipes belong to three species (M. humilis, M. grammopodia, and M. malenconii), where M. humilis or M. malenconii may resemble Bulliard’s original species. According to these results, Agaricus brevipes is therefore considered a nomen dubium. A lectotype of M. grammopodia is proposed herein.

Crepidotus stenocystis and C. brunnescens are morphologically similar species defined by globose to subglobose spores, the presence of clamp connections in all tissues and bottle-like or flask-like cheilocystidia. They are also similar in the pileal aspect which is hygrophanous, glabrous or white-fibrillose, at first white but becoming brownish with age. Each are described from separate continents and have only been reported from their respective continents, C. stenocystis from Europe and C. brunnescens from North America. The phylogenetic analysis of ITS and LSU nrDNA regions from original type material confirms the existence of two distinct species, C. brunnescens that is more closely related to C. malachioides , and C. stenocystis that is more closely related to C. applanatus . Crepidotus stenocystis differs from C. brunnescens by its more prominent spore ornamentation and longer cheilocystidia that are frequently narrowly utriform. Based on the studied material and published data, it seems that C. stenocystis is distributed throughout all of Europe and does not occur in North America, whereas C. brunnescens is only known from Michigan in the USA.

The current generally accepted concept of Russula maculata defines the species by yellow-brownish spots on the basidiomata, an acrid taste, a yellow spore print and a red pileus. This concept was tested using collections originating from various geographical areas mainly in Europe. Analyses of the ITS region suggested that there were three species within this broad concept. One of them, R. maculata , was identified based on the sequence from the epitype. Two other species, R. nympharum and R. sp., are described here as newly identified species. The European species R. maculata and R. nympharum grow in deciduous forests, are similar in their field aspect and are distinctly different in micro-morphological characteristics of spores, pleurocystidia and pileipellis. An Asian species, R. sp., is associated with pine and has smaller basidiomata and spores. These three species form the R. maculata complex and represent the sister clade to the R. globispora complex. This clade consists of species also characterized by a yellow-brownish context discolouration but with a different type of spore ornamentation. All of the other tested species had an acrid taste and yellow spore print but did not have a conspicuous yellow-brownish context discolouration and were placed in various unrelated clades.

In this second contribution on native American species in Russula subsect. Decolorantes , the authors focus on four additional species that exhibit a blackening context: two species from the West Coast, R. californiensis and R. occidentalis , and two species from the East Coast, R. rubescens and R. rubriceps . A key to the species in Decolorantes is provided. Russula californiensis and R. rubescens are lectotypified.