Holarctic Species in the Pluteus podospileus Clade: Description of Six New Species and Reassessment of Old Names.

We studied the taxonomy of Pluteus podospileus and similar species using morphological and molecular (nrITS, TEF1-α) data, including a detailed study of the type collections of P. inflatus var. alneus, Pluteus minutissimus f. major, and P. granulatus var. tenellus. Within the P. podospileus complex, we phylogenetically confirmed six species in Europe, five in Asia, and eight in North America. Based on our results, we recognize P. seticeps as a separate species occurring in North America, while P. podospileus is limited to Eurasia. We describe six new species and a new variety: P. absconditus, P. fuscodiscus, P. gausapatus, P. inexpectatus, P. millsii, and P. notabilis and its variety, P. notabilis var. insignis. We elevate Pluteus seticeps var. cystidiosus to species rank as Pluteus cystidiosus. Based on the holotype of P. inflatus var. alneus, collections of P. inflatus identified by Velenovský, and several modern collections, we resurrect the name P. inflatus. Based on molecular analyses of syntypes of Pluteus minutissimus f. major and a holotype of Pluteus granulatus var. tenellus, we synonymize them under P. inflatus. We also increase our knowledge about the morphology and distribution of P. cutefractus.

Resources of Iranian agarics (Basidiomycota) with an outlook on their antioxidant potential.

Agaric fungi are an important group of macromycetes with diverse ecological and functional properties, yet are poorly studied in many parts of the world. Here, we comprehensively analyzed 558 agaric species in Iran to reveal their resources of edible and poisonous species as well as their ecological guilds and luminescence potential. We also made a thorough survey of the antioxidant activity of the species. Phylogenetic relationships were reconstructed based on nuclear ribosomal LSU and ITS sequences. Our results reveal that agarics of Iran comprise about 189 edible, 128 poisonous, 254 soil saprotrophic, 172 ectomycorrhizal, 146 wood-inhabiting, 18 leaf/litter-inhabiting, 9 parasitic, and 19 luminescent species. Twenty percent of the Iranian agaric species possess antioxidant activity, phylogenetically distributed in four orders and 21 agaric families. About 5% of the antioxidant species can be considered strong antioxidants, many of which are also edible and could be utilized to develop functional foods. This is the first study combining phylogeny and antioxidant potential of agaric mushrooms in a large scale, and the obtained results would guide the selection of agaric taxa to be examined in the future for taxonomic revisions, biotechnological applications, and applied phylogeny studies.

Holarctic Species in the Pluteus romellii Clade. Five New Species Described and Old Names Reassessed.

We studied the taxonomy of Pluteus romellii, and morphologically similar Holarctic species in the /romellii clade of section Celluloderma, using morphological and molecular data (nrITS, TEF1-α). Pluteus romellii is lectotypified and epitypified and accepted as an exclusively Eurasian species. Pluteus lutescens and P. pallescens are considered synonyms of P. romellii. Pluteus fulvibadius is accepted as a related, but separate, North American species. Five species in the /romellii clade are described as new to science: two from North America (P. austrofulvus and P. parvisporus), one from Asia (P. parvicarpus), one from Europe (P. siccus), and one widely distributed across the Holarctic region (P. vellingae). Basidioma size, pileus color, lamellae color, basidiospore size, hymenial cystidia shape and size, habitat and geographical distribution help separate the species described here, but in some instances only molecular data allows for confident identification. The current status of P. californicus, P. melleipes, P. romellii var. luteoalbus, P. splendidus, P. sternbergii and P.sulphureus is discussed.

Emending Gymnopus sect. Gymnopus (Agaricales, Omphalotaceae) by including two new species from southern China.

Based on phylogenetic analyses, some newly studied Chinese mushroom specimens were found to represent two distinct species within the genus Gymnopus. Along with G.fusipes (sect. Gymnopus) they form a distinct clade with high support, although their macromorphological characters seem to be closer to members of Gymnopus sect. Levipedes or sect. Vestipedes (Collybiopsis). When examined in detail, their micromorphological characters, especially the type of pileipellis, support them as new members of G. sect. Gymnopus. Therefore, two new species, G.omphalinoides and G.schizophyllus, and the emended circumscription of sect. Gymnopus are proposed in this paper. Detailed morphological descriptions, colour photos, illustrations of the two new species, morphological comparisons with similar taxa and the molecular-phylogenetic analyses of the combined nrITS and nrLSU data are presented. A key to the known species of G. sect. Gymnopus is also presented.

Multilocus phylogeny and taxonomy of European Melanoleuca subgenus Melanoleuca.

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 galbuserae, M. fontenlae and M. acystidiata-Three New Species in Subgenus Urticocystis (Pluteaceae, Basidiomycota) with Comments on M. castaneofusca and Related Species.

Melanoleuca is one of the taxonomically most complicated genera of Agaricomycetes with several taxonomically lineages. The subgenus Urticocystis of the genus Melanoleuca contains species with either urticoid or absent cheilocystidia. In this paper, three new European species, Melanoleuca galbuserae, Melanoleuca fontenlae, and Melanoleuca acystidiata are described as new to science. Melanoleuca galbuserae, related to Melanoleuca stepposa and Melanoleuca tristis, was discovered in alpine grasslands in North Italy. The type specimens and recent collections of Melanoleuca angelesiana, Melanoleuca castaneofusca, Melanoleuca luteolosperma, Melanoleuca pseudopaedida, and Melanoleuca robertiana were sequenced and morphologically examined. Moreover, the related Melanoleuca microcephala and Melanoleuca paedida were included in morphological examination and DNA sequence analyses. All the species were delimited by macro- and micromorphological characters and the multigene phylogenetic analyses of a combined (ITS, rpb2, and tef1) dataset on the basis of the species tree estimation. In accordance with new molecular and morphological data, we suggest taxonomic reappraisal of M. pseudopaedida and M. robertiana, and M. fontenlae and M. acystidiata are proposed as new species. The differences between the type material of M. angelesiana from the USA and European M. angelesiana specimens are discussed.

Desarmillaria caespitosa, a North American vicariant of D. tabescens.

Desarmillaria caespitosa, a North American vicariant species of European D. tabescens, is redescribed in detail based on recent collections from the USA and Mexico. This species is characterized by morphological features and multilocus phylogenetic analyses using portions of nuc rDNA 28S (28S), translation elongation factor 1-alpha (tef1), the second largest subunit of RNA polymerase II (rpb2), actin (act), and glyceraldehyde-3-phosphate dehydrogenase (gpd). A neotype of D. caespitosa is designated here. Morphological and genetic differences between D. caespitosa and D. tabescens were identified. Morphologically, D. caespitosa differs from D. tabescens by having wider basidiospores, narrower cheilocystidia, which are often irregular or mixed (regular, irregular, or coralloid), and narrower caulocystidia. Phylogenetic analyses of five independent gene regions show that D. caespitosa and D. tabescens are separated by nodes with strong support. The new combination, D. caespitosa, is proposed.

Marasmioid and Gymnopoid Fungi of the Republic of Korea. 8. Gymnopus Section Levipedes.

Collections of Gymnopus sect. Levipedes from the Republic of Korea have been studied. Two new species, Gymnopus dryophiloides and G. brunneodiscus , are described based on their macro- and micromorphological and phylogenetic characteristics. Three other species, referred to as Gymnopus spp. 1, 2, and 3, are distinguished as separate taxa without formal descriptions. Taxonomic and phylogenetic positions of all taxa have been inferred and confirmed by analyses of ITS and LSU sequence data. Their detailed descriptions, illustrations and an identification key are provided.

New Cantharellus species from South Korea.

In this third contribution involving new Cantharellus species from South Korea, two new species are introduced. In addition, we document a first report of the recently described Japanese Cantharellus anzutake outside of Japan based on identical ITS sequence data. Cantharellus citrinus sp. nov. is introduced as a new member of subgenus Cinnabarini, to which the closely related Korean C. albovenosus and Chinese C. phloginus also belong. Cantharellus curvatus sp. nov. is introduced as a new member of subgenus Parvocantharellus, in which the Korean C. koreanus was recently placed. The respective placements of the new taxa are significantly supported by a phylogenetic analysis of sequences from the transcription elongation factor (tef-1).

Taxonomic update of Clitocybula sensu lato with a new generic classification.

The taxonomy and phylogeny of the hydropoid clade (genera Clitocybula s.l., Megacollybia, Leucoinocybe gen. nov., Hydropus, Trogia, Gerronema, Porotheleum and Lignomphalia gen. nov.) in Europe is studied using morphological and molecular approaches; the first three genera in detail including all known European species. Only two European species remain in Clitocybula s.str., Clitocybula lacerata and Clitocybula familia. The European C. lacerata is a species complex which should be treated as C. lacerata agg. at the current state of knowledge. A neotype originating from type area was designated to fix the application of the name. The presence of American species Clitocybula abundans in Europe is insufficiently proved. "Clitocybula dryadicola Ë® belongs to the genus Hydropus, and Clitocybula tilieti has an unclear systematic position. The results showed that Megacollybia and Leucoinocybe represent independent genera separated from Clitocybula. The genus Leucoinocybe is validly published with two European species, Leucoinocybe lenta and Leucoinocybe taniae. "Clitocybula flavoaurantia" proved to be conspecific with the latter species. The genus Lignomphalia is published for "Pseudoomphalina lignicola", a lignicolous omphalinoid species. The Indian "Clitocybula sulcata" is transferred to Leucoinocybe and "Clitocybula atrialba" to Gerronema. The first European records of Megacollybia marginata are published.

Megaphylogeny resolves global patterns of mushroom evolution.

Mushroom-forming fungi (Agaricomycetes) have the greatest morphological diversity and complexity of any group of fungi. They have radiated into most niches and fulfil diverse roles in the ecosystem, including wood decomposers, pathogens or mycorrhizal mutualists. Despite the importance of mushroom-forming fungi, large-scale patterns of their evolutionary history are poorly known, in part due to the lack of a comprehensive and dated molecular phylogeny. Here, using multigene and genome-based data, we assemble a 5,284-species phylogenetic tree and infer ages and broad patterns of speciation/extinction and morphological innovation in mushroom-forming fungi. Agaricomycetes started a rapid class-wide radiation in the Jurassic, coinciding with the spread of (sub)tropical coniferous forests and a warming climate. A possible mass extinction, several clade-specific adaptive radiations and morphological diversification of fruiting bodies followed during the Cretaceous and the Paleogene, convergently giving rise to the classic toadstool morphology, with a cap, stalk and gills (pileate-stipitate morphology). This morphology is associated with increased rates of lineage diversification, suggesting it represents a key innovation in the evolution of mushroom-forming fungi. The increase in mushroom diversity started during the Mesozoic-Cenozoic radiation event, an era of humid climate when terrestrial communities dominated by gymnosperms and reptiles were also expanding.

Fungal Systematics and Evolution: FUSE 5.

Thirteen new species are formally described: Cortinarius brunneocarpus from Pakistan, C. lilacinoarmillatus from India, Curvularia khuzestanica on Atriplex lentiformis from Iran, Gloeocantharellus neoechinosporus from China, Laboulbenia bernaliana on species of Apenes, Apristus, and Philophuga (Coleoptera, Carabidae) from Nicaragua and Panama, L. oioveliicola on Oiovelia machadoi (Hemiptera, Veliidae) from Brazil, L. termiticola on Macrotermes subhyalinus (Blattodea, Termitidae) from the DR Congo, Pluteus cutefractus from Slovenia, Rhizoglomus variabile from Peru, Russula phloginea from China, Stagonosporopsis flacciduvarum on Vitis vinifera from Italy, Strobilomyces huangshanensis from China, Uromyces klotzschianus on Rumex dentatus subsp. klotzschianus from Pakistan. The following new records are reported: Alternaria calendulae on Calendula officinalis from India; A. tenuissima on apple and quince fruits from Iran; Candelariella oleaginescens from Turkey; Didymella americana and D. calidophila on Vitis vinifera from Italy; Lasiodiplodia theobromae causing tip blight of Dianella tasmanica 'variegata' from India; Marasmiellus subpruinosus from Madeira, Portugal, new for Macaronesia and Africa; Mycena albidolilacea, M. tenuispinosa, and M. xantholeuca from Russia; Neonectria neomacrospora on Madhuca longifolia from India; Nothophoma quercina on Vitis vinifera from Italy; Plagiosphaera immersa on Urtica dioica from Austria; Rinodina sicula from Turkey; Sphaerosporium lignatile from Wisconsin, USA; and Verrucaria murina from Turkey. Multi-locus analysis of ITS, LSU, rpb1, tef1 sequences revealed that P. immersa, commonly classified within Gnomoniaceae (Diaporthales) or as Sordariomycetes incertae sedis, belongs to Magnaporthaceae (Magnaporthales). Analysis of a six-locus Ascomycota-wide dataset including SSU and LSU sequences of S. lignatile revealed that this species, currently in Ascomycota incertae sedis, belongs to Pyronemataceae (Pezizomycetes, Pezizales).

Considerations and consequences of allowing DNA sequence data as types of fungal taxa.

Nomenclatural type definitions are one of the most important concepts in biological nomenclature. Being physical objects that can be re-studied by other researchers, types permanently link taxonomy (an artificial agreement to classify biological diversity) with nomenclature (an artificial agreement to name biological diversity). Two proposals to amend the International Code of Nomenclature for algae, fungi, and plants (ICN), allowing DNA sequences alone (of any region and extent) to serve as types of taxon names for voucherless fungi (mainly putative taxa from environmental DNA sequences), have been submitted to be voted on at the 11th International Mycological Congress (Puerto Rico, July 2018). We consider various genetic processes affecting the distribution of alleles among taxa and find that alleles may not consistently and uniquely represent the species within which they are contained. Should the proposals be accepted, the meaning of nomenclatural types would change in a fundamental way from physical objects as sources of data to the data themselves. Such changes are conducive to irreproducible science, the potential typification on artefactual data, and massive creation of names with low information content, ultimately causing nomenclatural instability and unnecessary work for future researchers that would stall future explorations of fungal diversity. We conclude that the acceptance of DNA sequences alone as types of names of taxa, under the terms used in the current proposals, is unnecessary and would not solve the problem of naming putative taxa known only from DNA sequences in a scientifically defensible way. As an alternative, we highlight the use of formulas for naming putative taxa (candidate taxa) that do not require any modification of the ICN.

Insights into the phylogeny of Northern Hemisphere Armillaria: Neighbor-net and Bayesian analyses of translation elongation factor 1-α gene sequences.

Armillaria possesses several intriguing characteristics that have inspired wide interest in understanding phylogenetic relationships within and among species of this genus. Nuclear ribosomal DNA sequence-based analyses of Armillaria provide only limited information for phylogenetic studies among widely divergent taxa. More recent studies have shown that translation elongation factor 1-α (tef1) sequences are highly informative for phylogenetic analysis of Armillaria species within diverse global regions. This study used Neighbor-net and coalescence-based Bayesian analyses to examine phylogenetic relationships of newly determined and existing tef1 sequences derived from diverse Armillaria species from across the Northern Hemisphere, with Southern Hemisphere Armillaria species included for reference. Based on the Bayesian analysis of tef1 sequences, Armillaria species from the Northern Hemisphere are generally contained within the following four superclades, which are named according to the specific epithet of the most frequently cited species within the superclade: (i) Socialis/Tabescens (exannulate) superclade including Eurasian A. ectypa, North American A. socialis (A. tabescens), and Eurasian A. socialis (A. tabescens) clades; (ii) Mellea superclade including undescribed annulate North American Armillaria sp. (Mexico) and four separate clades of A. mellea (Europe and Iran, eastern Asia, and two groups from North America); (iii) Gallica superclade including Armillaria Nag E (Japan), multiple clades of A. gallica (Asia and Europe), A. calvescens (eastern North America), A. cepistipes (North America), A. altimontana (western USA), A. nabsnona (North America and Japan), and at least two A. gallica clades (North America); and (iv) Solidipes/Ostoyae superclade including two A. solidipes/ostoyae clades (North America), A. gemina (eastern USA), A. solidipes/ostoyae (Eurasia), A. cepistipes (Europe and Japan), A. sinapina (North America and Japan), and A. borealis (Eurasia) clade 2. Of note is that A. borealis (Eurasia) clade 1 appears basal to the Solidipes/Ostoyae and Gallica superclades. The Neighbor-net analysis showed similar phylogenetic relationships. This study further demonstrates the utility of tef1 for global phylogenetic studies of Armillaria species and provides critical insights into multiple taxonomic issues that warrant further study.

Phylogenetic studies in Psathyrella focusing on sections Pennatae and Spadiceae--new evidence for the paraphyly of the genus.

The sections Pennatae and Spadiceae were chosen to test the agreement of current infrageneric classifications of Psathyrella (Psathyrellaceae, Agaricales) with molecular phylogenetic data and to evaluate the systematic significance of relevant morphological characters. The ITS and partial LSU regions of nu-rDNA from 53 specimens representing 34 species of Psathyrella were sequenced and analysed with parsimony-based and model-based phylogenetic methods. According to our analyses, the sections Pennatae and Spadiceae are polyphyletic and distributed across the family Psathyrellaceae, which is divided into at least five major groups. The first one comprises most of the included Psathyrella species and, probably, the whole genus Coprinellus. The second group is made up of Psathyrella gossypina and P. delineata. The third clade consists of the genus Coprinopsis and includes Psathyrella aff. huronensis and P. marcescibilis. The fourth clade is composed of two sister groups, the subgenus Homophron and the genus Lacrymaria, and the fifth group represents the genus Parasola including Psathyrella conopilus. These results are in agreement with neither the current circumscription of the two subgenera, Psathyra and Psathyrella, nor with the pre-sent disposition of the Psathyrellaceae. Taxonomically important morphological characters in the genus Psathyrella show a high degree of homoplasy. Although these characters are useful for species delimitation, and in some cases for the circumscription of sections, they appear insufficient for a phylogenetically correct generic concept.

The genus Setulipes (Marasmiaceae) in Madagascar and the Mascarenes, including a key to other African taxa.

The authors report first records for the genus Setulipes in Madagascar, with the presence of Setulipes cf. hakgalensis and two new species, Setulipes funaliformis and S. moreaui, as well as a new species from Mauritius: S. mauritiensis. A key to these taxa, as well as to other African species, is supplied.

Crinipellis pedemontana sp. nov. (Agaricomycetes), a new basidiomycete from Italy.

Crinipellis pedemontana, a new agaric growing on stems of dead grasses in an Italian urban park, is described and illustrated. It fits in sect. Grisentinae on account of the green reaction with KOH and ammonia and its bright colored pileus. The unique combination of purple-violaceous tinges in the pileus and the presence of chlamydospores in the pileipellis characterize this species as new. Its taxonomic position within Crinipellis and the related genus Moniliophthora is discussed.