Ancestral predisposition toward a domesticated lifestyle in the termite-cultivated fungus Termitomyces.

The ancestor of termites relied on gut symbionts for degradation of plant material, an association that persists in all termite families.1,2 However, the single-lineage Macrotermitinae has additionally acquired a fungal symbiont that complements digestion of food outside the termite gut.3 Phylogenetic analysis has shown that fungi grown by these termites form a clade-the genus Termitomyces-but the events leading toward domestication remain unclear.4 To address this, we reconstructed the lifestyle of the common ancestor of Termitomyces using a combination of ecological data with a phylogenomic analysis of 21 related non-domesticated species and 25 species of Termitomyces. We show that the closely related genera Blastosporella and Arthromyces also contain insect-associated species. Furthermore, the genus Arthromyces produces asexual spores on the mycelium, which may facilitate insect dispersal when growing on aggregated subterranean fecal pellets of a plant-feeding insect. The sister-group relationship between Arthromyces and Termitomyces implies that insect association and asexual sporulation, present in both genera, preceded the domestication of Termitomyces and did not follow domestication as has been proposed previously. Specialization of the common ancestor of these two genera on an insect-fecal substrate is further supported by similar carbohydrate-degrading profiles between Arthromyces and Termitomyces. We describe a set of traits that may have predisposed the ancestor of Termitomyces toward domestication, with each trait found scattered in related taxa outside of the termite-domesticated clade. This pattern indicates that the origin of the termite-fungus symbiosis may not have required large-scale changes of the fungal partner.

Six new species and reports of Hydnum (Cantharellales) from eastern North America.

Five species of Hydnum have been generally recognized from eastern North America based on morphological recognition: H.albidum, H.albomagnum, H.repandum and varieties, H.rufescens, and H.umbilicatum. Other unique North American species, such as H.caespitosum and H.washingtonianum, are either illegitimately named or considered synonymous with European taxa. Here, seventeen phylogenetic species of Hydnum are detected from eastern North America based on a molecular phylogenetic survey of ITS sequences from herbarium collections and GenBank data, including environmental sequences. Based on current distribution results, sixteen of these species appear endemic to North America. Of these, six species are described as new: H.alboaurantiacum, H.cuspidatum, H.ferruginescens, H.subconnatum, H.subtilior, and H.vagabundum. Geographic range extensions and taxonomic notes are provided for five additional species recently described as new from eastern North America. A new name, H.geminum, is proposed for H.caespitosum Banning ex Peck, non Valenti. Overall, species of Hydnum are best recognized by a combination of morphological and molecular phylogenetic analyses. Taxonomic descriptions are provided for seventeen species, including epitype designations for H.albidum, H.albomagnum, and H.umbilicatum, taxa described more than 100 years ago, and molecular annotation of the isotype of H.washingtonianum. Photographs and a key to eastern North American Hydnum species are presented.

Four new species of Morchella from the Americas.

Morphological and molecular phylogenetic studies of true morels (Morchella) in North America, the Dominican Republic, Venezuela, Ecuador, and Peru led to the discovery of four undescribed species of Morchella. Two new species in the Elata clade, one from the Dominican Republic, initially distinguished by the informal designation Mel-18, and a newly discovered sister species from northern Arizona, are now recognized. Mel-18 is described as a novel phylogenetically distinct species, M. hispaniolensis. Its sister species from Arizona is described as M. kaibabensis, also recovered as an endophyte of Rocky Mountain juniper. Two additional species in the Esculenta clade, M. peruviana discovered in Peru and M. gracilis (previously reported as Mes-14) from the Dominican Republic, Venezuela, and Ecuador, are described as new. We also demonstrate that scanning electron microscopy (SEM) imaging of ascospores using rehydration/dehydration/critical point drying preparation techniques provides for enhanced resolution of spore wall surfaces, thereby increasing the number of morphological traits available to assess differences among otherwise closely related species.

A global view of Gyroporus: molecular phylogenetics, diversity patterns, and new species.

Gyroporus (Gyroporaceae, Boletales) is a highly diverse genus of poroid ectomycorrhizal mushrooms with a nearly worldwide distribution. Previous attempts to unravel the diversity within this genus proved difficult due to the presence of semicryptic species and ambiguous results from analysis of ribosomal RNA markers. In this study, we employ a combined morphotaxonomic and phylogenetic approach to delimit species and elucidate geographic and evolutionary patterns in Gyroporus. For phylogenetic analyses, the protein-coding genes atp6 (mitochondrial adenosine triphosphate [ATP] synthase subunit 6) and rpb2 (nuclear second largest subunit of RNA polymerase II) were selected based on their utility in studies of Boletales. We infer several distinct clades, most notably one corresponding to G. castaneus as a speciose Northern Hemisphere group, another unifying G. cyanescens and like entities, and a third group unifying G. longicystidiatus and a New World sister species. Also notable is the recovery of a sister relationship between the cyanescens and longicystidiatus clades. We formally describe five new species of Gyroporus, outline a number of provisional species, and briefly discuss distributional patterns. This study provides an important scaffold for future work on this well-known but poorly understood genus of fungi.

A new stipitate species of Crepidotus from India and Thailand, with notes on other tropical species.

A new Asian species of Crepidotus (Basidiomycota, Agaricales), C. asiaticus, is presented based on morphological and nuc rDNA internal transcribed spacer (ITS1-5.8S-ITS2 = ITS) and large subunit (28S) sequence data. This new species, found in India and Thailand, is characterized by the centrally stipitate medium-sized basidiomata, orange to reddish brown pileus, white to brownish orange lamellae, and white stipe. Based on morphology, C. asiaticus is similar to the neotropical C. thermophilus. However, the microscopic characters, especially the size and shape of the basidiospores, can be used to distinguish these two taxa, as well as their geographic distributions. Further, the phylogenetic position of C. asiaticus is unique based on ITS and 28S nuc rDNA sequences. Melanomphalia argipoda, described by Singer from Ecuador, is also a stipitate Crepidotus based on an ITS sequence of the type specimen, so the new combination is proposed here. Phylogenetically, the three species form a monophyletic group with the Asiatic C. asiaticus forming the sister lineage to the neotropical C. argipodus and C. thermophilus.

Sarcodon in the Neotropics I: new species from Guyana, Puerto Rico and Belize.

Four species of the ectomycorrhizal (ECM) genus Sarcodon (Bankeraceae, Thelephorales, Basidiomycota) are described as new to science. Sarcodon pakaraimensis sp. nov. is described from forests dominated by the ECM trees Pakaraimaea dipterocarpacea (Dipterocarpaceae) and Dicymbe jenmanii (Fabaceae subfam. Caesalpinioideae) in the Pakaraima Mountains of Guyana. Sarcodon portoricensis sp. nov. is described from lower montane wet forest within the El Yunque National Forest of Puerto Rico. Sarcodon quercophilus sp. nov. and Sarcodon umbilicatus sp. nov. are described from Quercus (Fagaceae) cloud forests within the Maya Mountains of Belize. The discovery of these species is significant given that the majority of the approximately 87 described Sarcodon species are north temperate or boreal in distribution and frequently associate with coniferous host plants; these constitute the most recent records for Sarcodon from the greater Neotropics. Each of the new species is morphologically consistent with accepted diagnostic characters for Sarcodon: pileate-stipitate stature, a dentate hymenophore, determinate basidiomatal development, fleshy, non-zonate context and brown, tuberculate basidiospores. DNA (ITS) sequence analysis corroborated the generic placement of S. pakaraimensis, S. portoricensis, S. quercophilus and S. umbilicatus and, along with morphological differences, supported their recognition as distinct species. Macromorphological, micromorphological, habitat and DNA sequence data from the nuc rDNA internal transcribed spacer region (ITS) are provided for each of the new species. A key to Neotropical Sarcodon species and similar extralimital taxa is provided.

Toward a stable classification of genera within the Entolomataceae: a phylogenetic re-evaluation of the Rhodocybe-Clitopilus clade.

Despite the recent molecular systematic analyses of the Entolomataceae (Agaricales, Basidiomycota), a robust classification of genera supported by morphological and phylogenetic evidence remains unresolved for this cosmopolitan family of pink-spored fungi. Here, a phylogenetic analysis for one of the two major clades (Rhodocybe-Clitopilus) was conducted using three nuclear protein-coding gene regions, the mitochondrial ATP synthase subunit 6 (atp6), the nuclear RNA polymerase subunit II (rpb2) and the nuclear translation elongation factor subunit 1-α (tef1). Five monophyletic groups are resolved with strong statistical support and a set of morphological features for delineation of genera is presented. In the revised classification proposed here, Clitopilus is retained, Rhodocybe is emended, two genera previously accepted as synonyms of Rhodocybe (Clitopilopsis and Rhodophana) are resurrected and Clitocella is described as new.

Cercopemyces crocodilinus, a new genus and species related to Ripartitella, is described from North America.

Cercopemyces is described as a new genus based on collections of a newly discovered agaricoid species from the arid Cercocarpus forests of Utah and Colorado. The new genus is near Ripartitella and Cystodermella based on nLSU, rpb1 and ITS molecular sequences but distinctly different from these taxa. The ornamented basidiospores and inflated cells in the scales of the pileus surface of Cercopemyces indicate a close relationship to some species of Ripartitella, and the molecular analyses support the sister group relationship. Morphologically Cercopemyces crocodilinus is reminiscent of the genus Amanita in the field, but the inamyloid, cyanophilic, ornamented basidiospores and lack of acrophysalidic hyphae in the trama indicate otherwise. A rare eastern USA species, Ripartitella ponderosa, is transferred to Cercopemyces based on morphological and molecular data.

The Entolomataceae of the Pakaraima Mountains of Guyana IV: new species of Calliderma, Paraeccilia and Trichopilus.

This paper is the fourth in a series documenting the Entolomataceae taxa (Agaricales, Basidiomycota) from Guyana. Six new species in three genera are described-Calliderma caeruleosplendens, Paraeccilia unicolorata, Trichopilus tibiiformis, T. fasciculatus, T. vividus, and T. luteolamellatus-occurring primarily in mixed tropical rainforests of the Potaro River Basin in the Pakaraima Mountains. Macromorphological, micromorphological and habitat data are provided for each. None of these genera had been reported from Guyana.

The Entolomataceae of the Pakaraima mountains of Guyana I: four new species of Entoloma s. str.

This paper constitutes the first documented accounts of Entolomataceae taxa (Agaricales, Basidiomycota) from Guyana. Entoloma olivaceocoloratum, Entoloma rugosostriatum, Entoloma fragilum and Entoloma illinita are described as new species from the Pakaraima Mountains, occurring in rainforests dominated by ectomycorrhizal Dicymbe and Aldina spp. Macromorphological, micromorphological and habitat data are provided for the new species.

A new genus of Boletaceae from eastern North America.

Bothia is described as a new genus in the Boletaceae based on Boletinus castanellus described by C.H. Peck from eastern North America. A widespread, occasionally encountered taxon, Bothia castanella possesses a combination of macro- and microscopic features that has prompted past placement in seven different genera. Yet, as a species it is readily recognizable with its chestnut brown, dry pileus, decurrent, pale brown hymenophore with radially elongated tubes, a short, sometimes eccentric, exannulate stipe, yellow brown spore deposit and constant association with Quercus. Phylogenetic analyses of large subunit rDNA and BLAST searches using the ITS region confirm the placement of B. castanella as a unique generic lineage in the Boletaceae.

Arthromyces and Blastosporella, two new genera of conidia-producing lyophylloid agarics (Agaricales, Basidiomycota) from the neotropics.

Two new genera encompassing three new species of lyophylloid agarics that produce conidia on the basidiomata are described. Arthromyces is a genus comprised of two very different arthrospore-producing mushroom species found in the Greater Antilles and Central America. Blastosporella is a monotypic genus with spherical balls of blastospores covering the pileus surface with age and is known from Hispaniola and Colombia. A key to the species of Arthromyces is included.

Contributions of rpb2 and tef1 to the phylogeny of mushrooms and allies (Basidiomycota, Fungi).

A phylogeny of the fungal phylum Basidiomycota is presented based on a survey of 160 taxa and five nuclear genes. Two genes, rpb2, and tef1, are presented in detail. The rpb2 gene is more variable than tef1 and recovers well-supported clades at shallow and deep taxonomic levels. The tef1 gene recovers some deep and ordinal-level relationships but with greater branch support from nucleotides compared to amino acids. Intron placement is dynamic in tef1, often lineage-specific, and diagnostic for many clades. Introns are fewer in rpb2 and tend to be highly conserved by position. When both protein-coding loci are combined with sequences of nuclear ribosomal RNA genes, 18 inclusive clades of Basidiomycota are strongly supported by Bayesian posterior probabilities and 16 by parsimony bootstrapping. These numbers are greater than produced by single genes and combined ribosomal RNA gene regions. Combination of nrDNA with amino acid sequences, or exons with third codon positions removed, produces strong measures of support, particularly for deep internodes of Basidiomycota, which have been difficult to resolve with confidence using nrDNA data alone. This study produces strong boostrap support and significant posterior probabilities for the first time for the following monophyletic groups: (1) Ustilaginomycetes plus Hymenomycetes, (2) an inclusive cluster of hymenochaetoid, corticioid, polyporoid, Thelephorales, russuloid, athelioid, Boletales, and euagarics clades, (3) Thelephorales plus the polyporoid clade, (4) the polyporoid clade, and (5) the cantharelloid clade. Strong support is also recovered for the basal position of the Dacrymycetales in the Hymenomycetidae and paraphyly of the Exobasidiomycetidae.

Major clades of Agaricales: a multilocus phylogenetic overview.

An overview of the phylogeny of the Agaricales is presented based on a multilocus analysis of a six-gene region supermatrix. Bayesian analyses of 5611 nucleotide characters of rpb1, rpb1-intron 2, rpb2 and 18S, 25S, and 5.8S ribosomal RNA genes recovered six major clades, which are recognized informally and labeled the Agaricoid, Tricholomatoid, Marasmioid, Pluteoid, Hygrophoroid and Plicaturopsidoid clades. Each clade is discussed in terms of key morphological and ecological traits. At least 11 origins of the ectomycorrhizal habit appear to have evolved in the Agaricales, with possibly as many as nine origins in the Agaricoid plus Tricholomatoid clade alone. A family-based phylogenetic classification is sketched for the Agaricales, in which 30 families, four unplaced tribes and two informally named clades are recognized.

Rhodocybe paurii, a new species from the Indian Himalaya.

A new species of Entolomataceae, Rhodocybe paurii, is described from Garhwal in the western Indian Himalaya. This species grows on wood in dense clusters and belongs to section Claudopodes Singer ex Baroni because of its pleurotoid habit and lack of hymenial pseudocystidia. It is distinguished from the other pleurotoid species in that section by its layered caespitose habit, a brown spore deposit and a tomentose pileus surface composed of a well-developed layer of hyaline, erect, filamentous hyphae. Phylogenetic analysis using nucleotide sequence data from the nuclear large ribosomal subunit gene indicates a close relationship between R. paurii and the type species of the genus, Rhodocybe caelata. This analysis also suggests a possible paraphyly of the genus Rhodocybe and supports monophyly of Entoloma sensu lato.

Scleroderma stellatum versus Scleroderma bermudense: the status of Scleroderma echinatum and the first record of Veligaster nitidum from the Virgin Islands.

The type of Scleroderma stellatum from Brazil exhibits a sharp echinulate, dark brown peridium, and the type of S. bermudense from Bermuda has a peridium that is loosely woven and fibrillose, whitish to pale brownish. These characters indicate two independent species. This information is contrary to that of Guzmán in 1970, who interpreted S. bermudense to be a synonym of S. stellatum based on the similar spores. Scleroderma echinatum from Borneo and Panama, as recently discussed by Guzmán and Ovrebo, also has an echinulate, dark brown peridium and is a synonym of S. stellatum. All these fungi have a stellate dehiscence. New records of S. bermudense from the Greater Antilles and Mexico's Pacific Coast, and Veligaster nitidum from Virgin Islands also are discussed.

New species of Psilocybe in the Caribbean, with an emendation of P. guilartensis.

Five new species of Psilocybe from the Caribbean are described: P. caribaea, P. egonii, P. subpsilocybioides, P. zapotecoantillarum and P. zapotecocaribaea. All except P. zapotecocaribaea, which is known only from Martinique, are native to Puerto Rico. Psilocybe guilartensis, previously described from Puerto Rico, is the most commonly collected species of Psilocybe in Puerto Rico. New information on morphology is provided for P. guilartensis, and an emendation of the species circumscription is presented.

One hundred and seventeen clades of euagarics.

This study provides a first broad systematic treatment of the euagarics as they have recently emerged in phylogenetic systematics. The sample consists of 877 homobasidiomycete taxa and includes approximately one tenth (ca. 700 species) of the known number of species of gilled mushrooms that were traditionally classified in the order Agaricales. About 1000 nucleotide sequences at the 5(') end of the nuclear large ribosomal subunit gene (nLSU) were produced for each taxon. Phylogenetic analyses of nucleotide sequence data employed unequally weighted parsimony and bootstrap methods. Clades revealed by the analyses support the recognition of eight major groups of homobasidiomycetes that cut across traditional lines of classification, in agreement with other recent phylogenetic studies. Gilled fungi comprise the majority of species in the euagarics clade. However, the recognition of a monophyletic euagarics results in the exclusion from the clade of several groups of gilled fungi that have been traditionally classified in the Agaricales and necessitates the inclusion of several clavaroid, poroid, secotioid, gasteroid, and reduced forms that were traditionally classified in other basidiomycete orders. A total of 117 monophyletic groups (clades) of euagarics can be recognized on the basis on nLSU phylogeny. Though many clades correspond to traditional taxonomic groups, many do not. Newly discovered phylogenetic affinities include for instance relationships of the true puffballs (Lycoperdales) with Agaricaceae, of Panellus and the poroid fungi Dictyopanus and Favolaschia with Mycena, and of the reduced fungus Caripia with Gymnopus. Several clades are best supported by ecological, biochemical, or trophic habits rather than by morphological similarities.