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ABSTRACTSuillus(order Boletales) is a diverse genus of epigeous, mushroom-forming fungi native to temperate forests across the Northern Hemisphere; however, some species are also present in areas where Pinaceae has been introduced in the Southern Hemisphere. Unlike the closely related genus Rhizopogon, there are no described hypogeous, sequestrate species of Suillus. Here, we describe Suillus hypogaeus, the first known species of the genus with hypogeous, sequestrate sporocarps. Collections were made on Marys Peak in Benton County, Oregon, USA, at an elevation of 800 m in forests dominated by Pseudotsuga menziesiivar. menziesii. The peridium is white, quickly staining pink to purple-reddish where bruised or cut. The gleba is pale yellow when young, becoming purple with maturity, and the basidiospores are obovoid, light yellow in KOH, and amyloid in Melzer’s reagent. Multilocus molecular phylogenetic analyses support the placement of S. hypogaeusamong the Larixspecialists in the spectabilis group of Suillus. Although Larixand Pseudotsugaare sister genera, Larixdoes not occur on Marys Peak or elsewhere in western Oregon. Suillus hypogaeus, therefore, represents both an independent origin of the hypogeous, sequestrate sporocarp within the Boletales and an independent host shift between Larixand Pseudotsugawithin the genus Suillus.

ABSTRACTIn the northwestern Himalayan mountains of India, the hypogeous sequestrate fungus Trappeindia himalayensisis harvested from forests dominated by the ectomycorrhizal tree Cedrus deodara(Himalayan cedar). This truffle has basidiospores that are ornamented with raised reticulation. The original description of Trappeindia himalayensissuggested that the gleba of this species is similar to young specimens of Scleroderma(Boletales), whereas its basidiospores are ornamented with raised reticulation, suggesting a morphological affinity to Leucogaster(Russulales) or Strobilomyces(Boletales). Given this systematic ambiguity, we have generated DNA sequence data from type material and other herbarium specimens and present the first molecular phylogenetic analysis of this unusual Cedrus-associated truffle. Despite the irregular ornamented basidiospore morphology, T. himalayensisis resolved within the genus Rhizopogon(Suillineae, Boletales) and represents a unique lineage that has not been previously detected. All known Rhizopogonspecies possess an ectomycorrhizal trophic mode, and because of its placement in this lineage, it is likely that Trappeindia himalayensisis an ectomycorrhizal partner of Cedrus deodara. This study highlights the importance of generating sequence data from herbarium specimens in order to identify fungal biodiversity and clarify the systematic relationships of poorly documented fungi.

ABSTRACTA new genus and eight new species, all with isaria-like phialides, are described in Cordycipitaceae from Thailand. The new genus, Samsoniella, is segregated from Akanthomycesbased on morphological and molecular evidence. Samsonielladiffers from Akanthomycesin producing orange cylindrical to clavate stromata with superficial perithecia and orange conidiophores with isaria-like phialides and white to cream conidia. A new combination for CBS 240.32, originally identified as Paecilomyces farinosus(Isaria farinosa), and CBS 262.58, originally identified as Penicillium alboaurantium, respectively, is made in Samsoniella. Two new species, Samsoniella aurantiaand S. inthanonensis, are described from lepidopteran larvae. Two new species of Cordyceps, C. blackwelliaeand C. lepidopterorum, were also found on coleopteran and lepidopteran larvae. Both produce isaria-like morphs with globose phialides and attenuated long necks and white mycelium in culture. The authors established a sexual-asexual link for Cordyceps javanica(= Isaria javanica) on lepidopteran larvae. Four new species, Akanthomyces kanyawimiae, A. sulphureus, A. thailandicus, and A. waltergamsii, were pathogenic on spiders, with some strains of A. kanyawimiaealso found on unidentified insect larvae. These four species of Akanthomycesoccur on the underside of leaves and produce white to cream white powdery conidia, whereas S. aurantiaand S. inthanonensiswere found in leaf litter and produce bright orange stromata and synnemata with white conidia. Another new combination, Akanthomyces ryukyuensis, is proposed. Phylogenetic analyses based on a combined data set comprising the nuc rDNA region encompassing the internal transcribed spacers 1 and 2 along with the 5.8S rDNA (ITS), nuc 28S rDNA (28S), partial sequences of translation elongation factor 1-α gene (TEF1), and the genes for RNA polymerase II largest (RPB1) and second-largest (RPB2) subunits strongly support the delimitation of these new species of Cordyceps, Akanthomyces, and in a new genus Samsoniellain Cordycipitaceae.

ABSTRACTHost specialization is common among parasitic fungi; however, there are examples when transitions in host specificity between disparately related hosts have occurred. Here, we examine the interkingdom host jump from insect pathogenicity and mycoparasitism in Tolypocladium. Previous phylogenetic inferences made using only a few genes and with poor support reconstructed an ancestral character state of insect pathogenesis, a transition to mycoparasitism, and reversions to insect pathogenesis. To further explore the directionality and genes underlying the transitions in host, we sequenced two additional species of Tolypocladium(T. capitatumand T. paradoxum) and used phylogenomics to compare two insect pathogens and two mycoparasites. Our whole-genome-scale analysis suggests that the diversification of Tolypocladiumspecies happened relatively quickly and that the truffle parasites form a monophyletic, derived lineage within the genus that is the result of a single ecological transition or host jump from insects to fungi. A significant amount of gene tree/species tree discordance occurs within the data set, and we infer this to be the product of both an historical hybridization event and incomplete lineage sorting that was likely because of the rapid diversification of the clade. Furthermore, comparative genomic analyses revealed a set of genes that are exclusive to the mycoparasitic species. These potentially mycoparasitic gene clusters were characterized by a reduced proportion of secreted proteins when compared with entomopathogen-enriched genes and involved the reshaping of the fungal secretome in the ecological context of mycoparasitism.

ABSTRACTThe end of the 20thcentury and the beginning of the 21stwill be remembered as the golden age of molecular systematics of Fungi. The development of the Polymerase Chain Reaction (PCR), automated sequencing, and explicit algorithms for inferring phylogenetic trees transformed fungal systematics into a hypothesis-testing discipline that would go on to serve as the foundation for comparative and evolutionary genomics. Many mycologists participated in this modernization of mycology and perhaps none more than Meredith Blackwell. She has played – and continues to play – a leadership role in transforming mycology into a modern science based on a robust understanding of organismal biology, development, testing and refinement of biologically meaningful hypotheses, and incorporation of emerging technologies to data collection and analysis. In this volume we recognize Meredith and her contributions to mycology with the publication of the first Festschrift issue in the 110 year history of Mycologia.

In the Amazon the only described species of Cordycepssensu stricto (Hypocreales, Cordycipitaceae) that parasitize insects of Orthopterida (orders Orthoptera and Phasmida) are Cordyceps locustiphilaand C. uleana. However, the type specimens for both taxa have been lost and the concepts of these species are uncertain. To achieve a more comprehensive understanding of the systematics of these species, collections of Cordycepsfrom the Amazon regions of Colombia, Ecuador and Guyana were subjected to morphological, ecological and molecular phylogenetic studies. Phylogenetic analyses were conducted on partial sequences of SSU, LSU, TEF, RPB1and RPB2nuclear loci. Two new species are proposed including C. diapheromeriphila, a parasite of Phasmida, and C. acridophila, a parasite of the superfamily Acridomorpha (Orthoptera), which is broadly distributed across the Amazon. For C. locustiphilaa lectotypification and an epitypification are made. Cordyceps locustiphilais host specific with Colpolopha(Acridomorpha: Romaleidae), and its distribution coincides with that of its host. The phylogenetic placement of these three species was resolved with strong support in the Beauveriaclade of Cordycepss. str. (Cordycipitaceae). This relationship and the morphological similarity of their yellow stromata with known teleomorphs of the clade, suggest that the holomorphs of these species may include Beauveriaor Beauveria-like anamorphs. The varying host specificity of the beauverioid Cordycepsspecies suggest the potential importance of identifying the natural host taxon before future consideration of strains for use in biological control of pest locusts.

Rhizopogonsubgenus Villosuliare the only members of the genus known to form an ectomycorrhizal relationship exclusively with Pseudotsuga. The specificity of this host relationship is unusual in that Rhizopogonis broadly associated with several tree genera within the Pinaceae and relationships with a host genus are typically distributed across Rhizopogonsubgenera. Naturally occurring specimens of R. subg. Villosulihave been described only from North American collections, and the unique host relationship with Pseudotsugais demonstrated only for Rhizopogonassociated with P. menziesii(Douglas-fir), the dominant species of Pseudotsugain North America. Species of Pseudotsugaare naturally distributed around the northern Pacific Rim, and Rhizopogonassociates of other Pseudotsugaspp. are not yet described. Here we present the results of field sampling conducted in P. japonicaforests throughout the Japanese archipelago and describe Rhizopogon togasawarianasp. nov., which occurs in ectomycorrhizal association with P. japonica. Placement of this new species within R. subg. Villosuliis supported by morphological and molecular phylogenetic analysis, and its implications to Pseudotsuga-Rhizopogonbiogeography are discussed.

Using spatial autocorrelation analysis, we examined the within-population genetic structure of Rhizopogon vinicolorand R. vesiculosus, two hypogeous ectomycorrhizal (EM) species that are sympatric sister taxa known to differ in their clonal structure. We collected 121 sporocarps and 482 tuberculate EM of both species from a 20 ha forest stand dominated by Douglas-fir (Pseudotsuga menziesii). Field collections were identified to species with restriction fragment length polymorphism analysis of the nuclear ribosomal internal transcribed spacer. Five and six microsatellite markers were used to characterize the genetic diversity of EM and sporocarp samples from R. vesiculosusand R. vinicolorrespectively. After correcting for genet structure, spatial autocorrelation analyses of the EM samples were used to test the null hypothesis that multilocus genotypes characterized from each species were randomly distributed within the study area. We detected positive and statistically significant fine-scale genetic structure up to 120 m within the R. vesiculosussample. In contrast, no spatial genetic structure was evident for R. vinicolor, indicating that the genotypes characterized for this species were randomly distributed throughout the study area. Differences in statistical power or the nuclear count of basidiospores are unlikely agents of the genetic patterns observed. Our results suggest that differences in reproductive output or competitive ability may act individually or in combination to create clusters of similar genotypes for R. vesiculosusthroughout the study area.

Cenococcumis a genus of ectomycorrhizal Ascomycota that has a broad host range and geographic distribution. It is not known to produce either meiotic or mitotic spores and is known to exist only in the form of hyphae, sclerotia and host-colonized ectomycorrhizal root tips. Due to its lack of sexual and asexual spores and reproductive structures, it has proven difficult to incorporate into traditional classification within Ascomycota. Molecular phylogenetic studies of ribosomal RNA placed Cenococcumin Dothideomycetes, but the definitive identification of closely related taxa remained elusive. Here we report a phylogenetic analysis of five nuclear loci (SSU, LSU, TEF1, RPB1, RPB2) of Dothideomycetes that placed Cenococcumas a close relative of the genus Gloniumof Gloniaceae (Pleosporomycetidae incertae sedis) with strong statistical support. Gloniumis a genus of saprobic Dothideomycetes that produces darkly pigmented, carbonaceous, hysteriate apothecia and is not known to be biotrophic. Evolution of ectomycorhizae, Cenococcumand Dothideomycetes is discussed.

Molecular phylogenetic analyses indicate that the monophyletic classes Orbiliomycetes and Pezizomycetes are among the earliest diverging branches of Pezizomycotina, the largest subphylum of the Ascomycota. Although Orbiliomycetes is resolved as the most basal lineage in some analyses, molecular support for the node resolving the relationships between the two classes is low and topologies are unstable. We provide ultrastructural evidence to inform the placement of Orbiliomycetes by studying an Orbilia, a member of the only order (Orbiliales) of the class. The truncate ascus apex in the Orbiliais thin-walled except at the margin, and an irregular wall rupture of the apex permits ascospore discharge. Ascus, ascogenous and non-ascogenous hyphae were simple septate, with septal pores plugged by unelaborated electron-dense, non-membranous occlusions. Globose Woronin bodies were located on both sides of the septum. Nuclear division was characterized by the retention of an intact nuclear envelope, and a two-layered disk-shaped spindle pole body. The less differentiated nature of the spore discharge apparatus and septal pore organization supports an earliest diverging position of Orbiliomycetes within the subphylum, while the closed nuclear division and disk-shaped spindle pole body are interpreted as ancestral state characters for Ascomycota.

Research in fungal phylogenetics and systematics progressed rapidly in the past decade due to advances in DNA sequencing technologies and analytical methods. A newfound wealth of sequence data acquired through community-wide initiatives has advanced the process of acquiring a stable phylogenetic classification of many fungal taxa. Financial support from the National Science Foundation Research Coordination Networks: a phylogeny for kingdom Fungi (Deep Hypha) for 5 y enabled more than 100 fungal systematists to assess the taxon sampling, molecular markers and analytical methods necessary to facilitate such a project. Later a second NSF program provided financial support for the Assembling the Fungal Tree of Life (AFTOL) project to accomplish much of the research. Deep Hypha may be viewed as an involved parent of AFTOL with a continuing role as coordinator of likeminded workers. Many questions posed at the beginning of the Deep Hypha project have been addressed, at least in part, although some details remain to be clarified. Many of the main branches of the fungal tree are stable and well supported, often as a result of multigene analyses that involved collaboration of many laboratories. More work is necessary, however, to resolve certain branching events near the base of the tree, as well as to reconstruct relationships in some terminal groups. The phylogenetic classification in this issue of Mycologiais a product of the AFTOL project and many other independent research initiatives, and it is an initial synthesis of a working classification designed to be used for all majorPUBLICations that require a phylogenetic classification of fungi.

Cordyceps cardinalis,a pathogen of lepidopteran larva, is described as a new species from the southern Appalachian Mountains of the eastern United States and from southeastern Japan. It is macroscopically similar to both C. militarisand C. pseudomilitaris,however, microscopic, cultural and molecular evidence support it being a phylogenetically distinct species. Cordyceps cardinalisis most similar to C. pseudomilitarisin the microscopic characters of non-disarticulating ascospore morphology and its host affinity for lepidopteran larvae, which contrast with the characters of C. militaris.Aspects of morphology, host association, phylogeny and biogeography are discussed.

We are re-addressing species concepts in the Rhizopogon vinicolorspecies complex (Boletales, Basidiomycota) using sequence data from the internal-transcribed spacer (ITS) region of the nuclear ribosomal repeat, as well as genotypic data from five microsatellite loci. The R. vinicolorspecies complex by our definition includes, but is not limited to, collections referred to as R. vinicolorSmith, R. diabolicusSmith, R. ochraceisporusSmith, R. parvulusSmith or R. vesiculosusSmith. Holo- and/or paratype material for the named species is included. Analyses of both ITS sequences and microsatellite loci separate collections of the R. vinicolorspecies complex into two distinct clades or clusters, suggestive of two biological species that subsequently are referred to as R. vinicolorsensu Kretzer et al and R. vesiculosussensu Kretzer et al. Choice of the latter names, as well as morphological characters, are discussed.

Rhizopogon(Basidiomycota, Boletales) is a genus of hypogeous fungi that form ectomycorrhizal associations mostly with members of the Pinaceae. This genus comprises an estimated 100+species, with the greatest diversity found in coniferous forests of the Pacific northwestern United States. Maximum parsimony analyses of 54 nuclear ribosomal DNA internal transcribed spacer (ITS) sequences including 27 Rhizopogonand 10 Suillusspecies were conducted to test sectional relationships in Rhizopogonand examine phylogenetic relationships with the closely related epigeous genus, Suillus. Sequences from 10 Rhizopogontype collections were included in these analyses. Rhizopogonand Suilluswere both monophyletic. Rhizopogonsection Rhizopogonis not monophyletic and comprised two clades, one of which consisted of two well supported lineages characterized by several long insertions. Rhizopogonsections Amylopogonand Villosuliformed well supported groups, but certain species concepts within these sections were unresolved. Four species from section Fulviglebaeformed a strongly supported clade within section Villosuli. Subgeneric taxonomic revisions are presented.

AbstractPhylogenetic relationships of the genus Ramariaand additional related taxa were examined through phylogenetic analysis of mitochondrial and nuclear ribosomal DNA sequence data. Related genera included Clathrus, Clavariadelphus, Gautieria, Gomphus, Hysterangium, Kaviniaand Pseudocolus.Outgroup genera included Bondarzewia, Favolus, Ganoderma, Lactariusand Russula.The nuclear large subunit ribosomal DNA (nuc LSU rDNA) (approx. 610 bp) from 78 collections including 34 Ramariaspecies was used to test generic, subgeneric and selected species concepts for Ramaria.The mitochondrial small subunit ribosomal DNA (mt SSU rDNA) (ca 470 bp) from 51 representative holobasidiomycetes including ten Ramariaspecies also was used to independently test the monophyly of the genus Ramaria.Parsimony analyses of both datasets indicated that the genus Ramariawas paraphyletic and that several morphologically distinct groups of holobasidiomycetes were derived from ramarioid ancestors. In the nuc LSU rDNA analyses, Gautieriawas nested among the terricolous Ramariaspecies and was closely related to R.subg Ramaria.The nuc LSU rDNA results also indicated that R.subg Laeticoloraand R.subg Lentoramariaformed paraphyletic grades. Ramariasubg Laeticolorawas paraphyletic due to the terminally derived R.subg Ramaria/Gautieriaclade and a nested Gomphus. Ramariasubg Lentoramariawas paraphyletic due to a nested Clavariadelphus, Kavinia, and R. abietinaof subg Echinoramaria.Both the nuc LSU rDNA and the mt SSU rDNA analyses demonstrated strong bootstrap support for the inclusion of the Phallales with the ingroup taxa, but the relationship of the Phallales to the Gomphales was indeterminate. Selected species concepts were also tested (e.g., R. amyloideaand R. celerivirescens, and R. araiosporaand R. stuntzii). These data rejected the monophyly of Ramariaand suggested that the ramarioid morphology and lignicolous substrate affinity were ancestral for the Gomphales.

AbstractA new species and genus of the gramini-colous Clavicipitaceae are described. Neoclaviceps monostipainfects individual florets in the same way as species of genus Clavicepsbut does not produce sclerotia. In culture it shows dimorphism in production of a yeast-like microcyclic conidiation (ephelidial) phase and a mycelial phase. Phylogenetic analysis of the 26S large subunit rDNA suggests that Neoclavicepsalong with genus Myriogenosporaare evolutionary intermediate between Clavicepsand Balansia.

AbstractThe phylogenetic relationship between Alpova, Chroogomphus, Gomphidius, Rhizopogon, Suillus, and Truncocolumellaand their placement in the Boletales was tested through maximum parsimony analyses of large subunit nuclear ribosomal DNA sequences. Taxon sampling included representatives of the genera Alpova, Boletellus, Boletus, Chroogomphus, Gomphidius, Melanogaster, Paragyrodon, Phylloporus, Rhizopogon, Suillus, Truncocolumella, Tylopilus, and Xerocomus, as well as species from the Agaricales, Russulales, and Polyporales. The order Boletales was strongly supported as monophyletic. Furthermore, within this order, two major groups emerged: the boletoid radiation and a Melanogastraceae/Paragyrodon sphaerosporusparaphyletic assemblage, and the suilloid radiation. The boletoid radiation and a Melanogastraceae/Paragyrodon sphaerosporusassemblage was strongly supported. Although the boletoid radiation was not strongly supported by bootstrap analysis, the clade did not collapse in the strict consensus topology. This clade comprised species from the genera Boletellus, Boletus, Phylloporus, Tylopilus, and Xerocomus.The Boletaceae sensu Singer was not monophyletic. Although species relationships within the boletoid radiation were not well resolved, the genus Boletusdid not appear to be monophyletic. Alpova diplophloeus, A. trappei, Melanogaster tuberiformis, and Paragyrodon sphaerosporusformed a paraphyletic assemblage basal to the boletoid radiation clade. The suilloid clade was strongly supported and included Alpova olivaceotinctus, Chroogomphus, Gomphidius, Rhizopogon, Suillusand Truncocolumella citrina.Species in the genus Alpovatherefore were grouped both within and outside the suilloid radiation. Within the suilloid radiation, Suilluswas inferred to be more closely related to Truncocolumella citrinaand the Gomphidiaceae than to Rhizopogon, suggesting a more distant relationship between Suillusand Rhizopogonthan previously hypothesized. Rhizopogonwas not supported as monophyletic, although its monophyly could not be rejected by the data. Alpova olivaceotinctusgrouped within the genus Rhizopogon, consistent with previous classifications, and is returned to that genus.

AbstractThe nonsexual lichenized family Siphulaceae is comprised of the obligately sterile genera, Siphulaand Thamnolia.These genera are currently placed in the order Lecanorales, apparently based solely on the character of lichenization since apothecia and other lichen reproductive structures are absent. Using the nucleotide characters of rDNA, we tested the phylogenetic position of these genera among the Ascomycota. Cladistic analyses of SSU nrDNA and partial LSU nrDNA did not support the recognition of the Siphulaceae. Rather, Siphulaand Thamnoliarepresent independent origins of nonsexual lichens among the Icmadophilaceae; a family that is currently classified in the order Helotiales. In addition to these molecular characters, the secondary chemistry of these genera also supports their affinity with the Icmadophilaceae. We propose taxonomically emending the Icmadophilaceae to include the genera Siphulaand Thamnolia.The phylogenetic relationships between the Icmadophilaceae and the Baeomycetaceae, the other family of lichenized fungi classified in the Helotiales, are discussed. Cladistic evidence is provided which demonstrates that the Baeomycetaceae is a distinct taxon from the Icmadophilaceae and the two families represent independent lichenization events. The phylogenetic relationships between these families and other higher taxa (e.g., families and orders) of the Ascomycota are also discussed.

AbstractBased on molecular studies using 1031 bp of the nuclear small subunit (SSU) gene and 594 bp of the nuclear large subunit (LSU) rDNA gene and using morphological characters, the genera Lulworthiaand Lindraare removed from Halosphaeriales and assigned to the new order Lulworthiales and new family Lulworthiaceae in the class Sordariomycetes. Eleven accepted species of Lulworthiaand 6 species of Lindraare listed.

AbstractThe termite pathogen Cordycepioideus bisporusis characterized by perithecia produced in a Cordyceps-like stroma, ascospores that are dark, thick-walled and 8–15 septate at maturity, and a Hirsutellaanamorph. The genus traditionally has been placed in the Hypocreales on the basis of purported apical paraphyses and immature ascospore features; however, phylogenetic analyses of partial sequences of the nuclear small and large subunit ribosomal DNA showed a close relationship between C. bisporusand clavicipitaceous ascomycetes, which is in agreement with a newer understanding of morphological characters of this species. Hirsutella thompsoniiwas the sister taxon to C. bisporus, and several insect parasitic species of Cordycepsalso showed close relationships to C. bisporusand H. thompsonii.The results of this study suggest strongly that the genus Cordycepioideusbelongs within the Clavicipitaceae, not with other hypocrealean species where the genus originally was placed.