Actualización del conocimiento del género Cyttaria Berk. (Cyttariales, Ascomycota) en Chile / Update of the knowledge of the genus Cyttaria Berk (Cyttariales, Ascomycota) in Chile

El género Cyttaria pertenece a la familia Cyttariaceae y sus especies son parásitos obligados de árboles del género Nothofagus. Se distribuye naturalmente en el hemisferio sur, encontrándose siete especies de Cyttaria presentes en nuestro país: C. berteroi, C. darwinii, C. espinosae, C. hariotii, C. hookeri, C. johowii y C. exigua, que comúnmente se conocen como "Digüeñes" y que tienen una gran importancia desde el punto de vista alimenticio, ya que han sido consumidos desde la prehistoria por pueblos originarios. En Chile se han realizado estudios sobre la taxonomía, ecología y propiedades medicinales de algunas especies del género Cyttaria y en esta revisión, se describen los caracteres macro y microscópicos e información relevante de las siete especies de este género presentes en nuestro país, de acuerdo a la literatura disponible y observaciones personales de la autora. Además, se presenta un resumen sobre los resultados de un estudio reciente de las propiedades bioactivas de las especies más consumidas en nuestro país. (AU)

The genus Cyttariabelongs to the family Cyttariaceae; its species are obligate parasites of trees of the genus Nothofagus. It is naturally distributed in the Southern Hemisphere, with seven species of Cyttariapresent in Chile: C. berteroi, C. darwinii, C. espinosae, C. hariotii, C. hookeri, C. johowiiand C. exigua, which are commonly known as "Digüeñes" and are of great nutritional importance, since they have been consumed from prehistory by native people. Studies have been carried out in Chile on the taxonomy, ecology and medicinal properties of some species of the genus Cyttaria. In this review, the macro and microscopic characteristics and relevant information of the seven species of this genus present in our country are described, according to the available literature and personal observations of the author. A summary of the results of a recent study of the bioactive properties of the most consumed species in our country is also presented. (AU)

Hymenogaster macmurphyi and Splanchnomyces behrii are sequestrate species of Xerocomellus from the western United States.

Hymenogaster is an ectomycorrhizal genus of brown-spored sequestrate fungi that is related to the mushroom-forming genera Hebeloma and Alnicola (Agaricales). However, because of difficulties in morphological taxonomy of sequestrate fungi, Hymenogaster has become a polyphyletic repository for a variety of unrelated brown-spored sequestrate species. During studies of ectomycorrhizal ecology and sequestrate fungal evolution in the western USA, we encountered specimens of a morphologically unique species. It was originally described as Hymenogaster macmurphyi, but our morphological and molecular analyses indicate that it is not closely related to Hymenogaster. Phylogenetic analyses of multiple gene regions indicate that H. macmurphyi is actually a member of the Boletineae (Boletales, Basidiomycota) and is nested within the epigeous genus Xerocomellus, distantly related to any of the other known genera of sequestrate Boletales. While examining additional herbarium collections, we came upon isotype material of Splanchnomyces behrii, which represents a closely related species. Here we document the morphology and phylogenetic affinities of these unusual sequestrate Boletineae and transfer both species to Xerocomellus as X. macmurphyi and X. behrii. During our study, we also noted that the sequestrate taxon Rhopalogaster transversarius is nested within the epigeous genus Suillus.

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

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

Phylogenetic assessment of Chromocyphellaceae (Agaricineae, Basidiomycota) and a new lamellate species of Chromocyphella.

Cyphelloid fungi represent a polyphyletic assemblage of reduced agarics, including the brown-spored family Chromocyphellaceae. In order to elucidate the phylogenetic position of Chromocyphellaceae, newly generated sequences of Chromocyphella were included in a multigene alignment of the Agaricineae and phylogenetically analyzed. The current analyses show that the Chromocyphella muscicola specimen used to phylogenetically place Chromocyphellaceae in its original description was misidentified and that the Chromocyphellaceae nests in the Hymenogastraceae, Chromocyphella being sister to Flammula. Chromocyphella is emended, including now a new species with lamellate and stipitate basidiomata, C. lamellata. The name Cymbella crouanii, type species of Chromocyphella, is lecto- and epitypified. Our analyses support a new origin of cyphelloid fungi. The shift to a cyphelloid basidioma from an agaric ancestor is suggested to have occurred in two evolutionary steps in Chromocyphella, an initial reduction in basidioma size and a subsequent loss of lamellae and stipe.

Revisión del género Macrolepiota Singer (Agaricales, Basidiomycota) en Chile / Revision of the genus Macrolepiota Singer (Agaricales, Basidiomycota) in Chile

El género Macrolepiota cuenta con 30 especies ampliamente distribuidas en el mundo, siendo éstas saprobiontes. En Chile se han documentado tres especies: M. procera, M. bonaerensis y M. rachodes, perteneciendo actualmente esta última al género Chlorophyllum. En esta revisión, se describen según la literatura los caracteres macromorfológicos y microscópicos de las tres especies mencionadas con anterioridad. Adicionalmente, se adaptó una clave dicotómica en base a la bibliografía para facilitar la determinación de las especies.

The Macrolepiota genus include 30 species widely distributed around the world, being these saprobionts. In Chile, three species have been documented: M. procera, M. bonaerensis y M. rachodes, at present the last specie currently belong to the genus Chlorophyllum. In this review, the macromorphological, and microscopic characteristics of the three species mentioned above are described according to the literature. In addition, a dichotomous key was adapted based on the bibliography to facilitate the determination of the species.

Typification and Characterization of Trametes multicolor (Agaricomycetes), a Perspective Species of Medicinal Mushrooms.

Nomenclature revision and enlarged taxonomical descriptions are still needed for some well-known species whose interpretation is complicated by many nomenclature or taxonomical problems. The polyporoid fungus widely known as Trametes ochracea (= Coriolus zonatus) belongs to such a problematic group. At the same time, recent data show that this species, like its sister species T. versicolor, seems to be a perspective subject for fungal biotechnology and pharmacology. This article is devoted to stabilizing the nomenclature of the species in question via lectotypification and epitypification of Boletus multicolor. It will clarify the name T. multicolor as applied to this species is nomenclaturally correct and useful, free of further problems. An expanded species description and cultural characterization of epitype materials are presented.

Nuclear populations of the multinucleate fungus of leafcutter ants can be dekaryotized and recombined to manipulate growth of nutritive hyphal nodules harvested by the ants.

We dekaryotized the multinucleate fungus Leucocoprinus gongylophorus, a symbiotic fungus cultivated vegetatively by leafcutter ants as their food. To track genetic changes resulting from dekaryotization (elimination of some nuclei from the multinuclear population), we developed two multiplex microsatellite fingerprinting panels (15 loci total), then characterized the allele profiles of 129 accessions generated by dekaryotization treatment. Genotype profiles of the 129 accessions confirmed allele loss expected by dekaryotization of the multinucleate fungus. We found no evidence for haploid and single-nucleus strains among the 129 accessions. Microscopy of fluorescently stained dekaryotized accessions revealed great variation in nuclei number between cells of the same vegetative mycelium, with cells containing typically between 3 and 15 nuclei/cell (average = 9.4 nuclei/cell; mode = 8). We distinguish four mycelial morphotypes among the dekaryotized accessions; some of these morphotypes had lost the full competence to produce gongylidia (nutritive hyphal-tip swellings consumed by leafcutter ants as food). In mycelial growth confrontations between different gongylidia-incompetent accessions, allele profiles suggest exchange of nuclei between dekaryotized accessions, restoring full gongylidia competence in some of these strains. The restoration of gongylidia competence after genetic exchange between dekaryotized strains suggests the hypothesis that complementary nuclei interact, or nuclear and cytoplasmic factors interact, to promote or enable gongylidia competence.

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

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

Sulfite determination by an inhibitor biosensor-based mushroom (Agaricus bisporus) tissue homogenate.

The aim of the study presented here is to develop a biosensor based on mushroom (Agaricus bisporus) tissue homogenate for sensitive and economical determination of sulfite in foods. The working principle of the biosensor is based on an inhibition effect of sulfite on polyphenol oxidases in mushroom. Mushroom tissue homogenate was immobilized by gelatin and glutaraldehyde on a Clark-type oxygen electrode. Some optimization studies related to the bioactive layer components and working conditions were identified. The biosensor was applied to the food samples. The biosensor reported here was successfully allowed to analyze sulfite, which was a food additive in real food samples.

Rhizomarasmius epidryas (Physalacriaceae): phylogenetic placement of an arctic-alpine fungus with obligate saprobic affinity to Dryas spp.

As a part of a large-scale biogeographical study we examined the evolutionary relationships and taxonomic position of Marasmius epidryas, one of the most typical circumpolar arctic-alpine fungi, characterized by a specific, saprobic affinity to dead tissues of Dryas spp. A phylogenetic analysis based on nLSU and RPB2 DNA regions unequivocally indicated the phylogenetic placement of this species within the Physalacriaceae. The Bayesian MCMCMC analysis as well as other inference methods tested (ML, NJ) revealed a well supported affinity of M. epidryas to Rhizomarasmius pyrrhocephalus, type species of a recently circumscribed genus, Rhizomarasmius. As a consequence, based on these results, we introduce a new combination, Rhizomarasmius epidryas (Kühner ex A. Ronikier) A. Ronikier and M. Ronikier. Thus our results demonstrate that neither the traditional taxonomic placement of the fungus in genus Marasmius nor the recent transfer into genus Mycetinis are phylogenetically correct. In contrast they support the importance of the third lineage of the polyphyletic Marasmius s. l., having evolutionary links with taxa forming the Physalacriaceae clade of agaricoid fungi. In addition the lineage of Rhizomarasmius was confirmed to be closely related to the representatives of Gloiocephala, comprising small, often narrowly specialized saprobic species previously also classified within Marasmius s. l.

Understanding the evolutionary processes of fungal fruiting bodies: correlated evolution and divergence times in the Psathyrellaceae.

Fruiting body evolution is one of the central topics in fungal evolutionary biology. A number of hypotheses have been developed to explain the contemporary diversity of fruiting body forms, but their evaluation has been hampered by the lack of well-sampled data sets and suitable statistical methods. Phylogenetic evidence of the physiological changes that accompany switches in fruiting body type is lacking, and very little is known about the age of major events of fruiting body evolution. Based on a new multigene phylogeny, by using Bayesian methods, we demonstrate the existence of correlation between a number of morphological features and switches from nondeliquescent to deliquescent (autodigesting) fruiting bodies in the mushroom family Psathyrellaceae. Our results show that switches in the anatomy of two types of spacer cells (cystidia and pseudoparaphyses) and basidia (bimorphic or monomorphic) as well as the structure of the mushroom cap follow the evolution of deliquescent fruiting bodies, which suggests strong functional linkage between these traits. We performed Bayes factor-based tests, referred hereafter to as evolutionary pathway test (EPT), to decide which of the correlated characters were gained first during evolution. The EPTs strongly suggest that deliquescence was gained first, followed after short waiting times by the other morphological features. Bayesian relaxed molecular clock analyses suggest that the various events of switching between fruiting body types occurred independently at various ages during the history of the family. The utility of two mushroom fossils (Archaemarasmius and Protomycena), the only ones with unambiguous taxonomic positions, for the calibration of agaric trees were also examined. Based on our results, we suggest that the evolutionary benefit of deliquescence may be prevention against desiccation via accelerated ontogeny of the fruiting body. Hypotheses regarding the functional significance of the correlated evolution are presented and discussed. Further, we argue that the changes in fruiting body types in mushrooms in general can be attributed to independent events (e.g., dispersal and adaptation) and not to particular geologic ages.

Taxonomy of Pluteus eugraptus and morphologically similar taxa.

The status of the taxa morphologically similar to Pluteus eugraptus (Basidiomycota, Agaricales) was investigated with morphological and molecular (ITS region) characters. This group of species belongs in Pluteus sect. Celluloderma based on morphological and molecular characters. Two species, Pluteus multiformis, from Spain and Pluteus eludens from Madeira, Portugal, Russia and USA, are described as new. Both species share pigmented cheilocystidia and a pileipellis composed of both clavate-spheropedunculate and elongated elements with P. eugraptus, but they can be separated based on the characteristics of the cystidia and pileipellis. Pluteus multiformis is characterized by the scarce pleurocystidia, clavate cheilocystidia and caulocystidia and highly polymorphic elements of the pileipellis. Pluteus eludens is characterized mainly by utriform pleurocystidia. Pluteus eugraptus is known only with certainty from the type collection (Sri Lanka), which has been re-examined here, and it is characterized by narrowly lageniform pleurocystidia. Phylogenetic analyses based on ITS region sequence data supported the separation of P. multiformis, P. eludens and an additional collection from Japan that likely represents the true P. eugraptus.

Lepiota in California: species with a hymeniform pileus covering.

Eight Lepiota species with a hymeniform pileus covering that are known in California are presented and discussed. Lepiota phaeoderma is described as new; this species is characterized by a dark pileus surface, which splits open into small patches around the umbo, and the absence of an annulus. Lepiota neophana, a species with an annulus and a closed brown pileus surface, and L. lilacea, with a conspicuous dark annulus, are both recorded for the first time west of the Rocky Mountains. The identity of L. neophana is reviewed and clarified based on morphological and molecular data. The type collection of L. rufipes was studied, and L. rufipes was placed in synonymy with Cystolepiota seminuda. A key to the species is given.

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 mushroom family Psathyrellaceae: evidence for large-scale polyphyly of the genus Psathyrella.

Psathyrella is the archetypal little brown mushroom genus with few easily discernable characters causing it to be considered a "clean-up" genus for other small brown-spored saprotrophic species found worldwide. While molecular studies have demonstrated that mushroom genera based on homoplastic morphological characters are artificial, the degree of phylogenetic heterogeneity contained within Psathyrella and Psathyrellaceae has never been appropriately addressed. For this study, 132 ribosomal sequences from approximately one-tenth of the known Psathyrella species worldwide, including representatives of most subgeneric subdivisions, and three closely related coprinoid genera (Parasola, Coprinopsis, Coprinellus) were evaluated using multiple phylogenetic methods, including likelihood, with Agaricaceae as the outgroup. Our results indicated that Psathyrella was polyphyletic. Conservatively, the genus can be separated into 11 clades of which five can be raised to generic status. Most species of Psathyrella, including its type species P. gracilis, formed a large clade with Coprinellus, which appeared to be derived from within Psathyrella. Generic limits of Parasola, Lacrymaria, and Coprinopsis should be reevaluated. Several taxa previously synonymized based on morphological features were phylogenetically distinct. Morphological features traditionally used to subdivide Psathyrella appeared to be mostly convergent (homoplasious) when traced upon the resulting phylogenies, although several had high RI values. These results were interpreted in light of the two major taxonomic treatments of Psathyrella and revealed substantial inconsistencies between the molecular- and morphology-derived inferences of relationships.

Wood-decay fungi in fine living roots of conifer seedlings.

The mycorrhizal basidiomycetes are known to have multiple, independent evolutionary origins from saprotrophic ancestors. To date, a number of studies have revealed functional resemblance of mycorrhizal fungi to free-living saprotrophs, but information on the ability of saprotrophic fungi to perform as mycorrhizal symbionts is scarce. Here, the objective was to investigate the ability of three wood-decay fungi, Phlebiopsis gigantea, Phlebia centrifuga and Hypholoma fasciculare, to colonize fine roots of conifer seedlings. For each fungus, mycorrhizal syntheses were attempted with Picea abies and Pinus sylvestris. After 24 wk, isolation of fungi and direct sequencing of fungal internal transcribed spacer (ITS) rDNA were carried out from healthy-looking surface-sterilized root tips that yielded both pure cultures and ITS sequences of each inoculated strain. Mycelial mantle of P. gigantea was frequently formed on root tips of P. abies, and microscopical examination has shown the presence of intercellular hyphae inside the roots. The results provide evidence of the ability of certain wood-decay fungi to colonise fine roots of tree seedlings.

Sumoylation of a meiosis-specific RecA homolog, Lim15/Dmc1, via interaction with the small ubiquitin-related modifier (SUMO)-conjugating enzyme Ubc9.

Sumoylation is a post-translational modification system that covalently attaches the small ubiquitin-related modifier (SUMO) to target proteins. Ubc9 is required as the E2-type enzyme for SUMO-1 conjugation to targets. Here, we show that Ubc9 interacts with the meiosis-specific RecA homolog, Lim15/Dmc1 in the basidiomycete Coprinus cinereus (CcLim15), and mediates sumoylation of CcLim15 during meiosis. In vitro protein-protein interaction assays revealed that CcUbc9 interacts with CcLim15 and binds to the C-terminus (amino acids 105-347) of CcLim15, which includes the ATPase domain. Immunocytochemistry demonstrates that CcUbc9 and CcLim15 colocalize in the nuclei from the leptotene stage to the early pachytene stage during meiotic prophase I. Coimmunoprecipitation experiments indicate that CcUbc9 interacts with CcLim15 in vivo during meiotic prophase I. Furthermore, we show that CcLim15 is a target protein of sumoylation both in vivo and in vitro, and identify the C-terminus (amino acids 105-347) of CcLim15 as the site of sumoylation in vitro. These results suggest that sumoylation is a candidate modulator of meiotic recombination via interaction between Ubc9 and Lim15/Dmc1.

Anamorphs of the Bolbitiaceae (Basidiomycota, Agaricales).

We describe and illustrate thallic conidiogenesis in 14 species of the Bolbitiaceae sensu Singer studied in culture. Conidiogenesis of 12 species is shown for the first time. Bolbitius vitellinus and the investigated species of Conocybe (C. albipes, C. appendiculata, C. magnicapitata, C. semiglobata, C. subovalis, C. subpubescens, C. sulcatipes and C. teneroides) possessed a similar mode of conidiogenesis. Species of both genera formed mostly coiled conidiogenous hyphae arising sympodially from differentiated conidiophores. The anamorphs of the Agrocybe species were not uniform and predominantly differed from those of Conocybe and Bolbitius. The conidia of Agrocybe dura, A. firma and A. praecox developed by the simple fragmentation of normally branched hyphae. Sympodially proliferating conidiophores occurred in Agrocybe arvalis and A. aegerita. Secretory cells of different size and shape were found in Agrocybe and in Conocybe. Our results corroborate a close phylogenetic relationship between Bolbitius and Conocybe as well as the polyphyly of the Bolbitiaceae as currently treated, which is consistent with recent molecular phylogenetic studies. Consequently we emend the family concept based on anamorphic characters.

Mycorrhizal synthesis of Lactarius indigo (Schw.) Fr. with five Neotropical pine species.

This paper describes for the first time the ectomycorrhiza synthesized between two Guatemalan strains of Lactarius indigo (Schw.) Fr. and the Neotropical species Pinus ayacahuite var. ayacahuite Ehren, P. hartwegii Lindl., P. oocarpa Schiede ex Schltdl. var oocarpa, P. pseudostrobus Lindl. and P. rudis Endl. The synthesis was carried out in a controlled growth chamber using plastic containers with peat moss-vermiculite substrate and mycelial inoculum. Mycorrhiza were obtained 25 days after inoculation. A description of the morphology, appearance and structure of mantle and Hartig net is given for each combination. Mycorrhiza were saffron to cinnamon greenish with age, with a net of saffron laticifers visible through outer mantle; orange latex secreted when injured. Cystidia-like emanating hyphae were observed on the mantle surface of young mycorrhiza. Plectenchymatous mantle with abundant interhyphal gelatinous material.