Molecular phylogenetic analyses have addressed the systematic position of several major Northern Hemisphere lineages of Pezizales but the taxa of the Southern Hemisphere remain understudied. This study focuses on the molecular systematics and taxonomy of Southern Hemisphere species currently treated in the genera Underwoodia and Gymnohydnotrya. Species in these genera have been identified as the monophyletic /gymnohydnotrya lineage, but no further research has been conducted to determine the evolutionary origin of this lineage or its relationship with other Pezizales lineages. Here, we present a phylogenetic study of fungal species previously described in Underwoodia and Gymnohydnotrya, with sampling of all but one described species. We revise the taxonomy of this lineage and describe three new species from the Patagonian region of South America. Our results show that none of these Southern Hemisphere species are closely related to Underwoodia columnaris, the type species of the genus Underwoodia. Accordingly, we recognize the genus Geomorium described by Spegazzini in 1922 for G. fuegianum. We propose the new family, Geomoriaceae fam. nov., to accommodate this phylogenetically and morphologically unique Southern Hemisphere lineage. Molecular dating estimated that Geomoriaceae started to diverge from its sister clade Tuberaceae c. 112 MYA, with a crown age for the family in the late Cretaceous (c. 67 MYA). This scenario fits well with a Gondwanan origin of the family before the split of Australia and South America from Antarctica during the Paleocene-Eocene boundary (c. 50 MYA).
The harvesting of morels is a vital economic activity for local communities in Chile because they are a significant commercial export for the country. Although many species of morels produce ascomata in the absence of fire, abundant ascomata production occurs among some Morchella species when triggered by fire. The intentional burning of Nothofagus forests in Southern Chile, as a means to increase morel production, has become a problem and has negatively impacted ecosystems. Information on the distribution of morels in South America is limited. Spegazzini (1909) described M. patagonica from Argentina and Gamundi et al. (2004) listed five Morchella species from Patagonia and surrounding areas including Spegazzini’s species. Recently Pildain et al. (2014) and Baroni et al. (2018) have examined diversity of Morchella species in South America and the Caribbean using molecular methods. To better understand which Morchella species are being commercially harvested in Southern Chile, molecular markers were used to identify collections of morels being harvested and/or sold commercially and determine their phylogenetic relationships. Morels were sampled from collections in the Fungarium of the Fundación Fungi, Chile (FFCL) and batches purchased from commercial harvesters and bulk gatherers in 2015 and 2016. DNA sequence from the EF1a, RPB1, ITS and RPB2 were obtained and used for phylogenetic analyses. This study will contribute to the knowledge of morels in South America and help to understand their phylogenetic relationships with other Morchella species found worldwide.
Species of Tympanis are well‐known pathogens in Holarctic forests, but we know lit‐ tle about their relationship to other genera in Tympanidaceae. The genus Myriodiscus, remarkable macroscopically and a possible pathogen on bamboo, has a complicated taxonomic history and has not been conclusively placed phylogenetically. Species of Myriodiscus have been described under two other generic names, Ascotremellopsis and Gelatinomyces, the latter being related to the pathogenic genus Collophorina. There has been no formal synonymy of these three bambusicolous genera or agree‐ ment on their systematic positions. We combine accurate micromorphology and DNA sequence data to show the link between Tympanis and Myriodiscus and reveal previously unrecognized features of the latter. These two genera show a type of ascus development unique in the Leotiomycetes. With this new data, we have re‐ solved past confusions over the identity of these fungi, determined their systematic position and have proposed the proper synonymies for Myriodiscus sparassoides and one new combination (M. conus).
The monotypic genus Biatorellina is currently considered a taxonomic synonym of Tryblidiopsis but has an obscure and complicated history. During the revision of the genus Tympanis a syntype of Biatorellina buchsii was fortuitously found and reviewed. Initially our revision led to the hypothesis that B. buchsii could be conspecific with Tympanis confusa. A bibliographic and morphological revision, together with a biometric study, was done using the syntype of B. buchsii and specimens of Tympanisconfusa to verify the identity of the genus Biatorellina. Our results show an overlap in the morphology, biometry, distribution and ecology of B. buchsii (≡ Tympanis buchsii) and Tympanis confusa. The identity and the placement of Biatorellina is resolved and B. buchsii is proposed as a synonym of Tympanis confus.
Fungi in the class Leotiomycetes are ecologically diverse, including mycorrhizas, endophytes of roots and leaves, plant pathogens, aquatic and aero-aquatic hyphomycetes, mammalian pathogens, and saprobes. These fungi are commonly detected in cultures from diseased tissue and from environmental DNA extracts. The identification of specimens from such character-poor samples increasingly relies on DNA sequencing. However, the current classification of Leotiomycetes is still largely based on morphologically defined taxa, especially at higher taxonomic levels. Consequently, the formal Leotiomycetes classification is frequently poorly congruent with the relationships suggested by DNA sequencing studies. Previous class-wide phylogenies of Leotiomycetes have been based on ribosomal DNA markers, with most of the published multi-gene studies being focussed on particular genera or families. In this paper we collate data available from specimens representing both sexual and asexual morphs from across the genetic breadth of the class, with a focus on generic type species, to present a phylogeny based on up to 15 concatenated genes across 279 specimens. Included in the dataset are genes that were extracted from 72 of the genomes available for the class, including 10 new genomes released with this study. To test the statistical support for the deepest branches in the phylogeny, an additional phylogeny based on 3156 genes from 51 selected genomes is also presented. To fill some of the taxonomic gaps in the 15-gene phylogeny, we further present an ITS gene tree, particularly targeting ex-type specimens of generic type species. A small number of novel taxa are proposed: Marthamycetales ord. nov., and Drepanopezizaceae and Mniaeciaceae fams. nov. The formal taxonomic changes are limited in part because of the ad hoc nature of taxon and specimen selection, based purely on the availability of data. The phylogeny constitutes a framework for enabling future taxonomically targeted studies using deliberate specimen selection. Such studies will ideally include designation of epitypes for the type species of those genera for which DNA is not able to be extracted from the original type specimen, and consideration of morphological characters whenever genetically defined clades are recognized as formal taxa within a classification.
Triblidiaceae is a family of uncommonly encountered, non-lichenized discomycetes. A recent classification circumscribed the family to include Triblidium (4 spp. and 1 subsp.), Huangshania (2 spp.) and Pseudographis (2 spp. and 1 var.). The apothecia of these fungi are persistent and drought-tolerant; they possess stromatic, highly melanized covering layers that open and close with fluctuations of humidity. Triblidialean fungi occur primarily on the bark of Quercus, Pinaceae and Ericaceae, presumably as saprobes. Though the type species of Huangshania is from China, these fungi are mostly known from collections originating from Western Hemisphere temperate and boreal forests. The higher-rank classification of triblidialean fungi has been in flux due in part to an overemphasis on ascospore morphology. Muriform ascospores are observed in species of Triblidium and in Pseudographis elatina. An intense, dark blue/purple ascospore wall reaction in iodine-based reagents is observed in species of Pseudographis. These morphologies have led, in part, to these genera being shuffled among unrelated taxa in Hysteriaceae (Dothideomycetes, Hysteriales) and Graphidaceae (Lecanoromycetes, Ostropales). Triblidiaceae has been placed within the monofamilial order Triblidiales (affinity Lecanoromycetes). Here, we demonstrate with a three-gene phylogenetic approach that triblidialean fungi are related to taxa in Rhytismatales (Leotiomycetes). We synonymize Triblidiales under Rhytismatales and emend Triblidiaceae to include Triblidium and Huangshania, with Pseudographis placed within Rhytismataceae. A history of Triblidiaceae is provided along with a description of the emended family. We discuss how the inclusion of triblidialean fungi in Rhytismatales brings some rarely observed or even unique ascospore morphologies to the order and to Leotiomycetes.
Ruhlandiella is a genus of exothecial, ectomycorrhizal fungi in the order Pezizales. Ascomata of exothecial fungi typically lack a peridium and are covered with a hymenial layer instead. Ruhlandiella species have nonoperculate asci and highly ornamented ascospores. The genus was first described by Hennings in 1903 to include the single species, R. berolinensis. Since then, mycologists have uncovered Ruhlandiella species in many locations around the globe, including Australia, Spain, Italy, and the USA. Currently, there are four recognized species: R. berolinensis, R. peregrina, R. reticulata, and R. truncata. All were found near Eucalyptus or Melaleuca trees of Australasian origin. Recently, we discovered two new species of Ruhlandiella in Nothofagaceae forests in South America. They regularly form mitotic spore mats directly on soil in the forests of Patagonia. Here, we formally describe these new species and construct the phylogeny of Ruhlandiella and related genera using a multilocus phylogenetic analysis. We also revise the taxonomy of Ruhlandiella and provide an identification key to accepted species of Ruhlandiella.
The class Laboulbeniomycetes comprises biotrophic parasites associated with arthropods and fungi. Two orders are currently recognized, Pyxidiophorales and Laboulbeniales. Herpomyces is an isolated genus of Laboulbeniales, with species that exclusively parasitize cockroaches (Blattodea). Here, we evaluate 39 taxa of Laboulbeniomycetes with a three-locus phylogeny (nrSSU, ITS, nrLSU) and propose a new order in this class. Herpomycetales accommodates a single genus, Herpomyces, with currently 26 species, one of which is described here based on morphological and molecular data. Herpomyces shelfordellae is found on Shelfordella lateralis cockroaches from Hungary, Poland, and the USA. We also build on the six-locus dataset from the Ascomycota Tree of Life paper (Schoch and colleagues, 2009) to confirm that Laboulbeniomycetes and Sordariomycetes are sister classes, and we apply laboulbeniomyceta as a rankless taxon for the now well-resolved node that describes the most recent common ancestor of both classes.
In this paper, new species and formae of the genus Gloeandromyces (Ascomycota, Laboulbeniales) are described and illustrated. These are: Gloeandromyces dickii sp. nov. on Trichobius joblingi from Nicaragua and Panama; G. pageanus f. alarum f. nov. on Tri. joblingi from Panama; G. pageanus f. polymorphus f. nov. on Tri. dugesioides and Tri. joblingi from Panama and Trinidad; and G. streblae f. sigmomorphus f. nov. on Tri. joblingi from Panama. Gloeandromyces pageanus on Tri. dugesioides from Panama as described in Nova Hedwigia 105 (2017) is referred to as G. pageanus f. pageanus. Support for these descriptions of species and formae comes from phylogenetic reconstruction of the large subunit ribosomal DNA and from the application of species delimitation methods (ABGD, bPTP, GMYC). Host specialization results in phylogenetic segregation by host species in both G. pageanus and G. streblae and this may represent a case of incipient speciation. A second mechanism driving diversity involves position induced morphological adaptations, leading to the peculiar morphotypes that are associated to growing on a particular position of the integument (G. pageanus f. alarum, G. streblae f. sigmomorphus).
Herbarium specimens represent important records of morphological and genetic diversity of plants that inform questions relevant to global change, including species distributions, phenology and functional traits. It is increasingly appreciated that plant microbiomes can influence these aspects of plant biology, but little is known regarding the historic distribution of microbes associated with plants collected in the pre-molecular age. If microbiomes can be observed reliably in herbarium specimens, researchers will gain a new lens with which to examine microbial ecology, evolution, species interactions. Here, we describe a method for accessing historical plant microbiomes from preserved herbarium specimens, providing a proof of concept using two plant taxa from the imperiled boreal biome (Andromeda polifolia and Ledum palustre subsp. groenlandicum, Ericaceae). We focus on fungal endophytes, which occur within symptomless plant tissues such as leaves. Through a three-part approach (i.e. culturing, cloning and next-generation amplicon sequencing via the Illumina MiSeq platform, with extensive controls), we examined endophyte communities in dried, pressed leaves that had been processed as regular herbarium specimens and stored at room temperature in a herbarium for four years. We retrieved only one endophyte in culture, but cloning and especially the MiSeq analysis revealed a rich community of foliar endophytes. The phylogenetic distribution and diversity of endophyte assemblages, especially among the Ascomycota, resemble endophyte communities from fresh plants collected in the boreal biome. We could distinguish communities of endophytes in each plant species and differentiate likely endophytes from fungi that could be surface contaminants. Taxa found by cloning were observed in the larger MiSeq dataset, but species richness was greater when subsets of the same tissues were evaluated with the MiSeq approach. Our findings provide a proof of concept for capturing endophyte DNA from herbarium specimens, supporting the importance of herbarium records as roadmaps for understanding the dynamics of plant-associated microbial biodiversity in the Anthropocene.This article is part of the theme issue 'Biological collections for understanding biodiversity in the Anthropocene'.
First published by Fries in 1825, the genus Sarea today comprises two accepted species of resinicolous discomycetes. Both species have a very broad range, with S. difformis reported from North America, Europe, and northwestern Africa, and S. resinae reported from North America, Europe, northern and central Africa, and central and eastern Asia. Both species have also been reported in southern hemisphere locations, such as New Zealand, on non-native trees. Both species also have a broad range of hosts in the Pinaceae, with S. difformis reported on Cedrus atlantica and both Sarea species reported on species of Pinus, Picea, Larix, Pseudotsuga, Abies and Tsuga. In addition, S. resinae has been reported on species in the Cupressaceae, including members of the genera Cupressus, Chamaecyparis,Juniperus and Taxodium. With few exceptions, specimens of each Sarea species share a very similar macro- and micromorphology, with specimens from multiple hosts fitting the specific concepts published by Hawksworth and Sherwood in 1981. Some molecular work has been done on the genus, but in almost all cases sequences are not associated with a vouchered herbarium specimen including the sexual morph. The objective of this study is to determine the degree of relatedness of geographically distant specimens collected in North America, Europe, and Macaronesia on different native and non-native host species. With permission, collections have been made of both species of Sarea from California, Georgia and the New England states in the USA, Northern and Southern Europe and Macaronesia. In addition to detailed measurements of the micromorphological features of specimens, ITS and LSU sequences have been generated using Sanger sequencing for analysis and comparison with published sequences. In contrast to the generally only slight morphological differences noted among specimens, ITS sequences from Europe, Asia, and North America not only differ by about 4% from each other, but also when submitted to phylogenetic analyses form multiple well-supported clades for each continent. These patterns are supported by similar analyses using the LSU sequences and ITS+LSU sequences. These clades also point to host specificity at the host family or genus level. In conclusion, the composition of the genus Sarea seems much more complicated than previously reported, with the possibility of multiple cryptic species in both accepted taxa; additional work must be done to further expand geographical and host range sampling of specimens to include in these analyses in order to approach a full picture of the diversity in Sarea.
The new genus Aotearoamyces is proposed to accommodate a single species that was repeatedly collected on fallen wood in Nothofagaceae forests of New Zealand and was previously misidentified as a Claussenomyces species. This monotypic genus belongs to Tympanidaceae, a recently erected family in Phacidiales. Aotearoamyces is differentiated from other Tympanidaceae by phragmospores that do not form conidia either in or outside the asci, an exciple of textura intricata with hyphae widely spaced and strongly gelatinized (plectenchyma), and apically flexuous, partly helicoid paraphyses. The asexual morph was studied in pure culture. Phylogenetic analyses of combined SSU, ITS and LSU sequences strongly support a sister relationship between the sexually typified Aotearoamyces and the asexually typified “Collophorina” paarla characterized morphologically by forming endoconidia, a feature not found in the genetically distinct type species of Collophorina. Based on our molecular results, we place the genus Epithamnolia in the Mniaecia lineage within Phacidiales.
Parasola is a genus of small, veil-less coprinoid mushrooms in the family Psathyrellaceae (Agaricales). The genus is not well documented in Asia, specifically in Pakistan. In this study we describe two new species Parasola glabra and P. pseudolactea from Pakistan, based on morphological and molecular data. Phylogeny based on three DNA regions: nuc rDNA region encompassing the internal transcribed spacers 1 and 2 along with the 5.8S rDNA (ITS), nuc 28S rDNA D1-D2 domains (28S) and translation elongation factor 1α gene (TEF1α) show that the new taxa are clustered in a clade formed by the members of section Parasola of genus Parasola. Parasola glabra with grayish pileus, slightly depressed pileal disc, lamellae separated from the stipe by pseudocollarium, basidiospores 14.5–16.5 × 9.5–11.5 × 8.0–10.5 µm, in front view broadly ovoid to oblong, some with rhomboidal outline, in side view ellipsoid, with eccentric germ-pore of 1.5 µm diameter. Parasola pseudolactea with yellowish brown to dull brown pileus, disc indistinctly umbonate, lamellae free, pseudocollarium absent, basidiospores 13.5–14.5 × 10.5–12.0 × 9.5–10.5 µm, in face view rounded triangular to heart shaped, rarely ovoid to subglobose, in side view ellipsoid to oblong, with eccentric germ-pore of 1.5 µm diam. In addition to these new species, P. auricoma and P.lilatincta were also studied. Morphological descriptions for the new species and comparison with known Parasola species are provided. Our observations highlight the diversity of Parasola in northern Pakistan and further document the need for additional systematic focus on the region’s fungi.
The genus Otidea was recently monographed and studied phylogenetically, but knowledge of the diversity and distribution of Otidea species in China is fragmentary. In this study, collections from China were examined morphologically and included in phylogenetic analyses. Using LSU, TEF1-α, and RPB2 new species were placed within previously recognized clades in the genus. The results agree with both Genealogical Concordance Phylogenetic Species Recognition (GCPSR) and genetic divergence as previously reported. Three new species, Otidea hanseniae, Otidea korfii and Otidea purpureogrisea are recognized based on phylogenetic reconstruction using ITS, LSU, TEF1- α and RPB2. Comments on some incompletely known species are added. With the discovery of these three new species, the genus Otidea in China proves to be more diverse than previously recognized.
The large genus Leucoagaricus (Basidiomycota) is poorly studied in Pakistan, where the northern parts of the country are considered hotspots for biodiversity. Based on morphological and molecular data, five new species are described: Leucoagaricus badius, L. lahorensiformis, L. pakistaniensis, L. sultanii and L. umbonatus. Descriptions and illustrations are given for the new taxa. A phylogeny based on morphology and four DNA regions, including the internal transcribed spacers (ITS1-5.8S-ITS2 = ITS) and D1-D2 domains of the 28S gene of the nuc rDNA), the gene for RNA polymerase II second largest subunit (RPB2) and the translation elongation factor 1α gene (TEF1-α), show that the new taxa are clustered in a clade representing Leucoagaricus section Rubrotincti subgenus Sericeomyces.