Historically, thallus-forming Laboulbeniomycetes, including the orders Laboulbeniales and Herpomycetales, were set apart because of their distinctive morphology and ecology. Although some biologists correctly interpreted these arthropod ectoparasites as fungi, even ascomycetes, others thought they were worms, red algae, or members of taxa described especially for them. Speculation on the evolution of the group involving red algae, the morphology-based Floridean Hypothesis, persisted deep into the 20th century, in part because valid alternatives were not presented. Although the distinctive features of Laboulbeniales clearly set them apart from other fungi, the difficulty was in the absence of characters grouping them among the fungi. Thaxter considered the Laboulbeniales to be ascomycetes, but he avoided phylogenetic discussions involved in the Floridean Hypothesis all of his life. Eventually, developmental studies of the life history of Pyxidiophora species, hyphal perithecial ascomycetes with 2-celled ascospores, revealed characters connecting Laboulbeniales to other ascomycetes. The distinctive morphological features of Laboulbeniales (absence of mycelium, a thallus developed from 2-celled ascospores by cell divisions in several planes, arthropod parasitism) can be best understood by comparison with Pyxidiophora. The development of a 3-dimensional thallus composed of true parenchyma occurs not only in Laboulbeniales, but also in Pyxidiophora species. The life history of arthropod ectoparasitism of Laboulbeniales as well as mycoparasitism and phoretic dispersal by arthropods of Pyxidiophora species can be explained by Tranzschel’s Law, originally applied to rust fungi. Molecular analyses including other arthropod-associated fungi have contributed to a better understanding of an enlarged class, Laboulbeniomycetes, which now includes a clade comprising Chantransiopsis, Tetrameronycha, and Subbaromyces. A two-locus phylogenetic tree highlights evolutionary and life history questions with regard to the placement of Herpomycetales as the first diverging lineage of the Laboulbeniomycetes. The sister group for all the Laboulbeniomycetes remains to be discovered.
Full citation: Beimforde, C., Schmidt, A.R., Rikkinen, J. and J.K. Mitchell. 2020. Sareomycetes cl. nov.: A new proposal for placement of the resinicolous genus Sarea (Ascomycota, Pezizomycotina). Fungal Systematics and Evolution.
Resinicolous fungi constitute a heterogeneous assemblage of fungi that live on fresh and solidified plant resins. The genus Sarea includes, according to current knowledge, two species, S. resinae and S. difformis. In contrast to other resinicolous discomycetes, which are placed in genera also including non-resinicolous species, Sarea species only ever fruit on resin. The taxonomic classification of Sarea has proven to be difficult and currently the genus, provisionally and based only on morphological features, has been assigned to the Trapeliales (Lecanoromycetes). In contrast, molecular studies have noted a possible affinity to the Leotiomycetes. Here we review the taxonomic placement of Sarea using sequence data from seven phylogenetically informative DNA regions including ribosomal (ITS, nucSSU, mtSSU, nucLSU) and protein-coding (rpb1, rpb2, mcm7) regions.
We combined available and new sequence data with sequences from major Pezizomycotina classes, especially Lecanoromycetes and Leotiomycetes, and assembled three different taxon samplings in order to place the genus Sarea within the Pezizomycotina. Based on our data, none of the applied phylogenetic approaches (Bayesian Inference, Maximum Likelihood and Maximum Parsimony) supported the placement of Sarea in the Trapeliales or any other order in the Lecanoromycetes. A placement of Sarea within the Leotiomycetes is similarly unsupported. Based on our data, Sarea forms an isolated and highly supported phylogenetic lineage within the "Leotiomyceta". From the results of our multilocus phylogenetic analyses we propose here a new class, order, and family, Sareomycetes, Sareales and Sareaceae in the Ascomycota to accommodate the genus Sarea. The genetic variability within the newly proposed class suggests that it is a larger group that requires further infrageneric classification.
Herpomyces periplanetae is an obligate biotroph of Periplaneta americana, the American cockroach. Its nearly cosmopolitan distribution is shaped by its globally invasive host and the international pet trade. Here, we report the draft genome sequence of H. periplanetae, based on a thallus from P. americana collected in Cambridge, Massachusetts
Geodina salmonicolor is shown to be a synonym of G. guanacastensis, the type and only species of the genus. Comparisons of ITS rDNA sequences of a paratype and two recent collections of G. guanacastensis with published ITS sequences of G. salmonicolor, from the Dominican Republic, show that these are nearly identical. When G. salmonicolor was erected no sequences of the type species were available. Morphological comparisons supports the conspecificity. Details regarding the description of G. salmonicolor are pointed out. A four-gene phylogeny places Geodina and Wynnea as a supported sister group to the rest of the Sarcoscyphaceae. Species in these genera share morphological traits of cyanophobic spore markings, dark angular outer excipular cells that give rise to hairs and the origin of several apothecia from a common basal stalk. Their occurrence on soil rather than on wood or plant material distinguish them from other Sarcoscyphaceae. Based on morphology, phylogenic relationships and trophic interactions we erect a new family, Wynneaceae, for Geodina and Wynnea.
Micraspis acicola was described more than 50 years ago to accommodate a phacidium-like fungus that caused a foliar disease of Picea mariana. After its publication, two more species were added, M. strobilina and M. tetraspora, all of them growing on Pinaceae in the Northern Hemisphere, but each species occupying a unique type of host tissue (needles, cones or wood). Micraspis is considered to be a member of class Leotiomycetes, but was originally placed in Phacidiaceae (Phacidiales), later transferred to Helotiaceae (Helotiales) and recently returned to Phacidiales but in a different family (Tympanidaceae). The genus remains poorly sampled, and hence poorly understood both taxonomically and ecologically. Here, we use morphology, cultures and sequences to provide insights into its systematic position in Leotiomycetes and its ecology. Our results show that the genus should not be included in Tympanidaceae or Phacidiaceae, and support the erection of a new family and order with a unique combination of morphological features supported by molecular data
Little is known about the diversity of Orbiliomycetes from Puerto Rico. Cantrell & Lodge (2008) compiled a list of the fungi from Puerto Rico, and only mentioned four species of Orbilia: O. andina, O. chysocoma, O. delicata and O. cf. gaillardii. During IMC11 in Puerto Rico, 2018, several collections of Orbiliomycetes were found in Juan Enrique Monagas Park during the Ascomycete workshop field trip. Some of these were sent to the Cornell and Farlow Herbaria. One collection caught the attention of Luis Quijada due to its interesting morphological features. This species is not related to any of the species reported by Cantrell & Lodge (2008). The morphology of the asci and ascospores of the sexual morph clearly indicates a relationship with Orbilia auricolor and related species (section Arthrobotrys). The morphology of the strongly mammiform paraphyses and the excipulum with large cortical cells with knob-like glassy caps had never before seen in this section. Our cultures produced an Arthrobotrys-like anamorph most similar to the anamorph of O. blumenaviensis (= A. vermicola), but the conidia are distinctly smaller and never more than 1-septate. Molecular data supports the placement of this Orbilia in series Arthrobotrys. Species in this series produce adhesive networks as trapping organs in the presence of nematodes, and this behavior was confirmed in cultures of this Orbilia. Our phylogenetic analysis shows this species as very distinct from O. blumenaviensis and supports it being new to science.
This paper presents 13 new records of Laboulbenialean parasites on ground beetles (Coleoptera, Carabidae) from the Boston Harbor Islands National Recreation Area in Massachusetts: Laboulbenia anoplogenii, L. casnoniae, L. clivinalis, L. egens, L. filifera, L. flagellata, L. inflata, L. macrotheca, L. pedicellata, L. terminalis, L. varia-bilis, L. vulgaris, and Peyritschiella geminata. Laboulbenia clivinalis and L. egens are new country records for the US. Moreover, we present additional localities for L. casnoniae, L. clivinalis, L. filifera, L. flagellata, L. inflata, L. pedicellata, L. variabilis, and L. vulgaris. The following new country records are presented: Laboulbenia clivinalis, L. filifera, and L. variabilis from Canada; L. flagellata from the Democratic Republic of the Congo; L. pedicellata from Ukraine; L. vulgaris from Croatia and Slovenia (and the first undoubtful record from Slovakia). Laboulbenia flagellata was found on 11 host species in the genera Agonum, Oxypselaphus, Patrobus, Platynus, and Pterostichus. Using this abundant material, we performed morphometrics to test the hypothesis that L. flagel-lata is a species complex. Specimens cannot be separated based on host genus (Agonum, Pterostichus). One parameter is significant between Pterostichus mutus and each of the 4 Agonum species after applying a strong Bonferroni P-value correction: H1T, the ratio of height of cell I (HC1) to total thallus length (TTL). In addition, we collected fresh material to be able to add a molecular phylogenetic component to test said hypothesis. We generated ITS and nrLSU ribosomal sequences of several species of Laboulbenia, including isolates of L. flagellata from multiple hosts. Phylogenetic inference of the concatenated dataset shows that L. flagellata isolates from 3 host species form 2 distinct clades, providing support for our hypothesis. We also show that L. coneglianensis is sepa-rate from L. flagellata, unequivocally ending a long-standing taxonomic debate. Finally, examination of Roland Thaxter’s 1891–1932 slides led to the designation of lectotypes for L. macrothecia, L. terminalis, and P. geminata.
A new species, Geopora ahmadii, is described and illustrated based on material from Punjab, Pakistan. This species is characterized by sessile, cup- to saucer-shaped, partly immersed apothecia with whitish to grayish hymenial surfaces; broad ellipsoid, mostly uniguttulate ascospores; and brown excipular hairs. Phylogenetic analyses of the nrDNA ITS region with maximum parsimony, maximum likelihood, and Bayesian inference methods reveal that G. ahmadii is distinct from other described Geopora species. A collection previously identified as Geopora arenosa from Rawalakot, Pakistan, likely represents a second locality of G. ahmadii.
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'.