Publications by authors named "Donald H Pfister"

42 Publications

Sareomycetes: more diverse than meets the eye.

IMA Fungus 2021 Mar 16;12(1). Epub 2021 Mar 16.

Farlow Reference Library and Herbarium of Cryptogamic Botany, Harvard University, 22 Divinity Avenue, Cambridge, MA, 02138, USA.

Since its resurrection, the resinicolous discomycete genus Sarea has been accepted as containing two species, one with black apothecia and pycnidia, and one with orange. We investigate this hypothesis using three ribosomal (nuITS, nuLSU, mtSSU) regions from and morphological examination of 70 specimens collected primarily in Europe and North America. The results of our analyses support separation of the traditional Sarea difformis s.lat. and Sarea resinae s.lat. into two distinct genera, Sarea and Zythia. Sarea as circumscribed is shown to conservatively comprise three phylospecies, with one corresponding to Sarea difformis s.str. and two, morphologically indistinguishable, corresponding to the newly combined Sarea coeloplata. Zythia is provisionally maintained as monotypic, containing only a genetically and morphologically variable Z. resinae. The new genus Atrozythia is erected for the new species A. klamathica. Arthrographis lignicola is placed in this genus on molecular grounds, expanding the concept of Sareomycetes by inclusion of a previously unknown type of asexual morph. Dating analyses using additional marker regions indicate the emergence of the Sareomycetes was roughly concurrent with the diversification of the genus Pinus, suggesting that this group of fungi emerged to exploit the newly-available resinous ecological niche supplied by Pinus or another, extinct group of conifers. Our phylogeographic studies also permitted us to study the introductions of these fungi to areas where they are not native, including Antarctica, Cape Verde, and New Zealand and are consistent with historical hypotheses of introduction.
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http://dx.doi.org/10.1186/s43008-021-00056-0DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7961326PMC
March 2021

Mortality of native and invasive ladybirds co-infected by ectoparasitic and entomopathogenic fungi.

PeerJ 2020 4;8:e10110. Epub 2020 Nov 4.

Agricultural Research Service, Southeastern Fruit and Tree Nut Research Laboratory, United States Department of Agriculture, Byron, GA, United States of America.

is an invasive alien ladybird in North America and Europe. Studies show that multiple natural enemies are using as a new host. However, thus far, no research has been undertaken to study the effects of simultaneous infection by multiple natural enemies on . We hypothesized that high thallus densities of the ectoparasitic fungus on a ladybird weaken the host's defenses, thereby making it more susceptible to infection by other natural enemies. We examined mortality of the North American-native and co-infected with and an entomopathogenic fungus-either or . Laboratory assays revealed that -infected individuals are more susceptible to entomopathogenic fungi, but does not suffer the same effects. This is in line with the enemy release hypothesis, which predicts that invasive alien species in new geographic areas experience reduced regulatory effects from natural enemies compared to native species. Considering our results, we can ask how affects survival when confronted by other pathogens that previously had little impact on .
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http://dx.doi.org/10.7717/peerj.10110DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7648450PMC
November 2020

Laboulbeniomycetes: Intimate Fungal Associates of Arthropods.

Annu Rev Entomol 2021 01 31;66:257-276. Epub 2020 Aug 31.

Farlow Reference Library and Herbarium of Cryptogamic Botany, Harvard University, Cambridge, Massachusetts 02138, USA; email:

Arthropod-fungus interactions involving the Laboulbeniomycetes have been pondered for several hundred years. Early studies of Laboulbeniomycetes faced several uncertainties. Were they parasitic worms, red algal relatives, or fungi? If they were fungi, to which group did they belong? What was the nature of their interactions with their arthropod hosts? The historical misperceptions resulted from the extraordinary morphological features of these oddly constructed ectoparasitic fungi. More recently, molecular phylogenetic studies, in combination with a better understanding of life histories, have clearly placed these fungi among filamentous Ascomycota (subphylum Pezizomycotina). Species discovery and research on the classification of the group continue today as arthropods, and especially insects, are routinely collected and examined for the presence of Laboulbeniomycetes. Newly armed with molecular methods, mycologists are poisedto use Laboulbeniomycetes-insect associations as models for the study of a variety of basic evolutionary and ecological questions involving host-parasite relationships, modes of nutrient intake, population biology, host specificity, biological control, and invasion biology. Collaboration between mycologists and entomologists is essential to successfully advance knowledge of Laboulbeniomycetes and their intimate association with their hosts.
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http://dx.doi.org/10.1146/annurev-ento-013020-013553DOI Listing
January 2021

New species of (Agaricales, Inocybaceae) from Pakistan revealed by morphology and multi-locus phylogenetic reconstruction.

MycoKeys 2020 10;69:1-31. Epub 2020 Jul 10.

Department of Botany, University of the Punjab, Lahore, Pakistan University of the Punjab Lahore Pakistan.

During fungal surveys between 2012 and 2014 in pine-dominated forests of the western Himalayas in Pakistan, several collections of (Agaricales, Inocybaceae) were made. These were documented, based on morphological and molecular data. During this work, three new species came to light, which are here formally described as , and . These species belong in the genus Matheny et al. (2019) = Pseudosperma clade Matheny (2005) = sect. Rimosae s.s. fideLarsson et al. (2009). Macro- and micro-morphological descriptions, illustrations and molecular phylogenetic reconstructions of the studied taxa are provided. The new species are differentiated from their close relatives by basidiospore size and colouration of basidiomata. Molecular phylogenetic relationships are inferred using ITS (ITS1-5.8S-ITS2), nrLSU and mtSSU sequence data. All three newly-described taxa likely share an ectomycorrhizal association with trees in the genus . In addition, five names are recombined in , and . These are , , , and .
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http://dx.doi.org/10.3897/mycokeys.69.33563DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7367896PMC
July 2020

Phylogenetic study documents different speciation mechanisms within the lineage in boreal and arctic environments of the Northern Hemisphere.

IMA Fungus 2019 7;10. Epub 2019 Jun 7.

Department of Cryptogams, Institute of Botany, Plant Science and Biodiversity Centre, Slovak Academy of Sciences, Dúbravská cesta 9, SK-845 23 Bratislava, Slovakia.

The lineage is a morphologically and phylogenetically well-defined group of ectomycorrhizal fungi occurring in various climatic areas. In this study we performed a multi-locus phylogenetic study based on collections from boreal, alpine and arctic habitats of Europe and Western North America, subalpine collections from the southeast Himalayas and collections from subtropical coniferous forests of Pakistan. European and North American collections are nearly identical and probably represent a single species named distributed from the Alps to the Rocky Mountains. Collections from the southeast Himalayas belong to two distinct species: sp. nov. from subtropical monodominant forests of and sp. nov. from subalpine mixed forests of and . The results suggest that speciation in this group is driven by a climate disjunction and adaptation rather than a host switch and geographical distance.
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http://dx.doi.org/10.1186/s43008-019-0003-9DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7325667PMC
June 2019

Laboulbeniomycetes: Evolution, natural history, and Thaxter's final word.

Mycologia 2020 Nov-Dec;112(6):1048-1059. Epub 2020 Mar 17.

Department of Organismic and Evolutionary Biology and Farlow Reference Library and Herbarium of Cryptogamic Botany, Harvard University , Cambridge, Massachusetts 02138.

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 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 . The development of a 3-dimensional thallus composed of true parenchyma occurs not only in Laboulbeniales, but also in species. The life history of arthropod ectoparasitism of Laboulbeniales as well as mycoparasitism and phoretic dispersal by arthropods of 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 , and . 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.
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http://dx.doi.org/10.1080/00275514.2020.1718442DOI Listing
March 2020

Draft Genome Sequence of the Globally Distributed Cockroach-Infecting Fungus Herpomyces periplanetae Strain D. Haelew. 1187d.

Microbiol Resour Announc 2020 Feb 6;9(6). Epub 2020 Feb 6.

Farlow Reference Library and Herbarium of Cryptogamic Botany, Harvard University, Cambridge, Massachusetts, USA.

is an obligate biotroph of , 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 , based on a thallus from collected in Cambridge, Massachusetts.
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http://dx.doi.org/10.1128/MRA.01458-19DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7005119PMC
February 2020

Placement of Triblidiaceae in Rhytismatales and comments on unique ascospore morphologies in Leotiomycetes (Fungi, Ascomycota).

MycoKeys 2019 18;54:99-133. Epub 2019 Jun 18.

Farlow Herbarium of Harvard University, 22 Divinity Avenue, Cambridge, MA, 02138, USA Harvard University Cambridge United States of America.

Triblidiaceae is a family of uncommonly encountered, non-lichenized discomycetes. A recent classification circumscribed the family to include (4 spp. and 1 subsp.), (2 spp.) and (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 , Pinaceae and Ericaceae, presumably as saprobes. Though the type species of 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 and in . An intense, dark blue/purple ascospore wall reaction in iodine-based reagents is observed in species of . 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 and , with 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.
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http://dx.doi.org/10.3897/mycokeys.54.35697DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6592975PMC
June 2019

Systematic study of truffles in the genus , with the description of two new species from Patagonia.

Mycologia 2019 May-Jun;111(3):477-492. Epub 2019 Apr 1.

a Department of Plant Pathology , University of Florida , Gainesville , Florida 32611.

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. species have nonoperculate asci and highly ornamented ascospores. The genus was first described by Hennings in 1903 to include the single species, . Since then, mycologists have uncovered species in many locations around the globe, including Australia, Spain, Italy, and the USA. Currently, there are four recognized species: , and . All were found near or trees of Australasian origin. Recently, we discovered two new species of 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 and related genera using a multilocus phylogenetic analysis. We also revise the taxonomy of and provide an identification key to accepted species of .
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http://dx.doi.org/10.1080/00275514.2019.1574490DOI Listing
April 2020

Birth of an order: Comprehensive molecular phylogenetic study excludes Herpomyces (Fungi, Laboulbeniomycetes) from Laboulbeniales.

Mol Phylogenet Evol 2019 04 6;133:286-301. Epub 2019 Jan 6.

Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA, USA.

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.
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http://dx.doi.org/10.1016/j.ympev.2019.01.007DOI Listing
April 2019

Overview of , including gen. nov. on .

IMA Fungus 2018 Jul 30;9:371-382. Epub 2018 Oct 30.

Department of Organismic and Evolutionary Biology, Harvard Herbarium, 22 Divinity Avenue, Cambridge MA 02138, United States of America.

The new genus is proposed to accommodate a single species that was repeatedly collected on fallen wood in forests of New Zealand and was previously misidentified as a species. This monotypic genus belongs to , a recently erected family in . is differentiated from other by phragmospores that do not form conidia either in or outside the asci, an exciple of 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 and the asexually typified " characterized morphologically by forming endoconidia, a feature not found in the genetically distinct type species of . Based on our molecular results, we place the genus in the lineage within .
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http://dx.doi.org/10.5598/imafungus.2018.09.02.08DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6317588PMC
July 2018

A novel proof of concept for capturing the diversity of endophytic fungi preserved in herbarium specimens.

Philos Trans R Soc Lond B Biol Sci 2018 11 19;374(1763). Epub 2018 Nov 19.

School of Plant Sciences, University of Arizona, Tucson, AZ 85721, USA.

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 ( and subsp 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'.
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http://dx.doi.org/10.1098/rstb.2017.0395DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6282087PMC
November 2018

Integrative taxonomy reveals hidden species within a common fungal parasite of ladybirds.

Sci Rep 2018 10 29;8(1):15966. Epub 2018 Oct 29.

Department of Organismic and Evolutionary Biology, Harvard University, 22 Divinity Avenue, Cambridge, Massachusetts, 02138, USA.

Our understanding of fungal diversity is far from complete. Species descriptions generally focus on morphological features, but this approach may underestimate true diversity. Using the morphological species concept, Hesperomyces virescens (Ascomycota, Laboulbeniales) is a single species with global distribution and wide host range. Since its description 120 years ago, this fungal parasite has been reported from 30 species of ladybird hosts on all continents except Antarctica. These host usage patterns suggest that H. virescens could be made up of many different species, each adapted to individual host species. Using sequence data from three gene regions, we found evidence for distinct clades within Hesperomyces virescens, each clade corresponding to isolates from a single host species. We propose that these lineages represent separate species, driven by adaptation to different ladybird hosts. Our combined morphometric, molecular phylogenetic and ecological data provide support for a unified species concept and an integrative taxonomy approach.
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http://dx.doi.org/10.1038/s41598-018-34319-5DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6206035PMC
October 2018

Laboulbeniales hyperparasites (Fungi, Ascomycota) of bat flies: Independent origins and host associations.

Ecol Evol 2018 Aug 24;8(16):8396-8418. Epub 2018 Jul 24.

Department of Organismic and Evolutionary Biology Farlow Reference Library and Herbarium of Cryptogramic Botany Harvard University Cambridge Massachusetts.

The aim of this study was to explore the diversity of ectoparasitic fungi (Ascomycota, Laboulbeniales) that use bat flies (Diptera, Hippoboscoidea) as hosts. Bat flies themselves live as ectoparasites on the fur and wing membranes of bats (Mammalia, Chiroptera); hence this is a tripartite parasite system. Here, we collected bats, bat flies, and Laboulbeniales, and conducted phylogenetic analyses of Laboulbeniales to contrast morphology with ribosomal sequence data. Parasitism of bat flies by Laboulbeniales arose at least three times independently, once in the Eastern Hemisphere () and twice in the Western Hemisphere (, ). We hypothesize that the genera and evolved independently from lineages of ectoparasites of true bugs (Hemiptera). We assessed phylogenetic diversity of the genus by considering the LSU rDNA region. Phenotypic plasticity and position-induced morphological adaptations go hand in hand. Different morphotypes belong to the same phylogenetic species. Two species, and , show divergence by host utilization. In our assessment of coevolution, we only observe congruence between the Old World clades of bat flies and Laboulbeniales. The other associations are the result of the roosting ecology of the bat hosts. This study has considerably increased our knowledge about bats and their associated ectoparasites and shown the necessity of including molecular data in Laboulbeniales taxonomy.
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http://dx.doi.org/10.1002/ece3.4359DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6145224PMC
August 2018

Considerations and consequences of allowing DNA sequence data as types of fungal taxa.

IMA Fungus 2018 Jun 24;9(1):167-175. Epub 2018 May 24.

Museum of Evolution, Uppsala University, Norbyvägen 16, 75236 Uppsala, Sweden.

Nomenclatural type definitions are one of the most important concepts in biological nomenclature. Being physical objects that can be re-studied by other researchers, types permanently link taxonomy (an artificial agreement to classify biological diversity) with nomenclature (an artificial agreement to name biological diversity). Two proposals to amend the International Code of Nomenclature for algae, fungi, and plants (ICN), allowing DNA sequences alone (of any region and extent) to serve as types of taxon names for voucherless fungi (mainly putative taxa from environmental DNA sequences), have been submitted to be voted on at the 11 International Mycological Congress (Puerto Rico, July 2018). We consider various genetic processes affecting the distribution of alleles among taxa and find that alleles may not consistently and uniquely represent the species within which they are contained. Should the proposals be accepted, the meaning of nomenclatural types would change in a fundamental way from physical objects as sources of data to the data themselves. Such changes are conducive to irreproducible science, the potential typification on artefactual data, and massive creation of names with low information content, ultimately causing nomenclatural instability and unnecessary work for future researchers that would stall future explorations of fungal diversity. We conclude that the acceptance of DNA sequences alone as types of names of taxa, under the terms used in the current proposals, is unnecessary and would not solve the problem of naming putative taxa known only from DNA sequences in a scientifically defensible way. As an alternative, we highlight the use of formulas for naming putative taxa (candidate taxa) that do not require any modification of the ICN.
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http://dx.doi.org/10.5598/imafungus.2018.09.01.10DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6048565PMC
June 2018

Influence of Elytral Color Pattern, Size, and Sex of Harmonia axyridis (Coleoptera, Coccinellidae) on Parasite Prevalence and Intensity of Hesperomyces virescens (Ascomycota, Laboulbeniales).

Insects 2018 Jun 15;9(2). Epub 2018 Jun 15.

Department of Organismic and Evolutionary Biology, Faculty of Arts and Sciences, Harvard University, Cambridge, MA 02138, USA.

is an invasive ladybird (Coleoptera, Coccinellidae) with the potential to outcompete native ladybird species in its invasive distribution area. It was introduced as a biological control agent in many countries but has also spread unintentionally in many others. (Ascomycota, Laboulbeniales) is a minute (200⁻400 µm in size) biotrophic fungus that infects over 30 species of ladybirds. The aim of this study was to evaluate whether the elytral color pattern, size, and sex of affect infection by . Coloration in has been linked to the presence of an antimicrobial alkaloid (harmonine). In fall 2016, we collected 763 individuals in Cambridge, Massaschusetts, of which 119 (16%) bore fruiting bodies. We analyzed 160 individuals (80 infected, 80 uninfected) concerning the intensity of infection by . Elytral sizes and coloration patterns were quantified using digital photography and analytical methods. Smaller ladybirds had a higher prevalence and higher intensity of parasitism. Additionally, male ladybirds bore more thalli compared to female ladybirds. Elytral color patterns had an effect on neither prevalence nor intensity of infection by Laboulbeniales in our dataset, although we found a slight trend to higher intensity of parasitism in more melanic males. This suggests that the development of Laboulbeniales might be affected by certain insect alkaloids.
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http://dx.doi.org/10.3390/insects9020067DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6023472PMC
June 2018

Lost and found: the Bermudan Donadinia seaveri found in North America, with comments on its juniper associates.

Mycologia 2018 Jan-Feb;110(1):215-221

a Farlow Reference Library and Herbarium and the Department of Organismic and Evolutionary Biology , Harvard University , 22 Divinity Ave., Cambridge , Massachusetts 02138.

Collections of a species referred to Sarcosomataceae (Pezizomycetes) from eastern North America were studied both morphologically and using nuc rDNA internal transcribed spacer (ITS) region (ITS1-5.8S-ITS2 = ITS) and approximately 800 bp from the 5' region of the nuc 28S rDNA (28S) to construct a phylogeny. The analyses indicate that these collections are Donadinia seaveri, a species previously known only from Bermuda. Because the associated tree, Juniperus bermudiana, has declined as a result of insect attack, it was thought that D. seaveri might be extinct. This work indicates that it is not extinct but is present in eastern North America. The species is described, new distributional records are given, and its association with the genus Juniperus is discussed.
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http://dx.doi.org/10.1080/00275514.2017.1409052DOI Listing
December 2018

Underexplored regions of Pakistan yield five new species of Leucoagaricus.

Mycologia 2018 Mar-Apr;110(2):387-400. Epub 2018 May 8.

a Harvard University Herbaria and Department of Organismic and Evolutionary Biology , Harvard University , Cambridge , Massachusetts 02138.

The large genus Leucoagaricus (Basidiomycota) is poorly studied in Pakistan, where the northern parts of the country are considered hot spots 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 1-α gene (TEF1-α), shows that the new taxa are clustered in a clade representing Leucoagaricus section Rubrotincti subgenus Sericeomyces.
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http://dx.doi.org/10.1080/00275514.2018.1439651DOI Listing
January 2019

The genus in Pakistan with the description of two new species.

MycoKeys 2018 27(30):41-60. Epub 2018 Feb 27.

Department of Botany, University of the Punjab, Lahore 54590, Pakistan.

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 and 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 1 gene () show that the new taxa are clustered in a clade formed by the members of section Parasola of genus . 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. 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, and were also studied. Morphological descriptions for the new species and comparison with known species are provided. Our observations highlight the diversity of in northern Pakistan and further document the need for additional systematic focus on the region's fungi.
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http://dx.doi.org/10.3897/mycokeys.30.21430DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5904494PMC
February 2018

Bulgariella pulla, a Leotiomycete of uncertain placement, with an uncommon type of ascus opening.

Mycologia 2017 14;109(6):900-911. Epub 2018 Mar 14.

a Farlow Herbarium, Harvard University , 22 Divinity Avenue, Cambridge , Massachusetts 02138.

Bulgariella pulla (Leotiomycetes) is redescribed with the addition of characters of the ascus, spores, and habitat that were previously unconsidered. The ascus dehiscence mechanism in Bulgariella is unusual among Leotiomycetes. In this genus, asci lack a pore and open by splitting to form valves. Phylogenetic analyses of partial sequences of translation elongation factor 1-α (TEF1-α), the second largest subunit of RNA polymerase II (RPB2), and the 18S and 28S nuc rRNA genes determined that Bulgariella belongs within Leotiomycetes but without conclusive assignment to an order or family. A comparison of the nuc rDNA internal transcribed spacers 1 and 2 plus the 5.8S gene (ITS) determined that Bulgariella isolates from the USA, Norway, and Sweden had 100% sequence similarity, and an isolate from Chile had 99.3% similarity with these isolates. These results support the proposition that these collections represent a single species, B. pulla. Bulgariella sphaerospora, a more recently described species, is confirmed as conspecific with B. pulla.
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http://dx.doi.org/10.1080/00275514.2017.1418590DOI Listing
November 2018

Richard Paul Korf (1925-2016).

Mycologia 2017 May-Jun;109(3):529-534. Epub 2017 Jul 31.

f State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences , No. 1 Bei-Chen-Xi Road, Chaoyang District, Beijing , 100101 , China.

"With poetry, the tune is in the words themselves-and once you begin to hear it, it will stay with you." Richard P. Korf, notes to his narration of John Brown's Body.
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http://dx.doi.org/10.1080/00275514.2017.1360126DOI Listing
May 2018

Parasites of parasites of bats: Laboulbeniales (Fungi: Ascomycota) on bat flies (Diptera: Nycteribiidae) in central Europe.

Parasit Vectors 2017 02 21;10(1):96. Epub 2017 Feb 21.

Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, Massachusetts, USA.

Background: Bat flies (Streblidae and Nycteribiidae) are among the most specialized families of the order Diptera. Members of these two related families have an obligate ectoparasitic lifestyle on bats, and they are known disease vectors for their hosts. However, bat flies have their own ectoparasites: fungi of the order Laboulbeniales. In Europe, members of the Nycteribiidae are parasitized by four species belonging to the genus Arthrorhynchus. We carried out a systematic survey of the distribution and fungus-bat fly associations of the genus in central Europe (Hungary, Romania).

Results: We encountered the bat fly Nycteribia pedicularia and the fungus Arthrorhynchus eucampsipodae as new country records for Hungary. The following bat-bat fly associations are for the first time reported: Nycteribia kolenatii on Miniopterus schreibersii, Myotis blythii, Myotis capaccinii and Rhinolophus ferrumequinum; Penicillidia conspicua on Myotis daubentonii; and Phthiridium biarticulatum on Myotis capaccinii. Laboulbeniales infections were found on 45 of 1,494 screened bat flies (3.0%). We report two fungal species: Arthrorhynchus eucampsipodae on Nycteribia schmidlii, and A. nycteribiae on N. schmidlii, Penicillidia conspicua, and P. dufourii. Penicillidia conspicua was infected with Laboulbeniales most frequently (25%, n = 152), followed by N. schmidlii (3.1%, n = 159) and P. dufourii (2.0%, n = 102). Laboulbeniales seem to prefer female bat fly hosts to males. We think this might be due to a combination of factors: female bat flies have a longer life span, while during pregnancy female bat flies are significantly larger than males and accumulate an excess of fat reserves. Finally, ribosomal DNA sequences for A. nycteribiae are presented.

Conclusions: We screened ectoparasitic bat flies from Hungary and Romania for the presence of ectoparasitic Laboulbeniales fungi. Arthrorhynchus eucampsipodae and A. nycteribiae were found on three species of bat flies. This study extends geographical and host ranges of both bat flies and Laboulbeniales fungi. The sequence data generated in this work contribute to molecular phylogenetic studies of the order Laboulbeniales. Our survey shows a complex network of bats, bat flies and Laboulbeniales fungi, of which the hyperparasitic fungi are rare and species-poor. Their host insects, on the other hand, are relatively abundant and diverse.
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http://dx.doi.org/10.1186/s13071-017-2022-yDOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5320862PMC
February 2017

A molecular and morphological re-examination of the generic limits of truffles in the tarzetta-geopyxis lineage - Densocarpa, Hydnocystis, and Paurocotylis.

Fungal Biol 2017 03 13;121(3):264-284. Epub 2017 Jan 13.

Department of Plant Biology, University of Minnesota, 1445 Gortner Avenue, St. Paul, MN 55108, USA; Department of Plant Pathology, University of Florida, Gainesville, FL 32611, USA. Electronic address:

Truffle species within the /tarzetta-geopyxis lineage share smooth, globose, hyaline spores, but differ in the amount of convolution of hymenia in ascomata. The relationships among truffle species in this lineage have historically been confused. Phylogenetic analyses of the ITS and 28S nuclear ribosomal DNA from recently collected members of the /tarzetta-geopyxis lineage from Asia, Austral Asia, North America, and South America prompted a reinvestigation of species and generic limits in the truffle genera Hydnocystis, Paurocotylis, and Stephensia. Our analyses support emendations of Hydnocystis and Paurocotylis, abandonment of Stephensia and the resurrection of the genus Densocarpa. Nomenclatural changes include the transfer of Stephensia bombycina to Hydnocystis, the transfer of Hydnocystis singeri and Stephensia bynumii to Paurocotylis, the reinstatement of Densocarpa for Stephensia shanori and transfer of Stephensia crocea to Densocarpa. This is the first detection of the genus Paurocotylis in the Americas. We describe three new species, Hydnocystis transitoria from North America, Paurocotylis patagonica from South America, and Paurocotylis watlingii from Australia. Our work highlights the unexplored diversity, morphological plasticity, and remaining taxonomic problems among truffles in the /tarzetta-geopyxis lineage.
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http://dx.doi.org/10.1016/j.funbio.2016.12.004DOI Listing
March 2017

Competing sexual-asexual generic names of and recommendations for use.

IMA Fungus 2016 Dec 23;7(2):285-288. Epub 2016 Nov 23.

Swedish Museum of Natural History, Department of Botany, P.O. Box 50007, SE-104 05 Stockholm, Sweden.

Following the change that eliminated dual naming of sexual and asexual morphs of fungi, generic names of have been evaluated to determine which of the competing names should be recommended for use. Evaluation is based on congruence of type species to determine if the names are congeneric and which name is most commonly cited as well as priority. In the six pairs of generic names were determined to compete. In all cases the older name, representing the sexual morph, is recommended for use, specifically rather than , rather than , rather than , rather than , rather than , and rather than .
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http://dx.doi.org/10.5598/imafungus.2016.07.02.08DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5159599PMC
December 2016

Mycorrhizal detection of native and non-native truffles in a historic arboretum and the discovery of a new North American species, Tuber arnoldianum sp. nov.

Mycorrhiza 2016 Oct 10;26(7):781-92. Epub 2016 Jun 10.

Department of Organismic and Evolutionary Biology, Farlow Herbarium, Harvard University, 22 Divinity Ave, Cambridge, MA, 02138-2020, USA.

During a study comparing the ectomycorrhizal root communities in a native forest with those at the Arnold Arboretum in Massachusetts (USA), the European species Tuber borchii was detected on the roots of a native red oak in the arboretum over two successive years. Since T. borchii is an economically important edible truffle native to Europe, we conducted a search of other roots in the arboretum to determine the extent of colonization. We also wanted to determine whether other non-native Tuber species had been inadvertently introduced into this 140-year-old Arboretum because many trees were imported into the site with intact soil and roots prior to the 1921 USDA ban on these horticultural practices in the USA. While T. borchii was not found on other trees, seven other native and exotic Tuber species were detected. Among the North American Tuber species detected from ectomycorrhizae, we also collected ascomata of a previously unknown species described here as Tuber arnoldianum. This new species was found colonizing both native and non-native tree roots. Other ectomycorrhizal taxa that were detected included basidiomycetes in the genera Amanita, Russula, Tomentella, and ascomycetes belonging to Pachyphlodes, Helvella, Genea, and Trichophaea. We clarify the phylogenetic relationships of each of the Tuber species detected in this study, and we discuss their distribution on both native and non-native host trees.
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http://dx.doi.org/10.1007/s00572-016-0713-4DOI Listing
October 2016

Fireworks under the microscope: a spectacular new species of Zodiomyces from the Thaxter collection.

Mycologia 2016 Jul-Aug;108(4):709-15. Epub 2016 Apr 7.

Farlow Reference Library and Herbarium of Cryptogamic Botany, Harvard University, 22 Divinity Avenue, Cambridge Massachusetts 02138

A new species of Zodiomyces (Ascomycota, Laboulbeniales) is described, Z. rhizophorus, parasitic on a hydrophilid beetle (Coleoptera, Hydrophilidae) from Trinidad. This species was discovered during the examination of the slides of Laboulbeniales made by Roland Thaxter. It is characterized by numerous long, slender, multicellular and multiseriate outgrowths at the base of the receptacle. Thaxter's outstanding illustrations have set a standard in the field of mycology; we provide a review of the methods he employed in the preparation of these illustrations.
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http://dx.doi.org/10.3852/15-148DOI Listing
January 2018

Bringing Laboulbeniales into the 21st century: enhanced techniques for extraction and PCR amplification of DNA from minute ectoparasitic fungi.

IMA Fungus 2015 Dec 4;6(2):363-72. Epub 2015 Nov 4.

Department of Organismic and Evolutionary Biology, Harvard University, 22 Divinity Avenue, Cambridge, MA 02138, USA;; Farlow Reference Library and Herbarium of Cryptogamic Botany, Harvard University, 22 Divinity Avenue, Cambridge, MA 02138, USA.

Laboulbeniales is one of the most peculiar orders of Ascomycota. These fungi are characterized by an ectoparasitic life-style on arthropods, determinate growth, lack of an asexual stage, high species richness, and intractability to culture. The order Laboulbeniales, sister to Pyxidiophorales, has only recently been assigned a separate class, the Laboulbeniomycetes, based on very few ribosomal DNA sequences. So far, DNA isolations and PCR amplifications have proven difficult. Here, we provide details of isolation techniques and the application of commercially available kits that enable efficient and reliable genetic analyses of these fungi. We provide 43 newly generated Laboulbeniales ribosomal DNA sequences, among which are the first published sequences for species in the genera Gloeandromyces, Herpomyces, Laboulbenia, Monoicomyces, and Polyandromyces. DNA extractions were possible using from 1 to 30 thalli from hosts preserved in ethanol (70-100 %). In two cases, we successfully isolated DNA from thalli on dried insect collections. Laboulbeniales molecular systematics could be substantially enhanced through these improved methods by allowing more complete sampling of both taxa and gene regions.
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http://dx.doi.org/10.5598/imafungus.2015.06.02.08DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4681260PMC
December 2015

Multigene molecular phylogeny and biogeographic diversification of the earth tongue fungi in the genera Cudonia and Spathularia (Rhytismatales, Ascomycota).

PLoS One 2014 1;9(8):e103457. Epub 2014 Aug 1.

Department of Plant Pathology, University of Florida, Gainesville, Florida, United States of America.

The family Cudoniaceae (Rhytismatales, Ascomycota) was erected to accommodate the "earth tongue fungi" in the genera Cudonia and Spathularia. There have been no recent taxonomic studies of these genera, and the evolutionary relationships within and among these fungi are largely unknown. Here we explore the molecular phylogenetic relationships within Cudonia and Spathularia using maximum likelihood and Bayesian inference analyses based on 111 collections from across the Northern Hemisphere. Phylogenies based on the combined data from ITS, nrLSU, rpb2 and tef-1α sequences support the monophyly of three main clades, the /flavida, /velutipes, and /cudonia clades. The genus Cudonia and the family Cudoniaceae are supported as monophyletic groups, while the genus Spathularia is not monophyletic. Although Cudoniaceae is monophyletic, our analyses agree with previous studies that this family is nested within the Rhytismataceae. Our phylogenetic analyses circumscribes 32 species-level clades, including the putative recognition of 23 undescribed phylogenetic species. Our molecular phylogeny also revealed an unexpectedly high species diversity of Cudonia and Spathularia in eastern Asia, with 16 (out of 21) species-level clades of Cudonia and 8 (out of 11) species-level clades of Spathularia. We estimate that the divergence time of the Cudoniaceae was in the Paleogene approximately 28 Million years ago (Mya) and that the ancestral area for this group of fungi was in Eastern Asia based on the current data. We hypothesize that the large-scale geological and climatic events in Oligocene (e.g. the global cooling and the uplift of the Tibetan plateau) may have triggered evolutionary radiations in this group of fungi in East Asia. This work provides a foundation for future studies on the phylogeny, diversity, and evolution of Cudonia and Spathularia and highlights the need for more molecular studies on collections from Europe and North America.
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http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0103457PLOS
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4118880PMC
April 2015

Placement of the genus Angelina within Rhytismatales and observations of Angelina rufescens.

Mycologia 2014 Jan-Feb;106(1):154-62

Farlow Herbarium of Harvard University, 22 Divinity Avenue, Cambridge, Massachusetts 02138.

Angelina rufescens is placed within the core clade of Rhytismatales (Leotiomycetes, Pezizomycotina, Ascomycota) based on analysis of LSU and mtSSU rDNA. The only species in the genus, it produces distinctive ascomata that reoccur annually on wood and on the remains of its own previous fructifications, forming dense conglomerations of interlocking longitudinally elongated apothecia with gray hymenia. Known collections and references of A. rufescens indicate that it is endemic to eastern and central United States. Morphological and cultural characters are described with notes on ascomata development. No mitospores were observed in field collections or in culture. Lectotypes are designated for Hysterium rufescens and its synonym Ascobolus conglomeratus. Angelina rufescens is illustrated here for the first time in the taxonomic literature.
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http://dx.doi.org/10.3852/13-174DOI Listing
April 2014

A phylogeny of the highly diverse cup-fungus family Pyronemataceae (Pezizomycetes, Ascomycota) clarifies relationships and evolution of selected life history traits.

Mol Phylogenet Evol 2013 May 9;67(2):311-35. Epub 2013 Feb 9.

Department of Organismic and Evolutionary Biology, Harvard University, 22 Divinity Ave., Cambridge, MA 02138, USA.

Pyronemataceae is the largest and most heterogeneous family of Pezizomycetes. It is morphologically and ecologically highly diverse, comprising saprobic, ectomycorrhizal, bryosymbiotic and parasitic species, occurring in a broad range of habitats (on soil, burnt ground, debris, wood, dung and inside living bryophytes, plants and lichens). To assess the monophyly of Pyronemataceae and provide a phylogenetic hypothesis of the group, we compiled a four-gene dataset including one nuclear ribosomal and three protein-coding genes for 132 distinct Pezizomycetes species (4437 nucleotides with all markers available for 80% of the total 142 included taxa). This is the most comprehensive molecular phylogeny of Pyronemataceae, and Pezizomycetes, to date. Three hundred ninety-four new sequences were generated during this project, with the following numbers for each gene: RPB1 (124), RPB2 (99), EF-1α (120) and LSU rDNA (51). The dataset includes 93 unique species from 40 genera of Pyronemataceae, and 34 species from 25 genera representing an additional 12 families of the class. Parsimony, maximum likelihood and Bayesian analyses suggest that Pyronemataceae is paraphyletic due to the nesting of both Ascodesmidaceae and Glaziellaceae within the family. Four lineages with taxa currently classified in the family, the Boubovia, Geopyxis, Pseudombrophila and Pulvinula lineages, form a monophyletic group with Ascodesmidaceae and Glaziellaceae. We advocate the exclusion of these four lineages in order to recognize a monophyletic Pyronemataceae. The genus Coprotus (Thelebolales, Leotiomycetes) is shown to belong to Pezizomycetes, forming a strongly supported monophyletic group with Boubovia. Ten strongly supported lineages are identified within Pyronemataceae s. str. Of these, the Pyropyxis and Otidea lineages are identified as successive sister lineages to the rest of Pyronemataceae s. str. The highly reduced (gymnohymenial) Monascella is shown to belong to Pezizomycetes and is for the first time suggested to be closely related to the cleistothecial Warcupia, as a sister group to the primarily apothecial Otidea. None of the lineages of pyronemataceous taxa identified here correspond to previous families or subfamily classifications. Ancestral character state reconstructions (ASR) using a Bayesian approach support that the ancestors of Pezizomycetes and Pyronemataceae were soil inhabiting and saprobic. Ectomycorrhizae have arisen within both lineages A, B and C of Pezizomycetes and are suggested to have evolved independently seven to eight times within Pyronemataceae s. l., whereas an obligate bryosymbiotic lifestyle has arisen only twice. No reversals to a free-living, saprobic lifestyle have happened from symbiotic or parasitic Pyronemataceae. Specializations to various substrates (e.g. burnt ground and dung) are suggested to have occurred several times in mainly saprobic lineages. Although carotenoids in the apothecia are shown to have arisen at least four times in Pezizomycetes, the ancestor of Pyronemataceae s. str., excluding the Pyropyxis and Otidea lineages, most likely produced carotenoids, which were then subsequently lost in some clades (- and possibly gained again). Excipular hairs were found with a high probability to be absent from apothecia in the deepest nodes of Pezizomycetes and in the ancestor of Pyronemataceae s. str. True hairs are restricted to the core group of Pyronemataceae s. str., but are also found in Lasiobolus (Ascodesmidaceae), the Pseudombrophila lineage and the clade of Chorioactidaceae, Sarcoscyphaceae and Sarcosomataceae. The number of gains and losses of true hairs within Pyronemataceae s. str., however, remains uncertain. The ASR of ascospore guttulation under binary coding (present or absent) indicates that this character is fast evolving and prone to shifts.
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http://dx.doi.org/10.1016/j.ympev.2013.01.014DOI Listing
May 2013