Publications

2014
Haelewaters, D., Schilthuizen, M. & Pfister, D.H., 2014. On Diphymyces (Laboulbeniales, Ascomycota) in Malaysian Borneo. Plant Ecology and Evolution , 147 , pp. 93-100.Abstract

Background - Laboulbeniales (Fungi, Ascomycota) are microscopic ectoparasites of Arthropoda. Since many Laboulbeniales are known to be host-specific, wet tropical areas with high arthropod diversity are likely to house a high diversity of Laboulbeniales, as well. In this paper, we reveal the presence of the genus Diphymyces I.I.Tav. in Malaysian Borneo. After fieldwork in disturbed and pristine tropical rainforest in Borneo, representatives of this genus were discovered that did not match the description of any of the fourteen known species.Methods - Insects were collected with dry pitfall traps baited with Limburger cheese. Fungal material was studied and described using morphology-based methods.Key results - A new species, Diphymyces sabahensis Haelew. & Pfister, is described and illustrated. Hosts for this species are recently described taxa in Ptomaphaginus Portevin, 1914 (Coleoptera, Leiodidae, Cholevinae, Ptomaphagini), representing a new host genus for Diphymyces. Other, morphologically distinctive records of Diphymyces were found; on one host specimen they co-occur with D. sabahensis. These divergent thalli are remarkable in their restricted occurrence on the metatibiae, and thus may represent a morphological variant of D. sabahensis. A review of all described species of Diphymyces, with hosts and geographical distribution, is also presented in tabulated form.

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Haelewaters, D., et al., 2014. Erratum to: Hesperomyces virescens (Fungi, Ascomycota, Laboulbeniales) attacking Harmonia axyridis (Coleoptera, Coccinellidae) in its native range. Chinese Science Bulletin , 59 , pp. 1482-1482.Abstract

This study intended to find data on obligate ec- toparasitic Laboulbeniales (Fungi, Ascomycota) on ChineseHarmonia axyridis (Coleoptera, Coccinellidae). After having screened over four thousand dried specimens of H. axyridisand close relatives, we present the first (historical) record of Chinese H. axyridis infected with Hesperomyces virescens. We suggest that H. virescens is a historically globally dis- tributed species and hypothesize that (native) infection was lost when H. axyridis was introduced in North America.

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Haelewaters, D., et al., 2014. Hesperomyces virescens (Fungi, Ascomycota, Laboulbeniales) attacking Harmonia axyridis (Coleoptera, Coccinellidae) in its native range. Chinese Science Bulletin , 59 , pp. 528-532.Abstract

This study intended to find data on obligate ec- toparasitic Laboulbeniales (Fungi, Ascomycota) on ChineseHarmonia axyridis (Coleoptera, Coccinellidae). After having screened over four thousand dried specimens of H. axyridisand close relatives, we present the first (historical) record of Chinese H. axyridis infected with Hesperomyces virescens. We suggest that H. virescens is a historically globally dis- tributed species and hypothesize that (native) infection was lost when H. axyridis was introduced in North America.

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Medardi, G., et al., 2014. Morphological and molecular study of Peziza emileia and P-howsei, two distinct taxa. Mycological Progress , 13 , pp. 1227-1234.Abstract

Peziza emileia Cooke and P. howsei Roze & Boud. are compared here; they are morphologically very similar, but the internal transcribed spacer (ITS) sequence is unique for each of the two species. Furthermore, since the holotype of P. emileia deposited in Kew (K) contains an unidentified fungus and the holotype of P. howsei in Paris (PC) no longer exists, we provide lectotypification and epitypification for both taxa.

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Ge, Z.W., et al., 2014. Multigene Molecular Phylogeny and Biogeographic Diversification of the Earth Tongue Fungi in the Genera Cudonia and Spathularia (Rhytismatales, Ascomycota). Plos One , 9.Abstract

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-1 alpha 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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Karakehian, J.M., LoBuglio, K.F. & Pfister, D.H., 2014. Placement of the genus Angelina within Rhytismatales and observations of Angelina rufescens. Mycologia , 106 , pp. 154-162.Abstract

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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2013
Pfister, D.H., et al., 2013. The Caloscyphaceae (Pezizomycetes, Ascomycota), with a new genus. Mycological Progress , 12 , pp. 667-674.Abstract

The family Caloscyphaceae with a single genus, Caloscypha, has been considered to include a single species, C. fulgens. Study of an overlooked second species, Caloscypha incarnata from North Africa and Italy, using SSU, LSU rDNA, and morphology allows placement of this species in a new genus, Kallistoskypha, in the Caloscyphaceae. This fungus is found in association with Eucalyptus species. The species was recently redescribed from Spain under the name Marcelleina parvispora. Caloscypha fulgens, the type species of the genus Caloscypha, shows sequence variation from across its range.

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Healy, R.A., et al., 2013. High diversity and widespread occurrence of mitotic spore mats in ectomycorrhizal Pezizales. Molecular Ecology , 22 , pp. 1717-1732.Abstract

Fungal mitospores may function as dispersal units and/ or spermatia and thus play a role in distribution and/or mating of species that produce them. Mitospore production in ectomycorrhizal (EcM) Pezizales is rarely reported, but here we document mitospore production by a high diversity of EcM Pezizales on three continents, in both hemispheres. We sequenced the internal transcribed spacer (ITS) and partial large subunit (LSU) nuclear rDNA from 292 spore mats (visible mitospore clumps) collected in Argentina, Chile, China, Mexico and the USA between 2009 and 2012. We collated spore mat ITS sequences with 105 fruit body and 47 EcM root sequences to generate operational taxonomic units (OTUs). Phylogenetic inferences were made through analyses of both molecular data sets. A total of 48 OTUs from spore mats represented six independent EcM Pezizales lineages and included truffles and cup fungi. Three clades of seven OTUs have no known meiospore stage. Mitospores failed to germinate on sterile media, or form ectomycorrhizas on Quercus, Pinus and Populus seedlings, consistent with a hypothesized role of spermatia. The broad geographic range, high frequency and phylogenetic diversity of spore mats produced by EcM Pezizales suggests that a mitospore stage is important for many species in this group in terms of mating, reproduction and/or dispersal.

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Bonito, G., et al., 2013. Historical Biogeography and Diversification of Truffles in the Tuberaceae and Their Newly Identified Southern Hemisphere Sister Lineage. Plos One , 8.Abstract

Truffles have evolved from epigeous (aboveground) ancestors in nearly every major lineage of fleshy fungi. Because accelerated rates of morphological evolution accompany the transition to the truffle form, closely related epigeous ancestors remain unknown for most truffle lineages. This is the case for the quintessential truffle genus Tuber, which includes species with socio-economic importance and esteemed culinary attributes. Ecologically, Tuber spp. form obligate mycorrhizal symbioses with diverse species of plant hosts including pines, oaks, poplars, orchids, and commercially important trees such as hazelnut and pecan. Unfortunately, limited geographic sampling and inconclusive phylogenetic relationships have obscured our understanding of their origin, biogeography, and diversification. To address this problem, we present a global sampling of Tuberaceae based on DNA sequence data from four loci for phylogenetic inference and molecular dating. Our well-resolved Tuberaceae phylogeny shows high levels of regional and continental endemism. We also identify a previously unknown epigeous member of the Tuberaceae - the South American cup-fungus Nothojafnea thaxteri (E.K. Cash) Gamundi. Phylogenetic resolution was further improved through the inclusion of a previously unrecognized Southern hemisphere sister group of the Tuberaceae. This morphologically diverse assemblage of species includes truffle (e.g. Gymnohydnotrya spp.) and non-truffle forms that are endemic to Australia and South America. Southern hemisphere taxa appear to have diverged more recently than the Northern hemisphere lineages. Our analysis of the Tuberaceae suggests that Tuber evolved from an epigeous ancestor. Molecular dating estimates Tuberaceae divergence in the late Jurassic (similar to 156 million years ago), with subsequent radiations in the Cretaceous and Paleogene. Intra-continental diversification, limited long-distance dispersal, and ecological adaptations help to explain patterns of truffle evolution and biodiversity.

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Haelewaters, D., et al., 2013. The Laboulbeniales of the Boston Harbor Islands. Phytopathology , 103 , pp. 54-54.Abstract

This paper, based on a recent comprehensive sampling of insects, is the rstreport of Laboulbeniales from the New England region since the 1930s. We present 7 new records of laboulbenialean parasites on Staphylinidae (rove beetles) and Coccinellidae (lady beetles) from the Boston Harbor Islands National Recreation Area. These are Clonophoro- myces nipponicus Terada & I.I. Tav., Hesperomyces virescens Thaxt., Ilyomyces cf. maireiF. Picard, Laboulbenia philonthi Thaxt., Peyritschiella protea Thaxt., Stichomyces conoso- matis Thaxt., and Teratomyces actobii Thaxt. One of these parasite species, C. nipponicus, has not been found previously outside of its type locality in Japan. Examination of Roland Thaxter’s 1891–1932 slides led to the designation of lectotypes for L. philonthi, P. protea,S. conosomatis, and T. actobii. The following synonymy is established: Teratomyces brevi- caulis Thaxt. = T. actobii. In addition, we discovered new localities for H. virescens (from Canada, Cuba, Guatemala, and Japan) and L. philonthi (from Canada, Grenada, Panama, Trinidad, and Venezuela).

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Hansen, K., et al., 2013. A phylogeny of the highly diverse cup-fungus family Pyronemataceae (Pezizomycetes, Ascomycota) clarifies relationships and evolution of selected life history traits. Molecular Phylogenetics and Evolution , 67 , pp. 311-335.Abstract

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-1 alpha (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. O 2013 Elsevier Inc. All rights reserved.

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Choi, Y.J., et al., 2013. Scutellinia (Pezizales) in Korea, with a new species and eight new records. Nova Hedwigia , 97 , pp. 457-476.Abstract

Eleven species of the genus Scutellinia (Pyronemataceae, Pezizales) are recognized in Korea by analysis of macro- and micro-morphological characteristics, substrates and geographical distributions. Eight species are recorded new to Korea, namely, S. ahmadiopsis, S. badio-berbis, S. colensoi, S. jilinensis, S. nigrohirtula, S. olivascens, S. setosa and S. patagonica. Based on the exceptional length of marginal hairs and tuberculate ascospore wall ornamentation, two Korean specimens occurring on wood are described as a new species. Infraspecific morphological variations among collections within S. scutellata and S. patagonica were found. These are tentatively treated as species complexes. The highly questionable occurrence of S. setosa in Asia was confirmed. The importance of characteristics of paraphyses for species delimitation is highlighted. Descriptions and taxonomic notes of the recognized species of Scutellinia are provided with a taxonomic key, illustrations and photographs of these species from Korea.

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Haelewaters, D. & Pfister, D.H., 2013. Two new species of Diphymyces (Fungi, Laboulbeniales) from Borneo. Phytopathology , 103 , pp. 54-54.
2012
Romero, A.I., LoBuglio, K.F. & Pfister, D.H., 2012. Rickiella edulis and its phylogenetic relationships within Sarcoscyphaceae. Kurtziana , (37) , pp. 79-89.Abstract

Rickiella edulis is reported from Argentina for the rst time and is documented with photographs of fresh specimens and molecular data. Previously the species was known as R. transiens (= Phillipsia transiens)and was reported from southern Brazil and Paraguay. Phylogenetic analyses based on SSU rDNA and LSU rDNA shows its placement in a monophyletic family, the Sarcoscyphaceae. The relationship ofRickiella, Phillipsia and Nanoscypha however could not be resolved from phylogenetic analyses of the ITS, SSU, and LSU rDNA sequences. The excipular tissue of Rickiella is shot through with regularly spaced channels and cavities. Because of this feature, the genus Rickiella is recognized as distinct from Phillipsia. Phillipsia and Nanoscypha are morphologically distinct but diversity within Phillipsiaremains a topic for further research. A new tribe in the Sarcoscyphaceae is proposed to accommodate the genus Wynnea.

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Medardi, G., et al., 2012. Clarification of Peziza fimeti with notes on P. varia collections on dung. Mycotaxon , 121 , pp. 465-476.Abstract

The smooth-spored species inhabiting dung, mainly of the Peziza fimeti group, were studied morphologically and through ITS sequence comparison. The results established that Peziza varia is also able to fruit on dung, clarifying a long-standing situation regarding two conflicting interpretations given in P. fimeti literature.

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Lantieri, A., Smith, M.E. & Pfister, D.H., 2012. A new species of Ruhlandiella (Pezizaceae) from Italy. Mycological Progress , 11 , pp. 509-513.Abstract

This paper describes the newly discovered species Ruhlandiella peregrina. Full description and illustrations of macro- and micromorphological features of the new taxon are provided. This species differs from other described species in ascus and ascospore size and in the crested and ridged ornamentation of ascospores. As is the case in two other similar species, the asci of R. peregrina do not becoming blue in iodine solutions.

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2010
Peterson, K.R., Pfister, D.H. & Bell, C.D., 2010. Cophylogeny and biogeography of the fungal parasite Cyttaria and its host Nothofagus, southern beech. Mycologia , 102 , pp. 1417-1425.Abstract

The obligate, biotrophic association among species of the fungal genus Cyttaria and their hosts in the plant genus Nothofagas often is cited as a classic example of cophylogeny and is one of the few cases in which the biogeography of a fungus is commonly mentioned or included in biogeographic analyses. In this study molecular and morphological data are used to examine hypotheses regarding the cophylogeny and biogeography of the 12 species of Cyttaria and their hosts, the 11 species of Nothofagas subgenera Lophozonia and Nothofagus. Our results indicate highly significant overall cophylogenetic structure, despite the fact that the associations between species of Cyttaria and Nothojagus usually do not correspond in a simple one to one relationship. Two major lineages of Cyttaria are confined to a single Nothofagus subgenus, a specificity that might. account for a minimum of two codivergences. We hypothesize other major codivergences. Numerous extinction also are assumed, as are an independent. parasite divergence followed by host switching to account for C. berteroi. Considering the historical association of Cyttaria and Nothofagus, our hypothesis may support the vicariance hypothesis for the trans-Antarctic distribution between Australasian and South American species of Cyttaria species hosted by subgenus Lophozonia. It also supports the hypothesis of transoceanic long distance dispersal to account for the relatively recent relationship between Australian and New Zealand Cyttaria species, which we estimate to have occurred 44.6-28.5 mya. Thus the history of these organisms is not only a reflection of the breakup of Gondwana but also of other events that have contributed to the distributions of many other southern hemisphere plants and fungi.

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Lantieri, A. & Pfister, D.H., 2010. A new species of Marcelleina from Italy. Mycotaxon , 111 , pp. 465-469.Abstract

Marcelleina mediterranea is described as a new species and is illustrated. It occurs on sandy soil among scattered mosses, in Southeast Sicily (Italy). It differs from other species in size and ornamentation of ascospores. Its ecology and taxonomical relationships are examined.

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Peterson, K.R. & Pfister, D.H., 2010. Phylogeny of Cyttaria inferred from nuclear and mitochondrial sequence and morphological data. Mycologia , 102 , pp. 1398-1416.Abstract

Cyttaria species (Leotiomycetes, Cyttariales) are obligate, biotrophic associates of Nothofagus (Hamamelididae, Nothofagaceae), the southern beech. As such Cyttaria species are restricted to the southern hemisphere, inhabiting southern South America (Argentina and Chile) and southeastern Australasia (southeastern Australia including Tasmania, and New Zealand). The relationship of Cyttaria to other Leotiomycetes and the relationships among species of Cyttaria were investigated with newly generated sequences of partial nucSSU, nucLSU and mitSSU rRNA, as well as TEE] sequence data and morphological data. Results found Cyttaria to be defined as a strongly supported clade. There is evidence for a close relationship between Cyttaria and these members of the Helotiales: Cordierites, certain Encoelia spp., Ionomidotis and to a lesser extent Chlorociboria. Order Cyttariales is supported by molecular data, as well as by the unique endostromatic apothecia, lack of chitin and highly specific habit of Cyttaria species. Twelve Cyttaria species are hypothesized, including all 11 currently accepted species plus an undescribed species that accommodates specimens known in New Zealand by the misapplied name C. gunnii, as revealed by molecular data. Thus the name C. gunnii sensu stricto is reserved for specimens occurring on N. cunninghamii in Australia, including Tasmania. Morphological data now support the continued recognition of C. septentrionalis as a species separate from C. gunnii. Three major clades are identified within Cyttaria: one in South America hosted by subgenus Nothofagus, another in South America hosted by subgenera Nothofagus and Lophozonia, and a third in South America and Australasia hosted by subgenus Lophozonia, thus producing a non-monophyletic grade of South American species and a monophyletic clade of Australasian species, including monophyletic Australian and New Zealand clades. Cyttaria species do not sort into clades according to their associations with subgenera Lophozonia and Nothofagus.

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LoBuglio, K.F. & Pfister, D.H., 2010. Placement of Medeolaria farlowii in the Leotiomycetes, and comments on sampling within the class. Mycological Progress , 9 , pp. 361-368.Abstract

Medeolaria farlowii, an ascomycetous parasite of Medeola virginiana, has been included as the only member of the family Medeolariaceae and order Medeolariales. Its assignment within the Ascomycota has been problematic due to the lack of distinctive ascomatal form and ascus morphology. Asci are formed in a loosely organized hymenium on hypertrophic portions of Medeola virginiana stems. Phylogenetic analyses of nuclear 5.8S, large subunit, and small subunit rDNA gene sequences place M. farlowii in the Leotiomycetes with parsimony, Bayesian, and maximum likelihood analyses, but its position within this class remains unresolved. In general, limited taxon and gene sampling in the Leotiomycetes hampers placement of taxa within this class. A survey of available sequence data in the Leotiomycetes is given, and the implication for phylogenetic studies within the class is discussed. Medeolaria farlowii should be treated as a monotypic genus in the monotypic order Medeolariales, class Leotiomycetes.

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