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Elaphomyces readii is presented as a new species to science based on morphological and molecular evidence. The new species is characterized mainly by the pale brown rugose ascomata, yellowish to reddish brown hues in the proximity to the gleba, dark purple gleba and ascospores 20–57 µm in diameter. Elaphomyces readii is morphologically similar to E. papillatus and E. muricatus but differs in the ascospore size and phylogenetical location. Further, E. readii is sold in traditional markets in the state of Oaxaca, southeastern Mexico as a medicinal fungus consumed to heal wounds, internal bleeding, and external injuries. Morphological descriptions, SEM ornamentation spores and nrITS-based phylogenetic analysis using maximum parsimony, maximum likelihood method and Bayesian inference support the proposal of this new species.

Elaphomyces readii Pérez-Moreno & de la Fuente, sp. nov. (Fig. 2). Mycobank no. 846810 Diagnosis: — Elaphomyces readii is mainly characterized by the peridium yellowish to reddish brown in the outer part, then whitish, with yellowish to reddish brown hues in the proximity to the gleba and ascospores ranging 20–57 µm in diameter, ornamented with small flat papillae, sometimes coalescing forming a subreticulum The macro- and micromorphological characteristics in conjunction with the nrITS-based phylogenetic analysis support the proposal of this new species. Holotype: — MEXICO. Oaxaca: Tlaxiaco City, state of Oaxaca, purchased in a traditional market, 17° 07’ 17. 21’’ N, 97° 34’ 97. 50’’ W, 2040 m a.s.l., 17 July 2016, Pérez-Moreno J. (2501-ITCV, holotype). Etymology: —This species is dedicated, as “ readii ”, in honor to Professor Sir David J. Read, eminent mycologist, Emeritus Professor of the University of Sheffield, England, Secretary of the Royal Society (London) and a giant scientist in the study of the mycorrhizal symbioses (like the giant spores of the new species, averaging 52.33 µm in diameter), with more than 55 years of experience contributing to the understanding of the ecophysiology of mycorrhizal symbioses, and was knighted in 2007 by the Queen of England for his outstanding scientific contributions. Description: — Ascomata 20–45 mm, globose to sub globose, slightly flattened when mature, sometimes lobulate, yellowish to reddish brown (5F4–5E4), dry, finely ornamented with flat papillae, crowded (2–3 papillae per mm), without mycelial strands. Peridium up to 1 to 1.5 mm thick, yellowish in the outer part, then whitish, yellowish to reddish brown near the gleba, firm, hard when dried, sometimes bruising reddish or orange when cut. Gleba pulverulent, white, then purple (18F8), dark purple (18F6) to blackish when mature, with some pale fibrils radially arranged, odor and taste fungoid. Peridium composed of three layers. Exoperidium 100–230 µm thick, composed of prostrate and horizontally arranged hyphae, 4–7 µm in diameter, hyaline or with golden content, septate, thin-walled, forming irregular pyramids of 400–650 µm. Mesoperidium 800–1200 µm thick, composed of interwoven tubulous hyphae, compacted, 3-5 µm in diameter, thin-walled. Endoperidium 250–450 µm thick, similar in structure to the mesoperidium, with tubulous interwoven hyphae, 5–15 µm in diameter, thick-walled, some with yellowish to reddish brown incrustations in the cell walls. Asci 45–62 × 38–42 µm, ellipsoid, rarely globose, hyaline, 6–8 spored, some with a pedicel up to 7 µm long, thick walled. Ascospores 20–57 µm (Average=52.33 µm, n=50), globose, with slender spines, crowded, reaching 3-5 µm long, sometimes converging in small ridges, forming a sub-reticulum or alveoli up to 5 µm in diameter, brown in 3% KOH, some with a big guttula, thick-walled. Habitat, habit, distribution: —Solitary to disperse. So far only known from the type community in Tlaxiaco municipality, growing under Pinus spp. from the beginning of July to the end of August, having a short distribution in the middle of the rainy season phenology. Specimens examined: — MEXICO. Oaxaca: Tlaxiaco Municipality, purchased in a traditional market, 17° 07’ 17. 21’’ N, 97° 34’ 97. 50’’ W, 2040 m, 20 August 2016, Pérez-Moreno J. (2501, 2502, 2503, 2504, 2505 ITCV)., Published as part of Fuente, Javier Isaac De La, Pérez-Moreno, Jesús, Martínez-Reyes, Magdalena, Ayala-Vásquez, Olivia, Martínez-González, César Ramiro & Aguirre-Acosta, Celia Elvira, 2023, Elaphomyces readii (Elaphomycetaceae, Eurotiomycetes), a new medicinal species of hypogeous fungus with biocultural importance from Mexico, pp. 241-250 in Phytotaxa 594 (4) on pages 245-247, DOI: 10.11646/phytotaxa.594.4.1, http://zenodo.org/record/7917021

Two new species of sequestrate fungi are described from south Mexico based on morphological and molecular evidences. Here we describe Elaphomyces castilloi characterized by the yellowish mycelial mat, dull blue gleba and ascospores of 9.7–11.5 µm; Entoloma secotioides is characterized by the secotioid basidiomata, sulcate, pale cream pileus, and basidiospores of 7–13 × 5–9 µm. Both species grow in montane cloud forest under Quercus sp. in the state of Chiapas, Mexico. Descriptions, photographs, and multilocus phylogeny for both species are presented.

The hypogeous, sequestrate ascomycete genus Elaphomyces is one of the oldest known truffle-like genera. Elaphomyces has a long history of consumption by animals in Europe and was formally described by Nees von Esenbeck in 1820 from Europe. Until recently most Elaphomyces specimens in North America were assigned names of European taxa due to lack of specialists working on this group and difficulty of using pre-modern species descriptions. It has recently been discovered that North America has a rich diversity of Elaphomyces species far beyond the four Elaphomyces species described from North America prior to 2012. We describe eight new Elaphomyces species (E. dalemurphyi, E. dunlapii, E. holtsii, E. lougehrigii, E. miketroutii, E. roodyi, E. stevemilleri and E. wazhazhensis) of eastern North America that were collected in habitats from Quebec, Canada south to Florida, USA, west to Texas and Iowa. The ranges of these species vary and with continued sampling may prove to be larger than we have established. Castellano has studied authentic material of all European Elaphomyces species published through 2016 and it is interesting to note that many Elaphomyces species from eastern North America have morphological similarities but with distinct morphological differences to a number of European Elaphomyces species.

The Eurotiales is a relatively large order of Ascomycetes with members frequently having positive and negative impact on human activities. Species within this order gain attention from various research fields such as food, indoor and medical mycology and biotechnology. In this article we give an overview of families and genera present in the Eurotiales and introduce an updated subgeneric, sectional and series classification for Aspergillus and Penicillium. Finally, a comprehensive list of accepted species in the Eurotiales is given. The classification of the Eurotiales at family and genus level is traditionally based on phenotypic characters, and this classification has since been challenged using sequence-based approaches. Here, we re-evaluated the relationships between families and genera of the Eurotiales using a nine-gene sequence dataset. Based on this analysis, the new family Penicillaginaceae is introduced and four known families are accepted: Aspergillaceae, Elaphomycetaceae, Thermoascaceae and Trichocomaceae. The Eurotiales includes 28 genera: 15 genera are accommodated in the Aspergillaceae (Aspergillago, Aspergillus, Evansstolkia, Hamigera, Leiothecium, Monascus, Penicilliopsis, Penicillium, Phialomyces, Pseudohamigera, Pseudopenicillium, Sclerocleista, Warcupiella, Xerochrysium and Xeromyces), eight in the Trichocomaceae (Acidotalaromyces, Ascospirella, Dendrosphaera, Rasamsonia, Sagenomella, Talaromyces, Thermomyces, Trichocoma), two in the Thermoascaceae (Paecilomyces, Thermoascus) and one in the Penicillaginaceae (Penicillago). The classification of the Elaphomycetaceae was not part of this study, but according to literature two genera are present in this family (Elaphomyces and Pseudotulostoma). The use of an infrageneric classification system has a long tradition in Aspergillus and Penicillium. Most recent taxonomic studies focused on the sectional level, resulting in a well-established sectional classification in these genera. In contrast, a series classification in Aspergillus and Penicillium is often outdated or lacking, but is still relevant, e.g., the allocation of a species to a series can be highly predictive in what functional characters the species might have and might be useful when using a phenotype-based identification. The majority of the series in Aspergillus and Penicillium are invalidly described and here we introduce a new series classification. Using a phylogenetic approach, often supported by phenotypic, physiologic and/or extrolite data, Aspergillus is subdivided in six subgenera, 27 sections (five new) and 75 series (73 new, one new combination), and Penicillium in two subgenera, 32 sections (seven new) and 89 series (57 new, six new combinations). Correct identification of species belonging to the Eurotiales is difficult, but crucial, as the species name is the linking pin to information. Lists of accepted species are a helpful aid for researchers to obtain a correct identification using the current taxonomic schemes. In the most recent list from 2014, 339 Aspergillus, 354 Penicillium and 88 Talaromyces species were accepted. These numbers increased significantly, and the current list includes 446 Aspergillus (32 % increase), 483 Penicillium (36 % increase) and 171 Talaromyces (94 % increase) species, showing the large diversity and high interest in these genera. We expanded this list with all genera and species belonging to the Eurotiales (except those belonging to Elaphomycetaceae). The list includes 1 187 species, distributed over 27 genera, and contains MycoBank numbers, collection numbers of type and ex-type cultures, subgenus, section and series classification data, information on the mode of reproduction, and GenBank accession numbers of ITS, beta-tubulin (BenA), calmodulin (CaM) and RNA polymerase II second largest subunit (RPB2) gene sequences.

Pseudotulostoma is a genus described for fungi with stipitate sporocarps that have an exposed gleba and a woody, volvate base. The two species that belong to this genus (P. volvatum and P. japonicum) form unusual epigeous ascomata that are atypical among the hypogeous members of the Elaphomycetaceae. The genus was first described from the Guiana Shield and was suggested to be restricted to rainforests dominated by the ectomycorrhizal tree Dicymbe corymbosa (Leguminosae-Detarioideae). Pseudotulostoma volvatum was also later described from Colombia in association with Pseudomonotes tropenbosii (Dipterocarpaceae- Monotoidea). Here we report a new occurrence of P. volvatum that is also the first occurrence of an ectomycorrhizal Ascomycota fungus in a native host plant in Brazil. The description includes images of the macroscopic and microscopic characteristics, a discussion of the distinctive features, and phylogenetic placement using the nLSU of this fungus among Elaphomyces species. This new collection (third known location) demonstrates that P. volvatum also occurs in a white-sand forest composed of the canopy tree Aldina heterophylla (Leguminosae-Papilionoideae). Thus, we provide additional information regarding P. volvatum that expands its known distribution.

The hypogeous ascomycete species, Elaphomyces decipiens Vittad., is reported as a new record from Turkey, based on the identification of the samples collected from Trabzon province. A brief description and photographs, related to the macroscopy and microscopy of the species, are provided., Toprak altı askomiset türü olan, Elaphomyces decipiens Vittad., Trabzon’dan toplanan örneklerin teşhis edilmesiyle, Türkiye’den yeni kayıt olarak rapor edilmiştir. Türün kısa bir betimlemesi ve makroskobi ve mikroskobisine ilişkin fotoğrafları verilmiştir.

We describe three new species of Elaphomyces from eastern North America. Of the three, Elaphomyces loebiae is the rarest, known only from North Carolina and South Carolina, and appears to associate primarily with ectomycorrhizal hardwoods but possibly also with conifers. Elaphomyces cibulae is widely distributed but disjunct from Florida, Mississippi, and North Carolina. Elaphomyces cibulae seems to primarily associate with Quercus species. Elaphomyces mitchelliae has the widest distribution of the three species, from Florida, Maryland, North Carolina, Virginia, and West Virginia, and appears to associate with either ectomycorrhizal hardwoods and/or conifers. In the course of comparing our new Elaphomyces species to previously described European species we discovered that E. persoonii var. minor is conspecific in all essential details with and thus a synonym of E. cyanosporus.

We describe five new species of Elaphomyces from Bartlett Experimental Forest, New Hampshire, USA (E. americanus, E. bartlettii, E. macrosporus, E. oreoides, and E. remickii) and revise the description of a sixth previously published species (E. verruculosus). Of the five new species, E. bartlettiii and E. remickii are only known from New Hampshire whereas E. americanus, E. macrosporus, and E. oreoides are widely distributed in eastern North America. Elaphomyces verruculosus is the most widespread and abundant Elaphomyces species in eastern North America with a distribution extending from eastern Canada south to northeastern Mexico. All six Elaphomyces species are putatively associated with Tsuga canadensis, a tree species in regional decline. For five of the six Elaphomyces species, we report partially consumed ascomata or rodent fecal samples containing spores, indicating that small mammals play a key role in dispersing these Elaphomyces species and that the Elaphomyces are an important part of the small mammals’ diet.

Thehypogeous Ascomycete species, Elaphomycesgranulatus isreported for the first time in Turkey. The morphological and microscopic charactersof the species recorded in Turkey are reported together with the localities ofcollection, and the photographs related to its macro and micromorphology areprovided., Toprakaltı Askomiset türü olan, Elaphomycesgranulatus Türkiye’den ilk kez rapor edilmiştir. Türkiye’den kaydedilentürün morfolojik ve mikroskobik özellikleri, toplanma lokaliteleri ile birlikteraporlanarak makro ve mikromorfolojisine ilişkin fotoğraflar verilmiştir.

First collections are reported for Elaphomyces species from Africa and Madagascar. On the basis of an ITS phylogeny, the authors question the monophyletic nature of family Elaphomycetaceae and of the genus Elaphomyces. The objective of this preliminary paper was not to propose a new phylogeny for Elaphomyces, but rather to draw attention to the very high dissimilarity among ITS sequences for Elaphomyces and to the unfortunate choice of species to represent the genus in most previous phylogenetic publications on Elaphomycetaceae and other cleistothecial ascomycetes. Our study highlights the need for examining the monophyly of this family and to verify the systematic status of Pseudotulostoma as a separate genus for stipitate species. Furthermore, there is an urgent need for an in-depth morphological study, combined with molecular sequencing of the studied taxa, to point out the phylogenetically informative characters of the discussed taxa.

Four species of the sequestrate fungal genus Elaphomyces are reported from New Zealand: Elaphomyces bollardii sp. nov. associated with Leptospermum spp. and Kunzea ericoides, E. luteicrustus sp. nov. associated with Nothofagus menziesii, E. putridus sp. nov. associated with Nothofagus spp., and an unnamed species associated with Nothofagus spp.

The sequestrate false truffles Elaphomyces favosus, E. iuppitercellus, and E. labyrinthinus spp. nov. are described as new to science from the Dja Biosphere Reserve, Cameroon. Elaphomyces adamizans sp. nov. is described as new from the Pakaraima Mountains of Guyana. The Cameroonian species are the first Elaphomyces taxa to be formally described from Africa, occurring in lowland Guineo-Congolian tropical rainforests dominated by the ectomycorrhizal (ECM) canopy tree Gilbertiodendron dewevrei (Fabaceae subfam. Caesalpinioideae). The Guyanese species is the third to be discovered in lowland tropical South America, occurring in forests dominated by the ECM trees Pakaraimaea dipterocarpacea (Dipterocarpaceae) and Dicymbe jenmanii (Fabaceae subfam. Caesalpinioideae). Macromorphological, micromorphological, habitat, and DNA sequence data are provided for each new species. Molecular and morphological data place these fungi in Elaphomycetaceae (Eurotiales, Ascomycota). Unique morphological features are congruent with molecular delimitation of each of the new species based on a phylogenetic analysis of the rDNA ITS and 28S loci across the Elaphomycetaceae. The phylogenetic analysis also suggests that a common ancestor is shared between some Elaphomyces species from Africa and South America, and that species of the stalked, volvate genus Pseudotulostoma may be nested in Elaphomyces.

Pseudotulostoma volvata gen. sp. nov. is described from the south-central Pakaraima Mountains of Guyana. Pseudotulostoma volvata is associated with ectomycorrhizal Dicymbe corymbosa trees (Caesalpiniaceae) and placed in the Ascomycota, Eurotiales, Elaphomycetaceae. Included are a description of the genus and species, illustrations of the macroscopic and microscopic features, and a discussion of the distinctive features and phylogenetic placement of this fungus.

Elaphomyces compleximurus sp. nov. and E. digitatus sp. nov. are described from the Pakaraima Mountains of Guyana. Macromorphological, micro- morphological, habitat and DNA sequence data are provided for each new species. This is the first report of Elaphomyces ascomata associated with ectomycor- rhizal members of the Fabaceae and also for the genus from the lowland South American tropics.

This dissertation is an investigation of the systematics, phylogeography, and ecology of a globally distributed fungal family, the Elaphomycetaceae. In Chapter 1, we assess the literature on fungal phylogeography, reviewing large-scale phylogenetics studies and performing a meta-data analysis of fungal population genetics. In particular, we examined the possible effects of asexuality, trophic niche, dispersal method, and ocean barriers on population structure. In Chapter 2, we examine the systematics and phylogeography of the Elaphomycetaceae, a family consisting of the truffle genus Elaphomyces and the stalked genus Pseudotulostoma, hypothesizing that the mammal-dispersed truffle would show evidence of dispersal limitation. Using DNA sequence data, we determined that Pseudotulostoma is derived from a lineage of Elaphomyces, indicating that Elaphomyces as currently defined is paraphyletic. The distribution of each subgenus of Elaphomyces is nearly global; representative species have been found on every continent save Africa and Antarctica. This biogeographic pattern does not follow the pattern expected by a scenario of continental vicariance. Dating analysis in BEAST confirmed that broadly distributed clades are, in most cases, too young for this pattern to be explained by continental vicariance, indicating that occasional long-distance dispersal has been a significant component in the biogeographic history of the Elaphomycetaceae. This finding contradicts our initial hypothesis that the mammal-dispersed truffles would be dispersal- limited. In Chapter 3, we investigate the role of Elaphomyces as a host for the fungal parasite Elaphocordyceps, a parasite derived from insect pathogens that attacks both insect larvae and Elaphomyces, its only fungal host. We examined the biogeography of Elaphocordyceps isolated from Elaphomyces specimens in order to test whether it, like its host, showed recent connections between the Southern and Northern Hemispheres. We also evaluated the pathogenicity of infection as determined by a visual rubric for the truffle gleba, the phylogenetic distribution of Elaphocordyceps species on its host, testing for seasonal, climate, and host-parasite effects. In Chapter 4, based on the phylogeographic pattern seen in Elaphomyces that resembles that of some air-dispersed fungi, we used theoretical and experimental methods to test whether Elaphomyces could be dispersed by air. We tested the capacity for air dispersal with an experimental test of passive air dispersal on the powdery spores of Elaphomyces morettii and found that these large spores could disperse over a short distance (10 m) in comparable numbers with the spores of the giant puffball Calvatia cyathiformis, which is known to be air-dispersed. The major findings of this thesis are that 1) fungi in general show high dispersal ability, but that trophic niche and dispersal mode may affect population structure, 2) that Pseudotulostoma, a stalked genus, is derived from the truffle Elaphomyces, 3) that the Elaphomycetaceae have experienced frequent long-distance dispersal despite 4) that the fitness of Elaphomyces as indicated by glebal development varies with host-parasite interactions based on species identity, but not with climate or season, and that 5) Elaphomyces spores, should they be released into the air, can remain in the air long enough to be dispersed long distances by the wind. The overall conclusion of this thesis is that, while Elaphomyces is clearly reliant on animal vectors for excavation and dispersal, its past history of long-distance dispersal and current spore trajectories indicate it can be passively air-dispersed as well.

Lycoperdon cervinum Linnaeus, Species Plantarum 2: 1183. 1753. "Habitat in Bohemia, Silesia. Vermelandia." RCN: 8512. Type not designated. Original material: [icon] in Micheli, Nov. Pl. Gen.: 220, t. 99, f. 4. 1729; [icon] in Bauhin & Cherler, Hist. Pl. Univ. 3(2): 851. 1651. Current name: Elaphomyces granulatus Fr. (Elaphomycetaceae)., Published as part of Jarvis, Charlie, 2007, Chapter 7: Linnaean Plant Names and their Types (part L), pp. 610-650 in Order out of Chaos. Linnaean Plant Types and their Types, London :Linnaean Society of London in association with the Natural History Museum on page 645, DOI: 10.5281/zenodo.291971

Lycoperdon tuber Linnaeus, Species Plantarum 2: 1183. 1753. "Habitat sub Terra." RCN: 8512. Type not designated. Original material: [icon] in Micheli, Nov. Pl. Gen.: 221, t. 102. 1729; [icon] in Mattioli, Comment. Dioscoridis: 544. 1565. Current name: Elaphomyces sp. (Elaphomycetaceae)., Published as part of Jarvis, Charlie, 2007, Chapter 7: Linnaean Plant Names and their Types (part L), pp. 610-650 in Order out of Chaos. Linnaean Plant Types and their Types, London :Linnaean Society of London in association with the Natural History Museum on page 646, DOI: 10.5281/zenodo.291971

Lycoperdon tuber Linnaeus, Species Plantarum 2: 1183. 1753. "Habitat sub Terra." RCN: 8512. Type not designated. Original material: [icon] in Micheli, Nov. Pl. Gen.: 221, t. 102. 1729; [icon] in Mattioli, Comment. Dioscoridis: 544. 1565. Current name: Elaphomyces sp. (Elaphomycetaceae).

Pseudotulostoma volvata (O. K. Mill. and T. W. Henkel) is a morphologically unusual member of the otherwise hypogeous Elaphomycetaceae due to its epigeous habit and exposed gleba borne on an elevated stalk at maturity. Field observations in Guyana indicated that P. volvata was restricted to rain forests dominated by ectomycorrhizal (EM) Dicymbe corymbosa (Caesalpiniaceae), suggesting an EM nutritional mode for the fungus. In this paper, we confirm the EM status of P. volvata with a combination of morphological, molecular, and mycosociological data. The EM status for P. volvata corroborates its placement in the ectotrophic Elaphomycetaceae.

The sequestrate ascomycete genus Elaphomyces is described and illustrated from Australia. The following thirteen new species are described: Elaphomyces aurantias, E. austrogranulatus, E. chlorocarpus, E. cooloolanus, E. coralloideus, E. laetiluteus, E. nothofagi, E. pedicellaris, E. queenslandicus, E. rugosisporus, E. suejoyceae, E. symeae, and E. timgroveii. A key is provided to all Elaphomyces species from Australia.

The study of hypogeous fungi has been neglected in Britain from the time of Berkeley & Broome until that of the present investigation. During the years 1948-53 some 700 collections have been made, mainly in the Bristol area, but also from other parts of England, Scotland, North Wales and Northern Ireland. These include members of the Phycomycetes (Endogonespp.), Ascomycetes (Elaphomycetaceae, Tuberales) and Basidiomycetes (Gasteromycetes). Some species were found sufficiently often to permit tentative conclusions to be drawn relating to the effect of weather and soil conditions on the production of fruit-bodies. Most of the species previously recorded in Britain have been collected and some new records made. Descriptions are given of all recorded British species, and most of these are illustrated by line drawings made from fresh material. Details of development are given for representative species and the probable relationships within the group and with other fungi are discussed.