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      True morels (Morchella, Pezizales) of Europe and North America: evolutionary relationships inferred from multilocus data and a unified taxonomy.

      Mycologia
      Mycological Society of America
      Ascomycota, nomenclature, Pezizomycetes, Morchellaceae, taxonomy

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          Abstract

          Applying early names, with or without original material, to genealogical species is challenging. For morels this task is especially difficult because of high morphological stasis and high plasticity of apothecium color and shape. Here we propose a nomenclatural revision of true morels (Morchella, Pezizales) from Europe and North America, based on molecular phylogenetic analyses of portions of the genes for RNA polymerase II largest subunit (RPB1) and second largest subunit (RPB2), translation elongation factor-1α (TEF1), the nuc rDNA region encompassing the internal transcribed spacers 1 and 2, along with the 5.8S rDNA (ITS), and partial nuc 28S rDNA D1-D2 domains (28S). The 107 newly sequenced collections were from both continents, including 48 types, together with previously published sequences. Names are applied to 30 of the 65 currently recognized genealogical species. Results of the present study revealed that the number of Morchella species in Europe (n = 21) is nearly identical to that in North America (n = 22). Only seven species were found on both continents, consistent with previous reports of high continental endemism within the genus. Presently it is not possible to tell whether the transoceanic disjunctions were due to human activities, migration across a Bering land bridge or long-distance dispersal. In an effort to stabilize the taxonomy, due in part to the recent publication of synonyms for 11 of the species, accepted names are presented together with their corresponding later synonyms. A new subclade that includes holotypes of M. castanea and M. brunneorosea is identified in sect. Morchella (Esculenta Clade). Lectotypes for Morchella deliciosa, M. eximia and M. tridentina are designated here, as well as epitypes for M. dunalii, M. eximia, M. purpurascens and M. vulgaris. Morchella conica was determined to be illegitimate, and further research is required to determine the identity of M. elata and M. inamoena.

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          Most cited references22

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          Phylogenetic species recognition and species concepts in fungi.

          The operational species concept, i.e., the one used to recognize species, is contrasted to the theoretical species concept. A phylogenetic approach to recognize fungal species based on concordance of multiple gene genealogies is compared to those based on morphology and reproductive behavior. Examples where Phylogenetic Species Recognition has been applied to fungi are reviewed and concerns regarding Phylogenetic Species Recognition are discussed.
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            Using RPB1 sequences to improve phylogenetic inference among mushrooms (Inocybe, Agaricales).

            An investigation of mushroom phylogeny using the largest subunit of RNA polymerase II gene sequences (RPB1) was conducted in comparison with nuclear ribosomal large subunit RNA gene sequences (nLSU) for the same set of taxa in the genus Inocybe (Agaricales, Basidiomycota). The two data sets, though not significantly incongruent, exhibit conflict among the placement of two taxa that exhibit long branches in the nLSU data set. In contrast, RPB1 terminal branch lengths are rather uniform. Bootstrap support is increased for clades in RPB1. Combined data sets increase the degree of confidence for several relationships. Overall, nLSU data do not yield a robust phylogeny when independently assessed by RPB1 sequences. This multigene study indicates that Inocybe is a monophyletic group composed of at least four distinct lineages-subgenus Mallocybe, section Cervicolores, section Rimosae, and subgenus Inocybe sensu Kühner, Kuyper, non Singer. Within subgenus Inocybe, two additional lineages, one composed of species with smooth basidiospores (clade I) and a second characterized by nodulose-spored species (clade II), are recovered by RPB1 and combined data. The nLSU data recover only clade I. The genera Astrosporina and Inocybella cannot be recognized phylogenetically. "Supersections" Cortinatae and Marginatae are not monophyletic groups.
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              Phylogeny and historical biogeography of true morels (Morchella) reveals an early Cretaceous origin and high continental endemism and provincialism in the Holarctic.

              True morels (Morchella, Ascomycota) are arguably the most highly-prized of the estimated 1.5 million fungi that inhabit our planet. Field guides treat these epicurean macrofungi as belonging to a few species with cosmopolitan distributions, but this hypothesis has not been tested. Prompted by the results of a growing number of molecular studies, which have shown many microbes exhibit strong biogeographic structure and cryptic speciation, we constructed a 4-gene dataset for 177 members of the Morchellaceae to elucidate their origin, evolutionary diversification and historical biogeography. Diversification time estimates place the origin of the Morchellaceae in the middle Triassic 243.63 (95% highest posterior density [HPD] interval: 169.35-319.89) million years ago (Mya) and the divergence of Morchella from its closest relatives in the early Cretaceous 129.61 (95% HPD interval: 90.26-173.16) Mya, both within western North America. Phylogenetic analyses identified three lineages within Morchella: a basal monotypic lineage represented by Morchella rufobrunnea, and two sister clades comprising the black morels (Elata Clade, 26 species) and yellow morels (Esculenta Clade, 16 species). Morchella possesses a Laurasian distribution with 37/41 species restricted to the Holarctic. All 33 Holarctic species represented by multiple collections exhibited continental endemism. Moreover, 16/18 North American and 13/15 Eurasian species appeared to exhibit provincialism. Although morel fruit bodies produce thousands of explosively discharged spores that are well suited to aerial dispersal, our results suggest that they are poorly adapted at invading novel niches. Morels also appear to have retained the ancestral fruit body plan, which has remained remarkably static since the Cretaceous. Published by Elsevier Inc.
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                Author and article information

                Journal
                25550303
                10.3852/14-166

                Ascomycota,nomenclature,Pezizomycetes,Morchellaceae,taxonomy

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