A Nomenclator for Ophiostomatoid Genera and Species in the Ophiostomatales and Microascales
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BIODIVERSITY SERIES 12: 245–322 A nomenclator for ophiostomatoid genera and species in the Ophiostomatales and Microascales Z. Wilhelm de Beer1, K.A. Seifert2, M.J. Wingfield1 1Department of Microbiology and Plant Pathology, Forestry and Agricultural Research Institute (FABI), University of Pretoria, Pretoria 0002, South Africa; 2Biodiversity (Mycology & Botany), Agriculture and Agri-Food Canada, 960 Carling Ave., Ottawa, Ontario K1A 0C6, Canada *Correspondence: Z. Wilhelm de Beer, [email protected] Abstract: In this updated nomenclator, the names of 646 species proposed for ophiostomatoid fungi are considered. The result is 397 accepted species in 12 genera classified in the Ophiostomatales and Microascales. The taxonomic status of each species was re-evaluated based on all published details and where available, phylogenetic inferences, and data on typification. The principle of single name nomenclature, as adopted by the 18th International Botanical Congress, Melbourne in July 2011, was applied to all genera and species. Based on these re-assessments, three genera were redefined:Graphilbum in the Ophiostomatales, and Graphium and Knoxdaviesia (= Gondwanamyces) in the Microascales. Species were reclassified as necessary, resulting in 30 new combinations and one new name. Ophiostoma s.l. now includes 134 accepted species, Ceratocystiopsis 16 species, Fragosphaeria two, Graphilbum eight, Raffaelea s.str. 13, and Leptographium s.l. (including Grosmannia) 94. A further 28 species could not be assigned to any of these six genera with certainty, and seven more species were invalidly described. In the Microascales, Ceratocystis contains 72 accepted species, Graphium nine, Knoxdaviesia nine, and Sphaeronaemella seven, while Cornuvesica and Custingophora are both monotypic. Three microascalean species of uncertain status and six invalidly published species remain. Type studies and evaluation of the literature allowed the remaining 144 species described in the classical concept of Graphium to be reconsidered. Finally, 19 species previously assigned to Ceratocystis, Ceratostomella, Leptographium, Ophiostoma, Raffaelea, Sphaeronaemella and/or Sporothrix, were excluded from both the Ophiostomatales and Microascales. Key words: anamorph taxonomy, Ascomycota, one fungus : one name, rDNA phylogeny, Sordariomycetes.. Taxonomic novelties: Cephalotrichum album (Costantin) Seifert comb. nov., Ceratocystiopsis neglecta (Kirschner & Oberw.) Z.W. de Beer & M.J. Wingf. comb. nov., Ceratocystis harringtonii Z.W. de Beer & M.J. Wingf. nom. nov., Graphilbum brunneocrinitum (E.F. Wright & Cain) Z.W. de Beer & M.J. Wingf. comb. nov., Graphilbum curvicolle (Olchow. & J. Reid) Z.W. de Beer & M.J. Wingf. comb. nov., Graphilbum fragrans (Math.-Käärik) Z.W. de Beer, Seifert & M.J. Wingf. comb. nov., Graphilbum microcarpum (Yamaoka & Masuya) Z.W. de Beer & M.J. Wingf. comb. nov., Graphilbum nigrum (R.W. Davidson) Z.W. de Beer & M.J. Wingf. comb. nov., Graphilbum rectangulosporium (R.W. Davidson) Z.W. de Beer & M.J. Wingf. comb. nov., Graphilbum tubicolle (Olchow. & J. Reid) Z.W. de Beer & M.J. Wingf. comb. nov., Grosmannia truncicola (R.W. Davidson) Z.W. de Beer & M.J. Wingf. comb. nov., Knoxdaviesia cecropiae (M. Kolařík) Z.W. de Beer & M.J. Wingf. comb. nov., Knoxdaviesia scolytodis (M. Kolařík) Z.W. de Beer & M.J. Wingf. comb. nov., Knoxdaviesia serotectus (Van der Linde & Jol. Roux) Z.W. de Beer & M.J. Wingf. comb. nov., Knoxdaviesia suidafrikana (Morgan-Jones & R.C. Sinclair) Z.W. de Beer & M.J. Wingf. comb. nov., Knoxdaviesia ubusi (Van der Linde & Jol. Roux) Z.W. de Beer & M.J. Wingf. comb. nov., Knoxdaviesia undulatistipes (Pinnoi) Z.W. de Beer & M.J. Wingf. comb. nov., Knoxdaviesia wingfieldii (Roets & Dreyer) Z.W. de Beer & M.J. Wingf. comb. nov., Leptographium obscurum (R.W. Davidson) Z.W. de Beer & M.J. Wingf. comb. nov., Leptographium rostrocylindricum (R.W. Davidson) Z.W. de Beer & M.J. Wingf. comb. nov., Leptographium verrucosum (Gebhardt, R. Kirschner & Oberw.) Z.W. de Beer & M.J. Wingf. comb. nov., Ophiostoma australiae (Kamgan, K. Jacobs & M.J. Wingf.) Z.W. de Beer & M.J. Wingf. comb. nov., Ophiostoma cupulatum (McNew & Harrington) Z.W. de Beer & M.J. Wingf. comb. nov., Ophiostoma denticulatum (R.W. Davidson) Z.W. de Beer & M.J. Wingf. comb. nov., Ophiostoma leucocarpum (R.W. Davidson) Z.W. de Beer & M.J. Wingf. comb. nov., Ophiostoma macrosporum (Francke-Grosm.) Z.W. de Beer & M.J. Wingf. comb. nov., Ophiostoma populicola (Olchow. & J. Reid) Z.W. de Beer, Seifert & M.J. Wingf. comb. nov., Ophiostoma tingens (Lagerb. & Melin) Z.W. de Beer & M.J. Wingf. comb. nov., Phaeostilbella nigra (Berk.) Seifert comb. nov., Sphaeronaemella betae (Delacr.) Z.W. de Beer & M.J. Wingf. comb. nov., Stilbocrea aterrima (Welw. & Curr.) Seifert comb. nov. INTRODUCTION genera in that order. Not all fungi in the Microascales are considered ophiostomatoid (De Beer et al. 2013). Therefore, we only consider Presented here is a nomenclatural implementation of the taxonomic genera of the three ophiostomatoid families in that order, the concepts for ophiostomatoid fungi proposed elsewhere in this Ceratocystidaceae, Gondwanamycetaceae and Graphiaceae. volume by De Beer et al. (2013), De Beer & Wingfield (2013) For the Microascales, we apply generic concepts as defined by B. and B. Wingfield et al. (2013). The nomenclator includes all Wingfieldet al. (2013) and De Beer et al. (2013). Sphaeronaemella genera and species currently classified in the Ophiostomataceae could not be placed with confidence in a microascalean family (De (Ophiostomatales), Ceratocystidaceae, Gondwanamycetaceae and Beer et al. 2013), but we included it in our list because several of its Graphiaceae (Microascales). In this listing, we follow the single name species were previously classified in Ceratocystis, and were thus principle accepted at the 18th International Botanical Congress held considered ophiostomatoid. in Melbourne during July 2011 (Hawksworth 2011, Hawksworth et In cases where a currently accepted genus includes the type al. 2011, Norvell 2011). These principles have been incorporated species of different genera, priority is given to the genus name into the newly named International Code of Nomenclature for that was described first. All other genus names are treated as its Algae, Fungi and Plants (ICN) and enforce equal status for taxa synonyms, irrespective of the morph they previously represented. (i.e. genera and species) whether they were originally described as either teleomorphic or anamorphic (McNeill et al. 2012). To maintain Species nomenclatural stability and avoid premature and unnecessary name changes, we followed recommendations explained by De Beer et al. All species are listed under the genus in which they are currently (2013), De Beer & Wingfield (2013) and B. Wingfield et al. (2013). treated in the accompanying papers by De Beer & Wingfield (2013) and B. Wingfield et al. (2013); this nomenclature has been applied Genera throughout this book. Three genera, Ophiostoma sensu lato (s.l.), Leptographium s.l., and Ceratocystis s.l., include species with In the nomenclator, we apply the generic concepts defined by De generic names differing from the genus where they are listed. This Beer & Wingfield (2013) for the Ophiostomatales, and include all reflects the taxonomic uncertainties surrounding these species, 245 DE BEER ET AL. usually the consequence of inadequate phylogenetic support Notes: Additional information, in most cases explaining and the need for additional multi-gene studies or more intensive synonymies and/or uncertainties surrounding the status of species, sampling. For example, Leptographium antibioticum is not part and the current placement of a species in a species complex based of Leptographium s.l., but of Ophiostoma s.l. based on rDNA on phylogenetic analyses. phylogenies (De Beer & Wingfield 2013), and is presently listed under the latter genus until its accurate phylogenetic classification can be assured. To improve searchability within these sensu lato A.1. ACCEPTED GENERA AND SPECIES IN THE genera, the species are alphabetically ordered by epithet, rather OPHIOSTOMATALES than their current genus name. Under the dual nomenclature system, previous monographs Genus concepts as defined by De Beer & Wingfield (2013) are and nomenclators distinguished between teleomorph and applied here. Under Ophiostoma and Leptographium, we list all anamorph binomials. These names now have equal status and species treated respectively in Ophiostoma s.l. and Leptographium we thus treat all names applied to any single species as equal s.l. as defined by De Beer & Wingfield (2013), following their synonyms. These synonyms are listed under the relevant species recommendations to ensure nomenclatural stability under the in order of priority. emended Code. Sections Ophiostoma Syd., In Sydow & Sydow, Annls mycol. 17: 43. 1919 [type species O. piliferum] The nomenclator is subdivided as follows: ?= Sporothrix Hektoen & C.F. Perkins, J. Exp. Med. 5: 80. 1900. Anamorphic synonym. [type species S. schenckii] A. The Ophiostomatales = Sporothrix section Sporothrix Weijman & de Hoog, Antonie van Leeuwenhoek 1. Accepted genera and species 51: 118. 1985. = Sporotrichopsis Guég., In De Beurmann & Gougerot, Archs Parasit. 15: 104. 2. Valid species of uncertain status 1911. Anamorphic synonym. [type species S. beurmannii nom. inval., Art. 36.1] 3. Invalidly published species = Dolichoascus Thibaut & Ansel, In Ansel & Thibaut, Compt. Rend. Hebd. B. The ophiostomatoid genera