Taxonomy and phylogeny of the Leptographium procerum complex, including Leptographium sinense sp. nov. and Leptographium longiconidiophorum sp. nov.
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Leptographium procerum (Ophiostomatales, Ascomycota) is a well-known fungal associate of pine root-infesting bark beetles and weevils, occurring in several countries of the world. The fungus is not a primary pathogen but has been associated with white pine root decline in the USA and with serious damage caused by the introduced red turpentine beetle (RTB) Dendroctonus valens in China. Several species closely related to L. procerum have been described during the past decade. The aim of this study was to reevaluate species boundaries in the L. procerum complex using multigene phylogenetic analyses and morphological comparisons. Phylogenetic analyses of seven gene regions (ITS2-LSU, actin, β-tubulin, calmodulin, translation elongation factor 1-α, and the mating type genes MAT1-1-3 and MAT1-2-1) distinguished between nine species in the complex. These included L. procerum, L. bhutanense, L. gracile, L. profanum, L. pini-densiflorae, L. sibiricum, L. sinoprocerum, as well as two new species described here as Leptographium sinense sp. nov. from Hylobitelus xiaoi on Pinus elliottii in China, and Leptographium longiconidiophorum sp. nov. from Pinus densiflora in Japan. Leptographium latens is reduced to synonymy with L. gracile, and an epitype is designated for L. procerum, because a living culture associated with the holotype of L. procerum did not exist. Amplification patterns of the mating type genes suggest that all known species in the L. procerum complex are heterothallic, although sexual states have not been observed for any of the species. The results also suggest that Eastern Asia is most probably the centre of species diversity for the L. procerum complex.
KeywordsBark beetle associates Epitype Leptographium Ophiostomatales Phylogeny Taxonomy
This study was initiated through the bilateral agreement between the Governments of South Africa and China, and we are grateful for the funding via projects 2012DFG31830 (International Science & Technology Cooperation Program of China), 2010KJCX015-03 (Forestry Science and Technology Innovation Project of Guangdong Province of China). We acknowledge members of Tree Protection and Cooperation Programme (TPCP), the National Research Foundation (NRF), the Department of Science and Technology (DST)/NRF, Center of Excellence in Tree Health Biotechnology (CTHB) and the University of Pretoria, Pretoria, South Africa. We also thank Mr. Runlei Chang for assistance with the fieldwork, and Ms. Yalin Fu for assistance with fungal isolations.
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