title: The Ophiostoma clavatum species complex: a newly defined group in the Ophiostomatales including three novel taxa.
abstract: Two species of blue-stain fungi with similar morphologies, Ophiostoma brunneo-ciliatum and Ophiostoma clavatum, are associates of bark beetles infesting Pinus spp. in Europe. This has raised questions whether they represent distinct taxa. Absence of herbarium specimens and contaminated or mistakenly identified cultures of O. brunneo-ciliatum and O. clavatum have accentuated the uncertainty regarding their correct identification. The aim of this study was to reconsider the identity of European isolates reported as O. brunneo-ciliatum and O. clavatum by applying DNA-based identification methods, and to provide appropriate type specimens for them. Phylogenetic analyses of the ITS, βT, TEF-1α and CAL gene sequences revealed that the investigated isolates represent a complex of seven cryptic species. The study confirmed that ITS data is insufficient to delineate species in some Ophiostoma species clusters. Lectotypes and epitypes were designated for O. clavatum and O. brunneo-ciliatum, and three new species, Ophiostoma brunneolum, Ophiostoma macroclavatum and Ophiostoma pseudocatenulatum, are described in the newly defined O. clavatum-complex. The other two species included in the complex are Ophiostoma ainoae and Ophiostoma tapionis. The results suggest co-evolution of these fungi in association with specific bark beetles. The results also confirm the identity of the fungus associated with the pine bark beetle Ips acuminatus as O. clavatum, while O. brunneo-ciliatum appears to be mainly associated with another pine bark beetle, Ips sexdentatus.
authors.affiliation: Department of Forest Sciences, University of Helsinki, P.O. Box 27, 00014, Helsinki, Finland. riikka.linnakoski@helsinki.fi.|Department of Microbiology and Plant Pathology, Forestry and Agricultural Biotechnology Institute (FABI), University of Pretoria, Pretoria, 0002, South Africa. riikka.linnakoski@helsinki.fi.
authors.forename: Riikka
authors.identifier: http://orcid.org/0000-0002-3294-8088
authors.initials: R
authors.lastname: Linnakoski
authors.affiliation: Department of Forest Pathology, Mycology and Tree Physiology, Institute of Forest Ecosystem Protection, University of Agriculture in Kraków, Al. 29 Listopada 46, 31-425, Kraków, Poland.
authors.forename: Robert
authors.initials: R
authors.lastname: Jankowiak
authors.affiliation: Department of Plant Pathology, Ohio State University, Columbus, OH, 43210, USA.|Dipartimento di Agronomia Animali Alimenti Risorse Naturali e Ambiente, Università di Padova, Agripolis, Legnaro, Padova, 35020, Italy.
authors.forename: Caterina
authors.initials: C
authors.lastname: Villari
authors.affiliation: Department of Forest and Soil Sciences, Institute of Forest Entomology, Forest Pathology and Forest Protection (IFFF), University of Natural Resources and Life Sciences, Vienna (BOKU), Hasenauerstraβe 38, 1190, Vienna, Austria.
authors.forename: Thomas
authors.initials: T
authors.lastname: Kirisits
authors.affiliation: Norwegian Institute of Bioeconomy Research, P.O. Box 115, 1431, Ås, Norway.
authors.forename: Halvor
authors.initials: H
authors.lastname: Solheim
authors.affiliation: Department of Microbiology and Plant Pathology, Forestry and Agricultural Biotechnology Institute (FABI), University of Pretoria, Pretoria, 0002, South Africa.
authors.forename: Z Wilhelm
authors.initials: ZW
authors.lastname: de Beer
authors.affiliation: Department of Microbiology and Plant Pathology, Forestry and Agricultural Biotechnology Institute (FABI), University of Pretoria, Pretoria, 0002, South Africa.
authors.forename: Michael J
authors.initials: MJ
authors.lastname: Wingfield
mesh_terms: D000818:Animals; D005075:Biological Evolution; D001517:Coleoptera / Q000382:microbiology; D004271:DNA, Fungal / Q000235:genetics; D021903:DNA, Ribosomal Spacer / Q000235:genetics; D005060:Europe; D055156:Ophiostoma / Q000145:classification* / Q000235:genetics / Q000254:growth & development / Q000302:isolation & purification; D010802:Phylogeny; D028223:Pinus / Q000382:microbiology; D016133:Polymerase Chain Reaction; D012340:RNA, Ribosomal, 5.8S / Q000235:genetics; D017422:Sequence Analysis, DNA; D013172:Spores, Fungal; D014197:Trees / Q000382:microbiology; D048889:Weevils / Q000382:microbiology
chemical_list: D004271:DNA, Fungal; D021903:DNA, Ribosomal Spacer; D012340:RNA, Ribosomal, 5.8S
keywords: Bark beetle-associated fungi; Ophiostoma brunneolum; Ophiostoma macroclavatum; Ophiostoma pseudocatenulatum; Ophiostomatoid fungi