{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,21]],"date-time":"2026-07-21T15:24:03Z","timestamp":1784647443190,"version":"3.55.0"},"reference-count":102,"publisher":"Public Library of Science (PLoS)","issue":"5","license":[{"start":{"date-parts":[[2023,5,8]],"date-time":"2023-05-08T00:00:00Z","timestamp":1683504000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000038","name":"Natural Sciences and Engineering Research Council of Canada","doi-asserted-by":"publisher","award":["RGPIN-2015-04376"],"award-info":[{"award-number":["RGPIN-2015-04376"]}],"id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000038","name":"Natural Sciences and Engineering Research Council of Canada","doi-asserted-by":"publisher","award":["RGPIN-2015-05279"],"award-info":[{"award-number":["RGPIN-2015-05279"]}],"id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS ONE"],"abstract":"<jats:p>Bark beetles (Coleoptera: Curculionidae; Scolytinae) are tree-infesting insects that consume subcortical tissues and fungi. Species capable of killing their host trees are most commonly associated with conifers, as very few bark beetle species infest and kill hardwood hosts directly. The alder bark beetle, <jats:italic>Alniphagus aspericollis<\/jats:italic>, is a hardwood-killing bark beetle that colonizes and kills red alder, <jats:italic>Alnus rubra<\/jats:italic>. Conifer-killing bark beetles have well-known associations with symbiotic ophiostomatoid fungi that facilitate their life histories, but it is unknown whether <jats:italic>A<\/jats:italic>. <jats:italic>aspericollis<\/jats:italic> has any fungal associates. This study was conducted to identify any consistent filamentous fungal associates of <jats:italic>A<\/jats:italic>. <jats:italic>aspericollis<\/jats:italic> and characterize the consistency of observed beetle\u2013fungus relationships. Beetles and gallery phloem samples were collected from seven sites throughout the Greater Vancouver region in British Columbia, Canada. Filamentous fungi were isolated from these samples and identified by DNA barcoding using the internal transcribed spacer (ITS) region and other barcode regions for resolution to the species-level for the most dominant isolates. The most common fungal associate was a previously undescribed <jats:italic>Neonectria major<\/jats:italic>-like fungus, <jats:italic>Neonectria<\/jats:italic> sp. nov., which was isolated from ~67% of adult beetles, ~59% of phloem samples, and ~94% of the beetle-infested trees. <jats:italic>Ophiostoma quercus<\/jats:italic> was isolated from ~28% of adult beetles, ~9% of phloem samples, and ~56% of infested trees and deemed a casual associate of <jats:italic>A<\/jats:italic>. <jats:italic>aspericollis<\/jats:italic>, while a putatively novel species of <jats:italic>Ophiostoma<\/jats:italic> was more infrequently isolated from <jats:italic>A<\/jats:italic>. <jats:italic>aspericollis<\/jats:italic> and its galleries. <jats:italic>Cadophora spadicis<\/jats:italic>, a new record for red alder, was rarely isolated and is probably coincidentally carried by <jats:italic>A<\/jats:italic>. <jats:italic>aspericollis<\/jats:italic>. Overall, <jats:italic>A<\/jats:italic>. <jats:italic>aspericollis<\/jats:italic> was only loosely associated with ophiostomatoid fungi, suggesting that these fungi have little ecological significance in the beetle\u2013tree interaction, while <jats:italic>Neonectria<\/jats:italic> sp. nov. may be a symbiote of <jats:italic>A<\/jats:italic>. <jats:italic>aspericollis<\/jats:italic> that is vectored by the beetle.<\/jats:p>","DOI":"10.1371\/journal.pone.0284393","type":"journal-article","created":{"date-parts":[[2023,5,8]],"date-time":"2023-05-08T18:29:41Z","timestamp":1683570581000},"page":"e0284393","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":6,"title":["Filamentous fungal associates of the alder bark beetle, Alniphagus aspericollis, including an undescribed species of Neonectria"],"prefix":"10.1371","volume":"18","author":[{"given":"Gervais Y. S.","family":"Lee","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8492-976X","authenticated-orcid":true,"given":"Debra L.","family":"Wertman","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1820-0759","authenticated-orcid":true,"given":"Allan L.","family":"Carroll","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Richard C.","family":"Hamelin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2023,5,8]]},"reference":[{"key":"pone.0284393.ref001","volume-title":"The bark beetles of Canada and Alaska: Coleoptera: Scolytidae","author":"DE Bright","year":"1976"},{"key":"pone.0284393.ref002","volume-title":"Great Basin Naturalist Memoirs No. 6","author":"SL Wood","year":"1982"},{"issue":"3","key":"pone.0284393.ref003","doi-asserted-by":"crossref","first-page":"293","DOI":"10.4039\/Ent116293-3","article-title":"Abundance of the fir engraver, Scolytus ventralis, and the Douglas-fir beetle, Dendroctonus pseudotsugae, following tree defoliation by the Douglas-fir tussock moth, Orgyia pseudotsugata","volume":"116","author":"LC Wright","year":"1984","journal-title":"Can Entomol"},{"key":"pone.0284393.ref004","first-page":"102","volume-title":"Beetle-pathogen interactions in conifer forests","author":"KF Raffa","year":"1993"},{"key":"pone.0284393.ref005","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1146\/annurev.ento.42.1.179","article-title":"Interactions among scolytid bark beetles, their associated fungi, and live host conifers","volume":"42","author":"TD Paine","year":"1997","journal-title":"Annu Rev Entomol"},{"key":"pone.0284393.ref006","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1146\/annurev.en.24.010179.002221","article-title":"Population dynamics of bark beetles","volume":"24","author":"RN Coulson","year":"1979","journal-title":"Annu Rev Entomol"},{"key":"pone.0284393.ref007","doi-asserted-by":"crossref","first-page":"484","DOI":"10.1086\/284497","article-title":"Mountain pine beetle outbreaks in the Rocky Mountains: regulators of primary productivity?","volume":"127","author":"WH Romme","year":"1986","journal-title":"Am Nat"},{"key":"pone.0284393.ref008","doi-asserted-by":"crossref","first-page":"65","DOI":"10.2307\/1551721","article-title":"Spruce beetles and fires in the nineteenth-century subalpine forests of western Colorado, USA","volume":"22","author":"WL Baker","year":"1990","journal-title":"Arc Alp Res"},{"key":"pone.0284393.ref009","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1016\/j.foreco.2006.02.038","article-title":"Spruce beetle outbreaks on the Kenai Peninsula, Alaska, and Kluane National Park and Reserve, Yukon Territory: relationship to summer temperatures and regional differences in disturbance regimes","volume":"227","author":"EE Berg","year":"2006","journal-title":"For Ecol Manage"},{"key":"pone.0284393.ref010","doi-asserted-by":"crossref","first-page":"987","DOI":"10.1038\/nature06777","article-title":"Mountain pine beetle and forest carbon: feedback to climate change","volume":"452","author":"WA Kurz","year":"2008","journal-title":"Nature"},{"issue":"6","key":"pone.0284393.ref011","doi-asserted-by":"crossref","first-page":"501","DOI":"10.1641\/B580607","article-title":"Cross-scale drivers of natural disturbances prone to anthropogenic amplification: the dynamics of bark beetle eruptions","volume":"58","author":"KF Raffa","year":"2008","journal-title":"Bioscience"},{"issue":"10","key":"pone.0284393.ref012","first-page":"2011","article-title":"The evolution of agriculture in beetles (Curculionidae: Scolytinae and Platypodinae)","volume":"55","author":"BD Farrell","year":"2001","journal-title":"Evolution"},{"key":"pone.0284393.ref013","doi-asserted-by":"crossref","first-page":"1187","DOI":"10.1093\/aesa\/63.4.1187","article-title":"Antagonism between Dendroctonus frontalis and the fungus Ceratocystis minor","volume":"63","author":"SJ Barras","year":"1970","journal-title":"Ann Entomol Soc Am"},{"key":"pone.0284393.ref014","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1111\/j.1365-2311.1983.tb00476.x","article-title":"Role of two phoretic mites in transmission of bluestain fungus, Ceratocystis minor","volume":"8","author":"JR Bridges","year":"1983","journal-title":"Ecol Entomol"},{"key":"pone.0284393.ref015","first-page":"83","article-title":"Symbiosis and competition: complex interactions among beetles, fungi and mites","volume":"30","author":"KD Klepzig","year":"2001","journal-title":"Symbiosis"},{"key":"pone.0284393.ref016","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1126\/science.167.3914.54","article-title":"Maxillary mycangium in the mountain pine beetle","volume":"167","author":"HS Whitney","year":"1970","journal-title":"Science"},{"key":"pone.0284393.ref017","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1016\/S0040-8166(71)80024-1","article-title":"Fine structure of the prothoracic mycangium, a chamber for the culture of symbiotic fungi, in the southern pine beetle, Dendroctonus frontalis","volume":"3","author":"GM Happ","year":"1971","journal-title":"Tissue Cell"},{"key":"pone.0284393.ref018","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1201\/9780203009918.ch7","volume-title":"Insect Symbioses","author":"DL Six","year":"2003a"},{"key":"pone.0284393.ref019","first-page":"183","volume-title":"Bark beetles in North American conifers","author":"HS Whitney","year":"1982"},{"key":"pone.0284393.ref020","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1146\/annurev-ento-120709-144839","article-title":"The role of phytopathogenicity in bark beetle-fungus symbioses: a challenge to the classic paradigm","volume":"56","author":"DL Six","year":"2011","journal-title":"Annu Rev Entomol"},{"issue":"6","key":"pone.0284393.ref021","doi-asserted-by":"crossref","first-page":"1393","DOI":"10.1093\/ee\/27.6.1393","article-title":"Effects of mycangial fungi and host tree species on progeny survival and emergence of Dendroctonus ponderosae (Coleoptera: Scolytidae)","volume":"27","author":"DL Six","year":"1998","journal-title":"Environ Entomol"},{"issue":"8","key":"pone.0284393.ref022","doi-asserted-by":"crossref","first-page":"2198","DOI":"10.1890\/0012-9658(2000)081[2198:NBOPFB]2.0.CO;2","article-title":"Nitrogen budgets of phloem-feeding bark beetles with and without symbiotic fungi","volume":"81","author":"MP Ayres","year":"2000","journal-title":"Ecology"},{"issue":"6","key":"pone.0284393.ref023","doi-asserted-by":"crossref","first-page":"1384","DOI":"10.1093\/ee\/36.6.1384","article-title":"Dietary benefits of fungal associates to an eruptive herbivore: potential implications of multiple associates on host population dynamics","volume":"36","author":"KP Bleiker","year":"2007","journal-title":"Environ Entomol"},{"issue":"8","key":"pone.0284393.ref024","doi-asserted-by":"crossref","first-page":"801","DOI":"10.1051\/forest\/2009066","article-title":"Stimulation of tree defenses by ophiostomatoid fungi can explain attack success of bark beetles in conifers","volume":"66","author":"F Lieutier","year":"2009","journal-title":"Ann For Sci"},{"issue":"3","key":"pone.0284393.ref025","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1046\/j.1365-3059.2002.00722.x","article-title":"Xylem sap flow of Norway spruce after inoculation with the blue-stain fungus Ceratocystis polonica","volume":"51","author":"T Kirisits","year":"2002","journal-title":"Plant Pathol"},{"issue":"6","key":"pone.0284393.ref026","doi-asserted-by":"crossref","first-page":"1599","DOI":"10.1007\/s00468-014-1068-0","article-title":"Sap flow-based quantitative indication of progression of Dutch elm disease after inoculation with Ophiostoma novo-ulmi","volume":"28","author":"J Urban","year":"2014","journal-title":"Trees"},{"issue":"7","key":"pone.0284393.ref027","doi-asserted-by":"crossref","first-page":"856","DOI":"10.1094\/PDIS-01-13-0056-FE","article-title":"Destructive tree diseases associated with ambrosia and bark beetles: black swan events in tree pathology","volume":"95","author":"RC Ploetz","year":"2013","journal-title":"Plant Dis"},{"key":"pone.0284393.ref028","first-page":"189","article-title":"Bark beetle\u2013fungal symbiosis: context dependency in complex associations","volume":"37","author":"KD Klepzig","year":"2004","journal-title":"Symbiosis"},{"key":"pone.0284393.ref029","doi-asserted-by":"crossref","first-page":"339","DOI":"10.3390\/insects3010339","article-title":"Ecological and evolutionary determinants of bark beetle\u2013fungus symbioses","volume":"3","author":"DL Six","year":"2012","journal-title":"Insects"},{"key":"pone.0284393.ref030","first-page":"245","article-title":"A nomenclator for ophiostomatoid genera and species in the Ophiostomatales and Microascales","volume":"12","author":"ZW de Beer","year":"2013","journal-title":"CBS Biodivers Ser"},{"issue":"2","key":"pone.0284393.ref031","doi-asserted-by":"crossref","first-page":"242","DOI":"10.2307\/3565273","article-title":"Why are there so few tree-killing bark beetles associated with angiosperms?","volume":"54","author":"CP Ohmart","year":"1989","journal-title":"Oikos"},{"issue":"5","key":"pone.0284393.ref032","first-page":"294","article-title":"Oak bark beetles\u2013important vectors of oak wilt","volume":"68","author":"CO Rexrode","year":"1970","journal-title":"J Forest"},{"key":"pone.0284393.ref033","first-page":"271","volume-title":"Invertebrate-microbial interactions: joint symposium of the British Mycological Society and the British Ecological Society. University of Exeter, Exeter, United Kingdom, September 1982","author":"JF Webber","year":"1984"},{"key":"pone.0284393.ref034","doi-asserted-by":"crossref","first-page":"126","DOI":"10.3832\/ifor1231-008","article-title":"Dutch elm disease and elm bark beetles: a century of association","volume":"8","author":"A Santini","year":"2015","journal-title":"Iforest"},{"key":"pone.0284393.ref035","volume-title":"The Scolytoidea of the Northwest","author":"WJ Chamberlin","year":"1958"},{"issue":"8","key":"pone.0284393.ref036","doi-asserted-by":"crossref","first-page":"870","DOI":"10.4039\/Ent101870-8","article-title":"Observations on the life history and habits of Alniphagus aspericollis (Coleoptera: Scolytidae) in southwestern British Columbia","volume":"101","author":"JH Borden","year":"1969","journal-title":"Can Entomol"},{"key":"pone.0284393.ref037","doi-asserted-by":"crossref","first-page":"586","DOI":"10.2307\/1307515","article-title":"Nitrogen fixation by actinomycete-nodulated angiosperms","volume":"28","author":"JG Torrey","year":"1978","journal-title":"BioScience"},{"key":"pone.0284393.ref038","first-page":"116","volume-title":"Silvics of North America","author":"CA Harrington","year":"1990"},{"key":"pone.0284393.ref039","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1139\/b91-056","article-title":"Endophytic fungi of red alder (Alnus rubra) leaves and twigs in British Columbia","volume":"69","author":"TN Sieber","year":"1991","journal-title":"Can J Bot"},{"issue":"1","key":"pone.0284393.ref040","doi-asserted-by":"crossref","first-page":"34","DOI":"10.3955\/046.089.0103","article-title":"Alder canopy dieback and damage in western Oregon riparian ecosystems","volume":"89","author":"L Sims","year":"2015","journal-title":"Northwest Sci"},{"issue":"16","key":"pone.0284393.ref041","doi-asserted-by":"crossref","first-page":"6241","DOI":"10.1073\/pnas.1117018109","article-title":"Nuclear ribosomal internal transcribed spacer (ITS) region as a universal DNA barcode marker for Fungi","volume":"109","author":"CL Schoch","year":"2012","journal-title":"Proc Natl Acad Sci USA"},{"key":"pone.0284393.ref042","first-page":"315","volume-title":"Protocols: a Guide to Methods and Applications","author":"TJ White","year":"1990"},{"key":"pone.0284393.ref043","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1111\/j.1365-294X.1993.tb00005.x","article-title":"ITS primers with enhanced specificity for basidiomycetes\u2013application to the identification of mycorrhizae and rusts","volume":"2","author":"M Gardes","year":"1993","journal-title":"Mol Ecol"},{"issue":"12","key":"pone.0284393.ref044","doi-asserted-by":"crossref","first-page":"1799","DOI":"10.1093\/oxfordjournals.molbev.a026092","article-title":"Phylogenetic relationships among ascomycetes: evidence from an RNA polymerase II subunit","volume":"16","author":"YJ Liu","year":"1999","journal-title":"Mol Biol Evol"},{"issue":"7","key":"pone.0284393.ref045","doi-asserted-by":"crossref","first-page":"664","DOI":"10.1007\/s11427-011-4184-8","article-title":"DNA barcoding of the fungal genus Neonectria and the discovery of two new species","volume":"54","author":"P Zhao","year":"2011","journal-title":"Sci China Life Sci"},{"issue":"3","key":"pone.0284393.ref046","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1080\/00275514.1999.12061051","article-title":"A method for designing primer sets for speciation studies in filamentous ascomycetes","volume":"91","author":"I Carbone","year":"1999","journal-title":"Mycologia"},{"key":"pone.0284393.ref047","first-page":"1417","article-title":"Phylogenetic relationships of Neonectria\/ Cylindrocarpon on Fagus in North America","volume":"84","author":"LA Castlebury","year":"2006","journal-title":"Can J Bot"},{"issue":"4","key":"pone.0284393.ref048","doi-asserted-by":"crossref","first-page":"1323","DOI":"10.1128\/aem.61.4.1323-1330.1995","article-title":"Development of primer sets designed for use with the PCR to amplify conserved genes from filamentous ascomycetes","volume":"61","author":"NL Glass","year":"1995","journal-title":"Appl Environ Microbiol"},{"key":"pone.0284393.ref049","first-page":"1","article-title":"Morphological and molecular data reveal cryptic speciation in Lasiodiplodia theobromae","volume":"28","author":"A Alves","year":"2008","journal-title":"Fungal Divers"},{"key":"pone.0284393.ref050","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1016\/j.funbio.2014.11.002","article-title":"Cadophora species associated with wood-decay of grapevine in North America","volume":"119","author":"R Travadon","year":"2015","journal-title":"Fungal Biol"},{"key":"pone.0284393.ref051","first-page":"95","article-title":"BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95\/98\/NT","volume":"41","author":"TA Hall","year":"1999","journal-title":"Nucleic Acids Symp Ser"},{"issue":"3","key":"pone.0284393.ref052","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1093\/sysbio\/syq010","article-title":"New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0","volume":"59","author":"S Guindon","year":"2010","journal-title":"Syst Biol"},{"key":"pone.0284393.ref053","first-page":"487","article-title":"Phialophora-like fungi associated with kiwifruit elephantiasis","volume":"90","author":"A Prodi","year":"2008","journal-title":"J Plant Pathol"},{"key":"pone.0284393.ref054","doi-asserted-by":"crossref","first-page":"139","DOI":"10.3114\/sim.2011.68.06","article-title":"Acremonium phylogenetic overview and revision of Gliomastix, Sarocladium, and Trichothecium","volume":"68","author":"RC Summerbell","year":"2011","journal-title":"Stud Mycol"},{"issue":"3","key":"pone.0284393.ref055","doi-asserted-by":"crossref","first-page":"532","DOI":"10.3852\/14-122","article-title":"Systematics of the Cosmospora viliuscula species complex","volume":"107","author":"CS Herrera","year":"2015","journal-title":"Mycologia"},{"key":"pone.0284393.ref056","doi-asserted-by":"crossref","first-page":"655","DOI":"10.1007\/s11557-011-0777-7","article-title":"Cylindrocarpon root rot: multi-gene analysis reveals novel species within the Ilyonectria radicicola species complex","volume":"11","author":"A Cabral","year":"2012","journal-title":"Mycol Prog"},{"issue":"4","key":"pone.0284393.ref057","doi-asserted-by":"crossref","first-page":"469","DOI":"10.1017\/S0953756203007445","article-title":"The Ophiostoma piceae complex in the Southern Hemisphere: a phylogenetic study","volume":"107","author":"ZW de Beer","year":"2003","journal-title":"Mycol Res"},{"key":"pone.0284393.ref058","first-page":"261","article-title":"Ophiostoma and Ceratocystiopsis spp. associated with two pine-infesting bark beetles in Chile","volume":"15","author":"XD Zhou","year":"2004","journal-title":"Fungal Divers"},{"key":"pone.0284393.ref059","doi-asserted-by":"crossref","first-page":"1475","DOI":"10.1016\/j.mycres.2008.06.007","article-title":"Fungi, including Ophiostoma karelicum sp. nov., associated with Scolytus ratzeburgi infesting birch in Finland and Russia","volume":"112","author":"R Linnakoski","year":"2008","journal-title":"Mycol Res"},{"key":"pone.0284393.ref060","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1007\/s11557-009-0594-4","article-title":"Delimitation of Ophiostoma quercus and its synonyms using multiple gene phylogenies","volume":"8","author":"JW Grobbelaar","year":"2009","journal-title":"Mycol Prog"},{"issue":"3","key":"pone.0284393.ref061","first-page":"285","article-title":"Ophiostoma species, including Ophiostoma borealis sp. nov., infecting wounds of native broad-leaved trees in Norway","volume":"31","author":"G Kamgan Nkuekam","year":"2010","journal-title":"Cryptogam, Mycol"},{"key":"pone.0284393.ref062","doi-asserted-by":"crossref","first-page":"111","DOI":"10.47371\/S10267-010-0081-4","article-title":"Discovery of Ophiostoma tsotsi on Eucalyptus wood chips in China","volume":"52","author":"JW Grobbelaar","year":"2011","journal-title":"Mycoscience"},{"key":"pone.0284393.ref063","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1071\/BT10231","article-title":"Ophiostoma species (Ophiostomatales, Ascomycota), including two new taxa on eucalypts in Australia","volume":"59","author":"G Kamgan Nkuekam","year":"2011","journal-title":"Aust J Bot"},{"key":"pone.0284393.ref064","first-page":"21","volume-title":"Ophiostomatoid fungi: expanding frontiers","author":"ZW de Beer","year":"2013"},{"key":"pone.0284393.ref065","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1080\/07060660409507116","article-title":"Fungi Canadenses No. 347. Pesotum fragrans","volume":"26","author":"K Jacobs","year":"2004","journal-title":"Can J Plant Pathol"},{"key":"pone.0284393.ref066","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1007\/s13225-009-0004-x","article-title":"Characterisation of synnematous bark beetle-associated fungi from China, including Graphium carbonarium sp. nov","volume":"40","author":"D Paciura","year":"2010","journal-title":"Fungal Divers"},{"key":"pone.0284393.ref067","unstructured":"Cootsona C. Identification and distribution of Neonectria major causing cankers on red alder (Alnus rubra). M.Sc. Thesis, College of Forest Resources, University of Washington. 2006."},{"issue":"2","key":"pone.0284393.ref068","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1017\/S0890037X00023289","article-title":"Biological control of red alder (Alnus rubra) with the fungus Nectria ditissima","volume":"9","author":"CE Dorworth","year":"1995","journal-title":"Weed Technol"},{"issue":"4","key":"pone.0284393.ref069","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1080\/07060669609500582","article-title":"Biological control of red alder (Alnus rubra) with indigenous pathogenic Ascomycotina","volume":"18","author":"CE Dorworth","year":"1996","journal-title":"Can J Plant Pathol"},{"key":"pone.0284393.ref070","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1007\/s00248-007-9251-0","article-title":"Geosmithia fungi are highly diverse and consistent bark beetle associates: evidence from their community structure in temperate Europe","volume":"55","author":"M Kola\u0159\u00edk","year":"2008","journal-title":"Microb Ecol"},{"key":"pone.0284393.ref071","doi-asserted-by":"crossref","first-page":"965","DOI":"10.1139\/w2012-069","article-title":"Identification and characterization of GEO1, a new class II hydrophobin from Geosmithia spp","volume":"58","author":"PP Bettini","year":"2012","journal-title":"Can J Microbiol"},{"key":"pone.0284393.ref072","doi-asserted-by":"crossref","first-page":"500","DOI":"10.1111\/j.1439-0329.2009.00626.x","article-title":"Factors influencing infection of Acacia mearnsii by the wilt pathogen Ceratocystis albifundus in South Africa","volume":"40","author":"RN Heath","year":"2010","journal-title":"For Pathol"},{"issue":"5","key":"pone.0284393.ref073","doi-asserted-by":"crossref","first-page":"1","DOI":"10.14214\/sf.1328","article-title":"Ophiostomatoid fungi and their roles in Quercus robur die-back in Tellermann forest, Russia","volume":"49","author":"NN Selochnik","year":"2015","journal-title":"Silva Fenn"},{"issue":"4","key":"pone.0284393.ref074","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1093\/forestry\/cpr018","article-title":"Decay, yield loss and associated fungi in stands of grey alder (Alnus incana) in Latvia","volume":"84","author":"N Arhipova","year":"2011","journal-title":"Forestry"},{"key":"pone.0284393.ref075","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1007\/978-1-4020-2241-8_10","volume-title":"Bark and Wood Boring Insects in Living Trees in Europe, a Synthesis","author":"T. Kirisits","year":"2007"},{"key":"pone.0284393.ref076","doi-asserted-by":"crossref","first-page":"9","DOI":"10.3767\/003158509X468038","article-title":"Ophiostoma denticiliatum sp. nov. and other Ophiostoma species associated with the birch bark beetle in southern Norway","volume":"23","author":"R Linnakoski","year":"2009","journal-title":"Persoonia"},{"key":"pone.0284393.ref077","doi-asserted-by":"crossref","first-page":"801","DOI":"10.1051\/forest:2006063","article-title":"Fungi associated with Tomicus piniperda in Poland and assessment of their virulence using Scots pine seedlings","volume":"63","author":"R. Jankowiak","year":"2006","journal-title":"Ann For Sci"},{"key":"pone.0284393.ref078","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1007\/s10310-003-0056-9","article-title":"Ophiostoma species associated with bark beetles infesting three Abies species in Nikko, Japan","volume":"9","author":"Y Yamaoka","year":"2004","journal-title":"J For Res"},{"key":"pone.0284393.ref079","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1007\/s00248-011-9841-8","article-title":"Spatial community structure of mountain pine beetle fungal symbionts across a latitudinal gradient","volume":"62","author":"AD Roe","year":"2011","journal-title":"Microb Ecol"},{"key":"pone.0284393.ref080","first-page":"211","article-title":"The biology of a species of Beauveria from the southern pine bark beetle","volume":"1","author":"JG Harrar","year":"1940","journal-title":"Proc W Va Acad Sci"},{"issue":"2","key":"pone.0284393.ref081","doi-asserted-by":"crossref","first-page":"188","DOI":"10.1016\/0022-2011(81)90074-4","article-title":"Comparative laboratory studies on three fungal pathogens of the elm bark beetle Scolytus scolytus: pathogenicity of Beauveria bassiana, Metarhizium anisopliae, and Paecilomyces farinosus to larvae and adults of S. scolytus","volume":"37","author":"JW Doberski","year":"1981","journal-title":"J Invertebr Pathol"},{"key":"pone.0284393.ref082","first-page":"149","article-title":"Valsalnicola oxystoma (Rehm) D.M. Walker & Rossman, comb. nov","volume":"128","author":"DM Walker","year":"2012","journal-title":"Fungal Planet"},{"issue":"4","key":"pone.0284393.ref083","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1080\/02827581.2012.670727","article-title":"Heart-rot and associated fungi in Alnus glutinosa stands in Latvia","volume":"27","author":"N Arhipova","year":"2012","journal-title":"Scand J Forest Res"},{"key":"pone.0284393.ref084","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1071\/AP04024","article-title":"Pathogenicity of fungi associated with a decay of kiwifruit","volume":"33","author":"S Di Marco","year":"2004","journal-title":"Australas Plant Pathol"},{"key":"pone.0284393.ref085","doi-asserted-by":"crossref","first-page":"2195","DOI":"10.1007\/s10482-018-1112-y","article-title":"Cadophora margaritata sp. nov. and other fungi associated with the longhorn beetles Anoplophora glabripennis and Saperda carcharias in Finland","volume":"111","author":"R Linnakoski","year":"2018","journal-title":"Antonie Van Leeuwenhoek"},{"key":"pone.0284393.ref086","unstructured":"Safranyik L, Whitney HS, Bleiker KP. A survey of microorganisms from the spruce beetle in central British Columbia. Information Report. Victoria: Pacific Forestry Centre, Canadian Forest Service; 2010. Report No.: BC-X-420. Available from: https:\/\/publications.gc.ca\/site\/eng\/9.566475\/publication.html"},{"key":"pone.0284393.ref087","doi-asserted-by":"crossref","first-page":"187","DOI":"10.1111\/j.1744-7348.1990.tb04206.x","article-title":"Biological control of Dutch elm disease: larvicidal activity of Trichoderma harzianum, T. polysporum and Scytalidium lignicola in Scolytus scolytus and S. multistriatus reared in artificial culture","volume":"117","author":"HK Jassim","year":"1990","journal-title":"Ann Appl Biol"},{"key":"pone.0284393.ref088","doi-asserted-by":"crossref","first-page":"636","DOI":"10.1111\/j.1365-2311.2006.00829.x","article-title":"Bacteria in oral secretions of an endophytic insect inhibit antagonistic fungi","volume":"31","author":"YJ Cardoza","year":"2006","journal-title":"Ecol Entomol"},{"issue":"4","key":"pone.0284393.ref089","doi-asserted-by":"crossref","first-page":"561","DOI":"10.14411\/eje.2014.070","article-title":"Fungi associated with the red-haired bark beetle, Hylurgus ligniperda (Coleoptera: Curculionidae) in the forest-steppe zone in eastern Ukraine","volume":"111","author":"K Davydenko","year":"2014","journal-title":"Eur J Entomol"},{"key":"pone.0284393.ref090","doi-asserted-by":"crossref","first-page":"184","DOI":"10.1111\/j.1439-0329.1990.tb01129.x","article-title":"Relative effectiveness of Scolytus scolytus, S. multistriatus and S. kirschi as vectors of Dutch elm disease","volume":"20","author":"JF Webber","year":"1990","journal-title":"Eur J Forest Pathol"},{"key":"pone.0284393.ref091","doi-asserted-by":"crossref","first-page":"1331","DOI":"10.1139\/x03-047","article-title":"A comparison of mycangial and phoretic fungi of individual mountain pine beetles","volume":"33","author":"DL Six","year":"2003b","journal-title":"Can J For Res"},{"key":"pone.0284393.ref092","doi-asserted-by":"crossref","first-page":"1815","DOI":"10.1139\/x03-107","article-title":"Fungi associated with the North American spruce beetle, Dendroctonus rufipennis","volume":"33","author":"DL Six","year":"2003","journal-title":"Can J For Res"},{"key":"pone.0284393.ref093","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1007\/s10341-014-0210-x","article-title":"Biology and control of the apple canker fungus Neonectria ditissima (syn. N. galligena) from a Northwestern European perspective","volume":"56","author":"RWS Weber","year":"2014","journal-title":"Erwerbs-Obstbau"},{"issue":"2","key":"pone.0284393.ref094","doi-asserted-by":"crossref","first-page":"325","DOI":"10.3852\/10-124","article-title":"Geosmithia morbida sp. nov., a new phytopathogenic species living in symbiosis with the walnut twig beetle (Pityophthorus juglandis) on Juglans in USA","volume":"103","author":"M Kola\u0159\u00edk","year":"2011","journal-title":"Mycologia"},{"key":"pone.0284393.ref095","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1094\/PDIS-91-3-0245","article-title":"Association of Ophiostoma novo-ulmi with Scolytus schevyrewi (Scolytidae) in Colorado","volume":"91","author":"WR Jacobi","year":"2007","journal-title":"Plant Dis"},{"key":"pone.0284393.ref096","first-page":"55","article-title":"First report of Ophiostoma karelicum from birch stands in Poland","volume":"59","author":"R. Jankowiak","year":"2011","journal-title":"Phytopathologia"},{"issue":"11","key":"pone.0284393.ref097","first-page":"510","article-title":"Transmission of fungus spores by bark beetles of the genus Scolytus with respect to the oak bark beetle (Scolytus intricatus Ratz.)","volume":"42","author":"P. \u0160r\u016ftka","year":"1996","journal-title":"Lesnictv\u00ed-Forestry"},{"key":"pone.0284393.ref098","first-page":"414","article-title":"The small oak barkbeetle transmits the oak wilt disease","volume":"56","author":"WD Buchanan","year":"1958","journal-title":"Journal of Forestry"},{"key":"pone.0284393.ref099","first-page":"213","article-title":"Ceratocystis manginecans sp. nov., causal agent of a destructive mango wilt disease in Oman and Pakistan","volume":"27","author":"M Van Wyk","year":"2007","journal-title":"Fungal Divers"},{"key":"pone.0284393.ref100","doi-asserted-by":"crossref","first-page":"381","DOI":"10.5248\/117.381","article-title":"Two new Ceratocystis species associated with mango disease in Brazil","volume":"117","author":"M Van Wyk","year":"2011","journal-title":"Mycotaxon"},{"key":"pone.0284393.ref101","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1007\/s10658-012-0081-7","article-title":"The mango sudden decline pathogen, Ceratocystis manginecans, is vectored by Hypocryphalus mangiferae (Coleoptera: Scolytinae) in Oman","volume":"135","author":"AO Al Adawi","year":"2013","journal-title":"Eur J Plant Pathol"},{"key":"pone.0284393.ref102","volume-title":"Insect transmission of plant diseases","author":"JG Leach","year":"1940"}],"container-title":["PLOS ONE"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0284393","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,5,8]],"date-time":"2023-05-08T18:30:38Z","timestamp":1683570638000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0284393"}},"subtitle":[],"editor":[{"given":"Simon Francis","family":"Shamoun","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2023,5,8]]},"references-count":102,"journal-issue":{"issue":"5","published-online":{"date-parts":[[2023,5,8]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pone.0284393","relation":{},"ISSN":["1932-6203"],"issn-type":[{"value":"1932-6203","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,5,8]]}}}