A taxonomic monograph of Nearctic Scolytus Geoffroy (Coleoptera, Curculionidae, Scolytinae)
Identifieur interne : 000275 ( Pmc/Corpus ); précédent : 000274; suivant : 000276A taxonomic monograph of Nearctic Scolytus Geoffroy (Coleoptera, Curculionidae, Scolytinae)
Auteurs : Sarah M. Smith ; Anthony I. CognatoSource :
- ZooKeys [ 1313-2989 ] ; 2014.
Abstract
The Nearctic bark beetle genus
Url:
DOI: 10.3897/zookeys.450.7452
PubMed: 25408617
PubMed Central: 4233402
Links to Exploration step
PMC:4233402Le document en format XML
<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">A taxonomic monograph of Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> Geoffroy (<named-content content-type="taxon-name"><named-content content-type="order">Coleoptera</named-content></named-content>, <named-content content-type="taxon-name"><named-content content-type="family">Curculionidae</named-content></named-content>, <named-content content-type="taxon-name"><named-content content-type="subfamily">Scolytinae</named-content></named-content>)</title><author><name sortKey="Smith, Sarah M" sort="Smith, Sarah M" uniqKey="Smith S" first="Sarah M." last="Smith">Sarah M. Smith</name><affiliation><nlm:aff id="A1">Department of Entomology, Michigan State University, Natural Science Building, room 243, 288 Farm Lane, East Lansing, MI 48824, United States of America</nlm:aff></affiliation></author><author><name sortKey="Cognato, Anthony I" sort="Cognato, Anthony I" uniqKey="Cognato A" first="Anthony I." last="Cognato">Anthony I. Cognato</name><affiliation><nlm:aff id="A1">Department of Entomology, Michigan State University, Natural Science Building, room 243, 288 Farm Lane, East Lansing, MI 48824, United States of America</nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">25408617</idno><idno type="pmc">4233402</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4233402</idno><idno type="RBID">PMC:4233402</idno><idno type="doi">10.3897/zookeys.450.7452</idno><date when="2014">2014</date><idno type="wicri:Area/Pmc/Corpus">000275</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000275</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">A taxonomic monograph of Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> Geoffroy (<named-content content-type="taxon-name"><named-content content-type="order">Coleoptera</named-content></named-content>, <named-content content-type="taxon-name"><named-content content-type="family">Curculionidae</named-content></named-content>, <named-content content-type="taxon-name"><named-content content-type="subfamily">Scolytinae</named-content></named-content>)</title><author><name sortKey="Smith, Sarah M" sort="Smith, Sarah M" uniqKey="Smith S" first="Sarah M." last="Smith">Sarah M. Smith</name><affiliation><nlm:aff id="A1">Department of Entomology, Michigan State University, Natural Science Building, room 243, 288 Farm Lane, East Lansing, MI 48824, United States of America</nlm:aff></affiliation></author><author><name sortKey="Cognato, Anthony I" sort="Cognato, Anthony I" uniqKey="Cognato A" first="Anthony I." last="Cognato">Anthony I. Cognato</name><affiliation><nlm:aff id="A1">Department of Entomology, Michigan State University, Natural Science Building, room 243, 288 Farm Lane, East Lansing, MI 48824, United States of America</nlm:aff></affiliation></author></analytic><series><title level="j">ZooKeys</title><idno type="ISSN">1313-2989</idno><idno type="eISSN">1313-2970</idno><imprint><date when="2014">2014</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><label>Abstract</label><p>The Nearctic bark beetle genus <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> Geoffroy was revised based in part on a molecular and morphological phylogeny. Monophyly of the native species was tested using mitochondrial (COI) and nuclear (28S, CAD, ArgK) genes and 43 morphological characters in parsimony and Bayesian phylogenetic analyses. Parsimony analyses of molecular and combined datasets provided mixed results while Bayesian analysis recovered most nodes with posterior probabilities >90%. Native hardwood- and conifer-feeding <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species were recovered as paraphyletic. Native Nearctic species were recovered as paraphyletic with hardwood-feeding species sister to Palearctic hardwood-feeding species rather than to native conifer-feeding species. The Nearctic conifer-feeding species were monophyletic. Twenty-five species were recognized. Four new synonyms were discovered: <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> LeConte, 1868 (= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic> Blackman, 1934; = <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content></italic> Blackman, 1934), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> Blackman, 1934 (= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">virgatus</named-content></named-content></italic> Bright, 1972; = <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic> Blackman, 1934). Two species were reinstated: <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> Blackman, 1934 and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic> Bright, 1972. 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<front><journal-meta><journal-id journal-id-type="nlm-ta">Zookeys</journal-id><journal-id journal-id-type="iso-abbrev">Zookeys</journal-id><journal-id journal-id-type="publisher-id">ZooKeys</journal-id><journal-title-group><journal-title>ZooKeys</journal-title></journal-title-group><issn pub-type="ppub">1313-2989</issn><issn pub-type="epub">1313-2970</issn><publisher><publisher-name>Pensoft Publishers</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">25408617</article-id><article-id pub-id-type="pmc">4233402</article-id><article-id pub-id-type="doi">10.3897/zookeys.450.7452</article-id><article-categories><subj-group subj-group-type="heading"><subject>Monograph</subject></subj-group></article-categories><title-group><article-title>A taxonomic monograph of Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> Geoffroy (<named-content content-type="taxon-name"><named-content content-type="order">Coleoptera</named-content></named-content>, <named-content content-type="taxon-name"><named-content content-type="family">Curculionidae</named-content></named-content>, <named-content content-type="taxon-name"><named-content content-type="subfamily">Scolytinae</named-content></named-content>)</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Smith</surname><given-names>Sarah M.</given-names></name><xref ref-type="aff" rid="A1">1</xref></contrib><contrib contrib-type="author"><name><surname>Cognato</surname><given-names>Anthony I.</given-names></name><xref ref-type="aff" rid="A1">1</xref></contrib></contrib-group><aff id="A1"><label>1</label>Department of Entomology, Michigan State University, Natural Science Building, room 243, 288 Farm Lane, East Lansing, MI 48824, United States of America</aff><author-notes><corresp>Corresponding author: Sarah M. Smith (<email xlink:type="simple">camptocerus@gmail.com</email>)</corresp><fn fn-type="edited-by"><p>Academic editor: M. Alonso-Zarazaga</p></fn></author-notes><pub-date pub-type="collection"><year>2014</year></pub-date><pub-date pub-type="epub"><day>29</day><month>10</month><year>2014</year></pub-date><issue>450</issue><fpage>1</fpage><lpage>182</lpage><history><date date-type="received"><day>7</day><month>3</month><year>2014</year></date><date date-type="accepted"><day>13</day><month>8</month><year>2014</year></date></history><permissions><copyright-statement>Sarah M. Smith, Anthony I. Cognato</copyright-statement><license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/4.0"><license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p></license></permissions><self-uri content-type="zoobank" xlink:type="simple" xlink:href="http://zoobank.org/6EAFB961-1C8C-4A88-BB84-CBCE13CDE663">http://zoobank.org/6EAFB961-1C8C-4A88-BB84-CBCE13CDE663</self-uri><abstract><label>Abstract</label><p>The Nearctic bark beetle genus <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> Geoffroy was revised based in part on a molecular and morphological phylogeny. Monophyly of the native species was tested using mitochondrial (COI) and nuclear (28S, CAD, ArgK) genes and 43 morphological characters in parsimony and Bayesian phylogenetic analyses. Parsimony analyses of molecular and combined datasets provided mixed results while Bayesian analysis recovered most nodes with posterior probabilities >90%. Native hardwood- and conifer-feeding <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species were recovered as paraphyletic. Native Nearctic species were recovered as paraphyletic with hardwood-feeding species sister to Palearctic hardwood-feeding species rather than to native conifer-feeding species. The Nearctic conifer-feeding species were monophyletic. Twenty-five species were recognized. Four new synonyms were discovered: <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> LeConte, 1868 (= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic> Blackman, 1934; = <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content></italic> Blackman, 1934), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> Blackman, 1934 (= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">virgatus</named-content></named-content></italic> Bright, 1972; = <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic> Blackman, 1934). Two species were reinstated: <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> Blackman, 1934 and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic> Bright, 1972. A diagnosis, description, distribution, host records and images were provided for each species and a key is presented to all species.</p></abstract><kwd-group><label>Keywords</label><kwd><named-content content-type="taxon-name"><named-content content-type="family">Scolytidae</named-content></named-content></kwd><kwd>bark beetle</kwd><kwd>fir engraver</kwd><kwd>banded elm bark beetle</kwd><kwd>lesser elm bark beetle</kwd><kwd>Douglas-fir engraver</kwd><kwd>hickory bark beetle</kwd><kwd>shot hole borer</kwd><kwd>large shot hole borer</kwd><kwd>hackberry engraver</kwd><kwd>taxonomy</kwd><kwd>revision</kwd><pmc-comment>PageBreak</pmc-comment>
</kwd-group></article-meta><notes><sec sec-type="Citation"><title>Citation</title><p>Smith SM, Cognato AI (2014) A taxonomic monograph of Nearctic <italic>Scolytus</italic> Geoffroy (Coleoptera, Curculionidae, Scolytinae). ZooKeys 450: 1–182. doi: <ext-link ext-link-type="doi" xlink:href="10.3897/zookeys.450.7452">10.3897/zookeys.450.7452</ext-link></p></sec></notes></front><body><sec sec-type="Introduction"><title>Introduction</title><p>This monograph presents a revision of the Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> Geoffroy, 1762 species (<named-content content-type="taxon-name"><named-content content-type="family">Curculionidae</named-content></named-content>: <named-content content-type="taxon-name"><named-content content-type="subfamily">Scolytinae</named-content></named-content>). The <named-content content-type="taxon-name"><named-content content-type="subfamily">Scolytinae</named-content></named-content> is comprised of approximately 6,000 species in 225 genera and 26 tribes (<xref rid="B1" ref-type="bibr">Alonso-Zarazaga and Lyal 2009</xref>). Termed ‘bark and ambrosia beetles’ this weevil subfamily is ubiquitous in forests worldwide. Many species contribute to the decomposition of dead vegetation while others are capable of causing substantial host mortality. True bark beetles, including <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic>, feed exclusively on phloem and cambium of living, declining, or dead trees and some species specialize on different plant parts from root to fruit and pith (<xref rid="B148" ref-type="bibr">Wood 1982</xref>). Beetle feeding on phloem and xylem hastens the decomposition of trees and the introduction of other xylophagous organisms (<xref rid="B126" ref-type="bibr">Stokland 2012</xref>). Some bark beetle species kill live trees by passively introducing fungi into otherwise healthy hosts as a by-product of phloem feeding and the galleries may girdle the host causing mortality (<xref rid="B148" ref-type="bibr">Wood 1982</xref>). At high density these species cause widespread destruction of economically valuable tree species, giving bark beetles their nefarious reputation as ecologically and economically destructive forest pests (<xref rid="B55" ref-type="bibr">Furniss and Carolin 1977</xref>). Other economically important bark beetle genera such as <italic><named-content content-type="taxon-name"><named-content content-type="genus">Dendroctonus</named-content></named-content></italic> Erichson, 1836 and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ips</named-content></named-content></italic> DeGeer, 1775 have received considerable attention and therefore have a reasonably solid understanding of the taxonomy (<xref rid="B151" ref-type="bibr">Wood and Bright 1992</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> is also important but has received considerably less attention.</p></sec><sec><title>Natural history</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> are distributed in the Holarctic, Oriental (Himalayan) and Neotropical regions and some Palearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species have been introduced around the world to the Nearctic, South Africa, Australia, New Zealand and temperate South America (<xref rid="B111" ref-type="bibr">Rosel and French 1975</xref>; <xref rid="B4" ref-type="bibr">Bain 1990</xref>; <xref rid="B151" ref-type="bibr">Wood and Bright 1992</xref>; <xref rid="B119" ref-type="bibr">Six et al. 2005</xref>; <xref rid="B150" ref-type="bibr">Wood 2007</xref>; <xref rid="B122" ref-type="bibr">Smith and Cognato 2013</xref>). In the Nearctic region, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species occur from the Atlantic to Pacific oceans and from the boundary of the Neotropical region to the northern limits of tree growth. In the Nearctic, native hardwood-feeders are generally found from the Atlantic coast to Texas and west to the foothills of the Rocky Mountains. With the exception of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic> (Swaine, 1910), conifer-feeding <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> are restricted to the occurrence of host trees in western mountain ranges including the Rocky Mountains. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic> has an expansive range from the east and west coasts of North America and from northern California and Colorado north to beyond the Arctic Circle. Invasive <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species are found throughout the US, northern Mexico and southern Canada (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic> (Bechstein, 1805), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> (Marsham, 1802), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic> (Müller, 1818), and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> Semenov, 1902).</p><p>In North America, <pmc-comment>PageBreak</pmc-comment><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species occur either in conifer or hardwood hosts (Table <xref ref-type="table" rid="T1">1</xref>). Members of the <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> conifer-feeding clade feed exclusively on <named-content content-type="taxon-name"><named-content content-type="family">Pinaceae</named-content></named-content> genera, including <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> Mill., <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content></named-content></italic> Mill., <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content></named-content></italic> D. Don. ex Loudon, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content></named-content></italic> Carrière, and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content></named-content></italic> Carrière, with the notable exception of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content></named-content></italic> L. (<xref rid="B151" ref-type="bibr">Wood and Bright 1992</xref>) (Table <xref ref-type="table" rid="T2">2</xref>). Members of the hardwood-feeding clade, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic> Say, 1824, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> Walsh, 1867, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic> Say, 1824 feed on the families <named-content content-type="taxon-name"><named-content content-type="family">Cannabaceae</named-content></named-content> (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content></named-content></italic> L.), <named-content content-type="taxon-name"><named-content content-type="family">Fagaceae</named-content></named-content> (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Fagus</named-content></named-content></italic> L., <italic><named-content content-type="taxon-name"><named-content content-type="genus">Quercus</named-content></named-content></italic> L.) and <named-content content-type="taxon-name"><named-content content-type="family">Juglandaceae</named-content></named-content> (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Carya</named-content></named-content></italic> Nutt.), respectively (Table <xref ref-type="table" rid="T1">1</xref>).</p><table-wrap id="T1" orientation="portrait" position="float"><label>Table 1.</label><caption><p>Host plant families for all Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic>.</p></caption><table frame="hsides" rules="all"><tbody><tr><th rowspan="1" colspan="1">Species</th><th rowspan="1" colspan="1"><named-content content-type="taxon-name"><named-content content-type="family">Cannabaceae</named-content></named-content></th><th rowspan="1" colspan="1"><named-content content-type="taxon-name"><named-content content-type="family">Fagaceae</named-content></named-content></th><th rowspan="1" colspan="1"><named-content content-type="taxon-name"><named-content content-type="family">Juglandaceae</named-content></named-content></th><th rowspan="1" colspan="1"><named-content content-type="taxon-name"><named-content content-type="family">Pinaceae</named-content></named-content></th><th rowspan="1" colspan="1"><named-content content-type="taxon-name"><named-content content-type="family">Rosaceae</named-content></named-content></th><th rowspan="1" colspan="1"><named-content content-type="taxon-name"><named-content content-type="family">Ulmaceae</named-content></named-content></th></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1">■</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr></tbody></table></table-wrap><table-wrap id="T2" orientation="portrait" position="float"><label>Table 2.</label><caption><p><named-content content-type="taxon-name"><named-content content-type="family">Pinaceae</named-content></named-content> host genus for conifer-feeding Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic>.</p></caption><table frame="hsides" rules="all"><tbody><tr><th rowspan="1" colspan="1">Species</th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content></named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content></named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content></named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content></named-content></italic></th></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic></td><td rowspan="1" colspan="1">■</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr></tbody></table></table-wrap><p>In the Nearctic region, six <pmc-comment>PageBreak</pmc-comment><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species have the potential to kill host trees and cause significant mortality of either conifers (<xref rid="B32" ref-type="bibr">Cibrián Tovar et al. 1995</xref>) or <italic><named-content content-type="taxon-name"><named-content content-type="genus">Carya</named-content></named-content></italic> spp. (<xref rid="B55" ref-type="bibr">Furniss and Carolin 1977</xref>). Two of these are the exotic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic>, both are key vectors of the Dutch elm disease fungi, pathogens that have killed millions of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> L. spp. trees in forest and urban areas across much of the US (<xref rid="B55" ref-type="bibr">Furniss and Carolin 1977</xref>). Mortality caused by various <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species is often sporadic and short-term, although some outbreaks locally affect thousands of acres a year (<xref rid="B55" ref-type="bibr">Furniss and Carolin 1977</xref>). Damage is most severe in times of environmental stress, which is mostly associated with drought and other insect infestations. For example, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> LeConte, 1876 killed approximately 72,843 hectares of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> spp. in California alone in a year (<xref rid="B55" ref-type="bibr">Furniss and Carolin 1977</xref>).</p><p>All Nearctic <pmc-comment>PageBreak</pmc-comment><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species are monogamous. Females select brood material and begin galley construction. Males walk across the host material in search of females. Mating occurs with the female in the entrance tunnel and the male on the bark. However, mating in twig crotches has been reported for <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> (<xref rid="B129" ref-type="bibr">Svihra and Clark 1980</xref>). The female creates an entrance tunnel at a 45° angle, boring through the bark to the cambium. From the entrance tunnel, she excavates a nuptial chamber and, depending on the species, one or two egg galleries in either direction from the entrance tunnel. The nuptial chamber and galleries are excavated in the cambium but also etch the sapwood. Females excavate egg niches on both sides of the egg galleries and a single egg is deposited in each niche and covered with boring dust. Adult males assist in removing frass and generally stay with the female until egg gallery construction is complete. The male then leaves the gallery and the female dies in the entrance tunnel with her abdomen projecting onto the bark surface. Larval galleries radiate away from the egg tunnels as they feed on phloem, also etching the sapwood. Once larvae mature, the prepupae burrow into the outer sapwood and pupate. The brood overwinters as pupae and adults emerge in the spring (<xref rid="B42" ref-type="bibr">Edson 1967</xref>). Upon emergence, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic>, and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> engage in maturation feeding at twig crotches and/or leaf petioles (<xref rid="B65" ref-type="bibr">Hoffman 1942</xref>; <xref rid="B5" ref-type="bibr">Baker 1972</xref>; <xref rid="B99" ref-type="bibr">Negrón et al. 2005</xref>). Feeding may also occur within small twigs in <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> Blackman, 1934 (reported as <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> LeConte, 1876 in <xref rid="B96" ref-type="bibr">McMullen and Atkins 1962</xref>).</p><p>Gallery shape is directly related to the resin system of the host genus. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content></named-content></italic> possess an elaborate system of resin ducts with vertical and radial ducts that are connected to each other (<xref rid="B87" ref-type="bibr">Lieutier 2004</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> galleries in these genera are consistently parallel to the grain of the wood. The vertical ducts are abundant in these tree genera and when <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> constructs a vertical egg gallery, both ducts are severed, and as the gallery is elongated, only vertical canals are severed. This minimizes exposure of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> to host resins (<xref rid="B87" ref-type="bibr">Lieutier 2004</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content></named-content></italic> lack resin canals and instead possess resin blisters. Galleries in these genera are quite variable and may be transverse, “V” or (“<bold>ε</bold>”) shaped depending on the species because there are no resin canals to avoid (<xref rid="B87" ref-type="bibr">Lieutier 2004</xref>).</p><p>Conifer-feeding <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species exhibit primary attraction to host volatiles rather than to pheromones produced by conspecifics (<xref rid="B89" ref-type="bibr">Macías-Sámano et al. 1998a</xref>). Attraction of hardwood-feeding species is not well understood but primary attraction seems probable for native <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic> and the invasive species <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> (<xref rid="B58" ref-type="bibr">Goeden and Norris 1964a</xref>; <xref rid="B79" ref-type="bibr">Kovach and Gorsuch 1985</xref>; <xref rid="B85" ref-type="bibr">Lee et al. 2010</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> exhibits secondary attraction and to 4-methyl-3-heptanol and multistriatin in combination with alpha-cubenene (<xref rid="B81" ref-type="bibr">Lanier et al. 1977</xref>). At endemic population levels, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> infest over mature, unthrifty or weakened standing trees, shaded out branches, recent logging slash, windthrown trees and fallen branches. During outbreaks, vigorous trees may be colonized by more aggressive species, including <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic> Wood, 1968 and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> LeConte, 1868 (<xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B55" ref-type="bibr">Furniss and Carolin 1977</xref>), and the secondary species <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> (Swaine, 1917), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> Blackman, 1934 [reported as <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic>] and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> LeConte, 1876 (<xref rid="B95" ref-type="bibr">McMullen and Atkins 1959</xref>; <xref rid="B136" ref-type="bibr">USDA 2004</xref>). While <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species are generally common in the forest, they are rarely abundant and seldom encountered by collectors. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species prefer fresh and moist host material. Sun-baked material is not preferred, but <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> galleries may be present in cooler, moister areas of the bark that are heavily shaded or on the underside of fallen trees and branches. Infested conifer branches and trees typically bear green needles while infested hardwoods have yellowing leaves (Smith, pers. obs.). The entrance can be identified by peeling off the bark flakes with a knife and searching for white to tan-colored boring dust or by close examination of the lateral and ventral sides of trunks and branches of smooth barked hosts. In <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content></named-content></italic>, the entrance tunnels of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudohylesinus</named-content></named-content></italic> Swaine, 1917 can be easily confused with those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic>, but <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> entrance tunnels are at a steeper angle and the boring dust is reddish rather than white. Some <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species, including <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic> LeConte, 1876 typically attack shaded out branches on standing trees. Yellowing needles are indicative of an infested tree. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> females typically conceal their entrance tunnels under bark flakes (common in <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content></named-content></italic>) or in rough patches of bark (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content></named-content></italic>).</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec><title>Sexual dimorphism</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species are sexually dimorphic. Sexually dimorphic structures are typically on the frons and epistoma (Fig. <xref ref-type="fig" rid="F1">1</xref>) and the abdominal ventrites and vary by clade (see clade discussions in results). When viewed laterally, males typically have a flattened, impressed frons and the female frons is always more strongly convex. The male frons is more strongly and coarsely longitudinally aciculate than the female and covered with longer, more abundant and dense erect setae. The epistomal process (when present) is more strongly developed in the male and less developed in the female. With the exception of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic>, the spines, tubercles, denticles, carinae and tumescence on the venter are more pronounced in males.</p><fig id="F1" orientation="portrait" position="float"><label>Figure 1.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> typical sexual dimorphism of the frons and epistoma as exhibited by <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic>. <bold>A</bold> male frons, frontal <bold>B</bold> male frontal oblique <bold>C</bold> female frons, frontal <bold>D</bold> female frontal oblique.</p></caption><graphic id="oo_33354.jpg" xlink:href="zookeys-450-001-g001"></graphic></fig><pmc-comment>PageBreak</pmc-comment>
</sec><sec><title>Systematics</title><p>The <named-content content-type="taxon-name"><named-content content-type="tribe">Scolytini</named-content></named-content> currently contains 213 species and six genera including <italic><named-content content-type="taxon-name"><named-content content-type="genus">Camptocerus</named-content></named-content></italic> Dejean, 1821, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ceratolepis</named-content></named-content></italic> Chapuis, 1869, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Cnemonyx</named-content></named-content></italic> Eichhoff, 1868, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Loganius</named-content></named-content></italic> Chapuis, 1869, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytopsis</named-content></named-content></italic> Blandford, 1896 and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> (<xref rid="B122" ref-type="bibr">Smith and Cognato 2013</xref>). With the exception of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Camptocerus</named-content></named-content></italic>, the tribe consists of cambium-feeding bark beetles. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species that feed exclusively on either hardwoods or conifers (<xref rid="B149" ref-type="bibr">Wood 1986</xref>). Twenty Nearctic and three Palearctic species feed on conifers. All other <named-content content-type="taxon-name"><named-content content-type="tribe">Scolytini</named-content></named-content> are restricted to angiosperm hosts (<xref rid="B151" ref-type="bibr">Wood and Bright 1992</xref>; <xref rid="B121" ref-type="bibr">Smith and Cognato 2010b</xref>; <xref rid="B122" ref-type="bibr">Smith and Cognato 2013</xref>).</p><p>A single apical, unarmed spine-like process that curves toward and extends beyond the process of the inner apical angle of the protibia and by a seven-segmented funicle readily distinguishes the tribe. All display a strongly sexually dimorphic head bearing hair-like setae, with the male frons variously excavated and female frons typically flat to convex. The eye is entire and the posterior area of head is subtruncate. The pronotum is unarmed with a costate lateral margin. In addition, the metapleural suture descends subventrally to the groove, receiving the groove on the costal margin of the elytra, then turns abruptly and parallels the groove near the metacoxal process (<xref rid="B144" ref-type="bibr">Wood 1972</xref>, <xref rid="B147" ref-type="bibr">1978</xref>, <xref rid="B148" ref-type="bibr">1982</xref>, <xref rid="B149" ref-type="bibr">1986</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> is perhaps one of the easiest bark beetle genera to recognize with its distinctive ascendant second abdominal ventrite, the slightly if at all declivous elytra and the depressed basal area of the elytra around the triangular scutellum (<xref rid="B148" ref-type="bibr">Wood 1982</xref>).</p></sec><sec><title>Taxonomic history</title><p>There are currently about 127 recognized <pmc-comment>PageBreak</pmc-comment><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species in the world (<xref rid="B151" ref-type="bibr">Wood and Bright 1992</xref>; <xref rid="B150" ref-type="bibr">Wood 2007</xref>; <xref rid="B78" ref-type="bibr">Knížek 2011</xref>; <xref rid="B104" ref-type="bibr">Petrov 2013</xref>; <xref rid="B122" ref-type="bibr">Smith and Cognato 2013</xref>). Twenty-five species occur in the Nearctic region, including four invasive Palearctic species. Several Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species were described as the North American continent was explored. General entomologists including <xref rid="B112" ref-type="bibr">Say (1824)</xref>, <xref rid="B109" ref-type="bibr">Riley (1867)</xref> and <xref rid="B137" ref-type="bibr">Walsh (1867)</xref> first described the eastern hardwood-feeders. <xref rid="B82" ref-type="bibr">LeConte (1868</xref>, <xref rid="B83" ref-type="bibr">1876</xref>) described the most common California species and species native to the interior mountain ranges. The prominent scolytine taxonomists <xref rid="B130" ref-type="bibr">Swaine (1910</xref>, <xref rid="B131" ref-type="bibr">1917</xref>), <xref rid="B10" ref-type="bibr">Blackman (1934)</xref>, <xref rid="B140" ref-type="bibr">Wood (1962</xref>, <xref rid="B142" ref-type="bibr">1967</xref>, <xref rid="B143" ref-type="bibr">1968</xref>) and <xref rid="B14" ref-type="bibr">Bright (1964</xref>, <xref rid="B16" ref-type="bibr">1972</xref>) also described several <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">californicus</named-content></named-content></italic><xref rid="B82" ref-type="bibr">LeConte (1868)</xref> was described from “California” but <xref rid="B10" ref-type="bibr">Blackman (1934)</xref> deemed this species a synonym of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">scolytus</named-content></named-content></italic> (Fabricius, 1775). Blackman also suggested that the specimen given to LeConte might have been incorrectly labeled with the ‘California’ locality. This species has been intercepted to the United States on numerous occasions but has never become established (<xref rid="B10" ref-type="bibr">Blackman 1934</xref>; <xref rid="B148" ref-type="bibr">Wood 1982</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">scolytus</named-content></named-content></italic> was therefore excluded from this monograph. <xref rid="B10" ref-type="bibr">Blackman (1934)</xref> and <xref rid="B29" ref-type="bibr">China (1962</xref>, <xref rid="B30" ref-type="bibr">1963</xref>) provide a detailed account of the taxonomic history of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> and outline the <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic>/<italic><named-content content-type="taxon-name"><named-content content-type="genus">Eccoptogaster</named-content></named-content></italic> Herbst, 1793 controversy, which occurred from the late 1800’s to the early 1900’s that resulted from intense debate on whether <xref rid="B57" ref-type="bibr">Geoffroy’s (1762)</xref> description was valid. <xref rid="B29" ref-type="bibr">China (1962</xref>, <xref rid="B30" ref-type="bibr">1963</xref>) outlines the International Commission on Zoological Nomenclature ruling that preserved the name <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> over <italic><named-content content-type="taxon-name"><named-content content-type="genus">Eccoptogaster</named-content></named-content></italic>.</p><p>Historically, there has been a great deal of uncertainty regarding the status of several <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species, particularly: <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> (Swaine, 1917); <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic> Blackman, 1934 and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content></italic> Blackman, 1934; <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic> Blackman, 1934; <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> Blackman, 1934 and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic>. <xref rid="B10" ref-type="bibr">Blackman (1934)</xref> and <xref rid="B148" ref-type="bibr">Wood (1982)</xref> formally revised the Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic>. A third revision by <xref rid="B42" ref-type="bibr">Edson (1967)</xref> was a M.S. thesis but was never published. <xref rid="B148" ref-type="bibr">Wood (1982)</xref> synonymized several species described by Blackman including <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic>. However, <xref rid="B42" ref-type="bibr">Edson (1967)</xref> recognized these species based on morphological, ecological and geographical differences. One of these species, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic> Blackman, was recently removed from synonymy because of such differences (<xref rid="B46" ref-type="bibr">Equihua-Martínez and Furniss 2009</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species are generally recognized based on differences in male morphology, specifically shape of the venter concavity or lack thereof, ascending angle and protrusion of the basal margin of the second abdominal ventrite, the placement and shape of the spine or lack thereof on the second ventrite, the presence or absence of a spine on the margin of ventrite 2 and 3 and frons aciculation and vestiture. Differences among species are generally subtle and substantial knowledge of the intraspecific variation is needed for accurate identifications. In addition, there is a paucity of diagnostic morphological characters for females; to date diagnostic characters are known only for females of hardwood-feeding species (<xref rid="B120" ref-type="bibr">Smith and Cognato 2010a</xref>). Association with a male or comparison to previously identified specimens is needed to determine the species of female specimens of conifer-feeders. Previous keys (<xref rid="B83" ref-type="bibr">LeConte 1876</xref>; <xref rid="B132" ref-type="bibr">Swaine 1918</xref>; <xref rid="B10" ref-type="bibr">Blackman 1934</xref>; <xref rid="B24" ref-type="bibr">Chamberlin 1939</xref>, <xref rid="B25" ref-type="bibr">1958</xref>; <xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>; <xref rid="B17" ref-type="bibr">Bright 1976</xref>; <xref rid="B148" ref-type="bibr">Wood 1982</xref>; <xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>) did not address the above deficiencies in species identification. In addition, previous authors were not explicit about their species concepts, raising doubts about species boundaries.</p><p>Unlike previous investigations (<xref rid="B10" ref-type="bibr">Blackman 1934</xref>; <xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B17" ref-type="bibr">Bright 1976</xref>; <xref rid="B148" ref-type="bibr">Wood 1982</xref>), this study incorporates molecular and morphological data in phylogenetic analyses, which provides a basis for assessing species boundaries. This is the first modern taxonomic treatment of any group of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species and will serve as the basis from which the rest of the genus will be revised. This regional monograph of the Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species provides a review of taxonomic characters, an assessment of their phylogenetic utility, an evaluation of intraspecific variation for each species, a key to both sexes and fully illustrates each species and diagnostic characters. We recognize 25 Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species.</p></sec><sec sec-type="materials"><title>Materials</title><sec sec-type="Morphology"><title>Morphology</title><p>This revision is based upon examination of 4,996 adult ingroup specimens and 447 outgroup specimens. Specimens were obtained from field collecting events, reared from host material, or borrowed from North American and European collections. Palearctic <pmc-comment>PageBreak</pmc-comment>species <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">intricatus</named-content></named-content></italic> (Ratzeburg, 1837), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laevis</named-content></named-content></italic> Chapuis, 1869, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">pygmaeus</named-content></named-content></italic> (Fabricius, 1787), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ratzeburgii</named-content></named-content></italic> E.W. Janson, 1856, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">scolytus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sinopiceus</named-content></named-content></italic> Tsai, 1962 and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sulcifrons</named-content></named-content></italic> Rey, 1892 were selected as outgroups. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">propinquus</named-content></named-content></italic> Blandford, 1896, a Neotropical species, was selected as the root taxon. These outgroup taxa were chosen based on the results of a large <named-content content-type="taxon-name"><named-content content-type="tribe">Scolytini</named-content></named-content> phylogenetic analysis (Smith et al. in prep.) which found Neotropical <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> sister to Holarctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic>. The following entomological collection abbreviations (most following <xref rid="B2" ref-type="bibr">Arnett et al. 1993</xref>) are referenced in the text. Names of the curators that prepared loans are listed in parentheses.</p><p><abbrev>ANSP<def><p> Academy of Natural Sciences, Philadelphia, Pennsylvania</p></def></abbrev>;</p><p><abbrev>AMNH<def><p> American Museum of Natural History, New York, New York (Lee Herman and Aaron Smith)</p></def></abbrev>;</p><p><abbrev>CASC<def><p> California Academy of Sciences, San Francisco, California (David Kavanaugh)</p></def></abbrev>;</p><p><abbrev>CSCA<def><p> California State Collection of Arthropods, Sacramento, California (Andrew Cline and Jacqueline Kishmirian)</p></def></abbrev>;</p><p><abbrev>CNCI<def><p> Canadian National Collection of Insects, Ottawa, Ontario, Canada (Hume Douglas, and Patrice Bouchard)</p></def></abbrev>;</p><p><abbrev>CSUC<def><p> C. P. Gillette Museum of Arthropod Biodiversity, Colorado State University, Fort Collins, Colorado (Boris Kondratieff)</p></def></abbrev>;</p><p><abbrev>CUIC<def><p> Cornell University Insect Collection, Cornell University, Ithaca, New York (James Liebherr)</p></def></abbrev>;</p><p><abbrev>DEBC<def><p> Donald E. Bright, Jr. Collection, Fort Collins, Colorado (Donald E. Bright, Jr.), to be housed at the CNCI</p></def></abbrev>;</p><p><abbrev>EMEC<def><p> Essig Museum of Entomology, University of California Berkeley, Berkeley, California (Cheryl Barr)</p></def></abbrev>;</p><p><abbrev>FMNH<def><p> Field Museum of Natural History, Chicago, Illinois (James Boone)</p></def></abbrev>;</p><p><abbrev>FSCA<def><p> Florida State Collection of Arthropods, Gainesville, Florida (Paul Skelley)</p></def></abbrev>;</p><p><abbrev>ISNB<def><p> Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium (Pol Limbourg)</p></def></abbrev>;</p><p><abbrev>MCZC<def><p> Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (Philip Perkins)</p></def></abbrev>;</p><p><abbrev>MSUC<def><p> Albert J. Cook Arthropod Research Collection, Michigan State University, East Lansing, Michigan (Gary Parsons)</p></def></abbrev>;</p><p><abbrev>NHMW<def><p> Naturhistorisches Museum Wien, Vienna, Austria (Harald Schillhammer)</p></def></abbrev>;</p><p><abbrev>OSAC<def><p> Oregon State Arthropod Collection, Oregon State University, Corvallis, Oregon (Christopher Marshall)</p></def></abbrev>;</p><p><abbrev>RJRC<def><p> Robert J. Rabaglia Collection, Annapolis, Maryland (Robert J. Rabaglia)</p></def></abbrev>;</p><p><abbrev>SBMN<def><p> Santa Barbara Museum of Nature, Santa Barbara, California (Michael Caterino)</p></def></abbrev>;</p><p><abbrev>SMEC<def><p> Snow Entomological Museum, Lawrence, Kansas (Zachary Falin)</p></def></abbrev>;</p><p><abbrev>THAC<def><p> Thomas H. Atkinson Collection, Austin, Texas (Thomas Atkinson)</p></def></abbrev>;</p><pmc-comment>PageBreak</pmc-comment>
<p><abbrev>WFBM<def><p> William F. Barr Entomological Collection, University of Idaho, Moscow, Idaho (Frank Merickel)</p></def></abbrev>;</p><p><abbrev>UMMZ<def><p> Museum of Zoology, University of Michigan, Ann Arbor, Michigan (Mark O’Brien)</p></def></abbrev>;</p><p><abbrev>USNM<def><p> National Museum of Natural History, Smithsonian Institution, Washington, DC (Including Stephen L. Wood Collection) (Natalia Vandenberg)</p></def></abbrev>;</p><p><abbrev>ZIFH<def><p> Zoologische Institut der Forsliche Hochschule, Eberswald, Germany</p></def></abbrev>;</p><p><abbrev>ZMBN<def><p> University Museum of Bergen, The Natural History Collections, Bergen, Norway (Bjarte Jordal)</p></def></abbrev>.</p><p>Additional distribution records were compiled from the following publications: <xref rid="B9" ref-type="bibr">Blackman 1922</xref>, <xref rid="B10" ref-type="bibr">1934</xref>; <xref rid="B40" ref-type="bibr">Doane et al. 1936</xref>; <xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>; <xref rid="B17" ref-type="bibr">Bright 1976</xref>; <xref rid="B55" ref-type="bibr">Furniss and Carolin 1977</xref>; <xref rid="B51" ref-type="bibr">Furniss and Johnson 1987</xref>; <xref rid="B56" ref-type="bibr">Gast et al. 1989</xref>; <xref rid="B151" ref-type="bibr">Wood and Bright 1992</xref>; <xref rid="B32" ref-type="bibr">Cibrián Tovar et al. 1995</xref>; <xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>; <xref rid="B54" ref-type="bibr">Furniss et al. 2002</xref>; <xref rid="B84" ref-type="bibr">Lee et al. 2006</xref>; <xref rid="B92" ref-type="bibr">Majka et al. 2007a</xref>,<xref rid="B93" ref-type="bibr">b</xref>; <xref rid="B67" ref-type="bibr">Humble et al. 2010</xref>; <xref rid="B120" ref-type="bibr">Smith and Cognato 2010a</xref>; <xref rid="B53" ref-type="bibr">Furniss and Kegley 2011</xref>; <xref rid="B86" ref-type="bibr">Lee et al. 2011</xref>). Plant nomenclature was verified using the Missouri Botanical Garden’s Tropicos database (<ext-link ext-link-type="uri" xlink:href="http://www.tropicos.org">www.tropicos.org</ext-link>).</p><p>Specimens were examined using either a Leica (Wetzlar, Germany) MZ125 or MZ16 compound microscope and illuminated with a SCHOTT (Mainz, Germany) 150W halogen light source (model ACE®1). Images were taken with a Visionary Digital Passport II system (Palmyra, VA) using a Canon EOS 5D Mark II, 58.0 mm Canon Macro photo lens, Canon Speedlite transmitter ST-E2, two Canon Speedlite 4303X II flashes and a Stack Shot (Cognisys, Inc, Kingsley, MI). Montage images were assembled using Helicon Focus Mac Pro 4.2.8 (Helicon Soft, Kharkov, Ukraine). Measurements were made using an ocular micrometer on the same microscope and light source as listed above and calibrated with ROK (Shenzhen, China) 150.0 mm digital calipers (model DC-122A) following the protocol of <xref rid="B121" ref-type="bibr">Smith and Cognato (2010b)</xref>. Measurements were taken from the specimen’s dorsal surface. Length was measured from the pronotum apex to the elytral apex. Width was measured at the widest part of the pronotum. Proportions are given as the ratio of length to width. A maximum of 50 specimens selected to encompass the known distribution were measured for each species. If important locality data such as state or county was missing from specimen labels, the information was inserted between square brackets in the material examined. Holotypes of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sulcatus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">californicus</named-content></named-content></italic> LeConte, 1868 were not physically examined. Type images were examined from the MCZC type database (<ext-link ext-link-type="uri" xlink:href="http://insects.oeb.harvard.edu/mcz/">http://insects.oeb.harvard.edu/mcz/</ext-link>) and the synonymy of these species was confirmed. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">californicus</named-content></named-content></italic> was not included in this revision because it is a synonym of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">scolytus</named-content></named-content></italic>, a species not established in the New World.</p></sec><sec sec-type="Taxonomic characters"><title>Taxonomic characters</title><p>External anatomical terminology followed <xref rid="B66" ref-type="bibr">Hopkins (1909)</xref> subsequently used by <xref rid="B113" ref-type="bibr">Schedl (1931)</xref>, <xref rid="B75" ref-type="bibr">Kaston (1936)</xref>, <xref rid="B42" ref-type="bibr">Edson (1967)</xref> and <xref rid="B148" ref-type="bibr">Wood (1982</xref>, <xref rid="B149" ref-type="bibr">1986</xref>, <xref rid="B150" ref-type="bibr">2007</xref>). Sculp<pmc-comment>PageBreak</pmc-comment>ture terminology followed <xref rid="B135" ref-type="bibr">Torre-Bueno (1989)</xref>. Several highly informative recently described <named-content content-type="taxon-name"><named-content content-type="tribe">Scolytini</named-content></named-content> characters were also scored (<xref rid="B121" ref-type="bibr">Smith and Cognato 2010b</xref>). Provisional morphological homology was assessed by similarity and relative positions of characters.</p><p>Characters were scored from both sexes unless otherwise noted. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> exhibits strong sexual dimorphism of the frons and abdominal venter; males display a wide array of morphological features, particularly on the venter, whereas females are more morphologically conserved. Consequently, characters were predominately male based. Male genitalia was found to be autapomorphic or extremely conserved in structure and thus characters were not scored. Characters and character state numbers correspond to data coded in the morphological data matrix for each taxon. The character matrix (Table <xref ref-type="table" rid="T3">3</xref>) was constructed and edited using the online database MX (<xref rid="B153" ref-type="bibr">Yoder et al. 2006</xref>). Character transformations were evaluated using MacClade 4.0 PPC (<xref rid="B91" ref-type="bibr">Maddison and Maddison 2000</xref>) and homology of characters and definitions of characters states were re-examined and modified if necessary.</p><table-wrap id="T3" orientation="portrait" position="float"><label>Table 3.</label><caption><p>Morphological character matrix of 43 characters for 37 species. Characters are described in Methods.</p></caption><table frame="hsides" rules="all"><tbody><tr><th rowspan="1" colspan="1"></th><th rowspan="1" colspan="43">Characters</th></tr><tr><th rowspan="1" colspan="1">Species</th><th rowspan="1" colspan="1">1</th><th rowspan="1" colspan="1">2</th><th rowspan="1" colspan="1">3</th><th rowspan="1" colspan="1">4</th><th rowspan="1" colspan="1">5</th><th rowspan="1" colspan="1">6</th><th rowspan="1" colspan="1">7</th><th rowspan="1" colspan="1">8</th><th rowspan="1" colspan="1">9</th><th rowspan="1" colspan="1">10</th><th rowspan="1" colspan="1">11</th><th rowspan="1" colspan="1">12</th><th rowspan="1" colspan="1">13</th><th rowspan="1" colspan="1">14</th><th rowspan="1" colspan="1">15</th><th rowspan="1" colspan="1">16</th><th rowspan="1" colspan="1">17</th><th rowspan="1" colspan="1">18</th><th rowspan="1" colspan="1">19</th><th rowspan="1" colspan="1">20</th><th rowspan="1" colspan="1">21</th><th rowspan="1" colspan="1">22</th><th rowspan="1" colspan="1">23</th><th rowspan="1" colspan="1">24</th><th rowspan="1" colspan="1">25</th><th rowspan="1" colspan="1">26</th><th rowspan="1" colspan="1">27</th><th rowspan="1" colspan="1">28</th><th rowspan="1" colspan="1">29</th><th rowspan="1" colspan="1">30</th><th rowspan="1" colspan="1">31</th><th rowspan="1" colspan="1">32</th><th rowspan="1" colspan="1">33</th><th rowspan="1" colspan="1">34</th><th rowspan="1" colspan="1">35</th><th rowspan="1" colspan="1">36</th><th rowspan="1" colspan="1">37</th><th rowspan="1" colspan="1">38</th><th rowspan="1" colspan="1">39</th><th rowspan="1" colspan="1">40</th><th rowspan="1" colspan="1">41</th><th rowspan="1" colspan="1">42</th><th rowspan="1" colspan="1">43</th></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic></td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic></td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">8</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">4</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">7</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">6</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic></td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">9</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">A</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic></td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic></td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">7</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">6</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic></td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">1</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic></td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">5</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">virgatus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">1</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">intricatus</named-content></named-content></italic></td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">8</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laevis</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">6</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">propinquus</named-content></named-content></italic></td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">pygmaeus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">6</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ratzeburgii</named-content></named-content></italic></td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">B</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">scolytus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">6</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sinopiceus</named-content></named-content></italic></td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sulcifrons</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">6</td></tr></tbody></table></table-wrap><p>Ecological characters including gallery pattern and host were scored. Character states were assigned based on a comprehensive literature review (e.g. <xref rid="B23" ref-type="bibr">Butovitsch 1929</xref>; <xref rid="B24" ref-type="bibr">Chamberlin 1939</xref>; <xref rid="B114" ref-type="bibr">Schedl 1948</xref>; <xref rid="B6" ref-type="bibr">Balachowsky 1949</xref>; <xref rid="B125" ref-type="bibr">Stark 1952</xref>; <xref rid="B25" ref-type="bibr">Chamberlin 1958</xref>; <xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>; <xref rid="B97" ref-type="bibr">Michalski 1973</xref>; <xref rid="B17" ref-type="bibr">Bright 1976</xref>; <xref rid="B55" ref-type="bibr">Furniss and Carolin 1977</xref>; <xref rid="B148" ref-type="bibr">Wood 1982</xref>; <xref rid="B3" ref-type="bibr">Atkinson and Equihua-Martínez 1986</xref>; <xref rid="B151" ref-type="bibr">Wood and Bright 1992</xref>; <xref rid="B105" ref-type="bibr">Pfeffer 1994</xref>; <xref rid="B32" ref-type="bibr">Cibrián Tovar et al. 1995</xref>; <xref rid="B18" ref-type="bibr">Bright and Skidmore 1997</xref>, <xref rid="B19" ref-type="bibr">2002</xref>; <xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>), and with field notes, and label data on pinned museum specimens.</p><p>Forty-three characters were used in this study (19 binary and 24 multistate). Ten morphological characters were coded from the head, two from the thorax, ten from the elytra, one from the metepimeron, and 17 from the venter. Three ecological characters were also coded. Consistency and retention index values from the morphological phylogeny (Fig. <xref ref-type="fig" rid="F2">2</xref>) and generated from MacClade are listed next to each character. Characters and states were scored as follows:</p><list list-type="order"><list-item><p><italic>Epistomal emargination</italic> (ci = 0.22; ri = 0.22): (0) weak; (1) moderate; (2) strong.</p></list-item><list-item><p><italic>Male epistomal process</italic> (ci = 0.25; ri = 0.63): (0) absent; (1) present.</p></list-item><list-item><p><italic>Male frons shape</italic> (ci = 0.25; ri = 0.45): (0) convex; (1) flat; (2) impressed.</p></list-item><list-item><p><italic>Male frons vestiture distribution</italic> (ci = 0.33; ri = 0.20): (0) glabrous; (1) uniform; (2) predominately on margins; (3) epistomal region.</p></list-item><list-item><p><italic>Male frons vestiture length</italic> (ci = 0.4; ri = 0.0): (0) less than width of eye; (1) equal to width of eye; (2) greater than width of eye.</p></list-item><list-item><p><italic>Male frons sculpturing</italic> (ci = 0.38; ri = 0.50): (0) coarsely aciculate; (1) weakly aciculate; (2) rugose-reticulate; (3) smooth.</p></list-item><list-item><p><italic>Male frons punctures</italic> (ci = 0.33; ri = 0.71): (0) small, fine; (1) small, coarse; (2) impunctate.</p></list-item><list-item><p><italic>Female frons shape</italic> (ci = 0.67; ri = 0.50): (0) flat; (1) impressed; (2) convex; (3) excavated.</p></list-item><list-item><p><italic>Female frons sculpturing</italic> (ci = 0.29; ri = 0.44): (0) coarsely aciculate; (1) finely aciculate; (2) reticulate.</p></list-item><list-item><p><italic>Female frons punctures</italic> (ci = 0.14; ri = 0.54): (0) small, fine; (1) small, coarse.</p></list-item><pmc-comment>PageBreak</pmc-comment>
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<list-item><p><italic>Pronotum length to width</italic> (ci = 0.50; ri = 0.50): (0) as long as wide (0.95–1.05); (1) wider than long.</p></list-item><list-item><p><italic>Metepimeron length</italic> (ci = 0.29; ri = 0.50): (0) less than half-length of metanepisternum; (1) half length of metanepisternum; (2) greater than half-length of metanepisternum.</p></list-item><list-item><p><italic>Elytral sides sub-parallel</italic> (<xref rid="B97" ref-type="bibr">Michalski 1973</xref>) (ci = 0.25; ri = 0.50); (0) on basal half only; (1) on apical half only.</p></list-item><list-item><p><italic>Interstrial setae</italic> (ci = 0.22; ri = 0.56); (0) glabrous; (1) sparse; (2) moderate.</p></list-item><list-item><p><italic>Interstrial impression</italic> (ci = 0.13; ri = 0.42): (0) not impressed; (1) faintly impressed.</p></list-item><list-item><p><italic>Interstrial width</italic> (ci = 0.29; ri = 0.44): (0) equal to striae; (1) twice width of striae; (2) more than twice the width of striae.</p></list-item><list-item><p><italic>Relative size of interstrial punctures</italic> (ci = 0.25; ri = 0.25): (0) equal to strial; (1) smaller than strial.</p></list-item><list-item><p><italic>Strial impression</italic> (ci = 0.40; ri = 0.70): (0) not impressed; (1) weakly impressed; (2) moderately impressed.</p></list-item><list-item><p><italic>Male elytral apex shape</italic> (ci = 0.25; ri = 0.67): (0) rounded; (1) subquadrate.</p></list-item><list-item><p><italic>Elytral apex emargination</italic> (ci = 0.38; ri = 0.76): (0) absent; (1) weak; (2) moderate; (3) strong.</p></list-item><list-item><p><italic>Apical margin of elytra serrate</italic> (ci = 0.20; ri = 0.33): (0) absent; (1) present.</p></list-item><list-item><p><italic>Punctures on elytral apical margin</italic> (ci = 0.20; ri = 0.53): (0) impunctate; (1) small, fine; (2) large, coarse.</p></list-item><list-item><p><italic>Venter appearance</italic> (<xref rid="B10" ref-type="bibr">Blackman 1934</xref>) (ci = 0.11; ri = 0.50): (0) smooth, shining; (1) shagreened, dull.</p></list-item><list-item><p><italic>Venter setae length</italic> (ci = 0.23; ri = 0.50): (0) less than 1 diameter of a puncture; (1) less than length of segment 3; (2) greater than length of segment 3; (3) glabrous.</p></list-item><list-item><p><italic>Female second ventrite</italic> (ci = 0.60; ri = 0.50): (0) unarmed; (1) apical; (2) basal; (3) medial.</p></list-item><list-item><p><italic>Suture between first and second ventrites</italic> (<xref rid="B148" ref-type="bibr">Wood 1982</xref>) (ci = 0.50; ri = 0.0): (0) clearly visible; (1) obsolete.</p></list-item><list-item><p><italic>Male first ventrite apical margin</italic> (ci = 0.42; ri = 0.68): (0) rounded, on vertical surface; (1) weakly elevated; (2) lip; (3) weakly produced; (4) strongly produced; (5) flush, not on vertical surface.</p></list-item><list-item><p><italic>Male second ventrite armature</italic> (ci = 0.36; ri = 0.42): (0) unarmed; (1) basal spine; (2) medial spine; (3) apical spine; (4) carina on basal half; (5) carina on apical half.</p></list-item><list-item><p><italic>Male second ventrite surface</italic> (ci = 0.33; ri = 0.67): (0) convex; (1) weakly concave; (2) strongly concave; (3) flat.</p></list-item><list-item><p><italic>Second ventrite punctures</italic> (ci = 0.25; ri = 0.63): (0) small, fine; (1) small, coarse.</p></list-item><list-item><p><italic>Male second ventrite setal tuft</italic> (ci = 1.00; ri = 0.0): (0) absent; (1) present.</p></list-item><list-item><p><italic>Second ventrite lateral spines</italic> (ci = 0.50; ri = 0.0): (0) absent; (1) present.</p></list-item><list-item><p><italic>Third ventrite lateral spines</italic> (ci = 0.50; ri = 0.0): (0) absent; (1) present.</p></list-item><list-item><p><italic>Fourth ventrite lateral spines</italic> (ci = 0.50; ri = 0.0): (0) absent; (1) present.</p></list-item><pmc-comment>PageBreak</pmc-comment>
<list-item><p><italic>Male fourth ventrite armed medially</italic> (ci = 0.17; ri = 0.29): (0) absent; (1) present.</p></list-item><list-item><p><italic>Male fifth ventrite</italic> (ci = 0.60; ri = 0.75): (0) unarmed – lacking carina; (1) midpoint of carina closer to apex; (2) midpoint of carina closer to base; (3) midpoint of carina equidistant from base and apex.</p></list-item><list-item><p><italic>Relative length of male fifth ventrite compared to third and fourth</italic> (ci = 0.15; ri = 0.48): (0) 5 larger; (1) 3+4 longer; (2) equal.</p></list-item><list-item><p><italic>Male fifth ventrite setal patch</italic> (ci = 0.33; ri = 0.50): (0) absent; (1) present.</p></list-item><list-item><p><italic>Male fifth ventrite with median depression</italic> (ci = 0.33; ri = 0.50): (0) absent; (1) present.</p></list-item><list-item><p><italic>Male metatibial setae</italic> (ci = 0.50; ri = 0.0): (0) not conspicuously longer than those of other tibiae; (1) much longer and more abundant than those of other tibiae.</p></list-item><list-item><p><italic>Mating system</italic> (ci = 1.00; ri = 0.0): (0) monogamy; (1) polygamy.</p></list-item><list-item><p><italic>Gallery type</italic> (ci = 0.50; ri = 0.70): (0) vertical; (1) transverse; (2) epsilon; (3) multi-branched.</p></list-item><list-item><p><italic>Host</italic> (ci = 0.73; ri = 0.73): (0) <named-content content-type="taxon-name"><named-content content-type="family">Fabaceae</named-content></named-content>; (1) <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content></named-content></italic>; (2) <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic>; (3) <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content></named-content></italic>; (4) <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content></named-content></italic>; (5) <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content></named-content></italic>; (6) <named-content content-type="taxon-name"><named-content content-type="family">Ulmaceae</named-content></named-content>; (7) <named-content content-type="taxon-name"><named-content content-type="family">Rosaceae</named-content></named-content>; (8) <named-content content-type="taxon-name"><named-content content-type="family">Fagaceae</named-content></named-content>; (9) <named-content content-type="taxon-name"><named-content content-type="family">Cannabaceae</named-content></named-content>; (A) <named-content content-type="taxon-name"><named-content content-type="family">Juglandaceae</named-content></named-content>; (B) <named-content content-type="taxon-name"><named-content content-type="family">Betulaceae</named-content></named-content>.</p></list-item></list><fig id="F2" orientation="portrait" position="float"><label>Figure 2.</label><caption><p>Morphological phylogeny of Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic>. Phylogram of one of 1016 most parsimonious trees (L = 262, CI = 0.332, RI = 0.539) generated for 37 taxa with 43 characters from a heuristic search of 1000 stepwise random additions with TBR in PAUP*. Numbers above the node indicate bootstrap values (>50). Bremer support values are listed below the node. The Nearctic hardwood and conifer clades are indicated in blue and green respectively. Non-monophyletic species are indicated in purple. * denotes species exotic to the Nearctic.</p></caption><graphic id="oo_33355.jpg" xlink:href="zookeys-450-001-g002"></graphic></fig></sec><sec sec-type="Taxa, DNA sequences and alignment"><title>Taxa, DNA sequences and alignment</title><p>We included 83 specimens (Table <xref ref-type="table" rid="T4">4</xref>) representing 32 <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species to reconstruct phylogenies using DNA sequences from mitochondrial and nuclear genes and morphology. The same Palearctic outgroup species used in the morphological analyses were included in the molecular dataset. As many Nearctic species as possible were included in the analysis. We were unable to collect fresh material for <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic>, which were therefore excluded from the molecular analyses. Sequences of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">intricatus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">scolytus</named-content></named-content></italic> were obtained from GenBank and included in the analyses. To assess mitochondrial cytochrome oxidase I (COI) intraspecific variation, we included 89 specimens (Table <xref ref-type="table" rid="T4">4</xref>) representing 30 <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species from as many populations as possible. Specimens were included in the four-gene phylogeny only if sequences were available for at least two genes.</p><table-wrap id="T4" orientation="portrait" position="float"><label>Table 4.</label><caption><p>Table of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> specimens sequenced, the collection locality, collector, host and GenBank accession numbers.</p></caption><table frame="hsides" rules="all"><tbody><tr><th rowspan="1" colspan="1">Species</th><th rowspan="1" colspan="1">Collection locality, date, collector and host</th><th rowspan="1" colspan="1">Matrix name</th><th rowspan="1" colspan="1">Extraction name</th><th rowspan="1" colspan="1">COI</th><th rowspan="1" colspan="1">28S</th><th rowspan="1" colspan="1">CAD</th><th rowspan="1" colspan="1">ArgK</th></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: California: Siskiyou Co.: Klamath National Forest, FR 41N16, <named-content content-type="dwc:verbatimCoordinates">N41°14.822', W122°53.562'</named-content>, 508ft, 28.VII.2010, S.M. Smith coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco abi 14</td><td rowspan="1" colspan="1">SMS 14</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909668">KJ909668</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923438">KJ923438</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923574">KJ923574</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923513">KJ923513</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Latah Co.: St. Joe National Forest, Moscow Mountain, <named-content content-type="dwc:verbatimCoordinates">N46°48.252', W116°49.811'</named-content>, 3834ft, S.M. Smith, A.R. Gillogly, M.M. Furniss coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco abi 15</td><td rowspan="1" colspan="1">SMS 15</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909669">KJ909669</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Bonner Co.: Kaniksu National Forest, FR 232, <named-content content-type="dwc:verbatimCoordinates">N48°20.609', W116°20.507'</named-content>, 2717ft, S.M. Smith, A.R. Gillogly coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco abi 16</td><td rowspan="1" colspan="1">SMS 16</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909670">KJ909670</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923439">KJ923439</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923575">KJ923575</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923514">KJ923514</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Oregon: Deschutes Co.: Deschutes National Forest, Black Butte Rd, Black Butte, <named-content content-type="dwc:verbatimCoordinates">N40°24.924', W121°38.323'</named-content>, 4212ft, 1.VIII.2010, S.M. Smith coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco abi 17</td><td rowspan="1" colspan="1">SMS 17</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909671">KJ909671</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic></td><td rowspan="1" colspan="1">Mexico: Nuevo Leon: San Antonio de las Alazanas: Santa Catarina, G. Cuellar coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco aze 161</td><td rowspan="1" colspan="1">SMS 161</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923440">KJ923440</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923576">KJ923576</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923515">KJ923515</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Pennsylvania: Cumberland Co.: Roadway Dr @ Schenider Dr., 29.V.2009, L.R. Donovall coll., Ex. Lindgren funnel trap</td><td rowspan="1" colspan="1">Sco fagi 1</td><td rowspan="1" colspan="1">Sco fagi 1</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909672">KJ909672</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923441">KJ923441</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Pennsylvania: Cumberland Co.: Roadway Dr @ Schenider Dr., 29.V.2009, L.R. Donovall coll., Ex. Lindgren funnel trap</td><td rowspan="1" colspan="1">Sco fagi 2</td><td rowspan="1" colspan="1">Sco fagi 2</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909673">KJ909673</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923442">KJ923442</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923577">KJ923577</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923516">KJ923516</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Colorado: Mesa Co.: Grand Mesa overlook, 8.VIII.2008, D.E. Bright, B.A. Barr coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> branches</td><td rowspan="1" colspan="1">Sco uni 1</td><td rowspan="1" colspan="1">Sco uni 1</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909736">KJ909736</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923496">KJ923496</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923625">KJ923625</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923561">KJ923561</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Colorado: Eastern Slope Rocky Mountains, 6.V.2010, D.E. Bright, B.A Barr, S.M. Smith coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco fis 36</td><td rowspan="1" colspan="1">SMS 36</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909737">KJ909737</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Colorado: Larimer Co.: Roosevelt National Forest, 9 km E Estes Park, Hwy 34, Big Thompson Canyon, <named-content content-type="dwc:verbatimCoordinates">N40°24.456', W105°24.565'</named-content>, 2143m, 5.V.2010, S.M. Smith, D.E. Bright, B.A. Barr coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco fis 37</td><td rowspan="1" colspan="1">SMS 37</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909738">KJ909738</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Boise Co.: Boise National Forest, Bogus Basin, NF275, <named-content content-type="dwc:verbatimCoordinates">N43°44.347', W116°07.099'</named-content>, 6042ft, 8.VIII.2010, S.M. Smith, A.R. Gillogly coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco fisi 38</td><td rowspan="1" colspan="1">SMS 38</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909739">KJ909739</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923498">KJ923498</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923627">KJ923627</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923563">KJ923563</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Latah Co.: Univeristy of Idaho experimental forest, <named-content content-type="dwc:verbatimCoordinates">N46°51.764', W116°43.786'</named-content>, 2855ft, 10.VIII.2010, S.M. Smith, A.R. Gillogly, M.M. Furniss coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco fis 41</td><td rowspan="1" colspan="1">SMS 41</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909742">KJ909742</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923501">KJ923501</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923630">KJ923630</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923566">KJ923566</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">intricatus</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sweden: Smaaland: Oskarshamn [GenBank]</td><td rowspan="1" colspan="1">Sco int BJ</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HQ883677">HQ883677</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HQ883589">HQ883589</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HQ883820">HQ883820</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HQ883909">HQ883909</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">intricatus</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sweden: Smaaland: Karlsburg, 15.VII.2005, B.H. Jordal coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Quercus</named-content></named-content></italic> sp.</td><td rowspan="1" colspan="1">Sco int 64</td><td rowspan="1" colspan="1">SMS 64</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923444">KJ923444</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923579">KJ923579</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923518">KJ923518</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laevis</named-content></named-content></italic></td><td rowspan="1" colspan="1">Denmark: Rødby Havn., 24.VIII.2005, J. Pedersen coll.</td><td rowspan="1" colspan="1">Sco laevis</td><td rowspan="1" colspan="1">Sco laevis</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923445">KJ923445</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923580">KJ923580</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923519">KJ923519</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Boundary Co.: Kaniksu National Forest, Robinson Lake campground, <named-content content-type="dwc:verbatimCoordinates">N48°58.385', W116°13.351'</named-content>, 2748ft, 11.VIII.2010, S.M. Smith, A.R. Gillogly coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco lar 2</td><td rowspan="1" colspan="1">SMS 2</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909676">KJ909676</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Boundary Co.: Kaniksu National Forest, Robinson Lake campground, <named-content content-type="dwc:verbatimCoordinates">N48°58.197', W116°13.068'</named-content>, 2748ft, 11.VIII.2010, S.M. Smith, A.R. Gillogly coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco lar 12</td><td rowspan="1" colspan="1">SMS 12</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909677">KJ909677</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923446">KJ923446</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923581">KJ923581</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923520">KJ923520</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Boundary Co.: Kaniksu National Forest, <named-content content-type="dwc:verbatimCoordinates">N48°39.185', W116°32.662'</named-content>, 5311ft, 12.VIII.2010, S.M. Smith, A.R. Gillogly coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco lar 13</td><td rowspan="1" colspan="1">SMS 13</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909678">KJ909678</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Pennsylvania: Lebanon Co.: Mt. Gretna, <named-content content-type="dwc:verbatimCoordinates">N40.242501 W76.462406</named-content>, IV-VIII.2009, S.E. Spichiger coll., Ex. Lindgren funnel trap</td><td rowspan="1" colspan="1">Sco mal 1</td><td rowspan="1" colspan="1">Sco mal 1</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923447">KJ923447</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923582">KJ923582</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923521">KJ923521</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic></td><td rowspan="1" colspan="1">Czech Republic: South Bohemia: Jindřichův Hradec env. Číměř; 23.VII.2011, M.Knížek lgt., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Prunus</named-content></named-content></italic> sp.</td><td rowspan="1" colspan="1">Sco mal 78</td><td rowspan="1" colspan="1">SMS 78</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909679">KJ909679</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923448">KJ923448</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923583">KJ923583</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923522">KJ923522</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Michigan: Kalamazoo Co.: Gourdneck Lake State Game Area, 6.VIII.2011, S.M. Smith, A.I. Cognato coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Prunus</named-content></named-content></italic> sp.</td><td rowspan="1" colspan="1">Sco mal 81</td><td rowspan="1" colspan="1">SMS 81</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909680">KJ909680</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923449">KJ923449</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923584">KJ923584</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923523">KJ923523</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Boundary Co.: Kaniksu National Forest, Robinson Lake campground, <named-content content-type="dwc:verbatimCoordinates">N48°58.385', W116°13.351'</named-content>, 2748ft, 11.VIII.2010, S.M. Smith, A.R. Gillogly coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco mon 33</td><td rowspan="1" colspan="1">SMS 33</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909681">KJ909681</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Soshone Co.: Coeur D’Alene National Forest, Placer Creek Rd <named-content content-type="dwc:verbatimCoordinates">N47°25.746', W115°53.601'</named-content>, 3547ft, 15.VIII.2010, S.M. Smith, A.R. Gillogly coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco mon 34</td><td rowspan="1" colspan="1">SMS 34</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909682">KJ909682</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923450">KJ923450</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Boise Co.: Boise Natioanl Forest, Bogus Basin, NF275, <named-content content-type="dwc:verbatimCoordinates">N43°44.347', W116°07.099'</named-content>, 6042ft, 8.VIII.2010, S.M. Smith, A.R. Gillogly coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco mon 35</td><td rowspan="1" colspan="1">SMS 35</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909683">KJ909683</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923451">KJ923451</ext-link></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923524">KJ923524</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic></td><td rowspan="1" colspan="1">Denmark: Storstrøm: Rødbyhavn, 24.VIII.2005, J. Pedersen coll.</td><td rowspan="1" colspan="1">Sco mul 65</td><td rowspan="1" colspan="1">SMS 65</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909685">KJ909685</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923453">KJ923453</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923586">KJ923586</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923525">KJ923525</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Michigan: Kalamazoo Co.: Gourdneck Lake State Game Area, 19.VI.2011, A.I. Cognato coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> sp.</td><td rowspan="1" colspan="1">Sco mul 82</td><td rowspan="1" colspan="1">SMS 82</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909686">KJ909686</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923454">KJ923454</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923587">KJ923587</ext-link></td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic></td><td rowspan="1" colspan="1">Russia: Moscow Oblast: Dolgoprudnly, <named-content content-type="dwc:verbatimCoordinates">N55°58.266', E37°30.191’</named-content> 570m, 28.VII.2011, A.V. Petrov, Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content><named-content content-type="species">laevis</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco mul 147</td><td rowspan="1" colspan="1">SMS 147</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909687">KJ909687</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923455">KJ923455</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923588">KJ923588</ext-link></td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Pennsylvania: Cumberland Co.: Roadway Dr @ Schenider Dr., 29.V.2009, L.R. Donovall coll., Ex. Lindgren funnel trap</td><td rowspan="1" colspan="1">Sco mut 1</td><td rowspan="1" colspan="1">Sco mut 1</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909688">KJ909688</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923456">KJ923456</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923589">KJ923589</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923526">KJ923526</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Michigan: Wayne Co.: Grosse Ile, <named-content content-type="dwc:verbatimCoordinates">N42.17060 W83.14496</named-content>, 7-20.VI.2007, R. Mech coll., Ex. Lindgren funnel trap</td><td rowspan="1" colspan="1">Sco mut 2</td><td rowspan="1" colspan="1">Sco mut 2</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909689">KJ909689</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923457">KJ923457</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Pennsylvania: Dauphin Co.: Harrisburg, <named-content content-type="dwc:verbatimCoordinates">N40.316325 W76.888783</named-content>, IV-VIII.2009, S.E. Spichiger coll., Ex. Lindgren Funnel trap</td><td rowspan="1" colspan="1">Sco mut 4</td><td rowspan="1" colspan="1">Sco mut 4</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909690">KJ909690</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: South Carolina: Columbia, 26.X.2011, W. Jones coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content></named-content></italic> sp.</td><td rowspan="1" colspan="1">Sco mut 169</td><td rowspan="1" colspan="1">SMS 169</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909691">KJ909691</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923458">KJ923458</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923590">KJ923590</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923527">KJ923527</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Arizona: Cochise Co.: Coronado National Forest, Chiricahua Mountains, <named-content content-type="dwc:verbatimCoordinates">N31°55.987', W109°16.331'</named-content>, 7022ft, 22.V.2010, S.M. Smith coll. Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco obe 18</td><td rowspan="1" colspan="1">SMS 18</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909692">KJ909692</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923459">KJ923459</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923591">KJ923591</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923528">KJ923528</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Arizona: Graham Co.: Coronado National Forest, Pinaleno Mountains, <named-content content-type="dwc:verbatimCoordinates">N32°37.702', W109°49.472'</named-content>, 7896ft, 23.V.2010, S.M. Smith coll. Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco obe 19</td><td rowspan="1" colspan="1">SMS 19</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909693">KJ909693</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Arizona: Coconino Co.: Arizona Snowbowl, <named-content content-type="dwc:verbatimCoordinates">N35°19.593', W111°42.681'</named-content>, 9032ft, 27.V.2010 S.M. Smith coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco obe 20</td><td rowspan="1" colspan="1">SMS 20</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909694">KJ909694</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: New Mexico: Torrance Co.: Cibola National Forest, W. Manzano, <named-content content-type="dwc:verbatimCoordinates">N34°37.226', W106°24.752'</named-content>, 7999ft, 11.V.2010, S.M. Smith, A.I. Cognato coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco obe 44</td><td rowspan="1" colspan="1">SMS 44</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909695">KJ909695</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923460">KJ923460</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923592">KJ923592</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923529">KJ923529</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Latah Co.: Univeristy of Idaho experimental forest, <named-content content-type="dwc:verbatimCoordinates">N46°51.372', W116°44.038'</named-content>, 2857ft, 10.VIII.2010, S.M. Smith, A.R. Gillogly, M.M. Furniss, Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco opa 21</td><td rowspan="1" colspan="1">SMS 21</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909696">KJ909696</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923461">KJ923461</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923593">KJ923593</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923530">KJ923530</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Boundary Co.: Kaniksu National Forest, Robinson Lake campground, <named-content content-type="dwc:verbatimCoordinates">N48°58.200', W116°13.067'</named-content>, 2696ft 13.VIII.2010, S.M. Smith, A.R. Gillogly, Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco opa 68</td><td rowspan="1" colspan="1">SMS 68</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909697">KJ909697</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923462">KJ923462</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Oregon: Jackson Co.: Rogue River National Forest, Rogue River gorge viewpoint, Hwy 62 1/4 mi N Union Creek Rd, <named-content content-type="dwc:verbatimCoordinates">N42°54.540', W122°26.733'</named-content>, 3489ft, 21.VIII.2010, S.M. Smith coll. Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco ore 5</td><td rowspan="1" colspan="1">SMS 5</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923463">KJ923463</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923594">KJ923594</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923531">KJ923531</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Oregon: Jackson Co.: Rogue River National Forest, Rogue River gorge viewpoint, Hwy 62 1/4 mi N Union Creek Rd, <named-content content-type="dwc:verbatimCoordinates">N42°54.540', W122°26.733'</named-content>, 3489ft, 21.VIII.2010, S.M. Smith coll. Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco ore 71</td><td rowspan="1" colspan="1">SMS 71</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909698">KJ909698</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923464">KJ923464</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923595">KJ923595</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923532">KJ923532</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Michigan: Livingston Co.: Howell, 431 Bishop Rd, <named-content content-type="dwc:verbatimCoordinates">N42.5076 W83.85698</named-content>, 25.VI.2009, R. Mech coll., Ex. Lindgren funnel trap</td><td rowspan="1" colspan="1">Sco pic 1</td><td rowspan="1" colspan="1">Sco mul 1</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909700">KJ909700</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Montana: Jefferson Co.: highway 15 N. of Butte, <named-content content-type="dwc:verbatimCoordinates">N46.2075 W112.3360</named-content>, 6013ft, 24.VII.2004, K.P. Dole coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content></named-content></italic> sp.</td><td rowspan="1" colspan="1">Sco pic</td><td rowspan="1" colspan="1">Sco pic 1</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909699">KJ909699</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923465">KJ923465</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923596">KJ923596</ext-link></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Latah Co.: University of Idaho Experimental Forest, <named-content content-type="dwc:verbatimCoordinates">N46°51.164', W116°44.838'</named-content>, 2939ft, S.M. Smith, A.R. Gillogly, M.M. Furniss coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">engelmanni</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco pic 7</td><td rowspan="1" colspan="1">SMS 7</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909701">KJ909701</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923466">KJ923466</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923597">KJ923597</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923533">KJ923533</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Valley Co.: Boise National Forest, Hwy 55, S. of Donnelly, <named-content content-type="dwc:verbatimCoordinates">N44°20.117', W116°02.698'</named-content>, 4816ft, 17.VIII.2010, S.M. Smith, A.R. Gillogly, Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">engelmannii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco pic 8</td><td rowspan="1" colspan="1">SMS 8</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909702">KJ909702</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: South Dakota: Lawrence Co.: Brownsville Rd near Leads, <named-content content-type="dwc:verbatimCoordinates">N44.2922 W103.7828</named-content>, 27.VII.2004, 5650ft, K.P. Dole coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco pic 73</td><td rowspan="1" colspan="1">SMS 73</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909703">KJ909703</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923467">KJ923467</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923598">KJ923598</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923534">KJ923534</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Wyoming: Carbon Co.: Medicine Bow National Forest, Snowy Mountains, WY130, Lake Marie, <named-content content-type="dwc:verbatimCoordinates">N41°19.965', W106°19.516’</named-content> 3208m, 26.VII.2011, S.M. Smith, D.E. Bright, B.A Barr coll., Emerged 1-5.IX.2011, Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">engelmannii</named-content></named-content></italic> branches</td><td rowspan="1" colspan="1">Sco pic 83</td><td rowspan="1" colspan="1">SMS 83</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909704">KJ909704</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923468">KJ923468</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923599">KJ923599</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923535">KJ923535</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: California: El Dorado Co.: El Dorado National Forest, Ise house resevoir, <named-content content-type="dwc:verbatimCoordinates">N38.50 W 120.22</named-content>, 1653m, 17.VI.2003, A.I. Cognato coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> sp.</td><td rowspan="1" colspan="1">Sco pra</td><td rowspan="1" colspan="1">Sco pra</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909709">KJ909709</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: California: Alpine Co.: Toiyabe National Forest, 11.6 miles E of Markleeville, <named-content content-type="dwc:verbatimCoordinates">N38°39.906', W119°38.540'</named-content>, 24.VII.2010, S.M. Smith coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">magnifica</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco pra 28</td><td rowspan="1" colspan="1">SMS 28</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909705">KJ909705</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: California: El Dorado Co.: El Dorado National Forest, nr. Ice House Resevoir <named-content content-type="dwc:verbatimCoordinates">N38°50.002', W120°21.160'</named-content>, 5413ft, 25.VII.2010, S.M. Smith coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco pra 29</td><td rowspan="1" colspan="1">SMS 29</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909706">KJ909706</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923469">KJ923469</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923600">KJ923600</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923536">KJ923536</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: California: Siskiyou Co.: Shasta Trinity National Forest, Mt. Shasta, <named-content content-type="dwc:verbatimCoordinates">N41°20.844', W122°16.691'</named-content>, 4892ft, S.M. Smith coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco pra 30</td><td rowspan="1" colspan="1">SMS 30</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909707">KJ909707</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923470">KJ923470</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923601">KJ923601</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923537">KJ923537</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: California: Siskiyou Co.: Klamath National Forest, FR 41N16, <named-content content-type="dwc:verbatimCoordinates">N41°14.822', W122°53.562'</named-content>, 5081ft, 28.VII.2010, S.M. Smith coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco pra 31</td><td rowspan="1" colspan="1">SMS 31</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909708">KJ909708</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Wyoming: Albany Co.: Snowy Mountains, Medicine Bow National Forest, Spruce campground, 6.IX.2010, D.E. Bright, B.A. Barr coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco pra 91</td><td rowspan="1" colspan="1">SMS 91</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909710">KJ909710</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923471">KJ923471</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923602">KJ923602</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923538">KJ923538</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">propinquus</named-content></named-content></italic></td><td rowspan="1" colspan="1">Mexico: Oaxaca: Huatulco, <named-content content-type="dwc:verbatimCoordinates">15.76234, -96.12885</named-content>, 41m, 23.VI.2009, T.H. Atkinson coll., THA 874, Ex. legume tree</td><td rowspan="1" colspan="1">Sco pro 1</td><td rowspan="1" colspan="1">Sco pro 1</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909711">KJ909711</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923472">KJ923472</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923603">KJ923603</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923539">KJ923539</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">propinquus</named-content></named-content></italic></td><td rowspan="1" colspan="1">Mexico: Oaxaca: Huatulco, <named-content content-type="dwc:verbatimCoordinates">15.76234, -96.12885</named-content>, 41m, 23.VI.2009, T.H. Atkinson coll., THA 874, Ex. legume tree</td><td rowspan="1" colspan="1">Sco pro 2</td><td rowspan="1" colspan="1">Sco pro 2</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909712">KJ909712</ext-link></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923604">KJ923604</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923540">KJ923540</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">pygmaeus</named-content></named-content></italic></td><td rowspan="1" colspan="1">Czech Republic: Moravia: Břeclav Kamci obora, 2004, K. Novakova coll.</td><td rowspan="1" colspan="1">Sco pyg 62</td><td rowspan="1" colspan="1">SMS 62</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909713">KJ909713</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923473">KJ923473</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923605">KJ923605</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923541">KJ923541</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">pygmaeus</named-content></named-content></italic></td><td rowspan="1" colspan="1">Denmark: Storstrøm: Rødbyhavn, 24.VIII.2005, J. Pedersen coll.</td><td rowspan="1" colspan="1">Sco pyg 67</td><td rowspan="1" colspan="1">SMS 67</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909714">KJ909714</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923474">KJ923474</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923606">KJ923606</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923542">KJ923542</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Pennsylvania: Cumberland Co.: Roadway Dr @ Schenider Dr., 24.VII.2009, L.R. Donovall coll., Ex. Lindgren funnel trap</td><td rowspan="1" colspan="1">Sco qua 1</td><td rowspan="1" colspan="1">Sco qua 1</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909715">KJ909715</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923475">KJ923475</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923607">KJ923607</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923543">KJ923543</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Pennsylvania: Cumberland Co.: Roadway Dr @ Schenider Dr., 24.VII.2009, L.R. Donovall coll., Ex. Lindgren funnel trap</td><td rowspan="1" colspan="1">Sco qua 2</td><td rowspan="1" colspan="1">Sco qua 2</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909716">KJ909716</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Pennsylvania: Lebanon Co.: Mt. Gretna, <named-content content-type="dwc:verbatimCoordinates">N40.242501 W76.462406</named-content>, IV-VIII.2009, S.E. Spichiger coll., Ex. Lindgren funnel trap</td><td rowspan="1" colspan="1">Sco qua 3</td><td rowspan="1" colspan="1">Sco qua 3</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909717">KJ909717</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Maryland: Ann Aruridel Co.: Annapolis, 26.V.2012, R.J. Rabaglia coll., Ex. Lindgren funnel trap</td><td rowspan="1" colspan="1">Sco qua 170</td><td rowspan="1" colspan="1">SMS 170</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909718">KJ909718</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923476">KJ923476</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923608">KJ923608</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923544">KJ923544</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ratzeburgii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Russia: Primorsky: Anisimovka, 12.VII.2008, B.H. Jordal coll.</td><td rowspan="1" colspan="1">Sco ratz 60</td><td rowspan="1" colspan="1">SMS 60</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909719">KJ909719</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923477">KJ923477</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923609">KJ923609</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923545">KJ923545</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Arizona: Pima Co.: Coronado National Forest, Santa Catalina Mountains, <named-content content-type="dwc:verbatimCoordinates">N32°24.529', W110°42.678'</named-content>, 7869ft, 22.V.2010, S.M. Smith coll. Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco ref 26</td><td rowspan="1" colspan="1">SMS 26</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909720">KJ909720</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923478">KJ923478</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923610">KJ923610</ext-link></td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Arizona: Cochise Co.: Coronado National Forest, Chiricahua Mountains, <named-content content-type="dwc:verbatimCoordinates">N31°54.915', W109°16.040'</named-content>, 8196ft, 20.V.2010, S.M. Smith coll. Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco ref 27</td><td rowspan="1" colspan="1">SMS 27</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909721">KJ909721</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923479">KJ923479</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923611">KJ923611</ext-link></td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Colorado: Larimer Co.: Roosevelt National Forest, 9 km E Estes Park, Hwy 34, Big Thompson Canyon, <named-content content-type="dwc:verbatimCoordinates">N40°24.456', W105°24.565'</named-content>, 2143m, 5.V.2010, S.M. Smith, D.E. Bright, B.A. Barr coll. Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco ref 32</td><td rowspan="1" colspan="1">SMS 32</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909722">KJ909722</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923480">KJ923480</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923612">KJ923612</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923546">KJ923546</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Colorado: Boulder Co.: Roosevelt National Forest, St. Vrain canyon, <named-content content-type="dwc:verbatimCoordinates">N40°10.072', W105°23.623'</named-content>, 2127m, 24.VII.2011, S.M. Smith, D.E. Bright, B.A. Barr coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco ref 85</td><td rowspan="1" colspan="1">SMS 85</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909684">KJ909684</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923452">KJ923452</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923585">KJ923585</ext-link></td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Arizona: Graham Co.: Coronado National Forest, Pinaleno Mountains, <named-content content-type="dwc:verbatimCoordinates">N32°37.702', W109°49.472'</named-content>, 7896ft, 23.V.2010, S.M. Smith coll. Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco rob 3</td><td rowspan="1" colspan="1">SMS 3</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909723">KJ909723</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923481">KJ923481</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923613">KJ923613</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923547">KJ923547</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: New Mexico: Taos Co.: Carson National Forest, Agua Piedra campground, Hwy 75, <named-content content-type="dwc:verbatimCoordinates">N36°07.960', W105°31.828’</named-content> 8477 ft, 13.V.2010, S.M. Smith, A.I. Cognato coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco rob 43</td><td rowspan="1" colspan="1">SMS 43</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909724">KJ909724</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923482">KJ923482</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923614">KJ923614</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923548">KJ923548</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Arizona: Cochise Co.: Coronado National Forest, Chiricahua Mountains, <named-content content-type="dwc:verbatimCoordinates">N31°54.665', W109°16.336'</named-content>, 2445m, 5.VIII.2012, S.M. Smith, A.I. Cognato coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco rob 178</td><td rowspan="1" colspan="1">SMS 178</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909725">KJ909725</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923483">KJ923483</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923615">KJ923615</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923549">KJ923549</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">Hungary: Győr-Moson-Sopron: Sopron, 2003, F. Lakatos coll.</td><td rowspan="1" colspan="1">Sco rug 66</td><td rowspan="1" colspan="1">SMS 66</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923484">KJ923484</ext-link></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923550">KJ923550</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">Morocco: 30 km S. Asni, Tizi’n’Test, 18.IV.2002, B.H. Jordal coll,. Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Prunus</named-content><named-content content-type="species">dulcis</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco rug 72</td><td rowspan="1" colspan="1">SMS 72</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923485">KJ923485</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923616">KJ923616</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923551">KJ923551</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">Czech Republic: South Bohemia, Jindřichův Hradec env. Číměř; 23.VII.2011, M.Knížek coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Prunus</named-content></named-content></italic> sp.</td><td rowspan="1" colspan="1">Sco rug 79</td><td rowspan="1" colspan="1">SMS 79</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909726">KJ909726</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923486">KJ923486</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923617">KJ923617</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923552">KJ923552</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">Iran: Guilan: Khoshkestalkh, <named-content content-type="dwc:verbatimCoordinates">N37°27'01” E49°42'06"</named-content>, 17.VII.2011, S. Amini coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Malus</named-content></named-content></italic> sp.</td><td rowspan="1" colspan="1">Sco rug 165</td><td rowspan="1" colspan="1">SMS 165</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923487">KJ923487</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923618">KJ923618</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923553">KJ923553</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Missouri: Saint Louis Co.: Maryland Heights, 22.V.2008, Ex. Lindgren funnel trap</td><td rowspan="1" colspan="1">Sco sch 2</td><td rowspan="1" colspan="1">Sco sch 2</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923488">KJ923488</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923619">KJ923619</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923554">KJ923554</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">scolytus</named-content></named-content></italic></td><td rowspan="1" colspan="1">Denmark: NEJ, Tofte Skov [GenBank]</td><td rowspan="1" colspan="1">Sco sco BJ</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HQ883678">HQ883678</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HQ883590">HQ883590</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HQ883821">HQ883821</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HQ883910">HQ883910</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sinopiceus</named-content></named-content></italic></td><td rowspan="1" colspan="1">China: Qinghai: Mai Xiu Forest Preserve, <named-content content-type="dwc:verbatimCoordinates">N35°16.288', E101°55.904'</named-content>, 2927m, 21.V.2008, A.I. Cognato coll., Ex. Lindgren funnel trap</td><td rowspan="1" colspan="1">Sco sin 1</td><td rowspan="1" colspan="1">Sco sin 1</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909728">KJ909728</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923489">KJ923489</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923620">KJ923620</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923555">KJ923555</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sinopiceus</named-content></named-content></italic></td><td rowspan="1" colspan="1">China: Sichuan: Highway 213 near Zhangla, 9.VII.2004, A.I. Cognato coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">purpurea</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco sin 70</td><td rowspan="1" colspan="1">SMS 70</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909729">KJ909729</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923490">KJ923490</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923556">KJ923556</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: California: Alpine Co.: Toiyabe National Forest, 11.6 miles E of Markleeville, <named-content content-type="dwc:verbatimCoordinates">N38°39.906', W119°38.540'</named-content>, 24.VII.2010, S.M. Smith coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">magnifica</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco sub 6</td><td rowspan="1" colspan="1">SMS 6</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909730">KJ909730</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923491">KJ923491</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923621">KJ923621</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923557">KJ923557</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Boundary Co.: Kaniksu National Forest, Robinson Lake campground, <named-content content-type="dwc:verbatimCoordinates">N48°58.197', W116°13.068'</named-content>, 2748ft 13.VIII.2010, S.M. Smith, A.R. Gillogly, Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco sub 51</td><td rowspan="1" colspan="1">SMS 51</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909731">KJ909731</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923492">KJ923492</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923558">KJ923558</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sulcifrons</named-content></named-content></italic></td><td rowspan="1" colspan="1">Russia: Moscow Oblast: Dolgoprudnly, <named-content content-type="dwc:verbatimCoordinates">N55°58.266', E37°30.191'</named-content>, 570m, 28.VII.2011, A.V. Petrov coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content><named-content content-type="species">laevis</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco sul 146</td><td rowspan="1" colspan="1">SMS 146</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909732">KJ909732</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923493">KJ923493</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923622">KJ923622</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923559">KJ923559</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Oregon: Hood River Co.: Mount Hood National Forest, Sherwood Campground, Hwy 35, <named-content content-type="dwc:verbatimCoordinates">N45°19.278', W121°37.104'</named-content>, 4293ft, 2.VIII.2010, S.M. Smith coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco tsu 9</td><td rowspan="1" colspan="1">SMS 9</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909733">KJ909733</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Soshone Co.: St. Joe National Forest, <named-content content-type="dwc:verbatimCoordinates">N47°00.790', W116°12.359'</named-content>, 3192 ft, 15.VIII.2010, S.M. Smith, A.R. Gillogly coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco tsu 10</td><td rowspan="1" colspan="1">SMS 10</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909734">KJ909734</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923494">KJ923494</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923623">KJ923623</ext-link></td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Boundary Co.: Kaniksu National Forest, <named-content content-type="dwc:verbatimCoordinates">N48°40.911', W116°34.345'</named-content>, 4353ft, 12.VIII.2010, S.M. Smith, A.R. Gillogly coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco tsu 11</td><td rowspan="1" colspan="1">SMS 11</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909735">KJ909735</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923495">KJ923495</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923624">KJ923624</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923560">KJ923560</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Oregon: Deschutes Co.: Deschutes National Forest, Black Butte Rd, Black Butte, <named-content content-type="dwc:verbatimCoordinates">N40°24.924', W121°38.323'</named-content>, 4212ft, 1.VIII.2010 S.M. Smith coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco fis 1</td><td rowspan="1" colspan="1">SMS 1</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909674">KJ909674</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923443">KJ923443</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923578">KJ923578</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923517">KJ923517</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Oregon: Hood River Co.: Mount Hood National Forest, Sherwood Campground, Hwy 35, <named-content content-type="dwc:verbatimCoordinates">N45°19.278', W121°37.104'</named-content>, 4293ft, 2.VIII.2010, S.M. Smith coll. Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco uni 39</td><td rowspan="1" colspan="1">SMS 39</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909740">KJ909740</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923499">KJ923499</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923628">KJ923628</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923564">KJ923564</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Oregon: Jackson Co.: Rogue River National Forest, NF 60, <named-content content-type="dwc:verbatimCoordinates">N42°53.926', W122°19.177'</named-content>, 4547ft, 30.VII.2010, S.M. Smith coll. Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco uni 40</td><td rowspan="1" colspan="1">SMS 40</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909741">KJ909741</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923500">KJ923500</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923629">KJ923629</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923565">KJ923565</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Oregon: Klamath Co.: Deschutes National Forest, NF 4672, <named-content content-type="dwc:verbatimCoordinates">N43°30.474', W121°52.147'</named-content>, 4923ft, 31.VII.2010, S.M. Smith coll. Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco uni 42</td><td rowspan="1" colspan="1">SMS 42</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909743">KJ909743</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923502">KJ923502</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923631">KJ923631</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923567">KJ923567</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Oregon: Marion Co.: Williamette National Forest,Breitenbush Rd, <named-content content-type="dwc:verbatimCoordinates">N44°47.200', W121°56.557'</named-content>, 2645ft, 1.VIII.2010, S.M. Smith coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco uni 151</td><td rowspan="1" colspan="1">SMS 151</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909744">KJ909744</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923503">KJ923503</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923632">KJ923632</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923568">KJ923568</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Arizona: Cochise Co.: Coronado National Forest, Chiricahua Mountains, 14.VII.2009, J. Hulcr coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco ven 1</td><td rowspan="1" colspan="1">Sco sp 1</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909745">KJ909745</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Boundary Co.: Kaniksu National Forest, Robinson Lake campground, <named-content content-type="dwc:verbatimCoordinates">N48°58.385', W116°13.351'</named-content>, 2748ft, 11.VIII.2010, S.M. Smith, A.R. Gillogly coll. Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco ven 22</td><td rowspan="1" colspan="1">SMS 22</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909746">KJ909746</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923504">KJ923504</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923633">KJ923633</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923569">KJ923569</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Adams Co.: Payette National Forest <named-content content-type="dwc:verbatimCoordinates">N45°00.225', W116°09.651'</named-content>, 6025ft, 17.VIII.2010, S.M. Smith, A.R. Gillogly coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco ven 45</td><td rowspan="1" colspan="1">SMS 45</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909747">KJ909747</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Benewah Co.: McCroskey State Park, <named-content content-type="dwc:verbatimCoordinates">N47°04.801', W116°54.960'</named-content>, 3744ft,11.VIII.2010, S.M. Smith, A.R. Gillogly coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco ven 46</td><td rowspan="1" colspan="1">SMS 46</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909748">KJ909748</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923505">KJ923505</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Idaho: Boundary Co.: Kaniksu National Forest, Robinson Lake campground, <named-content content-type="dwc:verbatimCoordinates">N48°58.200', W116°13.067'</named-content>, 2696ft 13.VIII.2010, S.M. Smith, A.R. Gillogly, Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco ven 47</td><td rowspan="1" colspan="1">SMS 47</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909749">KJ909749</ext-link></td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: New Mexico: Torrance Co.: Cibola National Forest, W. Manzano, <named-content content-type="dwc:verbatimCoordinates">N34°37.325', W106°24.642'</named-content>, 8026ft, 11.V.2010, S.M. Smith, A.I. Cognato coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco ven 48</td><td rowspan="1" colspan="1">SMS 48</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909750">KJ909750</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923506">KJ923506</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923634">KJ923634</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923570">KJ923570</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: California: El Dorado Co.: El Dorado National Forest, nr. Ice House Resevoir <named-content content-type="dwc:verbatimCoordinates">N38°50.002', W120°21.160'</named-content>, 5413ft, 25.VII.2010, S.M. Smith coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco ven 49</td><td rowspan="1" colspan="1">SMS 49</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909751">KJ909751</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923507">KJ923507</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923635">KJ923635</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923571">KJ923571</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Arizona: Coconino Co.: Arizona Snowbowl, <named-content content-type="dwc:verbatimCoordinates">N36°24.381', W112°05.619'</named-content>, 8923ft, 30.V.2010 S.M. Smith, K. Bush coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco ven 50</td><td rowspan="1" colspan="1">SMS 50</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909752">KJ909752</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923508">KJ923508</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923636">KJ923636</ext-link></td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">virgatus</named-content></named-content></italic></td><td rowspan="1" colspan="1">Mexico: Nuevo Leon: San Antonio de las Alazanas: Santa Catarina, G. Cuellar coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco vir 162</td><td rowspan="1" colspan="1">SMS 162</td><td rowspan="1" colspan="1">-</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923509">KJ923509</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923637">KJ923637</ext-link></td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: Arizona: Cochise Co.: Coronado National Forest, Chiricahua Mountains, <named-content content-type="dwc:verbatimCoordinates">N31°55.360', W109°15.702'</named-content>, 7882ft, 20.V.2010, S.M. Smith coll. Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco wick 24</td><td rowspan="1" colspan="1">SMS 24</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909753">KJ909753</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923510">KJ923510</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923638">KJ923638</ext-link></td><td rowspan="1" colspan="1">-</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: New Mexico: Otero Co.: Lincoln National Forest, Apache Point observatory, <named-content content-type="dwc:verbatimCoordinates">N32°47.046', W105°48.841'</named-content>, 9116ft, 16.V.2010, S.M. Smith coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco wick 25</td><td rowspan="1" colspan="1">SMS 25</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909754">KJ909754</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923511">KJ923511</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923639">KJ923639</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923572">KJ923572</ext-link></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic></td><td rowspan="1" colspan="1">USA: New Mexico: Colfax Co.: Highway 64 near Ute Park, <named-content content-type="dwc:verbatimCoordinates">N36°33'9.26” W105°07'6.74"</named-content>, 2242m, 24.VI.2004, A.I. Cognato, S.A. Stephens coll., Ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content><named-content content-type="species">ponderosa</named-content></named-content></italic></td><td rowspan="1" colspan="1">Sco wick 74</td><td rowspan="1" colspan="1">SMS 74</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ909755">KJ909755</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923512">KJ923512</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923640">KJ923640</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KJ923573">KJ923573</ext-link></td></tr></tbody></table></table-wrap><p>DNA was extracted from freshly collected specimens preserved in 200 proof ethanol and from pinned specimens killed in sawdust impregnated with ethyl acetate and immediately pinned. Specimens were dissected by removing the head and thorax from the abdomen. DNA extractions were performed on the head and thorax using a Qiagen DNEasy blood and tissue kit (Hilden, Germany) following manufacturer protocols except for the DNA elution procedure, which consisted of a single elution of 100–200 µl of buffer AE depending on the size of the specimen, with specimens measuring less than 5.0 mm in length receiving 100 µl and those greater than 5.0 mm in length receiving 200 µl. After the extraction process was completed, the body parts were rinsed in 70% ethanol, glued onto a mounting board, pinned, labeled and were vouchered in the MSUC. The resulting puri<pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment>fied DNA was used to amplify partial gene regions of COI, D2 region of nuclear ribosomal 28S, CAD, and Argenine Kinase (ArgK) using the PCR primers listed in Table <xref ref-type="table" rid="T5">5</xref>. The COI barcoding primers LCO 1490 and HCO 2198 did not consistently amplify scolytine COI sequences, which necessitated the construction of the degenerate scolytine specific primers, 1495b and rev750. These primers worked best for <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> specimens and have proven to be effective on a broad array of scolytine taxa (Cognato et al. unpublished). The 1495b and rev750 primers were used to create the <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> specific F215 and Rev453 primers, respectively for 5–20 years or older pinned specimens. All PCR cocktails consisted of a total volume of 25 µl and included 14.25–17.25 µl ddH<sub>2</sub>O, 2.5 µl 10X PCR buffer (Qiagen), 1.0 µl 25 mM MgCl<sub>2</sub> (Qiagen), 0.5µl dNTP mix (Qiagen), 2–5 µl DNA temfig, 0.25 µl HotStar Taq DNA polymerase (Qiagen). PCR reactions were performed on a thermal cycler (PTC-2000, MJ Research, Waltham, MA, USA or MyCylcer Thermocycler, BioRad, Hercules, CA, USA) under the following conditions: one cycle for 15 min at 95 °C, 40 cycles for 30 (COI)–45 (28S, CAD, ArgK) sec at 94 °C, 45 sec at 50–58 °C (see Table <xref ref-type="table" rid="T5">5</xref> for specific annealing temperatures), 1 min at 72 °C and a final elongation cycle of 5 min at 72 °C. PCR products were cleaned using ExoSAP-IT (USB Corp., Cleveland, OH, USA) and following the manufacturer protocols. Cleaned PCR products were then prepared for sequencing. Each reaction contained 3.5 µl of cleaned PCR product, 1 µl 33 pM/µl sequencing primers (identical to those used in PCR), and 7.5 µl of ddH<sub>2</sub>O. Samples were sequenced in the Michigan State University Research Technology Support Facility using a BigDye Terminator v 1.1 (Applied Biosystems, Foster City, CA, USA) and visualized using an ABI 3730 Genetic Analyzer (Applied Biosystems).</p><table-wrap id="T5" orientation="portrait" position="float"><label>Table 5.</label><caption><p>Table of PCR primers and the annealing temperatures used for the amplification of gene sequences.</p></caption><table frame="hsides" rules="all"><tbody><tr><th rowspan="1" colspan="1">Gene</th><th rowspan="1" colspan="1">Primer</th><th rowspan="1" colspan="1">Primer sequence</th><th rowspan="1" colspan="1">Annealing Temp °C</th><th rowspan="1" colspan="1">BP analyzed</th><th rowspan="1" colspan="1">First Cited</th></tr><tr><td rowspan="1" colspan="1">COI</td><td rowspan="1" colspan="1">LCO 1490</td><td rowspan="1" colspan="1">5’-GGTCAACAAATCATAAAGATATTGG-3’</td><td rowspan="1" colspan="1">50</td><td rowspan="1" colspan="1">615</td><td rowspan="1" colspan="1"><xref rid="B64" ref-type="bibr">Hebert et al. 2003</xref></td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">HCO 2198</td><td rowspan="1" colspan="1">5’-TAAACTTCAGGGTGACCAAAAAATCA-3’</td><td rowspan="1" colspan="1">50</td><td rowspan="1" colspan="1">615</td><td rowspan="1" colspan="1"><xref rid="B64" ref-type="bibr">Hebert et al. 2003</xref></td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">1495b</td><td rowspan="1" colspan="1">5’-AACAAATCATAAAGATATTGGRAC-3’</td><td rowspan="1" colspan="1">50</td><td rowspan="1" colspan="1">615</td><td rowspan="1" colspan="1">This study</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">rev 750</td><td rowspan="1" colspan="1">5’-GAAATTATNCCAATTCCTGG-3’</td><td rowspan="1" colspan="1">50</td><td rowspan="1" colspan="1">615</td><td rowspan="1" colspan="1">This study</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">ScoCOI F215</td><td rowspan="1" colspan="1">5’-CCCCCGACATAGCTTTCCC-3’</td><td rowspan="1" colspan="1">50</td><td rowspan="1" colspan="1">271</td><td rowspan="1" colspan="1">This study</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">ScoCOI Rev453</td><td rowspan="1" colspan="1">5’-TATTTGATCGAACTTTATTCC-3’</td><td rowspan="1" colspan="1">50</td><td rowspan="1" colspan="1">271</td><td rowspan="1" colspan="1">This study</td></tr><tr><td rowspan="1" colspan="1">28S</td><td rowspan="1" colspan="1">3665</td><td rowspan="1" colspan="1">5’-AGACAGAGTTCAAGAGTACGTG-3’</td><td rowspan="1" colspan="1">55</td><td rowspan="1" colspan="1">402–496</td><td rowspan="1" colspan="1"><xref rid="B8" ref-type="bibr">Belshaw and Quicke 1997</xref></td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">4068</td><td rowspan="1" colspan="1">5’-TTGGTCCGTGTTTCAAGACGGG-3’</td><td rowspan="1" colspan="1">55</td><td rowspan="1" colspan="1">402–496</td><td rowspan="1" colspan="1"><xref rid="B8" ref-type="bibr">Belshaw and Quicke 1997</xref></td></tr><tr><td rowspan="1" colspan="1">CAD</td><td rowspan="1" colspan="1">apCADforB2</td><td rowspan="1" colspan="1">5’-TGGAARGARGTBGARTACGARGTGGYCG-3’</td><td rowspan="1" colspan="1">56 or 58</td><td rowspan="1" colspan="1">690</td><td rowspan="1" colspan="1"><xref rid="B41" ref-type="bibr">Dole et al. 2010</xref></td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">apCADfor4</td><td rowspan="1" colspan="1">5’-TGGAARGARGTBGARTACGARGTGGTYCG-3’</td><td rowspan="1" colspan="1">56 or 58</td><td rowspan="1" colspan="1">472</td><td rowspan="1" colspan="1"><xref rid="B37" ref-type="bibr">Danforth et al. 2006</xref></td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">apCADrev1mod</td><td rowspan="1" colspan="1">5’-GCCATYRCTCBCCTACRCTYTTCAT-3’</td><td rowspan="1" colspan="1">56 or 58</td><td rowspan="1" colspan="1">472 or 690</td><td rowspan="1" colspan="1"><xref rid="B37" ref-type="bibr">Danforth et al. 2006</xref></td></tr><tr><td rowspan="1" colspan="1">ArgK</td><td rowspan="1" colspan="1">ArgKforB2</td><td rowspan="1" colspan="1">5’-GAYTCCGGWATYGGWATCTAYGCTCC-3’</td><td rowspan="1" colspan="1">56 or 58</td><td rowspan="1" colspan="1">692</td><td rowspan="1" colspan="1"><xref rid="B41" ref-type="bibr">Dole et al. 2010</xref></td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">ArgKrevB2</td><td rowspan="1" colspan="1">5’-GTATGYTCMCCRCGRGTACCACG-3’</td><td rowspan="1" colspan="1">56 or 58</td><td rowspan="1" colspan="1">692</td><td rowspan="1" colspan="1"><xref rid="B41" ref-type="bibr">Dole et al. 2010</xref></td></tr></tbody></table></table-wrap><p>Sequences were received as chromatograms and the sense and antisense strands were compiled using Sequencher (Ann Arbor, MI) to trim sequences, examine for ambiguities and create consensus sequences. Sequences were blasted in GenBank to examine for paralogous copies and other potential problems including contamination and pseudogenes. Sequences for COI, CAD and ArgK were aligned using SE-AL v.2.0a11 Carbon (<ext-link ext-link-type="uri" xlink:href="http://tree.bio.ed.ac.uk/software/seal/">http://tree.bio.ed.ac.uk/software/seal/</ext-link>). Sequence length variation was only observed in 28S. Sequences of 28S were manually aligned in SE-AL using a scolytine-specific secondary structure model (<xref rid="B73" ref-type="bibr">Jordal et al. 2008</xref>). Nexus files are available at <ext-link ext-link-type="uri" xlink:href="http://www.scolytid.msu.edu">http://www.scolytid.msu.edu</ext-link>.</p></sec></sec><sec sec-type="methods"><title>Methods</title><sec sec-type="Species concept"><title>Species concept</title><p>Species are hypotheses of unique evolutionary entities, tested by monophyly (<pmc-comment>PageBreak</pmc-comment><italic>sensu</italic><xref rid="B138" ref-type="bibr">Wheeler and Platnick 2000</xref>). These monophyletic groups of individuals were named if they were diagnosable by synapomorphies or a unique combination of homoplastic characters. If diagnostic characters were not found, the clade retained the original species name or was synonymized with the sister clade. Therefore, given our species level morphological and molecular phylogenies, species revisions were based on monophyly of multiple individuals from disjunct populations.</p></sec><sec sec-type="Morphology"><title>Morphology</title><p><italic>Parsimony phylogenetic analysis.</italic> A phylogeny was reconstructed using the criterion of parsimony implemented in PAUP*4.0 b10 PPC (<xref rid="B133" ref-type="bibr">Swofford 2002</xref>). A heuristic search with 1,000 stepwise random additions with tree bisection-reconnection (TBR) for 37 taxa (25 ingroup, 12 outgroup) was performed. Characters were unordered and equally weighted. Bootstrap values were calculated by performing 1,000 pseudoreplicates with simple additions in PAUP*. Bremer support values were calculated by creating a constraint tree in TreeRot v.2 (<xref rid="B123" ref-type="bibr">Sorenson 1999</xref>) and analyzed in PAUP* using a heuristic search with 100 addition-sequence replicates.</p></sec><sec sec-type="Molecular"><title>Molecular</title><p><italic>Likelihood phylogenetic analyses.</italic> We analyzed the molecular and morphological datasets using Bayesian estimation of phylogeny with MrBayes 3.2.2 (<xref rid="B110" ref-type="bibr">Ronquist et al. 2012</xref>) on the CIPRES Science Gateway (<xref rid="B116" ref-type="bibr">Schwartz 2010</xref>). The Bayesian analysis consisted of a combined molecular and morphological dataset (82 taxa). The dataset was divided into 11 partitions by gene and codon position for COI, CAD and ArgK, and one partition each for 28s and morphology. We selected the best model for each data partition using MrModeltest (<xref rid="B100" ref-type="bibr">Nylander 2004</xref>). The GTR+I+Γ (general time reversible with a proportion of invariant sites and a gamma-shaped distribution of rate variation across sites) model selected by AIC was found to have the optimal fit for the gene partitions and Γ (gamma) was chosen for the morphology partition. Taxa that were unable to be sequenced were included in the morphology partition of the combined dataset.</p><p>Four Metropolis-Coupled Markov Chain Monte Carlo searches (3 heated, 1 cold) were performed for 10 million generations with sampling as described for the molecular dataset. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic>, and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic> were included in this analysis with only morphological characters. All parameters reached stability at 500,000 generations and the standard deviation of split frequencies between runs was 0.006983. Bayesian posterior probabilities of clades were calculated by a majority-rule consensus of those trees after the burn-in (75,000 trees).</p><pmc-comment>PageBreak</pmc-comment>
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<p><italic>Parsimony phylogenetic analysis.</italic> Intraspecific (Table <xref ref-type="table" rid="T6">6</xref>) and interspecific (Table <xref ref-type="table" rid="T7">7</xref>) differences for 83 taxa were generated by computing pairwise distances for each gene in PAUP*. A single gene phylogeny was reconstructed using COI using a heuristic search with 100 stepwise random additions with TBR for 89 taxa (79 ingroup, 10 outgroup). Bootstrap values were calculated by performing 1,000 pseudoreplicates with simple additions in PAUP*. Bremer support values were calculated by creating a constraint tree in TreeRot v.2 and analyzed in PAUP* using a heuristic search with 100 addition-sequence replicates.</p><table-wrap id="T6" orientation="portrait" position="float"><label>Table 6.</label><caption><p>Intraspecific differences among genes expressed as the proportion of sites differing between sequences.</p></caption><table frame="hsides" rules="all"><tbody><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="8"><bold>Gene</bold></td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="2"><bold>COI</bold></td><td rowspan="1" colspan="2"><bold>28S</bold></td><td rowspan="1" colspan="2"><bold>CAD</bold></td><td rowspan="1" colspan="2"><bold>ArgK</bold></td></tr><tr><td rowspan="1" colspan="1"><bold>Species</bold></td><td rowspan="1" colspan="1"><bold>Range</bold></td><td rowspan="1" colspan="1"><bold>Average</bold></td><td rowspan="1" colspan="1"><bold>Range</bold></td><td rowspan="1" colspan="1"><bold>Average</bold></td><td rowspan="1" colspan="1"><bold>Range</bold></td><td rowspan="1" colspan="1"><bold>Average</bold></td><td rowspan="1" colspan="1"><bold>Range</bold></td><td rowspan="1" colspan="1"><bold>Average</bold></td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic></td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic></td><td rowspan="1" colspan="1">0–0.0231</td><td rowspan="1" colspan="1">0.0183</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0–0.0058</td><td rowspan="1" colspan="1">0.0034</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">intricatus</named-content></named-content></italic></td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic></td><td rowspan="1" colspan="1">0–0.0043</td><td rowspan="1" colspan="1">0.0020</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0.0043</td><td rowspan="1" colspan="1">0.0043</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0433</td><td rowspan="1" colspan="1">0.0433</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0–0.0030</td><td rowspan="1" colspan="1">0.0020</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic></td><td rowspan="1" colspan="1">0–0.0017</td><td rowspan="1" colspan="1">0.0011</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0–0.0016</td><td rowspan="1" colspan="1">0.0033</td><td rowspan="1" colspan="1">0–0.0027</td><td rowspan="1" colspan="1">0.0009</td><td rowspan="1" colspan="1">0–0.0127</td><td rowspan="1" colspan="1">0.0078</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0034–0.0105</td><td rowspan="1" colspan="1">0.0079</td><td rowspan="1" colspan="1">0–0.0074</td><td rowspan="1" colspan="1">0.0049</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0081–0.0195</td><td rowspan="1" colspan="1">0.0135</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0.0021</td><td rowspan="1" colspan="1">0.0021</td><td rowspan="1" colspan="1">0.0014</td><td rowspan="1" colspan="1">0.0014</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic></td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0.0085</td><td rowspan="1" colspan="1">0.0085</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic></td><td rowspan="1" colspan="1">0–0.0163</td><td rowspan="1" colspan="1">0.0100</td><td rowspan="1" colspan="1">0–0.0025</td><td rowspan="1" colspan="1">0.0008</td><td rowspan="1" colspan="1">0–0.00850</td><td rowspan="1" colspan="1">0.0050</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic></td><td rowspan="1" colspan="1">0–0.0148</td><td rowspan="1" colspan="1">0.0077</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0–0.0044</td><td rowspan="1" colspan="1">0.0024</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">propinquus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0.0063</td><td rowspan="1" colspan="1">0.0063</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">pygmaeus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0.0000</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0016–0.0049</td><td rowspan="1" colspan="1">0.0032</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0.0000</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0.0014</td><td rowspan="1" colspan="1">0.0014</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ratzeburgii</named-content></named-content></italic></td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0–0.0130</td><td rowspan="1" colspan="1">0.0070</td><td rowspan="1" colspan="1">0–0.0049</td><td rowspan="1" colspan="1">0.0012</td><td rowspan="1" colspan="1">0–0.0023</td><td rowspan="1" colspan="1">0.0003</td><td rowspan="1" colspan="1">0–0.0043</td><td rowspan="1" colspan="1">0.0029</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0033–0.0537</td><td rowspan="1" colspan="1">0.0358</td><td rowspan="1" colspan="1">0–0.0025</td><td rowspan="1" colspan="1">0.0017</td><td rowspan="1" colspan="1">0–0.0043</td><td rowspan="1" colspan="1">0.0029</td><td rowspan="1" colspan="1">0.0015–0.00612</td><td rowspan="1" colspan="1">0.0038</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">0–0.0074</td><td rowspan="1" colspan="1">0.0050</td><td rowspan="1" colspan="1">0.0065–0.0192</td><td rowspan="1" colspan="1">0.0128</td><td rowspan="1" colspan="1">0–0.0274</td><td rowspan="1" colspan="1">0.0150</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic></td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">scolytus</named-content></named-content></italic></td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sinopiceus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0157</td><td rowspan="1" colspan="1">0.0157</td><td rowspan="1" colspan="1">0.0123</td><td rowspan="1" colspan="1">0.0123</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0158</td><td rowspan="1" colspan="1">0.0158</td><td rowspan="1" colspan="1">0.0049</td><td rowspan="1" colspan="1">0.0049</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">0.0101</td><td rowspan="1" colspan="1">0.0101</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sulcifrons</named-content></named-content></italic></td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0016–0.0147</td><td rowspan="1" colspan="1">0.0103</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0016–0.0282</td><td rowspan="1" colspan="1">0.0121</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0–0.0023</td><td rowspan="1" colspan="1">0.0015</td><td rowspan="1" colspan="1">0–0.0029</td><td rowspan="1" colspan="1">0.0007</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic></td><td rowspan="1" colspan="1">0–0.0181</td><td rowspan="1" colspan="1">0.0089</td><td rowspan="1" colspan="1">0–0.0025</td><td rowspan="1" colspan="1">0.0010</td><td rowspan="1" colspan="1">0–0.0043</td><td rowspan="1" colspan="1">0.0022</td><td rowspan="1" colspan="1">0–0.0043</td><td rowspan="1" colspan="1">0.0022</td></tr></tbody></table></table-wrap><table-wrap id="T7" orientation="portrait" position="float"><label>Table 7.</label><caption><p>Interspecific differences among genes expressed as the proportion of sites differing among sister taxa.</p></caption><table frame="hsides" rules="all"><tbody><tr><th rowspan="1" colspan="1"></th><th rowspan="1" colspan="8">Gene</th></tr><tr><th rowspan="1" colspan="1"></th><th rowspan="1" colspan="2">COI</th><th rowspan="1" colspan="2">28S</th><th rowspan="1" colspan="2">CAD</th><th rowspan="1" colspan="2">ArgK</th></tr><tr><th rowspan="1" colspan="1">Species</th><th rowspan="1" colspan="1">Range</th><th rowspan="1" colspan="1">Average</th><th rowspan="1" colspan="1">Range</th><th rowspan="1" colspan="1">Average</th><th rowspan="1" colspan="1">Range</th><th rowspan="1" colspan="1">Average</th><th rowspan="1" colspan="1">Range</th><th rowspan="1" colspan="1">Average</th></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic></td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">0–0.0025</td><td rowspan="1" colspan="1">0.0020</td><td rowspan="1" colspan="1">0.0047–0.0088</td><td rowspan="1" colspan="1">0.0067</td><td rowspan="1" colspan="1">0.0119–0.0133</td><td rowspan="1" colspan="1">0.0128</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.1317–0.1382</td><td rowspan="1" colspan="1">0.1361</td><td rowspan="1" colspan="1">0.0176–0.0251</td><td rowspan="1" colspan="1">0.0201</td><td rowspan="1" colspan="1">0.0171–0.208</td><td rowspan="1" colspan="1">0.0190</td><td rowspan="1" colspan="1">0.0267</td><td rowspan="1" colspan="1">0.0267</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.1789–0.1821</td><td rowspan="1" colspan="1">0.1801</td><td rowspan="1" colspan="1">0.0528</td><td rowspan="1" colspan="1">0.0528</td><td rowspan="1" colspan="1">0.0531–0.0533</td><td rowspan="1" colspan="1">0.0532</td><td rowspan="1" colspan="1">0.0533–0.0549</td><td rowspan="1" colspan="1">0.0541</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic></td><td rowspan="1" colspan="1">0–0.0297</td><td rowspan="1" colspan="1">0.0143</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0–0.0128</td><td rowspan="1" colspan="1">0.0051</td><td rowspan="1" colspan="1">0.0029–0.0072</td><td rowspan="1" colspan="1">0.0051</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic></td><td rowspan="1" colspan="1">0–0.0.028</td><td rowspan="1" colspan="1">0.0109</td><td rowspan="1" colspan="1">0.0025–0.0049</td><td rowspan="1" colspan="1">0.0031</td><td rowspan="1" colspan="1">0.0065–0.0234</td><td rowspan="1" colspan="1">0.0096</td><td rowspan="1" colspan="1">0.0130–0.0159</td><td rowspan="1" colspan="1">0.0145</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0331–0.0520</td><td rowspan="1" colspan="1">0.043</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0–0.0149</td><td rowspan="1" colspan="1">0.0074</td><td rowspan="1" colspan="1">0–0.0072</td><td rowspan="1" colspan="1">0.0043</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic></td><td rowspan="1" colspan="1">0–0.0179</td><td rowspan="1" colspan="1">0.0151</td><td rowspan="1" colspan="1">0.0025–0.0052</td><td rowspan="1" colspan="1">0.0032</td><td rowspan="1" colspan="1">0.0043–0.0149</td><td rowspan="1" colspan="1">0.0138</td><td rowspan="1" colspan="1">0.013–0.0145</td><td rowspan="1" colspan="1">0.01375</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0397–0.0520</td><td rowspan="1" colspan="1">0.046</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0</td><td rowspan="1" colspan="1">0.0043–0.0093</td><td rowspan="1" colspan="1">0.007</td><td rowspan="1" colspan="1">0–0.0043</td><td rowspan="1" colspan="1">0.0024</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0390–0.0396</td><td rowspan="1" colspan="1">0.0393</td><td rowspan="1" colspan="1">0.0025</td><td rowspan="1" colspan="1">0.0025</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">0.0043–0.0046</td><td rowspan="1" colspan="1">0.0045</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0765–0.0846</td><td rowspan="1" colspan="1">0.0798</td><td rowspan="1" colspan="1">0.0025–0.0049</td><td rowspan="1" colspan="1">0.0031</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">0.0072–0.0087</td><td rowspan="1" colspan="1">0.0077</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0380–0.0413</td><td rowspan="1" colspan="1">0.0398</td><td rowspan="1" colspan="1">0.0025</td><td rowspan="1" colspan="1">0.0025</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">N/A</td><td rowspan="1" colspan="1">0.0185</td><td rowspan="1" colspan="1">0.0185</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.1626–0.1691</td><td rowspan="1" colspan="1">0.1659</td><td rowspan="1" colspan="1">0.0500–0.0575</td><td rowspan="1" colspan="1">0.0525</td><td rowspan="1" colspan="1">0.0488–0.0532</td><td rowspan="1" colspan="1">0.0510</td><td rowspan="1" colspan="1">0.0522–0.0552</td><td rowspan="1" colspan="1">0.0537</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0934–0.1089</td><td rowspan="1" colspan="1">0.1036</td><td rowspan="1" colspan="1">0.0025</td><td rowspan="1" colspan="1">0.0025</td><td rowspan="1" colspan="1">0.0021</td><td rowspan="1" colspan="1">0.0021</td><td rowspan="1" colspan="1">0.0052–0.0103</td><td rowspan="1" colspan="1">0.0089</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0098–0.0179</td><td rowspan="1" colspan="1">0.0131</td><td rowspan="1" colspan="1">0.0000</td><td rowspan="1" colspan="1">0.0000</td><td rowspan="1" colspan="1">0.0064–0.0142</td><td rowspan="1" colspan="1">0.0103</td><td rowspan="1" colspan="1">0.0155–0.0169</td><td rowspan="1" colspan="1">0.0162</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.0791–0.1041</td><td rowspan="1" colspan="1">0.0902</td><td rowspan="1" colspan="1">0.0025–0.0099</td><td rowspan="1" colspan="1">0.0066</td><td rowspan="1" colspan="1">0.0150–0.0194</td><td rowspan="1" colspan="1">0.0172</td><td rowspan="1" colspan="1">0.0089–0.01231</td><td rowspan="1" colspan="1">0.0107</td></tr><tr><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic> vs. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sinopiceus</named-content></named-content></italic></td><td rowspan="1" colspan="1">0.1260–0.1366</td><td rowspan="1" colspan="1">0.1320</td><td rowspan="1" colspan="1">0.0099–0.0149</td><td rowspan="1" colspan="1">0.0120</td><td rowspan="1" colspan="1">0.0336–0.0414</td><td rowspan="1" colspan="1">0.0370</td><td rowspan="1" colspan="1">0.0135–0.0245</td><td rowspan="1" colspan="1">0.0204</td></tr></tbody></table></table-wrap></sec><sec sec-type="Distribution mapping"><title>Distribution mapping</title><p>Distribution maps were created in ArcMap (RockWare; Golden, Colorado, USA) by Thomas Atkinson (University of Texas) and are a combination of specimens examined as part of this study (black) and reported material examined by Atkinson and records gleaned from literature (white). A database of Atkinson’s localities is available at <ext-link ext-link-type="uri" xlink:href="http://www.barkbeetles.info">www.barkbeetles.info</ext-link>.</p></sec><sec sec-type="Terminology"><title>Terminology</title><p>There are some traditionally used terms and characters in <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> literature that require clarification. The term ‘venter’ is regularly utilized. In general, the term refers to the entire ventral surface of an organism. However in <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> it only pertains to the abdominal ventral concavity, specifically ventrites 2–5. The length of ventrite 5 compared to ventrite 3 and 4 is another common misleading character in <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> literature. Rather than measuring the entire length of segment 5, only the distance from the basal margin to the subapical carinate ridge is measured and compared to the combined lengths of ventrites 3 and 4. The following <italic>Glossary</italic> (modified from <xref rid="B66" ref-type="bibr">Hopkins 1909</xref>; <xref rid="B135" ref-type="bibr">Torre-Bueno 1989</xref>; <xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B63" ref-type="bibr">Harris 1979</xref>) summarizes the meanings ascribed to some terms used in this monograph:</p><p><bold><italic>General</italic></bold></p><p><italic>Apical</italic>: referring to a point at or close to the apex or tip.</p><p><italic>Basal</italic>: referring to a point at or closest to the main body.</p><p><italic>Impressed</italic>: a depression in a surface, typically referring the elytral striae or frons.</p><p><italic>Concave</italic>: appearing hollowed out.</p><p><italic>Convex</italic>: appearing rounded.</p><p><italic>Vestiture</italic>: the surface covering composed of setae. In <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> the setae are either long or short.</p><p><bold><italic>Head</italic></bold> (Fig. <xref ref-type="fig" rid="F3">3</xref>)</p><fig id="F3" orientation="portrait" position="float"><label>Figure 3.</label><caption><p>Terminology of the head (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> male).</p></caption><graphic id="oo_33356.jpg" xlink:href="zookeys-450-001-g003"></graphic></fig><p><italic>Aciculate</italic>: referring to longitudinal groves or scratches on the frons that can appear coarse as if made by a knife or fine as if lightly scratched with a needle.</p><p><italic>Epistomal process</italic>: a raised, sinuate process composed of a median and two lateral sections apically fringed with thick, long bristles that cover the median epistomal area just above the mandibles.</p><pmc-comment>PageBreak</pmc-comment>
<p><italic>Frons</italic>: region of the head from just above the epistoma to a point that is just dorsal to the inner apices of the eyes.</p><p><italic>Inner apices of eyes</italic>: the innermost mesal margins of the eyes as viewed frontally.</p><p><italic>Strigate</italic>: having narrow, transverse lines in the cuticle.</p><p><italic>Vertex</italic>: the top of the head, above the eyes.</p><p><bold><italic>Elytra</italic></bold> (Figs <xref ref-type="fig" rid="F4">4</xref>–<xref ref-type="fig" rid="F5">5</xref>)</p><fig id="F4" orientation="portrait" position="float"><label>Figure 4.</label><caption><p>Habitus terminology (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> male), lateral.</p></caption><graphic id="oo_33357.jpg" xlink:href="zookeys-450-001-g004"></graphic></fig><fig id="F5" orientation="portrait" position="float"><label>Figure 5.</label><caption><p>Habitus terminology (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> male), dorsal.</p></caption><graphic id="oo_33358.jpg" xlink:href="zookeys-450-001-g005"></graphic></fig><p><italic>Apex</italic>: end of a structure that is distal to the base.</p><p><italic>Disc</italic>: the central upper surface of the elytra between the elytral bases and the sloped declivity.</p><p><italic>Striae</italic>: punctures in rows, which may or may not be impressed to make grooves.</p><p><italic>Interstriae</italic>: longitudinal spaces along the elytra between the striae, which is not as impressed and bear smaller punctures.</p><p><italic>Corrugated</italic>: with alternate ridges and channels, referring to the appearance of the elytral striae and interstriae.</p><p><italic>Sutural dehiscence</italic>: the central notch at the apical margin of the elytra.</p><p><bold><italic>Abdomen</italic></bold> (Fig. <xref ref-type="fig" rid="F6">6</xref>)</p><fig id="F6" orientation="portrait" position="float"><label>Figure 6.</label><caption><p>Abdominal venter terminology (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic> female).</p></caption><graphic id="oo_33359.jpg" xlink:href="zookeys-450-001-g006"></graphic></fig><p><italic>Carina</italic>: a ridge-like or keel-shaped projection of the exoskeleton.</p><p><italic>Cusp</italic>: a slight projection or elevation along a margin; refers to the apical margin of ventrite 1 in a few species.</p><p><italic>Denticle</italic>: a small tooth.</p><p><italic>Opaque</italic>: appearing dull in luster; referring to a surface that reflects little light.</p><p><italic>Produced</italic>: refers to a part of the exoskeleton that is extended, lengthened, or elevated.</p><p><italic>Punctate</italic>: set with fine impressed points appearing as pinpricks.</p><p><italic>Punctulate</italic>: minute punctures.</p><p><italic>Rugose</italic>: appearing wrinkled.</p><p><italic>Shagreened</italic>: covered with a closely set roughness and appearing similar to sharkskin.</p><p><italic>Shiny</italic>: appearing glossy or bright in luster; refers to a surface that appears polished and reflects light well.</p><pmc-comment>PageBreak</pmc-comment>
<pmc-comment>PageBreak</pmc-comment>
<p><italic>Spine</italic>: an elongate projection of the exoskeleton that is longer than its basal width.</p><p><italic>Ventrite</italic>: In <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> there are five visible ventrites. They are numbered from anterior to posterior with ventrite 1 closest to the head and ventrite 5 closest to the elytral apex.</p><p><italic>Subapical carinate ridge</italic>: a carinate ridge on ventrite 5 occurring just before the apical margin.</p><p><italic>Tubercle</italic>: a small knob-like or rounded protuberance of the exoskeleton.</p><p><italic>Tumescence</italic>: a swelling of the exoskeleton.</p><p><italic>Venter</italic>: the undersurface of the abdomen, in <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> this pertains to the five visible ventrites.</p><p><bold><italic>Gallery pattern</italic></bold> (Fig. <xref ref-type="fig" rid="F7">7</xref>)</p><fig id="F7" orientation="portrait" position="float"><label>Figure 7.</label><caption><p>Gallery pattern terminology (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic>).</p></caption><graphic id="oo_33360.jpg" xlink:href="zookeys-450-001-g007"></graphic></fig><p><italic>Adult gallery</italic>: the composite tunnel produced by the adult female includes 1–2 egg galleries, the egg niches and the nuptial chamber (if present).</p><pmc-comment>PageBreak</pmc-comment>
<p><italic>Egg gallery</italic>: a single extension of the adult gallery from the nuptial chamber (if present) along which eggs are deposited in niches.</p><p><italic>Egg niche</italic>: notches along the sides of the egg gallery excavated by females in which a single egg is deposited.</p><p><italic>Larval mine</italic>: the excavation tunnel produced by a larva as it feeds.</p><p><italic>Nuptial chamber</italic>: an enlarged area or short extension of the adult gallery at the base of the entrance tunnel that may be used for mating or as a turning niche.</p><p><italic>Pupation chamber</italic>: an ovoid or circular excavation at the end of a larval mine in which pupation occurs.</p></sec></sec><sec><title>Phylogenetic results</title><sec sec-type="Morphology"><title>Morphology</title><p>The morphological phylogeny was poorly resolved and few synapomorphic characters were found (Fig. <xref ref-type="fig" rid="F2">2</xref>). This reflects the morphologically similarity of many taxa. The morphological phylogeny recovered two clades: native hardwood species (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic>) and conifer species (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic>, and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic>). The outgroup taxa were poorly resolved except for <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sinopiceus</named-content></named-content></italic>, which were recovered as sister taxa. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> was recovered as a distinct lineage separate from <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic>, of which it was considered a synonym. The relationship among <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content></italic> was unresolved and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">virgatus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic> were strongly supported as monophyletic.</p></sec><sec sec-type="Mitochondrial COI"><title>Mitochondrial COI</title><p>The parsimony analysis of mitochondrial COI produced 2,958 most parsimonious trees with a length of 1,434 steps and 606 of 616 characters were parsimony informative (Fig. <xref ref-type="fig" rid="F8">8</xref>). There was a high degree of synapomorphy and low amount of homoplasy in the phylogeny (CI = 0.356, RI = 0.809).</p><fig id="F8" orientation="portrait" position="float"><label>Figure 8.</label><caption><p>Mitochondrial COI phylogeny of Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic>. Phylogram of one of 2958 most parsimonious trees (L = 1431, CI = 0.356, RI = 0.809) generated from a heurtistic search of 1000 stepwise random additions with TBR in PAUP*. Numbers above the node indicate bootstrap values (>50). Bremer support values are listed below the node. Non-monophyletic species are indicated in red. Mean intraspecific nucleotide differences (listed next to each taxon) and mean interspecific nucleotide differences (right of solid line) are given. Taxa highlighted in red were not recovered as monophyletic.</p></caption><graphic id="oo_33361.jpg" xlink:href="zookeys-450-001-g008"></graphic></fig></sec><sec sec-type="Combined analyses"><title>Combined analyses</title><p>Bayesian analysis of the combined dataset recovered the native Nearctic <pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> as paraphyletic with native species found in two clades: native hardwood and conifer feeders (Fig. <xref ref-type="fig" rid="F9">9</xref>). Bayesian analysis of the combined dataset (Fig. <xref ref-type="fig" rid="F9">9</xref>) recovered the monophyletic groups observed in the morphological phylogeny and the native species were recovered in two clades: native hardwood-feeders (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic>) and a second clade of conifer-feeders (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic>, and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic>) sister to <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic>.</p><fig id="F9" orientation="portrait" position="float"><label>Figure 9.</label><caption><p>Bayesian tree found by analysis of data using combined molecular and morphological datasets. St. dev. = 0.006983. Numbers above the nodes are Bayesian posterior probabilities. Taxa in purple were not recovered as monophyletic. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> was a synonym of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic>. * denotes species exotic to the Nearctic.</p></caption><graphic id="oo_33362.jpg" xlink:href="zookeys-450-001-g009"></graphic></fig></sec></sec><sec><title>Taxonomic treatment</title><p>Based on the results of our phylogenetic analyses, 25 species occur in the Nearctic. Several species were not recovered as monophyletic and had less than 1% difference in all genes sequences (Table <xref ref-type="table" rid="T6">6</xref>). These species are thus placed in synonymy with the oldest described species. The following synonyms were discovered: <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> (= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic>; = <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content></italic>), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> (= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">virgatus</named-content></named-content></italic>; = <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> was recovered as monophyletic and is thus removed from synonymy with <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic>. Further detail regarding taxonomic changes is found in the remarks section for each species.</p><sec sec-type="A checklist of the Nearctic Scolytus species"><title>A checklist of the Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species</title><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic> Wood, 1967</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic> Bright, 1964</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> Walsh, 1867</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> Blackman, 1934, valid species</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic> Wood, 1968</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> Blackman, 1934</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic> (Bechstein, 1805) – Introduced</styled-content></p><p><styled-content style="padding-left:60px; text-indent:-20px; display:block;">= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sulcatus</named-content></named-content></italic> LeConte, 1868</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> (Swaine, 1917)</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> (Marsham, 1802) – Introduced</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic> Wood, 1968</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic> Say, 1824</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic> Wood, 1962</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic> Blackman, 1934</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic> (Swaine, 1910)</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> LeConte, 1876</styled-content></p><p><styled-content style="padding-left:60px; text-indent:-20px; display:block;">= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic> Blackman, 1934, syn. n.</styled-content></p><p><styled-content style="padding-left:60px; text-indent:-20px; display:block;">= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content></italic> Blackman, 1934, syn. n.</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic> Say, 1824</styled-content></p><p><styled-content style="padding-left:60px; text-indent:-20px; display:block;">= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">carya</named-content></named-content></italic> Riley, 1867</styled-content></p><p><styled-content style="padding-left:60px; text-indent:-20px; display:block;">= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">caryae</named-content></named-content></italic> Walsh, 1867</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> Blackman, 1934</styled-content></p><p><styled-content style="padding-left:60px; text-indent:-20px; display:block;">= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">virgatus</named-content></named-content></italic> Bright, 1972, syn. n.</styled-content></p><p><styled-content style="padding-left:60px; text-indent:-20px; display:block;">= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic> Blackman, 1934, syn. n.</styled-content></p><pmc-comment>PageBreak</pmc-comment>
<p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic> Blackman, 1934</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic> (Müller, 1818) – Introduced</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> Semenov, 1902 – Introduced</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic> Bright, 1972, valid species</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic> LeConte, 1876</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> (Swaine, 1917)</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> LeConte, 1876</styled-content></p><p><styled-content style="padding-left:60px; text-indent:-20px; display:block;">= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sobrinus</named-content></named-content></italic> Blackman, 1934</styled-content></p><p><styled-content style="padding-left:40px; text-indent:-20px; display:block;"><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> LeConte, 1868</styled-content></p></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></title><p><named-content content-type="taxon-authority">Geoffroy, 1762</named-content></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content><named-content content-type="comment"> Geoffroy, 1762: 309</named-content></p></list-item></list><sec sec-type="treatment-Type species"><title>Type species.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Bostrichus</named-content><named-content content-type="species">scolytus</named-content></named-content></italic> Fabricius, 1775.</p><p>See <xref rid="B1" ref-type="bibr">Alonso-Zarazaga and Lyal (2009)</xref> for complete taxonomic history.</p></sec><sec sec-type="treatment-Diagnostic characters"><title>Diagnostic characters.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> is easily distinguished from all other Nearctic scolytines by the unarmed protibia with a single curved uncus at the outer apical angle, by the flattened antennal club with 0–1 septate procurved sutures, by the seven-segmented funicle, by the slightly declivous elytra, the depressed scutellar notch and scutellum and by the abruptly ascending abdominal ventrites 2–5 (except <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic>).</p></sec><sec sec-type="treatment-Description"><title>Description</title><p><bold>(modified from <xref rid="B148" ref-type="bibr">Wood 1982</xref>).</bold> Length 1.7–6.0 mm, 1.7–2.9 times as long as wide. Color variable, red-brown to black. Teneral adults are often light brown.</p><p>Head visible from above. Frons flattened to convex, sexually dimorphic (discussed below for each species group or clade). Eye elongate, sinuate to shallowly emarginated, finely faceted. Antennal scape club shaped, shorter than four funicle segments; funicle seven-segmented; club larger than funicle, flattened, oval to obovate, minutely pubescent and with 3 strongly procurved sutures, suture 1 partially to completely septate and with or without a surface groove. Pronotum large, wider than long; lateral and basal margins marked by a fine elevated line; disc finely punctate. Scutellum triangular, deeply depressed below elytral surface. Elytra depressed around scutellum and along basal one-fifth to half-length of elytral suture. Elytra wider than pronotum, flattened; striae punctate; interstriate punctate, with or without setae. Venter either gradually ascending from apical margin of ventrite 1 to elytral apex, or ventrite 2 abruptly ascending, sexually dimorphic, with or without tubercules and/or carinae; remaing ventrite usually unarmed, may have various armature or elevated margins, especially in males. Procoxae narrowly separated. Protibia rectangular; sides straight, nearly parallel, without denticles; outer distal angle produced into a curved uncus.</p></sec><sec sec-type="treatment-Species descriptions"><title>Species descriptions.</title><p>Species are redescribed and treated alphabetically by clade or group beginning with the introduced group, native hardwood clade and conifer clade. Common names listed are the official common names of the Entomological Society of America and Canada.</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="treatment-Introduced species group"><title>Introduced species group.</title><p>The introduced species group (1.9–4.2 mm long) (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic>) is not monophyletic and consists of a morphologically diverse group of Palearctic species that encompasses three formerly recognized subgenera (<xref rid="B23" ref-type="bibr">Butovitsch 1929</xref>; <xref rid="B6" ref-type="bibr">Balachowsky 1949</xref>). This diversity is reflected in the sexual dimorphism exhibited by the group. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> exhibit the typical frons dimorphism as discussed above. The venter lateral teeth are identical in both sexes of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> and the spine on ventrite 2 is smaller in the female compared to the male. Sexual dimorphism of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic> is much more subtle and differentiating the sexes can be challenging. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic> males have the medial area of the frons and epistomal regions impressed and the lateral margins of the epistoma are lightly covered with more abundant erect setae than the female. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic> males have the medial area of the frons and epistoma slightly more impressed and the frons bears longer and more abundant, erect setae.</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></title><p><named-content content-type="taxon-authority">(Bechstein, 1805)</named-content></p><p><xref ref-type="fig" rid="F10">Fig. 10</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Bostrichus</named-content><named-content content-type="species">mali</named-content></named-content><named-content content-type="comment"> Bechstein, 1805: 882.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content><named-content content-type="comment"> (Bechstein, 1805): <xref rid="B44" ref-type="bibr">Eichhoff 1881</xref>: 41.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sulcatus</named-content></named-content><named-content content-type="comment"> LeConte, 1868: 167. <xref rid="B21" ref-type="bibr">Brown 1950</xref>: 203.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content><named-content content-type="comment"> For complete taxonomic history see <xref rid="B151" ref-type="bibr">Wood and Bright (1992)</xref>.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Both sexes of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic> are distinguished by having ventrite 2 at an oblique angle to ventrite 1, by the aciculate frons, with most setae found just above the epistoma and on the lateral epistomal margins, by the smooth, shining appearance of the pronotum and elytra, and by the weakly rounded and smooth elytral apex.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>3.2–4.1 mm long (mean = 3.7 mm; n = 10); 2.1–2.5 times as long as wide. Head, pronotum and abdominal venter dark red-brown, legs and antennae light brown, elytra brown to red-brown. Pronotum typically darker than elytra.</p><p><italic>Head</italic>. Epistoma weakly, broadly emarginate; epistomal process absent; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing convex when viewed laterally, moderately transversely impressed just above epistoma, longitudinally impressed near median line; densely, finely longitudinally aciculate-punctate; aciculations converging at epistoma; punctures small, coarse; sparsely covered by long, fine, erect, yellow hair-like setae, these as long as width of midpoint of eye, more abundant along epistoma. Antennal scape short, elongate; club flattened, irregularly ovoid, setose with partial septum, two arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to weakly rounded, smooth apex; apex weakly emarginate at suture. Margin of apical edge bearing small, fine punctures. Disc smooth, shining; interstriae weakly impressed, more than twice width of striae, interstrial punctures uniseriate, smaller than those of striae, bearing sparse, long, semi-erect yellow hair-like setae (may be abraded); striae weakly impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron greater than half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 weakly elevated above base of ventrite 2. Ventrite 2 nearly at an oblique angle to ventrite 1; surface smooth, shining, finely punctate; punctures small, fine, shallow; surface convex, unarmed; setae erect, short, about half of ventrite 3 length; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 unarmed; length of ventrite 5 equal to combined lengths of ventrites 3 and 4; setal patch absent; median depression present.</p><fig id="F10" orientation="portrait" position="float"><label>Figure 10.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33363.jpg" xlink:href="zookeys-450-001-g010"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>3.0–4.2 mm long (mean = 3.7 mm; n = 10); 2.0–2.5 times as long as wide. Similar to male except epistoma feebly emarginate, frons more strongly convex when viewed laterally, weakly aciculate, setae sparser, shorter, less than width of eye.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>111.</p><p><italic>Type material.</italic> Syntypes <italic><named-content content-type="taxon-name"><named-content content-type="genus">Bostrichus</named-content><named-content content-type="species">mali</named-content></named-content></italic> Bechstein (location unknown). Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sulcatus</named-content></named-content></italic> LeConte: male, labeled “[pink disc = Middle States (Md., Del., N.Y., N.J., Pa., Conn.?, R.I.?)]; type 969" (MCZC).</p><p><italic>Non-type material.</italic><bold>CANADA:</bold><bold><italic>ONTARIO</italic>:</bold> Hamilton, 14–21.VI.1981, M. Sanborne (CNCI-2), 23.VII.1980, ex. malaise trap (CNCI-1). Owen Sound, 27.V.[19]65, K.E. Stewart, ex. elm [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> sp.] (CNCI-1). Vineland Station, 8.VI.1949, W.L. Putnam, ex. apple [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Malus</named-content></named-content></italic> sp.] (CNCI-2). <bold><italic>QUEBEC</italic>:</bold> Cantic, 11.VIII.1945, W.J. Brown, ex. apple [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Malus</named-content></named-content></italic> sp.] (CNCI-2). Cap-Saint-Ignace, VII.1970 (CNCI-1). Dunham, 2.IX.1998, Vignoble, L’Orpailleur, ECORC/CRDHAg-Cord (DEBC-1). Napierville, 2 mi N.E., 13.VIII.1945, W.J. Brown (CNCI-1). Noyan, 0.75 mi W., 15.VIII.1945, W.J. Brown (CNCI-1). <bold>UNITED STATES:</bold><bold><italic>CONNECTICUT</italic>:</bold> [<italic>Fairfield Co.</italic>]: Greenwich, 22.VII.1933, F.J. Dillaway, ex. in plum [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Prunus</named-content></named-content></italic> sp.] (USNM-2). <italic>New Haven Co.</italic>: New Haven, 25.VI.1956, C.W. O’Brien (EMEC-1). <bold><italic>MAINE</italic>:</bold> [<italic>Androscoggin Co.</italic>]: Livermore Falls, 5.VIII.1975, ex. plum [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Prunus</named-content></named-content></italic> sp.] (USNM-1). <bold><italic>MARYLAND</italic>:</bold><italic>Montgomery Co.</italic>: Ashton, 4 mi S.W., 31.V.1986, G.F. & J.F. Hevel, ex. malaise trap (USNM-1). <bold><italic>MASSACHUSETTS</italic>:</bold><italic>Worchester Co.</italic>: 16.II.[19]53, ex. indoors (USNM-2). <bold><italic>MICHIGAN</italic>:</bold><italic>Allegan Co.</italic>: Fennville area, 30.VI.2003, P. McGhee, ex. apple trees [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Malus</named-content></named-content></italic> sp.] (MSUC-6). <italic>Genesee Co.</italic>: Richfield County Park, <named-content content-type="dwc:verbatimCoordinates">N43°100610, W-83°55810</named-content>, 16.VI.2008, R. Mech, PI Anthony Cognato (MSUC-1). <italic>Ingham Co.</italic>: Lansing, 2 mi N., E. State Rd, <named-content content-type="dwc:verbatimCoordinates">N42.7842°, W84.5362°</named-content>, 261 m, 2-18.VI.2007, PI Anthony Cognato, ex. Lindgren trap with ipslure (MSUC-1). <italic>Kalamazoo Co.</italic>: Gourdneck Lake State Game Area, 16.VII.2011, S.M. Smith, A.I. Cognato, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Prunus</named-content></named-content></italic> sp. (MSUC-2). <italic>Oakland Co.</italic>: Farmington Hills, <named-content content-type="dwc:verbatimCoordinates">N42°27.668', W83°25.579'</named-content>, 2.VII.2004, B. Sullivan, ex. Lindgren funnel with multistriatus lure (MSUC-2). <italic>Saginaw Co.</italic>: St. Charles, 25.VI.1968, J.G. Truchan, ex. rotary trap (MSUC-1), 25.VI.1969 (MSUC-1). <italic>Wayne Co.</italic>: 20.VI.1960, G. Steyskai (USNM-2). <bold><italic>NEW JERSEY</italic>:</bold> [<italic>Essex Co.</italic>]: Maplewood, 7.VI.[19]34, D. Fivaz, ex. on elm [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> sp.] (USNM-12). [<italic>Morris Co.</italic>]: Chatham, 25.III.[19]34, W.D. Buchanan (USNM-1). <bold><italic>NEW YORK</italic>:</bold><italic>Albany Co.</italic>: near Rensselaerville, Huyck Preserve, 3-10.VII.1967, R. & J. Matthews, ex. window pane trap (CNCI-1). [<italic>Suffolk Co.</italic>]: Cutchogue, [19]45, Tuthill, ex. in apple [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Malus</named-content></named-content></italic> sp.] (USNM-26). [<italic>Tompkins Co.</italic>]: Groton, 24.V.1942, N.M. Downie (FMNH-1), 23.VI.1946 (FMNH-2). [<italic>Westchester Co.</italic>]: Armonk, 5.VI.[19]35, H. Dietrich, ex. apple [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Malus</named-content></named-content></italic> sp.] (CNCI-2). <bold><italic>OHIO</italic>:</bold><italic>Medina Co.</italic>: 15.VI.[19]62, C.L. Griswold (DEBC-4). <bold><italic>PENNSYLVANIA</italic>:</bold><italic>Luzerne Co.</italic>: Nanticoke, 20.IX.[19]60, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Malus</named-content></named-content></italic> sp. (CNCI-1). Nuangola, 12.V-23.VIII.1983, S. & J. Peck, ex. forest intercept (CNCI-1). <bold><italic>VERMONT</italic>:</bold><italic>Rutland Co.</italic>: Castleon, 18-19.VI.1989, H.V. Weems (FSCA-1). <bold><italic>WASHINGTON, D.C.</italic>:</bold> 5.IV.1983, E.R. Hodges, ex. on sweater of collector (USNM-1). <bold>Additional specimens:</bold><italic>Czech Republic</italic> (MSUC-21) and <italic>Italy</italic> (MSUC-1).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: Ontario, Quebec. UNITED STATES: Connecticut, Maine, Maryland, Massachusetts, Michigan, New Jersey, New York, Ohio, Pennsylvania, Vermont, Washington, D.C., Wisconsin (Fig. <xref ref-type="fig" rid="F11">11</xref>).</p><fig id="F11" orientation="portrait" position="float"><label>Figure 11.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33364.jpg" xlink:href="zookeys-450-001-g011"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Malus</named-content></named-content></italic> spp. (apple), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Prunus</named-content></named-content></italic> spp. (cherry), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> spp. (elm), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pyrus</named-content></named-content></italic> spp. (pear) and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Sorbus</named-content></named-content></italic> spp. (mountain ash).</p></sec><sec sec-type="treatment-Common name"><title>Common name.</title><p>Large shothole borer.</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p>This species colonizes dying and weakened limbs of its host as well as fresh slash. Typical host material is 15.0–38.0 cm in diameter but branches as small as 8.0 cm are colonized (<xref rid="B103" ref-type="bibr">Pechuman 1938</xref>).</p><p>Adult galleries are somewhat variable and are either parallel or at a slight oblique angle to the grain of the wood and consisting of a nuptial chamber and a single egg gallery. The adult gallery strongly scores the sapwood and ranges in size from 3.5–6.0 cm in length. Egg niches are constructed along the gallery and score the sapwood. Six to 100 eggs may be laid along the egg galleries with the typical gallery having 40 eggs. Larval mines lightly score the sapwood and radiate perpendicular to the egg gallery. Larval galleries later meander often at an oblique angle to the grain of wood, forming a fan shaped pattern. Pupation occurs within the sapwood and broods overwinter as larvae or adults. The following year, adults emerge and feed at twig crotches before selecting host material (<xref rid="B103" ref-type="bibr">Pechuman 1938</xref>; <xref rid="B6" ref-type="bibr">Balachowsky 1949</xref>; <xref rid="B5" ref-type="bibr">Baker 1972</xref>; <xref rid="B148" ref-type="bibr">Wood 1982</xref>). In New York, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic> has one generation per year although there are two generations per year in Europe (<xref rid="B103" ref-type="bibr">Pechuman 1938</xref>).</p></sec><sec sec-type="treatment-Collection notes"><title>Collection notes.</title><p>The senior author collected this species from an 8.0 cm diameter <italic><named-content content-type="taxon-name"><named-content content-type="genus">Prunus</named-content></named-content></italic> sp. branch that had broken during a recent windstorm. Females constructed gallery entrances beneath large flakes of bark on the sides and bottom surfaces of the branch. Specimens were infesting the same limbs as <italic><named-content content-type="taxon-name"><named-content content-type="genus">Phloeotribus</named-content><named-content content-type="species">liminaris</named-content></named-content></italic> (Harris, 1852).</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>This species is native to the Palearctic region and was first detected in New York in 1868 when it was described as <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sulcatus</named-content></named-content></italic> LeConte. <xref rid="B82" ref-type="bibr">LeConte (1868)</xref> noted in his description that the species most strongly resembled the Palearctic species <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic> and did not mention the collection date of his specimens. Interestingly, no other specimens were collected until 1933, 65 years after the initial discovery from southeastern New York, northern New Jersey and western Connecticut. It is likely that like LeConte’s <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">californicus</named-content></named-content></italic> (= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">scolytus</named-content></named-content></italic>), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic> was collected in 1868 but populations never became established. The later collections may be the result of multiple introduction events (<xref rid="B103" ref-type="bibr">Pechuman 1938</xref>).</p><p><xref rid="B21" ref-type="bibr">Brown (1950)</xref> recognized that LeConte’s species was morphologically and behaviorally identical to that of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic> and placed <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sulcatus</named-content></named-content></italic> into synonymy. The native range of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic> is Europe, Central Asia, Siberia, the Russian Far East, North Africa (<xref rid="B97" ref-type="bibr">Michalski 1973</xref>; <xref rid="B78" ref-type="bibr">Knížek 2011</xref>).</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></title><p><named-content content-type="taxon-authority">(Marsham, 1802)</named-content></p><p><xref ref-type="fig" rid="F12">Fig. 12</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Ips</named-content><named-content content-type="species">multistriatus</named-content></named-content><named-content content-type="comment"> Marsham, 1802: 54.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content><named-content content-type="comment"> (Marsham, 1802). Chapuis and Candeze 1853: 577.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">javanus</named-content></named-content><named-content content-type="comment"> Chapuis, 1869: 56. <xref rid="B115" ref-type="bibr">Schedl 1954</xref>: 137.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content><named-content content-type="comment"> For complete taxonomic history see <xref rid="B151" ref-type="bibr">Wood and Bright (1992)</xref>.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Both sexes of the species are distinguished by the presence of lateral teeth on the apical margins of ventrites 2–4 and by a median conical spine on the basal margin of ventrite 2.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>2.2–3.9 mm long (mean = 2.81 mm; n = 20); 2.0–2.6 times as long as wide. Head, pronotum, and abdominal venter dark red-brown, legs light brown, antennae yellow-brown, elytra usually dark red-brown but may be brown. Color not uniform and pronotal and elytral surfaces frequently contain patches of red-brown mixed with dark red-brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly, broadly emarginate; epistomal process absent; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing flattened when viewed laterally; moderately, coarsely, longitudinally aciculate-punctate; aciculations converging at epistoma; punctures small, coarse; smoderately, uniformly covered by long, fine, yellow erect hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, thinner on apical half, irregularly ovoid, setose with partial septum, two sharply arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, smooth apex; apex entire at suture. Margin of apical edge bearing small, fine punctures. Disc smooth, shining; interstriae weakly impressed, more than twice width of striae, interstrial punctures uniseriate, smaller than those of striae, bearing sparse, long, semi-erect yellow hair-like setae (may be abraded); striae weakly impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron greater than half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 weakly elevated above base of ventrite 2. Ventrite 2 nearly perpendicular to ventrite 1; surface smooth, shining, finely punctate, punctures small, coarse; surface flattened; basal margin armed with a long, smooth, conical spine with it’s base extending from basal margin to half length of segment; lateral margins of ventrites 2–4 armed with lateral tooth. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 greater than combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F12" orientation="portrait" position="float"><label>Figure 12.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33365.jpg" xlink:href="zookeys-450-001-g012"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>2.4–3.3 mm long (mean = 2.95 mm; n = 20); 2.2–2.7 times as long as wide. Similar to male except epistoma feebly entire, frons strongly convex when viewed laterally, weakly aciculate, setae sparser, shorter, less than the width of eye.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>165.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">javanus</named-content></named-content></italic> Chapuis: male, labeled “Java, Solier, Dejean” (ISNB).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>CANADA:</bold><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><bold><italic>ALBERTA</italic>:</bold> Calgary, 19.VII.[19]94, T. Reichardt, ex. pheromone trap no. EBB XI (CNCI-2). <bold><italic>ONTARIO</italic>:</bold> St. Catherines, 22.VI.1961, Kelton, Brumpton (CNCI-1). Toronto, 1970, ex. reared on elm [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> sp.] (CNCI-1). Queenston, 26.VI.[19]50, ex. elm [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> sp.] (CNCI-1). <bold><italic>QUEBEC</italic>:</bold> Montreal, 14.VII.1977, E.J. Kiteley (CNCI-1); Île de Montreal, Île de Sainte Hélène, 12.VIII.1985, L. Lariviere (CNCI-1). <bold>MEXICO:</bold><bold><italic>Chihuahua</italic>:</bold> Cuidad Juarez, II.1987, I.C. Rodriguez (CNCI-2). <bold>UNITED STATES:</bold><bold><italic>CALIFORNIA</italic>:</bold> [<italic>Alameda Co.</italic>]: Berkeley, 23.V.1962, ex. elm [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> sp.] (EMEC-2). <italic>Contra Costa Co.</italic>: Antioch [Dunes] National Wildlife Refuge, 10.X.[19]91, J.A. Powell (EMEC-1). <italic>Fresno Co.</italic>: 11.I.1950, N.J. Smith (SBMN-2). <italic>Inyo Co.</italic>: Big Pine, 4000 ft, V.1971, D. Guiliani, ex. black light (CASC-1), VII.1971 (CASC-1). <bold><italic>COLORADO</italic>:</bold><italic>Mesa Co.</italic>: Grand Junction, X.1979, D. Leatherman, ex. European ash [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Fraxinus</named-content></named-content></italic> sp.] (CSUC-1). <italic>Weed Co.</italic>: Briggsdale, 1.IX.198[sic!], D. Leatherman (CSUC-1). <bold><italic>GEORGIA</italic>:</bold><italic>Clarke Co.</italic>: Whitehall forest, 17-24.IX.1976, R. Turnbow, ex. window trap F-8 (CNCI-1). <bold><italic>ILLINOIS</italic>:</bold><italic>Alexander Co.</italic>: Horseshoe Lake, 28.IX.1968, T.E. Brooks (CNCI-4). <bold><italic>KENTUCKY</italic>:</bold><italic>Green Co.</italic>: 9.VII.1941, C. Cook (MSUC-1). <bold><italic>MASSACHUSETTS</italic>:</bold> [<italic>Hampshire Co.</italic>]: Northampton, 10.VIII.1974, E.J. Kiteley (MSUC-1). [<italic>Suffolk Co.</italic>]: Forest Hills [= Boston], 3.VI.[?], W.M. Mann (USNM-1). <bold><italic>MICHIGAN</italic>:</bold><italic>Antrim Co.</italic>: Eastport, ca. 3 mi N., <named-content content-type="dwc:verbatimCoordinates">N45°08'30", W85°22'50"</named-content>, 3–5.VII.2003, F.W. Stehr, ex. UV light (MSUC-1). <italic>Cass Co.</italic>: Cassopolis, 2.VI.2007, A.D. Smith (MSUC-3). <italic>Genesee Co.</italic>: Richfield County Park, <named-content content-type="dwc:verbatimCoordinates">N43.100610°, W-83.55810°</named-content>, ex. Lindgren with EtOH (MSUC-1). <italic>Ingham Co.</italic>: Lansing, 2 mi N., E. State Rd, <named-content content-type="dwc:verbatimCoordinates">N42.7842°, W84.5361°</named-content>, 261 m, 16-30.VII.2007, PI A.I. Cognato (MSUC-1); G.L. Parsons, ex. UV light. & white lights (MSUC-1). East Lansing, 1.X.1957, R.C. Fox (MSUC-1), 2.X.1957 (MSUC-5), 3.X.1957 (MSUC-7), 4.X.1957 (MSUC-1), 5.X.1957 (MSUC-3), 6.X.1957 (MSUC-4), 10.X.1957 (MSUC-1), 15.X.1957 (MSUC-5); 25.VI.1981, R. Fischer (MSUC-1); <named-content content-type="dwc:verbatimCoordinates">N42°11.320', W84°27.867'</named-content>, 258 m, 16.IX.2011, S.M. Smith, A.I. Cognato, I.A. Cognato, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> sp. (MSUC-7); Michigan State University campus, 17.VI.2011, D.G. McCullough (MSUC-3). <italic>Kalamazoo Co.</italic>: Gourdneck Lake State Game Area, 19.VI.2011, A.I. Cognato, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> sp. (MSUC-21). <italic>Kalkaska Co.</italic>: T27N R7W S18, 28.XII.1986, P. Waclawski, ex. basement wood (MSUC-1), 20.XII.1987 (MSUC-1), 3.I.1987 (MSUC-5). <italic>Lapeer Co.</italic>: Potter Lake, 19.VI.1967, Brivio (MSUC-2). <italic>Macomb Co.</italic>: East of Memphis, 3.V.1964, C. Brivio (MSUC-1), 17.VI.1965 (MSUC-1), 21.VIII.[19]66 (MSUC-2), 23.VIII.1969 (MSUC-1), 30.VIII.1969 (MSUC-1). <italic>Saginaw Co.</italic>: St. Charles, 30.V.1969, J.H. Truchan, ex. rotary trap at 6 ft height level (MSUC-2), 11.VI.1969, ex. rotary trap at 12 ft height level (MSUC-1), 16.VI.1969 (MSUC-3). <italic>Sanilac Co.</italic>: Port Sanilac, 20.VI.1986, Brivio (MSUC-1). <bold><italic>MONTANA</italic>:</bold><italic>Choteau Co.</italic>: Great Falls, VII-VIII.1977, S. Kohler, ex. caught in flight, sticky trap, multilure bait (CNCI-1). <italic>Gallatin Co.</italic>: Bozeman, VII-VIII.1977, S. Kohler, ex. caught in flight, sticky trap, multilure bait (CNCI-1). <bold><italic>NEW YORK</italic>:</bold> [<italic>Westchester Co.</italic>]: Yonkers, VIII.1935, H. Dietrich, ex. elm [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> sp.] (CNCI-3). <bold><italic>OKLAHOMA</italic>:</bold> [<italic>Oklahoma Co.</italic>]: Jones, 13.VI.1957, D. Alexander (USNM-1). <bold><italic>TEXAS</italic>:</bold> [<italic>El Paso Co.</italic>]: El Paso, 045483, 17.VII.[19]94, lot 94 07975, ex. Mexico–in log of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Prunus</named-content></named-content></italic> sp. (USNM-10). <bold><italic>WYOMING</italic>:</bold><italic>Carbon Co.</italic>: Medicine Bow National Forest, VI-VIII.1999, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Polyphorus</named-content><named-content content-type="species">volvatus</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Cryptoporus</named-content><named-content content-type="species">volvatus</named-content></named-content></italic> (Peck) Shear] (CSUC-1). <bold>Additional specimens:</bold><italic>Czech Republic</italic> (MSUC-2), <italic>Italy</italic> (MSUC-8) and <italic>Russia</italic> (MSUC-22).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: Alberta, Manitoba, New Brunswick, Nova Scotia, Ontario, Quebec, Saskatchewan. MEXICO: Aguascalientes, Chihuahua. UNITED STATES: Alabama, Arizona, Arkansas, California, Colorado, Connecticut, Florida, Georgia, Idaho, Illinois, Indiana, Kansas, Kentucky, Louisiana, Maine, Maryland, Massachusetts, Michigan, Missouri, Montana, Nebraska, Nevada, New Hampshire, New Jersey, New Mexico, New York, North Carolina, North Dakota, Ohio, Oklahoma, Oregon, Pennsylvania, Rhode Island, South Carolina, South Dakota, Tennessee, Texas, Utah, Vermont, Virginia, Washington, D.C., West Virginia, Wisconsin, Wyoming (Fig. <xref ref-type="fig" rid="F13">13</xref>).</p><fig id="F13" orientation="portrait" position="float"><label>Figure 13.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33366.jpg" xlink:href="zookeys-450-001-g013"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p>All native and introduced <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> spp. including <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content><named-content content-type="species">americana</named-content></named-content></italic> L. (American elm) and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Zelkova</named-content><named-content content-type="species">serrata</named-content></named-content></italic> (Thunb.) Makino.</p></sec><sec sec-type="treatment-Common names"><title>Common names.</title><p>Smaller European elm bark beetle and the European elm bark beetle.</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> colonizes cut, stressed, weakened and diseased elm trees (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> spp.) (<xref rid="B148" ref-type="bibr">Wood 1982</xref>). It seldom attacks healthy and vigorous trees (<xref rid="B17" ref-type="bibr">Bright 1976</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> females produce an aggregation pheromone to aggregate conspecifics to host trees. The pheromone bouquet is composed of three components: (-)-4-methyl-3-heptanol, (-)-2,4-dimethyl-5-ethyl-6,8-dioxabicylo[3.2.1]octane (α)-multistriatin and (-)-α-cubebene (<xref rid="B102" ref-type="bibr">Pearce et al. 1975</xref>).</p><p>Mating can occur either as the female is undergoing maturation feeding in twig crotches or within the gallery as described for the genus (<xref rid="B129" ref-type="bibr">Svihra and Clark 1980</xref>). The adult gallery is excavated parallel to the grain of the wood and consists of a single egg gallery; a nuptial chamber is not constructed. The adult gallery ranges in size from 2.5–5.0 cm in length. Egg niches are constructed along the gallery and score the sap<pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment>wood. Twenty-four to 96 eggs may be laid singly along the egg gallery. Larval mines lightly score the sapwood and radiate perpendicular to the egg gallery. Larval galleries later meander often at an oblique angle to the grain of wood, forming a fan shaped pattern. Larvae construct pupal chambers in the bark (<xref rid="B17" ref-type="bibr">Bright 1976</xref>).</p><p>There are one to one-half generations per year in Canada and three in the southern US (<xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>). In Canada, adults emerge in June and July and feed at twig crotches of healthy trees for 7–10 days before selecting host material (<xref rid="B25" ref-type="bibr">Chamberlin 1958</xref>; <xref rid="B5" ref-type="bibr">Baker 1972</xref>; <xref rid="B129" ref-type="bibr">Svihra and Clark 1980</xref>). The brood from these adults either emerges in August or September or overwinters as larvae.</p><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> is the principal vector of the Dutch elm disease fungus <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ophiostoma</named-content><named-content content-type="species">ulmi</named-content></named-content></italic> (Buisman) Melin & Nannf in North America. This beetle vectored disease killed 50–75% of the elms population in northeastern North America prior to the 1930s (<xref rid="B13" ref-type="bibr">Bloomfield 1979</xref>). Adults become covered in fungal spores upon emergence from brood material. Adults inoculate elms with the Dutch elm disease fungus as they feed in twig crotches (<xref rid="B129" ref-type="bibr">Svihra and Clark 1980</xref>). This feeding activity leaves wounds in the bark that allow spores to be transferred from the beetle’s cuticle to the tree tissues (<xref rid="B17" ref-type="bibr">Bright 1976</xref>).</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>This species is native to the Palearctic region and is primarily distributed throughout Europe but also occurs in Iran and Algeria (<xref rid="B78" ref-type="bibr">Knížek 2011</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> was first detected in North America in 1909 from elm trees on the Harvard University campus in Massachusetts (<xref rid="B26" ref-type="bibr">Chapman 1910</xref>).</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></title><p><named-content content-type="taxon-authority">(Müller, 1818)</named-content></p><p><xref ref-type="fig" rid="F14">Fig. 14</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Bostrichus</named-content><named-content content-type="species">rugulosus</named-content></named-content><named-content content-type="comment"> Müller, 1818: 247.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content><named-content content-type="comment"> (Müller, 1818): <xref rid="B44" ref-type="bibr">Eichhoff 1881</xref>: 41.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content><named-content content-type="comment"> For complete taxonomic history see <xref rid="B151" ref-type="bibr">Wood and Bright (1992)</xref>.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Both sexes of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic> are distinguished by the rounded elytral apices, by the serrate and deeply emarginated elytral apex, and by the oblique ventrite 2.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>1.9–2.6 mm long (mean = 2.4 mm; n = 10); 2.1–2.7 times as long as wide. Color red-brown to dark red brown, antenna yellow-brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly, broadly emarginate; epistomal process absent; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing convex when viewed laterally, slightly transversely impressed just above epistoma; moderately, finely, longitudinally aciculate-punctate; aciculations converging at epistoma; punctures small, coarse; moderately, uniformly covered by long, fine, yellow erect hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, irregularly ovoid, setose with partial septum, two arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on basal half, narrowing to strongly rounded, moderately serrate apex; apex strongly emarginated at suture. Margin of apical edge bearing large, coarse punctures. Disc glabrous, smooth, shining; interstriae not impressed and equal in width to striae, interstrial punctures uniseriate, equal in size to those of striae, bearing moderately abundant short, semi-erect yellow-brown hair-like setae; striae not impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron less than half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 rounded, flush with base of ventrite 2. Ventrite 2 nearly at an oblique angle to ventrite 1; surface shagreened, dull finely punctate; punctures small, coarse, shallow; surface flattened, unarmed; setae abundant, erect, long, greater than length of segment 3; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 greater than combined lengths of ventrites 3 and 4; setal patch absent, median depression present.</p><fig id="F14" orientation="portrait" position="float"><label>Figure 14.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33367.jpg" xlink:href="zookeys-450-001-g014"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>2.0–3.1 mm long (mean = 2.5 mm; n = 10); 2.1–2.8 times as long as wide. Similar to male except frontal setae sparser, shorter, less than width of eye.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>102.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Syntypes: (ISNB, ZIFH). None examined.</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>CANADA:</bold><bold><italic>BRITISH COLUMBIA</italic>:</bold> Vancouver, 5.VII.1975, N.M. Downie (FMNH-2). <bold><italic>NOVA SCOTIA</italic>:</bold> Middleton, 11.VI.1938 (CNCI-2). <bold><italic>ONTARIO</italic>:</bold> Brimley, 17.VII.[19]21 (CNCI-1), 30.VII.[19]22 (CNCI-1). Ottawa, 21.VIII.1999, A.T. Howden, ex. wild grape [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Vitis</named-content></named-content></italic> sp.] (CNCI-1). Simcoe, 3.IX.1931, J.A. Hall (CNCI-2). Smith Falls, V.1940, H.S. Fleming (CNCI-1). <bold>UNITED STATES:</bold><bold><italic>ARIZONA</italic>:</bold> [<italic>Cochise Co.</italic>]: Green Canyon, 26.IX.[19]98, Sequeira and Jordal, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Fraxinus</named-content></named-content></italic> sp. (ZMBN-1). <bold><italic>CALIFORNIA</italic>:</bold> [<italic>Los Angeles Co.</italic>]: San Marino, 25.VI.[19]42, G.P. Mackenzie (FMNH-1). [<italic>Orange Co.</italic>]: Santa Ana, 1.IV.1943, L.R. Gillogly, ex. bait traps (USNM-1). [<italic>Placer Co.</italic>]: Penryn, 1.X.1929, A.T. McClay (FMNH-2). <bold><italic>DELAWARE</italic>:</bold> [<italic>Kent Co.</italic>]: Camdem, XII.1940, J.M. Amos (FMNH-1). <bold><italic>INDIANA</italic>:</bold><italic>Madison Co.</italic>: 10.VIII.1937, ex. apricot [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Prunus</named-content><named-content content-type="species">armeniaca</named-content></named-content></italic>] (FMNH-2). <italic>Porter Co.</italic>: Jackson Township, Maple Knoll Farm, 6.IV.[19]60, C.C. Gregg (FMNH-10). <italic>Tippecanoe Co.</italic>: 7.VI.1956, N.M. Downie (FMNH-2), 1.XI.1978 (FMNH-1). <bold><italic>MICHIGAN</italic>:</bold><italic>Allegan Co.</italic>: Fennville area, 30.VI.2003, P. McGhee, ex. on apple trees [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Malus</named-content></named-content></italic> sp.] (MSUC-1). [<italic>Cass Co.</italic>]: Edwardsburg, 30.VIII.1928 (MSUC-6). [<italic>Ingham Co.</italic>]: [East Lansing], Agriculture College [= Michigan State University campus], 1.I.1917 (MSUC-3). <italic>Isabella Co.</italic>: 16.IX.[19]55, R.R. Dreisbach (MSUC-1). [<italic>Kent Co.</italic>]: Grand Rapids, 17.X.[19]11 (MSUC-6). [<italic>Lenawee Co.</italic>]: Adrian, 20.VII.1900 (MSUC-7). <italic>Livingston Co.</italic>: Howell, 214 Inverness St, <named-content content-type="dwc:verbatimCoordinates">N42.61678°, W84.92810°</named-content>, 23.V-7.VI.2007, R. Mech, ex. Lindgren trap with EtOH + alpha (MSUC-1). <italic>Midland Co.</italic>: 24.VII.[19]46, R.R. Dreisbach (MSUC-1), 14.IX.[19]56 (MSUC-1). <italic>Oakland Co.</italic>: A.W. Andrews (MSUC-1); 28.VI.[19]47, B. Summerville (MSUC-1). [<italic>Washtenaw Co.</italic>]: Manchester, 21.V.1913 (MSUC-4), 31.V.1913 (MSUC-7). <bold><italic>NEW YORK</italic>:</bold><italic>Onondaga Co.</italic>: Syracuse, X.1987, R.J. Rabaglia, ex. mountain ash [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Sorbus</named-content></named-content></italic> sp.] (RJRC-1). [<italic>Tompkins Co.</italic>]: Ithaca, ex. peach [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Prunus</named-content></named-content></italic> sp.] (CASC-1). [<italic>Ulster Co.</italic>]: Oliverea, 20.VI.[19]18 (USNM-1). <italic>Wayne Co.</italic>: 26.VI.1950, Shumaker (USNM-1), 8.VIII.[19]51 (USNM-1). [<italic>Unspecified County</italic>]: (CASC-2). <bold><italic>PENNSYLVANIA</italic>:</bold><italic>Allegheny Co.</italic>: Upper St. Clair Township, 28.VIII.[19]59 (EMEC-1). [<italic>Philadelphia Co.</italic>]: Philadelphia, 20.V.[18]98 (USNM-1). <bold><italic>UTAH</italic>:</bold> [<italic>Utah Co.</italic>]: Vineyard, 6.IX.[19]23, T. Spalding (USNM-1). <bold><italic>WASHINGTON</italic>:</bold> [<italic>Whitman Co.</italic>]: Steptoe Butte State Park, 9.VII.1971, N.M. Downie (FMNH-1). <bold><italic>WASHINGTON, D.C.</italic>:</bold> 26.VII.1943, L.J. Bottimer, ex. on <italic><named-content content-type="taxon-name"><named-content content-type="genus">Prunus</named-content></named-content></italic> sp. (CNCI-2). <bold>ADDITIONAL SPECIMENS:</bold><italic>Brazil</italic> (MSUC-21), <italic>Czech Republic</italic> (MSUC-1), <italic>Hungary</italic> (MSUC-1), <italic>Morocco</italic> (MSUC-1) and <italic>Italy</italic> (MSUC-2).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: British Columbia, New Brunswick, Nova Scotia, Ontario, Prince Edward Island. Greenland. MEXICO: Durango. UNITED STATES: Arizona, Arkansas, California, Colorado, Connecticut, Delaware, Florida, Georgia, Idaho, Illinois, Indiana, Iowa, Kansas, Kentucky, Louisiana, Maryland, Massachusetts, Michigan, Mississippi, Missouri, Montana, New Jersey, New York, North Carolina, Ohio, Oregon, Pennsylvania, South Carolina, Texas, Utah, Virginia, Washington, Washington D.C, West Virginia, Wisconsin (Fig. <xref ref-type="fig" rid="F15">15</xref>).</p><fig id="F15" orientation="portrait" position="float"><label>Figure 15.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33368.jpg" xlink:href="zookeys-450-001-g015"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p>Stone and pome fruit trees including <italic><named-content content-type="taxon-name"><named-content content-type="genus">Malus</named-content></named-content></italic> spp. (apple), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pyrus</named-content></named-content></italic> spp. (pear), and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Prunus</named-content></named-content></italic> spp. (cherry) and is less common in <italic><named-content content-type="taxon-name"><named-content content-type="genus">Crataegus</named-content></named-content></italic> (hawthorn), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Sorbus</named-content></named-content></italic> (mountain ash) and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> spp. (elm).</p></sec><sec sec-type="treatment-Common name"><title>Common name.</title><p>Shothole borer.</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic> primarily attacks and kills small twigs and branches but may kill whole trees through the process of excavating adult galleries and larval feeding. Twig injury is the result of feeding activity at the base of the bud. Attacks can begin near an injury or on a healthy tree (<xref rid="B24" ref-type="bibr">Chamberlin 1939</xref>; <xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>).</p><p>The adult gallery is parallel to the grain of the wood and solely consists of an egg gallery; a nuptial chamber is not constructed. The adult gallery strongly scores the cambium, lightly scores the sapwood and ranges in size from 1.0–5.0 cm in length (Bright and Stark 1976; <xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>). Eggs are laid singly in larval niches on both sides of the egg gallery. Larval mines first radiate perpendicular to the egg gallery against the grain of the wood and later turn parallel to the grain (<xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>). The larval stage typically lasts one month. Pupation occurs in the bark for summer emergence. If the brood is unable to complete development during the warmer months, larvae will burrow 1.0–2.0 cm deep within the sapwood to overwinter. The following year, adults emerge and feed at twig crotches before selecting host material (<xref rid="B5" ref-type="bibr">Baker 1972</xref>). There are one to four generations per year depending on locality, with more generations occurring in warmer climates (<xref rid="B31" ref-type="bibr">Chittenden 1898</xref>; <xref rid="B5" ref-type="bibr">Baker 1972</xref>; <xref rid="B148" ref-type="bibr">Wood 1982</xref>).</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>This species is native to the Palearctic region and was first detected in North America in 1877 from New York (<xref rid="B31" ref-type="bibr">Chittenden 1898</xref>). The native range of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic> encompasses Europe, North Africa, Asia Minor and middle Asia to Zabaikalye (Russia) (<xref rid="B97" ref-type="bibr">Michalski 1973</xref>; <xref rid="B78" ref-type="bibr">Knížek 2011</xref>).</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></title><p><named-content content-type="taxon-authority">Semenov, 1902</named-content></p><p><xref ref-type="fig" rid="F16">Fig. 16</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content><named-content content-type="comment"> Semenov, 1902: 265.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content><named-content content-type="comment"> For complete taxonomic history see <xref rid="B151" ref-type="bibr">Wood and Bright (1992)</xref>.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>The <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> male most strongly resembles that of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic> because in both species the spine on the second ventrite never attains the apical margin. Both sexes can be distinguished from <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic> by the subapical carina on ventrite 5 located just before end of segment. Males are further differentiated from those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic> by the laterally compressed spine with a bulbous apex, which frequently has a longitudinal groove and by the banded appearance of the elytra. The female is further differentiated from that of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic> by the banded pattern of the elytra and by the low median carina (variable and may be absent) that does not touch either margin on the second ventrite.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>2.8–3.5 mm long (mean = 3.2 mm; n = 10); 2.0–2.2 times as long as wide. Color red-brown to dark red brown. Elytra of most specimens appears banded with a characteristic dark brown band on reddish colored elytra, but may also be solely dark brown or red-brown without a band. Antennae yellow-brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly, broadly emarginated; epistomal process absent; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing impressed when viewed laterally; moderately, coarsely reticulate punctate to weakly longitudinally aciculate; punctures dense, small, coarse; moderately covered by long, fine, yellow, erect, hair-like setae, these longer than width midpoint of eye, setae on lateral and dorsal margins longer, thicker, incurved. Antennal scape short, elongate; club flattened, thinner on apical half, irregularly ovoid, setose with partial septum and two arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger, coarser, deeper and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on basal half, narrowing to moderately rounded, smooth apex; apex weakly emarginated at suture. Margin of apical edge bearing small, fine punctures. Scutellar notch densely covered in recumbent white setae (may be abraded). Disc glabrous, smooth, shining; interstriae not impressed, equal in width to striae; interstrial punctures uniseriate, equal in size to those of striae, bearing sparse short, semi-erect yellow hair-like setae; striae weakly impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 flush with base of ventrite 2. Ventrite 2 nearly perpendicular to ventrite 1; surface shagreened, dull, finely punctate; punctures small, fine, shallow; surface convex; densely covered with semi-recumbent, long setae, as long as length of ventrite 3 or less; surface armed with median laterally compressed spine (rarely absent) that has its base close to basal margin but does not touch it; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 greater than combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F16" orientation="portrait" position="float"><label>Figure 16.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33369.jpg" xlink:href="zookeys-450-001-g016"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>2.3–3.6 mm long (mean = 3.2 mm; n = 10); 1.7–2.4 times as long as wide. Similar to male except epistoma feebly emarginated, frons flattened when viewed laterally, weakly longitudinally aciculate, setae sparser, shorter, less than width of eye; weakly transversely impressed just above epistoma and longitudinally impressed in median area. Apical margin of ventrite 1 weakly elevated above base of ventrite 2. Second ventrite armed with low median carina (variable and may be absent) that does not touch either margin.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>24.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype, male (location unknown).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>UNITED STATES:</bold><bold><italic>CALIFORNIA</italic>:</bold><italic>Los Angeles Co.</italic>: Whittier, Whittier Fertilizer, 7.IX.2006, A. Sanchez, ex. Lindgren funnel + ETOH & alpha pinene (DEBC-1). <bold><italic>COLORADO</italic>:</bold><italic>Adams Co.</italic>: Aurora, wood recycler, <named-content content-type="dwc:verbatimCoordinates">N39°807', W104°994'</named-content>, 9.VII.2003, P. McPherren, ex. funnel trap with Ips lure (MSUC-11), 15.IV.2003 (MSUC-1), 26.IV.2003 (MSUC-1), 2.V.2003 (MSUC-1), 29.V.2003 (MSUC-1). <bold><italic>MICHIGAN</italic>:</bold><italic>Ingham Co.</italic>: <named-content content-type="dwc:verbatimCoordinates">42.736501 -84.464670</named-content>, 258 m, 16.IX.2011, S.M. Smith, A.I. Cognato, I.A. Cognato, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> sp. (MSUC-1). <italic>Wayne Co.</italic>: Trenton-Woodhaven, 26.VII.2004, T. Dutton, ex. Lindgren funnel alpha-pinene Trap WY4 (MSUC-1). <bold><italic>UTAH</italic>:</bold><italic>Weber Co.</italic>: Ogden, Ogden Nature Center, 13.VII.2003, ex. funnel trap with alpha pinene & ethanol lure (MSUC-6).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: Alberta, British Columbia, Manitoba, Ontario, Saskatchewan. UNITED STATES: Arizona, California, Colorado, Connecticut, Delaware, Idaho, Illinois, Indiana, Kansas, Maryland, Michigan, Minnesota, Missouri, Montana, Nebraska, Nevada, New Jersey, New Mexico, Ohio, Oklahoma, Oregon, Pennsylvania, South Dakota, Texas, Utah, Virginia, Washington, Wyoming (Fig. <xref ref-type="fig" rid="F17">17</xref>).</p><fig id="F17" orientation="portrait" position="float"><label>Figure 17.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33370.jpg" xlink:href="zookeys-450-001-g017"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> spp. including <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content><named-content content-type="species">americana</named-content></named-content></italic> L. (American elm), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content><named-content content-type="species">pumila</named-content></named-content></italic> L. (Siberian elm), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content><named-content content-type="species">thomasii</named-content></named-content></italic> Sarg. (rock elm) and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content><named-content content-type="species">procera</named-content></named-content></italic> Salisb. (English elm).</p></sec><sec sec-type="treatment-Common name"><title>Common name.</title><p>Banded elm bark beetle.</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p>This species prefers to attack drought stressed elms (<xref rid="B99" ref-type="bibr">Negrón et al. 2005</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> locate a suitable host via host tree volatiles (<xref rid="B85" ref-type="bibr">Lee et al. 2010</xref>).</p><p>The adult gallery consists solely of a single egg gallery parallel with grain of the wood; a nuptial chamber is not constructed (<xref rid="B84" ref-type="bibr">Lee et al. 2006</xref>). The adult gallery <pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment>strongly scores the sapwood. Egg niches are constructed along the gallery and score the sapwood. Twenty to 120 eggs are laid along the egg galleries (<xref rid="B85" ref-type="bibr">Lee et al. 2010</xref>). Larval mines lightly score the sapwood and radiate perpendicular to the egg gallery. Larval galleries later meander often at an oblique angle to the grain of wood, forming a fan shaped pattern. Pupation occurs in the outer bark and broods overwinter as mature larvae or pupae (<xref rid="B84" ref-type="bibr">Lee et al. 2006</xref>). In California, flight occurs from April-September or October. In Nevada, Utah, Wyoming, Colorado, Kansas and Utah, flight occurs from May to September (<xref rid="B86" ref-type="bibr">Lee et al. 2011</xref>). There are two to three generations per year (<xref rid="B86" ref-type="bibr">Lee et al. 2011</xref>). Development from egg to adult takes 30–45 days (<xref rid="B99" ref-type="bibr">Negrón et al. 2005</xref>). Upon emergence, adults feed at twig crotches prior to selecting host material (<xref rid="B99" ref-type="bibr">Negrón et al. 2005</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> is a less effective vector of the Dutch elm disease fungus <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ophiostoma</named-content><named-content content-type="species">ulmi</named-content></named-content></italic> in North America, especially in Rocky Mountain states (<xref rid="B68" ref-type="bibr">Jacobi et al. 2013</xref>).</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>This species is native to the Palearctic where it occurs from western Russia and Uzbekistan and east to China, Mongolia and Korea (<xref rid="B97" ref-type="bibr">Michalski 1973</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> was first detected in North America in 2003. The first specimens were collected in Colorado and Utah. By 2005, it was recorded from 21 states suggesting that it had been present for many years before its initial detection (<xref rid="B99" ref-type="bibr">Negrón et al. 2005</xref>; <xref rid="B80" ref-type="bibr">LaBonte 2010</xref>). Subsequent review of bark beetle survey collections revealed the earlier occurrence of this species in Denver, Colorado in 1994 and Clovis, New Mexico in 1998 (<xref rid="B84" ref-type="bibr">Lee et al. 2006</xref>).</p><p>In areas where populations of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> are both present, the abundance of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> is decreasing to the point where this once common species is difficult to find (<xref rid="B99" ref-type="bibr">Negrón et al. 2005</xref>; <xref rid="B85" ref-type="bibr">Lee et al. 2010</xref>). This competitive displacement of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> by <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> is likely the result of differences in fecundity, generation time, and emergence. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> produces larger broods that may overwinter as pupae, have a quicker development period, have an earlier flight, and exhibit rapid, strong aggregation to host kairomones as compared to <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> (<xref rid="B85" ref-type="bibr">Lee et al. 2010</xref>).</p><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> is a highly morphologically variable species. <xref rid="B80" ref-type="bibr">LaBonte (2010)</xref> provides an excellent discussion regarding intraspecific variation exhibited within <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic>. There is considerable variation observed in the shape and appearance of spine on the second ventrite. The male second ventrite spine is typically well developed with a blunt apex that is broader than the base and appears triangular when laterally viewed. In females the spine is variously reduced and may even be absent. In most males, the spine is closest to the apical margin of the second ventrite. In some males the base of the spine is closer to basal margin. Most individuals exhibit the characteristic dark brown band on reddish colored elytra, which is the derivative of the species’.</p></sec><sec sec-type="treatment-Common name"><title>Common name.</title><p>Occasionally individuals have been encountered with unicolorous dark brown or reddish elytra. Additional variation was also observed in the coloration of the pronotum. The pronotum can vary from almost entirely dark brown with reddish coloration along the margins (common form) to the entire dorsal surface being reddish with dark brown margins (uncommon form) (<xref rid="B80" ref-type="bibr">LaBonte 2010</xref>).</p><pmc-comment>PageBreak</pmc-comment>
<pmc-comment>PageBreak</pmc-comment>
</sec></sec><sec sec-type="Native Hardwood clade"><title>Native Hardwood clade</title><p>The native hardwood clade (2.2–5.6 mm long) (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic>, and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic>) is monophyletic. The sexual dimorphism of the native hardwood clade differs from the introduced group and the conifer clade as both the males and females have an impressed frons that is strongly longitudinally aciculate, except <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> in which the males have a faintly longitudinally aciculate frons and the female frons is granulate punctate. This group arguably has the strongest degree of sexual dimorphism with <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic> each possessing a strongly excavated second ventrite numerous spines and teeth or a large setal patch on ventrite 5 respectively. The abdominal venter of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> is identical in both sexes.</p><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></title><p><named-content content-type="taxon-authority">Walsh, 1867</named-content></p><p><xref ref-type="fig" rid="F18">Fig. 18</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content><named-content content-type="comment"> Walsh, 1867: 58.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Both sexes of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> are distinguished from <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic> by the rugose-reticulate frons that is covered by uniformly distributed setae.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>3.3–5.5 mm long (mean = 4.45 mm; n = 20); 1.96–2.75 times as long as wide. Color dark red-brown to black, antenna light brown. Pronotum typically same color as elytra.</p><p><italic>Head.</italic> Epistoma weakly, broadly emarginated; epistomal process absent; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing flattened when viewed laterally, slightly transversely impressed just above epistoma and along median line; rugose-reticulate, strongly punctate; punctures small, coarse; moderately, uniformly covered by long, fine, yellow erect hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, thinner on apical half, irregularly ovoid, setose with partial septum, two arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctulate, punctures moderately abundant, larger, coarse and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to weakly rounded, smooth apex; apex weakly emarginated at suture. Margin of apical edge bearing small, fine punctures. Disc smooth, shining; interstriae not impressed, more than twice width of striae, punctures uniseriate, smaller than those of striae, punctures bearing short recumbent yellow setae slightly longer than size of a puncture (may be abraded); striae moderately impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 weakly elevated above base of ventrite 2. Ventrite 2 nearly perpendicular to ventrite 1; surface smooth, shining, finely punctate; punctures large, fine, shallow; surface flattened; setae semi-recumbent, short, about half length of segment 3 or less; apical margin unarmed; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 unarmed; length of ventrite 5 greater than combined lengths of ventrites 3 and 4; setal patch absent, median depression present.</p><fig id="F18" orientation="portrait" position="float"><label>Figure 18.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33371.jpg" xlink:href="zookeys-450-001-g018"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>3.9–5.6 mm long (mean = 4.65 mm; n = 20); 1.9–2.8 times as long as wide. Similar to male except epistoma feebly emarginated, epistomal process absent, frons convex when viewed laterally, weakly longitudinally strigate-punctate, setae shorter, less than width of eye and sparse; weakly transversely impressed just above epistoma and between inner apices of eyes. Second ventrite unarmed.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>74.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Neotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> Walsh: male, labeled “Galesburg, Illinois, Liebeck Collection” (MCZC). Neotype designated <xref rid="B121" ref-type="bibr">Smith and Cognato 2010b</xref>: 36.</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>CANADA:</bold><bold><italic>ONTARIO</italic>:</bold> Point Pelee National Park, 27.IX.1989, K. Dunster, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content><named-content content-type="species">tenuifolia</named-content></named-content></italic> (CNCI-9). <bold>UNITED STATES:</bold><bold><italic>MISSISSIPPI</italic>:</bold> [<italic>Sharkey Co.</italic>]: Rolling Fork, VIII.1976, J.D. Solomon, ex. Nuttall oak [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Quercus</named-content></named-content></italic> sp.] (USNM-1). <bold><italic>PENNSYLVANIA</italic>:</bold><italic>Cumberland Co.</italic>: Roadway Dr @ Schneider Dr, <named-content content-type="dwc:verbatimCoordinates">40.229030°N, 77.111580°W</named-content>, 26.VI.2009, L.R. Donovall, (MSUC-25), 29.V.2009, ex. Lindgren-alpha pinene/EtOH (MSUC-4), 29.V.2009, ex. Lindgren-EtOH (MSUC-1). <italic>Dauphin Co.</italic>: Wildwood on Industrial Rd, <named-content content-type="dwc:verbatimCoordinates">40.316325°N, 76.888783°W</named-content>, 6.VIII.2009, S.-E. Spichiger, ex. Lindgren-EtoH (MSUC-2). <italic>Lancaster Co.</italic>: 7031 Elizabethtown Rd, <named-content content-type="dwc:verbatimCoordinates">40.182583°N, 76.498783°W</named-content>, 23.VII.2009, ex. Lindgren-BEBB/EtOH (MSUC-1). <italic>York Co.</italic>: 400 Mundis Rd, <named-content content-type="dwc:verbatimCoordinates">40.030170°N, 76.705330°W</named-content>, 10.VI.2009, S. Rebert, ex. Lindgren-alpha/EtOH (MSUC-4). <bold><italic>TEXAS</italic>:</bold> [<italic>Colorado Co.</italic>]: Columbus, [18]88 (MSUC-4, USNM-15). [<italic>Unspecified county</italic>]: (USNM-3). Fort Worth, 31.VII.[19]12, ex. bred from hackberry [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content></named-content></italic> sp.] (USNM-4).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: Ontario. UNITED STATES: Illinois, Kansas, Mississippi, Ohio, Pennsylvania, Texas (Fig. <xref ref-type="fig" rid="F19">19</xref>).</p><fig id="F19" orientation="portrait" position="float"><label>Figure 19.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33372.jpg" xlink:href="zookeys-450-001-g019"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> L. (common hackberry), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content><named-content content-type="species">tenuifolia</named-content></named-content></italic> Nutt. (dwarf hackberry), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content><named-content content-type="species">laevigata</named-content></named-content></italic> Willd. var. <italic>texana</italic> Sarg. (Texan sugarberry), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Fagus</named-content><named-content content-type="species">grandifolia</named-content></named-content></italic> Ehrh. (American beech), and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Quercus</named-content></named-content></italic> spp. (oak).</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p>The gallery has been reported as very confused and undecipherable. Larvae were reported boring in the wood. In addition, the species did not appear to colonize healthy trees (<xref rid="B101" ref-type="bibr">Packard 1890</xref>). <xref rid="B120" ref-type="bibr">Smith and Cognato (2010a)</xref> summarized all information known about this species.</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>This species is very rarely encountered. Most museum collections occurred in the early 1900’s and most recent collections have been from Lindgren funnel traps. There are many gaps in the known distribution of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> but the species likely occurs throughout the eastern US associated with its host trees.</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></title><p><named-content content-type="taxon-authority">Say, 1824</named-content></p><p><xref ref-type="fig" rid="F20">Fig. 20</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content><named-content content-type="comment"> Say, 1824: 182.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic> are differentiated from other <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species by the presence of long, erect hair-like setae on the elytral interstriae and on the abdominal venter. The male is further distinguished by a pair of strongly elevated areas on the basal two-thirds of ventrite 5, each densely covered with abundant fine, long hair-like setae.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>2.2–5.3 mm long (mean = 3.65 mm; n = 20); 1.75–2.6 times as long as wide. Head, pronotum, legs and abdominal venter dark red-brown, antennae yellow-brown, elytra usually dark red-brown but occasionally red-brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma moderately, broadly emarginated; epistomal process absent; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing strongly flattened when viewed laterally; densely, finely, longitudinally aciculate; aciculations converging at epistoma; impunctate; setae on lateral and dorsal margins covered by long, thick, incurved, yellow erect hair-like setae, these longer than width of midpoint of eye, median areas covered with sparse, shorter and thinner setae. Antennal scape short, elongate; club flattened, irregularly ovoid, setose with partial septum, three very sharply arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; smooth, shining, punctures on disc fine, shallow, moderately abundant, larger, coarser and dense laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, smooth apex; apex entire at suture. Margin of apical edge bearing small, fine punctures. Disc reticulate, shining; interstriae faintly impressed, equal to width of striae, interstrial punctures large, uniseriate, equal in size to those of striae, bearing moderately abundant, long, semi-erect, yellow hair-like setae (may be abraded); striae moderately impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 weakly elevated above base of ventrite 2. Ventrite 2 nearly perpendicular to ventrite 1; surface reticulate, shining, densely, coarsely punctured; punctures large, coarse; surface flattened; setae moderately abundant, longer than length of ventrite 3; apical margin unarmed; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 unarmed; length of ventrite 5 greater than combined lengths of ventrites 3 and 4; median depression absent; pair of strongly elevated areas on basal two-thirds, each densely covered with abundant fine, long hair-like setae; apical third strongly impressed.</p><fig id="F20" orientation="portrait" position="float"><label>Figure 20.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33373.jpg" xlink:href="zookeys-450-001-g020"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>3.1–5.1 mm long (mean = 4.0 mm; n = 20); 1.96–2.45 times as long as wide. Similar to male except epistoma feebly emarginated, epistomal process absent, frons less strongly flattened when viewed laterally, finely, narrowly longitudinally aciculate, setae shorter, less than width of eye and less abundant; weakly transversely impressed between inner apices of eyes. Second ventrite unarmed. Ventrite 5 without a pair of strongly elevated areas on basal two-thirds or setal patches.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>214.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: male, Missouri (ANSP, lost).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>CANADA:</bold><bold><italic>ONTARIO</italic>:</bold> Pelee Island, 29.VI.1940, W.J. Brown (CNCI-1). Point Pelee National Park, 23.VI.[19]31, W.J. Brown (CNCI-1); 27.IX.1989, K. Dunster, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content><named-content content-type="species">tenuifolia</named-content></named-content></italic> (CNCI-1), 2.VIII.1990 (CNCI-1). <bold>UNITED STATES:</bold><bold><italic>COLORADO</italic>:</bold><italic>Jefferson Co.</italic>: Evergreen, Upper Bear Creek Rd, 11.IX.1980, D. Leatherman, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CSUC-1). <bold><italic>ILLINOIS</italic>:</bold><italic>Alexander Co.</italic>: Pleasant Valley, 15.VI.1979, B.C. Weber, ex. Trap 6, Ht. 2 (USNM-1), 22.VI.1979, ex. Trap 14, Ht. 4 (USNM-1). <italic>Champaign Co.</italic>: 28.X.1960, ex. hackberry [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content></named-content></italic> sp.] (EMEC-4). <italic>Sangamon Co.</italic>: Springfield, 29.V.2003, C. Helm (CUIC-1). <italic>Stephenson Co.</italic>: Freeport, 4.VII.[19]17 (USNM-1). <bold><italic>INDIANA</italic>:</bold><italic>Tippecanoe Co.</italic>: 6.VI.1971, N.M. Downie, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (USNM-6). <bold><italic>IOWA</italic>:</bold> [<italic>Story Co.</italic>]: Ames, 18.VIII.1926, H.H. Harris (DEBC-2), 1.IV.1936 (DEBC-1), 22.V.1939, E. Polderboer (DEBC-1); 22.V.1939, C. Vocom (USNM-2); E. Snead (USNM-1). <bold><italic>KANSAS</italic>:</bold><italic>Douglas Co.</italic>: Lawrence, 5 mi N.E., Kansas University Natural History Research Station, 9.VII.1982, D.H. Wahl (CNCI-1). Lawrence, 10.VI.[19]20, M.W. Blackman (USNM-8); 5.IX.1950, S.L. Wood (USNM-4). <italic>Riley Co.</italic>: G.A. Dean (DEBC-1); Popenoe (DEBC-1). [<italic>Unspecified County</italic>]: Manhattan, 1.VI.[19]29, F. Kruger (DEBC-1); 5.VI.[19]29, T.N. Winburn (DEBC-1); 8.V.1968, G. Hevel (USNM-1). <bold><italic>KENTUCKY</italic>:</bold><italic>Christian Co.</italic>: 15.VI.1960, J.M. Campbell (CNCI-1). <bold><italic>LOUISANA</italic>:</bold><italic>Jefferson Parish</italic>: New Orleans, 4.XII.1975, emerged II.1976, S.G. Wellso, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content><named-content content-type="species">laevigata</named-content></named-content></italic> wood (MSUC-13). <italic>Saint Bernard Parish</italic>: J.N. Knull, ex. reared from <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content><named-content content-type="species">mississippiensis</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content><named-content content-type="species">tenuifolia</named-content></named-content></italic>] (USNM-3). <bold><italic>MARYLAND</italic>:</bold> [<italic>Cecil Co.</italic>]: Port Deposit, 26.VI.1977, D. Jump (USNM-1), 13.VII.1977 (USNM-1), 9.VI.1979 (USNM-2), 3.VII.1979 (USNM-1). <italic>Montgomery Co.</italic>: Bethesda, 25.V.1981, W.E. Steiner (USNM-1). Plummers Island, II-III.1912, E.A. Schwarz, ex. branch of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content></named-content></italic> sp. (USNM-30). <bold><italic>MICHIGAN</italic>:</bold><italic>Ingham Co.</italic>: East Lansing, Agriculture College [Michigan State University] (MSUC-8); S21 T4N R1W, VII.1970, S.G. Wellso (MSUC-12). East Lansing, IV.1972, D.K. Young (MSUC-4); 15.VI.1970, S.G. Wellso, ex. emerged from <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (MSUC-7). Okemos, 12.VIII.1969, S.G. Wellso (MSUC-1), 5.VI.1976 (MSUC-1); 28.V.1970, S.G. Wellso, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (MSUC-2), 10.VIII.1969 (MSUC-1). <italic>Kalamazoo Co.</italic>: Gourdneck Lake State Game Area, 19.VI.2011, A.I. Cognato, ex. funnel trap with EtOH (MSUC-1). <italic>Wayne Co.</italic>: Grosse Ile, Pke [sic! = Parke] lane, <named-content content-type="dwc:verbatimCoordinates">N42.17060°, W84.14496°</named-content>, 23.V–7.VI.2007, R. Mech, ex. Lindgren trap with ipslure (MSUC-1), ex. Lindgren trap with EtOH + alpha (MSUC-1), 20.VI-6.VII.2007, ex. Lindgren trap with ipslure (MSUC-1), 7-20.VI.2007 (MSUC-5). <bold><italic>MISSISSIPPI</italic>:</bold> [<italic>Unspecified County</italic>]: 19.V.1920, M.W. Blackman (CNCI-1). <bold><italic>MISSOURI</italic>:</bold> [<italic>Boone Co.</italic>]: Columbia, 24.IV.1954, P.J. Spangler (USNM-1). <bold><italic>NEBRASKA</italic>:</bold><italic>Dixon Co.</italic>: Ponca, Ponca State Park, <named-content content-type="dwc:verbatimCoordinates">42.607161°N, -96.73223°W</named-content>, 23.VIII.2007, T.P. Miller, ex. funnel trap with PSB alpha-pinene (MSUC-1). <italic>Knox Co.</italic>: Crofton, Lewis and Clarke Lake, <named-content content-type="dwc:verbatimCoordinates">42.8321983°N -97.575555°W</named-content>, 5.IX.2007, T.P. Miller, ex. funnel trap with Sirex (MSUC-1). [<italic>Lancaster Co.</italic>]: Lincoln, 29.VIII.[19]53, R. Roselle, ex. elm [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Ulmus</named-content></named-content></italic> sp.] (USNM-1). <italic>Sarpy Co.</italic>: Fontenelle Forest, <named-content content-type="dwc:verbatimCoordinates">41.171478°N -95.9068166°W</named-content>, 30.V.2006, N. Haxton, ex. funnel trap with ethanol lure (MSUC-1), 7.VII.2006 (MSUC-1). <bold><italic>NEW JERSEY</italic>:</bold> [<italic>Burlington Co.</italic>]: Riverton, 12.II.1934, Wadley, ex. on Hackberry [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content></named-content></italic> sp.] (USNM-1). [<italic>Essex Co.</italic>]: Newark (USNM-1). [<italic>Somerset Co.</italic>]: North Branch, 9.III.1937, C.H. Hoffman, ex. from gallery made in honey locust [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Gleditsia</named-content><named-content content-type="species">triacanthos</named-content></named-content></italic>] (USNM-1). <bold><italic>NEW YORK</italic>:</bold> [<italic>Tompkins Co.</italic>]: Ithaca, 18.VI.[19]59, ex. in <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (CUIC-11). <bold><italic>NORTH CAROLINA</italic>:</bold><italic>Mecklenburg Co.</italic>: Charlotte, 15-20.VI.2006, J.F. Cornell, ex. FIT in vacant lot (MSUC-1). <bold><italic>NORTH DAKOTA</italic>:</bold> [<italic>Ransom Co.</italic>]: Mcleod, 5 mi N.W., 4.VII.1968, H.F. Howden (CNCI-1). <bold><italic>OHIO</italic>:</bold> [<italic>Franklin Co.</italic>]: Columbus (USNM-1); 11.V.[19]14 (DEBC-2). [<italic>Hamilton Co.</italic>]: Cincinnati, 6.VI.[?] (USNM-1), 24.VI.[?] (USNM-1), 9.VII.[?] (USNM-1). <bold><italic>OKLAHOMA</italic>:</bold><italic>Latimer Co.</italic>: V.1982, K. Stephan (USNM-2). <bold><italic>PENNSYLVANIA</italic>:</bold> [<italic>Allegheny Co.</italic>]: Allegheny [= Pittsburgh], 7.VIII.[18]95 (CUIC-12). [<italic>Dauphin Co.</italic>]: Harrisburg, T.H. Hubbell (CUIC-1). Hummelstown, 26.VI.[19]37, J.N. Knull (CUIC-6). <bold><italic>TEXAS</italic>:</bold><italic>Brazos Co.</italic>: College Station, 13.IV.1964, S.G. Wellso (MSUC-1), 26.IV.1964 (MSUC-1). [<italic>Unspecified County</italic>]: Dallas, Hopk. U.S. 9929-X, F.C. Bishop, ex. hackberry [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content></named-content></italic> sp.] (USNM-5). <bold><italic>VIRGINIA</italic>:</bold><italic>Clarke Co.</italic>: Boyne, 2 mi S., U[niversity of] Virginia Blandy Experimental Farm, 8-18.VI.1990, D.R. Smith, ex. malaise trap (USNM-1), 19-30.VI.1990 (USNM-1), 1-12.VII.1990 (USNM-1). <italic>Essex Co.</italic>: Dunnsville, 1 mi S.E., <named-content content-type="dwc:verbatimCoordinates">37°52’N, 76°48’W</named-content>, 29.V-9.VI.1993, D.R. Smith, ex. malaise trap (USNM-1); 24.VI-9.VII.1992 (USNM-1). <italic>Fairfax Co.</italic>: Mount Vernon, 27.VI.1915, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Cornus</named-content><named-content content-type="species">stricta</named-content></named-content></italic> (USNM-1). [<italic>Unspecified County</italic>]: Falls Church, 4.III.1921, E.A. Chapin (USNM-1). <bold><italic>WASHINGTON, D.C.</italic>:</bold> 21.V.1908 (MSUC-5). <bold><italic>WEST VIRGINIA</italic>:</bold><italic>Morgan Co.</italic>: near Great Cacapon, 25.V.1985, G.F. & J.F. Hevel (USNM-2). <italic>Wood Co.</italic>: Hopk. U.S. 6675, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content></named-content></italic> (USNM-1).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: Ontario. UNITED STATES: Colorado, Florida, Iowa, Illinois, Indiana, Kansas, Kentucky, Louisiana, Maryland, Michigan, Mississippi, Missouri, Nebraska, New Jersey, New York, North Carolina, North Dakota, Ohio, Oklahoma, Pennsylvania, South Carolina, Tennessee, Texas, Virginia, Washington, D.C., West Virginia (Fig. <xref ref-type="fig" rid="F21">21</xref>).</p><fig id="F21" orientation="portrait" position="float"><label>Figure 21.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33374.jpg" xlink:href="zookeys-450-001-g021"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> L. (common hackberry), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content><named-content content-type="species">tenuifolia</named-content></named-content></italic> Nutt. (dwarf hackberry), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Celtis</named-content><named-content content-type="species">laevigata</named-content></named-content></italic> Willd. (sugar hackberry) and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Gleditsia</named-content><named-content content-type="species">triacanthos</named-content></named-content></italic> L. (honeylocust).</p></sec><sec sec-type="treatment-Common name"><title>Common name.</title><p>Hackberry engraver.</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic> prefers to colonize dead, dying or felled trees and the broken, dead and dying branches of live hackberry (<xref rid="B9" ref-type="bibr">Blackman 1922</xref>; <xref rid="B40" ref-type="bibr">Doane et al. 1936</xref>; <xref rid="B17" ref-type="bibr">Bright 1976</xref>).</p><p>The biology of this species is poorly understood. Adult galleries are parallel to the grain of the wood and consist of a single egg gallery; a nuptial chamber is not constructed. The adult gallery strongly scores the sapwood and ranges in size from 2.5–5.0 cm in length (<xref rid="B9" ref-type="bibr">Blackman 1922</xref>). Larval mines first radiate perpendicular to the egg <pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment>gallery and later meander, giving the galleries a tortuous appearance (<xref rid="B9" ref-type="bibr">Blackman 1922</xref>; <xref rid="B17" ref-type="bibr">Bright 1976</xref>). Larvae construct pupal chambers in burrows 2.0–3.0 cm deep within the sapwood (<xref rid="B17" ref-type="bibr">Bright 1976</xref>; Smith and Cognato, pers. obs.). There are two generations per year in the southeastern US and one generation per year in the northeastern US (<xref rid="B9" ref-type="bibr">Blackman 1922</xref>; <xref rid="B5" ref-type="bibr">Baker 1972</xref>).</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>The holotype of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic> has been lost (<xref rid="B148" ref-type="bibr">Wood 1982</xref>), however, <xref rid="B112" ref-type="bibr">Say’s (1824)</xref> description is unambiguous as to the characteristics of this species.</p><p><xref rid="B148" ref-type="bibr">Wood (1982)</xref> reports <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic> as having a transverse gallery that is perpendicular to the grain of the wood. Other authors, including <xref rid="B9" ref-type="bibr">Blackman (1922)</xref>, <xref rid="B5" ref-type="bibr">Baker (1972)</xref>, and <xref rid="B17" ref-type="bibr">Bright (1976)</xref> report this species as having a longitudinal gallery that is parallel to the grain of the wood. After examining the literature and photographs provided by T.H. Atkinson, it appears that Wood erred in his description and the gallery is parallel to the grain of the wood.</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></title><p><named-content content-type="taxon-authority">Say, 1824</named-content></p><p><xref ref-type="fig" rid="F22">Fig. 22</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content><named-content content-type="comment"> Say, 1824: 182.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">carya</named-content></named-content><named-content content-type="comment"> Riley, 1867: 68. <xref rid="B83" ref-type="bibr">LeConte 1876</xref>: 371.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">caryae</named-content></named-content><named-content content-type="comment"> Walsh, 1867: 58. <xref rid="B83" ref-type="bibr">LeConte 1876</xref>: 371.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>The <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic> male is easily distinguished by the autapomorphic features of the abdominal venter which include: the apical margin of ventrite 3 armed by three acute spines (two lateral and one medial), the apical margin of ventrite 4 armed by one median tooth, ventrite 1 apically descending, ventrite 2 deeply concave, with the basal margin produced and bearing a median tubercle. The female is distinguished by the flattened and moderately, finely longitudinally aciculate frons, bearing long, fine, incurved setae on the lateral and dorsal margins.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>2.8–4.8 mm long (mean = 4.0 mm; n = 20); 1.8–2.25 times as long as wide. Color red-brown to dark red-brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly emarginated; epistomal process strongly produced, moderately elevated, smooth, shining; median area above mandibles bearing a dense patch of long, yellow, hair-like setae. Frons appearing strongly flattened when viewed laterally; strongly, densely, coarsely, aciculate-punctate; aciculations converging at epistoma; punctures small, coarse; surface moderately covered by long, fine, yellow, erect hair-like setae, these longer than width of midpoint of eye, setae on lateral and dorsal margins longer, thicker, incurved. Antennal scape short, elongate; club flattened, irregularly ovoid, setose with partial septum, two arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, golden setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to weakly rounded, serrate apex; apex entire at suture. Margin of apical edge bearing large, coarse punctures.. Disc smooth, shining; interstriae weakly impressed, twice width of striae, punctures large, uniseriate, smaller than those of striae, interstrial punctures bearing sparse, long, semi-erect yellow hair-like setae (may be abraded); striae moderately impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 descending, strongly, acutely produced; ventrite 2 deeply concave, basal margin produced, bearing median tubercle. Ventrite 2 nearly perpendicular to ventrite 1; surface shagreened, dull, finely, obscurely punctate with small, fine, shallow punctures; apical margin armed with broad median denticle, occasionally absent. Apical margin of ventrite 3 armed by three acute spines (two lateral and one medial); apical margin of ventrite 4 armed by one median tooth. Ventrite 5 carinate ridge closer to basal margin of segment; length of ventrite 5 less than combined lengths of ventrites 3 and 4; median depression absent, apical half of segment pubescent.</p><fig id="F22" orientation="portrait" position="float"><label>Figure 22.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33375.jpg" xlink:href="zookeys-450-001-g022"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>2.9–5.0 mm long (mean = 3.9 mm; n = 20); 1.8–2.4 times as long as wide. Similar to male except epistoma feebly emarginated, frons less strongly flattened when viewed laterally, moderately and finely aciculate, setae shorter, less than width of eye and uniformly distributed. Apical margin of ventrite 1 weakly elevated above base of ventrite 2. Ventrite 2 surface flattened, nearly perpendicular to ventrite 1; setae erect, short, about half length of segment 3. Venter unarmed.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>143.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic> Say: male, Missouri (ANSP, lost). Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">carya</named-content></named-content></italic> Riley, location unknown. Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">caryae</named-content></named-content></italic> Walsh, location unknown.</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>CANADA:</bold><bold><italic>ONTARIO</italic>:</bold> Queenston, 15.VII.1938, D.F. Patterson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Carya</named-content><named-content content-type="species">ovata</named-content></named-content></italic> (CNCI-2); Z17 E464 N4820, 20.VII.2004, Seaforth, ex. handpicked from eastern white pine [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content><named-content content-type="species">strobus</named-content></named-content></italic>], 04-5-0265 (CNCI-1). <bold>UNITED STATES:</bold><bold><italic>GEORGIA</italic>:</bold> 1.IX.[19]45 (CASC-4). <bold><italic>ILLINOIS</italic>:</bold> [<italic>Cook Co.</italic>]: Edgebrook (CNCI-2). [<italic>Unspecified County</italic>]: Willow Springs, 21.VIII.[19]04 (FMNH-1), 13.V.[19]05, A.B Wolcott (FMNH-4). <bold><italic>KANSAS</italic>:</bold> [<italic>Shawnee Co.</italic>]: Topeka, 16.VI.[?], Popenoe (USNM-1). <bold><italic>MASSACHUSSETS</italic>:</bold> [<italic>Unspecified County</italic>]: (CASC-1). <bold><italic>MICHIGAN</italic>:</bold> [<italic>Ingham Co.</italic>]: East Lansing, 15.VII.1932, (MSUC-4). <italic>Wayne Co.</italic>: Detroit, 18.VIII.1902 (MSUC-3). [<italic>Unspecified County</italic>]: Lansing, 9.IX.1929 (MSUC-36). <bold><italic>MISSOURI</italic>:</bold><italic>Dent Co.</italic>: 28.VIII.1973, M.P. Rolling (USNM-1), 31.VIII.1973 (USNM-3). [<italic>Unspecified County</italic>]: (FMNH-1). <bold><italic>MINNESOTA</italic>:</bold> [<italic>Sherburne Co.</italic>]: Elk River, 8.VII.1959, E.J. Kingsley (CNCI-2). <bold><italic>Mississippi</italic>:</bold> [<italic>Madison Co.</italic>]: Canton, 16.VI.[19]04 (CUIC-1). [<italic>Oktibbeha Co.</italic>]: [Starkville], Agriculture College of Mississippi [= Mississippi State University], 15.IV.1922, F.M. Hull (CUIC-1). <bold><italic>New jersey</italic>:</bold> [<italic>Middlesex Co.</italic>]: Dunellen (CUIC-1). <bold><italic>NEW YORK</italic>:</bold><italic>Onondaga Co.</italic>: 10.VI.1942, N.M. Downie (FMNH-2), 14.VII.1946 (FMNH-1). Syracuse, C.J. Drake, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Hickoria</named-content><named-content content-type="species">glabra</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Carya</named-content><named-content content-type="species">glabra</named-content></named-content></italic>] (USNM-1). [<italic>Orange Co.</italic>]: Highland Falls, 20.VI.1920, F. Schott (CUIC-1). Middletown (CUIC-1). [<italic>Queens Co.</italic>]: Long Island Aqueduct, 14.VII.1912 (MSUC-4). [<italic>Tompkins Co.</italic>]: Groton, 7.VII.1946, N.M. Downie (FMNH-1). Ithaca, 4.VIII.1928, P.P. Babiy (CUIC-1). [<italic>Westchester Co.</italic>]: Mount Vernon, VII.1913, ex. from hickory [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Carya</named-content></named-content></italic> sp.] (CASC-14). Yonkers, 28.V.1935, P.A. Readio, H. Dietrich, ex. taken on air trap (CUIC-1). [<italic>Unspecified County</italic>]: New York City, 15.V.1912, (CNCI-1), 13.V.1912 (CUIC-1). <bold><italic>north carolina</italic>:</bold> [<italic>Buncombe Co.</italic>]: Asheville, Bent Creek, 17.VI.[19]29 (FMNH-1). <bold><italic>Pennsylvania</italic>:</bold> [<italic>Allegheny Co.</italic>]: Allegheny [= Pittsburgh] (FMNH-3), 24.VI.[18]93 (CUIC-1). [<italic>Armstrong Co.</italic>]: Ford City, 28.VIII.[19]11 (USNM-1). [<italic>Blair Co.</italic>]: Tyrone, VII.[19]12, lot 367, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Fraxinus</named-content><named-content content-type="species">alba</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Fraxinus</named-content><named-content content-type="species">americana</named-content></named-content></italic>] (CASC- 7). <italic>Cumberland Co.</italic>: Roadway Dr @ Schneider Dr, <named-content content-type="dwc:verbatimCoordinates">40.229030°N, -77.111580°W</named-content>, 26.VI.2009, L.R. Donovall (MSUC-1). [<italic>Dauphin Co.</italic>]: Harrisburg, 19.IV.1911 (CASC-1), VI.1911 (CASC-3); 1.III.[19]11, emerged 7.IV.[19]11, Champlain (CASC-12); Hopk. U.S. 10935-E, 13-14.VII.[?], W.S. Fisher, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Hicoria</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Carya</named-content></named-content></italic> sp.] (CUIC-1). Hummelstown, 20.VI.[19]15 (CUIC-1). Linglestown, 8.VI.1912, W.S. Fisher (CNCI-1, CUIC-2). [<italic>Philadelphia Co.</italic>]: Angora [= Philadelphia], IX.[19]15, H.A. Kaeber, ex. Hickory bark [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Carya</named-content></named-content></italic> sp.] (USNM-7). [<italic>Westmoreland Co.</italic>]: Jeannette, H.G. Klages (CASC-2). <bold><italic>TENNESSEE</italic>:</bold> [<italic>Hamilton Co.</italic>]: Chattanooga, 2.VI.[19]19, Leach (FMNH-1).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: Ontario, Quebec. UNITED STATES: Alabama, Connecticut, Delaware, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky, Louisiana, Maryland, Massachussets, Michigan, Minnesota, Mississippi, Missouri, New Jersey, New York, North Carolina, Ohio, Pennsylvania, South Carolina, Ohio, Pennsylvania, South Carolina, Tennessee, Texas, Virginia, West Virginia, Washington, D.C., Wisconsin (Fig. <xref ref-type="fig" rid="F23">23</xref>).</p><fig id="F23" orientation="portrait" position="float"><label>Figure 23.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33376.jpg" xlink:href="zookeys-450-001-g023"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p>Primary hosts: <italic><named-content content-type="taxon-name"><named-content content-type="genus">Carya</named-content></named-content></italic> spp. (hickory) including <italic><named-content content-type="taxon-name"><named-content content-type="genus">Carya</named-content><named-content content-type="species">illinoinensis</named-content></named-content></italic> (Wangenh.) K. Koch (pecan). Incidental hosts: <italic><named-content content-type="taxon-name"><named-content content-type="genus">Juglans</named-content><named-content content-type="species">cinerea</named-content></named-content></italic> L. (butternut).</p></sec><sec sec-type="treatment-Common name"><title>Common name.</title><p>Hickory bark beetle.</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic> is one of the most destructive pests of hardwoods in the US and the most important pest of hickory (<xref rid="B40" ref-type="bibr">Doane et al. 1936</xref>; <xref rid="B5" ref-type="bibr">Baker 1972</xref>). The species generally attacks and kills single trees, or solely treetops. However, outbreaks can develop during periods of drought, killing large tracts of hickory. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic> kills its host by mass attack in which a multitude of broods develop in the phloem and cambium, effectively girdling the host tree (<xref rid="B9" ref-type="bibr">Blackman 1922</xref>).</p><p>Adult galleries are parallel to the grain of the wood and deeply score the sapwood; a nuptial chamber is not constructed. The adult gallery is short (2.5–5.0 cm), and consists of a single egg gallery (<xref rid="B9" ref-type="bibr">Blackman 1922</xref>). Eggs are deposited singly in niches on each side of the egg gallery with 20–60 niches per gallery (<xref rid="B9" ref-type="bibr">Blackman 1922</xref>). After the eggs have been laid, the female constructs a postovipositional feeding tunnel parallel to the egg gallery (<xref rid="B61" ref-type="bibr">Goeden and Norris 1965b</xref>). Larval mines are excavated in the cambium. From the egg gallery, the larval mines are first perpendicular to the grain of the wood and then gradually turn and diverge creating a fan shaped appearance. The larvae bore into the inner bark to overwinter. Larvae pupate the following spring and emerge as adults the following summer (<xref rid="B9" ref-type="bibr">Blackman 1922</xref>). Upon emergence, adults feed at twig crotches <pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment>and leaf petioles before selecting a host (<xref rid="B5" ref-type="bibr">Baker 1972</xref>; <xref rid="B59" ref-type="bibr">Goeden and Norris 1964b</xref>). There is one generation per year in the north with larvae completing their development in March and April and emergence in May. There may be two generations per year in the south with the brood overwintering as larvae (<xref rid="B40" ref-type="bibr">Doane et al. 1936</xref>). See <xref rid="B58" ref-type="bibr">Goeden and Norris (1964a</xref>,<xref rid="B59" ref-type="bibr">b</xref>, <xref rid="B60" ref-type="bibr">1965a</xref>,<xref rid="B61" ref-type="bibr">b</xref>) for more information regarding the biology of this species.</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>The holotype of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic> is lost (<xref rid="B148" ref-type="bibr">Wood 1982</xref>), however <xref rid="B112" ref-type="bibr">Say’s (1824)</xref> description is unambiguous as to the characteristics of this species. The holotype of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">caryae</named-content></named-content></italic> Walsh was likely deposited in Walsh’s type collection, which was housed in the Chicago Academy of Science Museum. This collection burned in the Great Chicago Fire and the holotypes were lost (<xref rid="B118" ref-type="bibr">Sheppard 2004</xref>; J. Colby, pers. comm.).</p></sec></sec></sec><sec sec-type="Conifer clade"><title>Conifer clade</title><p>The conifer clade (1.7–5.9 mm) (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic>, and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic>) is monophyletic. The sexual dimorphism exhibited by species in the clade follows the general pattern outlined above except for the shape of the frons, which is quite variable. In some species, the frons is convex in both sexes with the female frons more strongly convex than that of the male (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic>, and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> males have a flattened and impressed frons when viewed from a lateral profile and that of the female is convex. In the remaining species, the male frons appears flattened when viewed from a lateral profile. Galleries of the conifer clade are in Figures <xref ref-type="fig" rid="F24">24</xref> and <xref ref-type="fig" rid="F25">25</xref>.</p><fig id="F24" orientation="portrait" position="float"><label>Figure 24.</label><caption><p>Galleries of conifer-feeding <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic>, from <xref rid="B42" ref-type="bibr">Edson 1967</xref>.</p></caption><graphic id="oo_33377.jpg" xlink:href="zookeys-450-001-g024"></graphic></fig><fig id="F25" orientation="portrait" position="float"><label>Figure 25.</label><caption><p>Galleries of conifer-feeding <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic>, from <xref rid="B42" ref-type="bibr">Edson 1967</xref>.</p></caption><graphic id="oo_33378.jpg" xlink:href="zookeys-450-001-g025"></graphic></fig><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></title><p><named-content content-type="taxon-authority">Wood, 1967</named-content></p><p><xref ref-type="fig" rid="F26">Fig. 26</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content><named-content content-type="comment"> Wood, 1967: 120.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic> are distinguished by the characteristics of the elytral apex with the apical margin produced between interstriae 1 and 2, deeply emarginated at interstriae 3, produced on interstriae 4 and deeply emarginated at stria 4.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>5.0 mm long (mean = 5.0 mm; n = 2); 2.5 times as long as wide. Body dark red-brown to black, antennae light brown, legs light brown to dark red-brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma broadly, deeply emarginated; epistomal process weakly developed, low; median area above mandibles bearing dense patch of long yellow hair-like setae. Frons appearing flattened when viewed laterally, slightly transversely impressed just above epistoma; surface moderately, coarsely longitudinally aciculate-punctate; aciculations converging at epistoma; punctures large, coarse and more dense medially above epistoma; surface moderately and uniformly covered by long, fine, yellow erect hair-like setae, setae longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, almost subquadrate, setose with partial septum, two arcuate sutures visible.</p><p><italic>Pronotum</italic> as long as wide; apical margin broadly rounded, area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow and moderately abundant, larger, more abundant laterally and on apical constriction; apical, anterolateral and lateral margins bearing moderately abundant, erect, yellow hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to rounded, serrate apex; apical margin of elytra produced between interstriae 1 and 2, deeply emarginated at interstria 3, produced on interstria 4 and deeply emarginated at stria 4; apical margin of elytral apices bearing large, coarse punctures; apex moderately emarginated at suture. Disc smooth, shining; interstriae bearing short, sparse yellow hair-like setae spaced by length of a setae or less; interstriae weakly impressed, more than twice width of striae, interstrial punctures uniseriate, smaller than those of striae; striae moderately impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 weakly elevated above base of ventrite 2. Ventrite 2 nearly perpendicular to ventrite 1; surface rugose, coarsely, deeply punctate; covered with abundant erect setae that are greater than length of segment 3; surface convex; armed medially with laterally compressed spine with base extending from apical margin to ¾ of segment, apex rounded, shining, impunctate; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge closer to basal margin of segment; length of ventrite 5 greater than combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F26" orientation="portrait" position="float"><label>Figure 26.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33379.jpg" xlink:href="zookeys-450-001-g026"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>4.6–5.9 mm long (mean = 5.23 mm; n = 4); 2.3–2.5 times as long as wide. Similar to male except epistoma feebly and broadly emarginated, epistomal process absent, frons convex when viewed laterally, weakly longitudinally aciculate, setae sparse, shorter, less than width of eye; frons weakly transversely impressed on lateral margins above epistoma and at vertex; median line on frons raised. Apical margin of elytra slightly produced between interstriae 1 and 2, emarginated at interstria 3, slightly produced on interstria 4 and emarginated at stria 4. Second ventrite unarmed.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>11.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic> Wood: female, labeled “27 mi E. Morelia, Mich., VI-4-1965, Mexico, 8,000 ft, S.L. Wood, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">religiosa</named-content></named-content></italic>” (USNM). Paratypes: <bold>MEXICO:</bold><bold><italic>Michoacán</italic>:</bold> 27 mi E. Morelia, 14.VI.1965, S.L. Wood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">religiosa</named-content></named-content></italic> (USNM-3).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>MEXICO:</bold><bold><italic>DURANGO</italic>:</bold> Potreyo del Sauto y Cebadillas, 19.IV.[19]90, J. Tulio Mendez M., 935-A, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (THAC-4). <bold><italic>Nuevo León</italic>:</bold> Santa Catarina, San Antonio de las Alazanas, 14.XII.2011, Gerardo Cuellar, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-3).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>MEXICO: Durango, Michoacán, Nuevo León (Fig. <xref ref-type="fig" rid="F27">27</xref>).</p><fig id="F27" orientation="portrait" position="float"><label>Figure 27.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33380.jpg" xlink:href="zookeys-450-001-g027"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">religiosa</named-content></named-content></italic> (Kunth) Schltdl. & Cham. (sacred fir), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">durangensis</named-content></named-content></italic> Martínez (Durango fir) and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (Mirb.) Franco (Douglas fir). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> species are likely the preferred host.</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><pmc-comment>PageBreak</pmc-comment>
<pmc-comment>PageBreak</pmc-comment>
<p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic> is an uncommon species and little is known regarding its biology. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic> colonizes both standing trees and felled trees (<xref rid="B142" ref-type="bibr">Wood 1967</xref>; <xref rid="B32" ref-type="bibr">Cibrián Tovar et al. 1995</xref>). The treetops attacked by <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic> exhibit red foliage. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic> has been collected feeding in the cambium of large <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> trees, although it has also been collected from <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content></named-content></italic>. Because of limited host records, it is unknown whether the primary host is <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> or <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content></named-content></italic>. Phylogenetic placement of the species suggests <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> (Fig. <xref ref-type="fig" rid="F4">4</xref>) because the species belongs to a clade of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> feeding taxa. The adult gallery is transverse across the grain of the wood with a central nuptial chamber, similar in appearance to that of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> (see <xref rid="B32" ref-type="bibr">Cibrián Tovar et al. 1995</xref>, fig 132). Larval mines are in the cambium and do not score the wood. Pupation occurs in the cambium (<xref rid="B142" ref-type="bibr">Wood 1967</xref>; <xref rid="B32" ref-type="bibr">Cibrián Tovar et al. 1995</xref>).</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>This is the first description of a male for this species.</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></title><p><named-content content-type="taxon-authority">Bright, 1964</named-content></p><p><xref ref-type="fig" rid="F28">Figs 28</xref><xref ref-type="fig" rid="F29">–29</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content><named-content content-type="comment"> Bright, 1964: 167.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>The <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic> male is easily distinguished by the endemic distribution in the Santa Lucia range of California, and by the presence of a median denticle on the apical margin of ventrite 4, occasionally median denticles may also be present on the apical margins of ventrites 2 and 3. The female is distinguished by its distribution and differentiated from the <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> female by the presence of a strongly developed and distinct epistomal process and larger size.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>3.0–4.0 mm long (mean = 3.45 mm; n = 15); 1.7–2.5 times as long as wide. Head, pronotum and abdominal venter dark red-brown, antennae light brown, legs dark red-brown to light brown, elytra light red-brown. Pronotum darker than elytra.</p><p><italic>Head.</italic> Epistoma moderately, acutely emarginated; epistomal process strongly produced, elevated; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing flattened when viewed laterally, not impressed; moderately, finely, longitudinally aciculate, coarsely punctate; aciculations converging at epistoma; punctures small, coarse; moderately, uniformly covered by long, fine, yellow erect hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, ovoid, setose with partial septum, two arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctulate, punctures moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on basal half, narrowing to subquadrate, smooth apex; apex entire at suture. Margin of apical edge bearing large, coarse punctures. Disc glabrous, smooth, shining; interstriae not impressed, more than twice width of striae, punctures uniseriate, smaller than those of striae; striae weakly impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 strongly produced, elevated above base of ventrite 2, ventrite base 2 appearing impressed. Ventrite 2 nearly perpendicular to ventrite 1; surface glabrous, shagreened, dull, finely, obscurely punctate; punctures small, fine and shallow; surface flattened. Apical margins of ventrites 2 and 3 may be armed with median denticle, median denticle always present on apical margin of ventrite 4; lateral margins of ventrites 2–4 unarmed. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 equal to combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F28" orientation="portrait" position="float"><label>Figure 28.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33381.jpg" xlink:href="zookeys-450-001-g028"></graphic></fig><fig id="F29" orientation="portrait" position="float"><label>Figure 29.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic> venters (from <xref rid="B42" ref-type="bibr">Edson 1967</xref>).</p></caption><graphic id="oo_33382.jpg" xlink:href="zookeys-450-001-g029"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>3.4–4.6 mm long (mean = 3.7 mm; n = 15); 2.2–2.5 times as long as wide. Similar to male except epistoma less strongly emarginated, epistomal process less strongly produced and elevated, frons convex when viewed laterally, weakly longitudinally aciculate, setae sparser, shorter, less than width of eye; weakly medially impressed between inner apices of eyes. Ventrites unarmed.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>95.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic> Bright: male, labeled “Calif: Monterey Co, Cone Peak, 6-29-[19]63, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">bracteata</named-content></named-content></italic>, C.J. Wray Collector” (CASC). Allotype, female, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic>, identical data as holotype. Paratypes: <bold>UNITED STATES:</bold><bold><italic>California</italic>:</bold><italic>Monterey Co.</italic>: Cone Peak, 29.VI.[19]63, C.J. Wray, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">bracteata</named-content></named-content></italic> (CASC-6, CNCI-20, EMEC-39, USNM-8). Carmel Valley, 15 mi S., 22.VII.[19]63, C.J. Wray, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">bracteata</named-content></named-content></italic>, (CASC-4).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>UNITED STATES:</bold><bold><italic>California</italic>:</bold><italic>Monterey Co.</italic>: Carmel Valley, 15 mi S., 23.VII.[19]63, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">bracteata</named-content></named-content></italic> (CASC-2); 30.VII.1964 (CASC-2). Cone Peak, 29.VI.[19]63, C.J. Wray, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">bracteata</named-content></named-content></italic> (CASC-1, USNM-12). Williams Canyon, Los Padres National Forest, 24.IV.1992, D.E. Bright, G. Ferrell, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">bracteata</named-content></named-content></italic> limbs (CNCI-1).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>UNITED STATES: California (Fig. <xref ref-type="fig" rid="F30">30</xref>).</p><fig id="F30" orientation="portrait" position="float"><label>Figure 30.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33383.jpg" xlink:href="zookeys-450-001-g030"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">bracteata</named-content></named-content></italic> (D. Don) Poit. (bristlecone fir / Santa Lucia fir).</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p>Little is known regarding the biology of this uncommon and narrowly geographically restricted species. The host, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">bracteata</named-content></named-content></italic>, is restricted to the Santa Lucia Mountains of California. The tree is distributed in small patches in deep, moist canyon bottoms as well as dry, rocky slopes and ledges within Los Padres National Forest in Monterey County, especially near Cone Peak and Church Creek (<xref rid="B62" ref-type="bibr">Griffin and Critchfield 1972</xref>; <xref rid="B128" ref-type="bibr">Sullivan 1993</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic> has been collected feeding in the bole of large standing trees (<xref rid="B14" ref-type="bibr">Bright 1964</xref>) but is also reported from larger limbs and fresh slash (<xref rid="B42" ref-type="bibr">Edson 1967</xref>).</p><p>The adult gallery somewhat resembles an expanded ‘E’ (Fig. <xref ref-type="fig" rid="F24">24</xref>). The gallery is parallel to the grain of the wood and consists of a central nuptial chamber and two egg galleries, one below and one above the nuptial chamber. Each egg gallery ends in a pronounced hook. Galleries strongly score the cambium and lightly score the sapwood. The adult gallery averages 2.5–12.0 cm in length (<xref rid="B14" ref-type="bibr">Bright 1964</xref>; <xref rid="B42" ref-type="bibr">Edson 1967</xref>). Egg niches are closely spaced on both sides of the maternal gallery (<xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>). Larval mines are first perpendicular to the egg gallery and later turn to form a fan shaped pattern before terminating at pupation chambers in the sapwood (<xref rid="B42" ref-type="bibr">Edson 1967</xref>).</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>Considerable variation is observed in the presence of median denticles on the apical margins of male ventrites 2 and 3. Frequently males only have a median denticle on the apical margin of ventrite 4 as those on 2 and 3 may or may not be present in the male. This species is related to <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic> based on morphological characters (Fig. <xref ref-type="fig" rid="F2">2</xref>). Both of these species also feed on true firs (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> spp.) and have the apical margin of male ventrite 1 strongly apically produced.</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></title><p><named-content content-type="taxon-authority">Blackman, 1934</named-content><named-content content-type="taxon-status">valid sp.</named-content></p><p><xref ref-type="fig" rid="F29">Figs 29</xref><xref ref-type="fig" rid="F31">, 31</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content><named-content content-type="comment"> Blackman, 1934: 25.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> is very morphologically similar to <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic>. Males of are distinguished from those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> by the frons flattened when viewed laterally, never deeply impressed, by the moderately abundant frontal setae (compared to dense) and by the host genus, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content></named-content></italic>. Males are distinguished from those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> by the following combination of characters: abdominal venter shiny in luster, the base of the ventrite 2 spine extends from the apical margin to three-quarters the length of the segment and geographical distribution east of the Rocky Mountains. The female is distinguished from that of both species by the shining luster of ventrite 2.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>2.2–2.8 mm long (mean = 2.4 mm; n = 15); 2.1–2.8 times as long as wide. Head, antennae, pronotum, and abdominal venter dark red-brown, elytra and legs yellow-brown to light brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly emarginated; epistomal process present, moderately developed, low; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing flattened when viewed laterally from epistoma to vertex, slightly transversely impressed just above epistoma to inner apices of eyes; moderately, coarsely, longitudinally aciculate-punctate; aciculations converging at epistoma; punctures large, sparse and coarse; moderately, uniformly covered by long, fine, yellow, erect, hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, irregularly ovoid, setose with partial septum, two broadly arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, smooth apex; apex moderately emarginated at suture. Margin of apical edge bearing small, fine punctures. Disc smooth, shining; interstriae not impressed, twice width of striae, punctures uniseriate, smaller than those of striae; punctures bearing short, sparse, recumbent, yellow setae slightly longer than size of a puncture (may be abraded); striae weakly impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron less than half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 rounded, 2 marked by weak carina. Ventrite 2 nearly perpendicular to ventrite 1; surface smooth, shining, finely punctate; punctures small, fine, shallow; covered with sparse setae less than length of segment 3; surface convex; apical margin armed with laterally compressed, median spine with base extending from apical margin to ¾ length of segment, apex rounded; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge equidistant between basal and apical margins of segment; length of ventrite 5 less than combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F31" orientation="portrait" position="float"><label>Figure 31.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33384.jpg" xlink:href="zookeys-450-001-g031"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>2.2–3.5 mm long (mean = 4.65 mm; n = 15); 2.1–2.5 times as long as wide. Similar to male except epistoma feebly emarginated, epistomal process weakly developed, frons convex when viewed laterally, weakly longitudinally aciculate, setae sparser, shorter, less than width of eye; weakly transversely impressed between epistoma and inner apices of eyes. Second ventrite apical margin armed with acute median denticle, with base extending from apical margin to half-length of segment.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>297.</p></sec><sec sec-type="treatment-Type specimens"><title>Type specimens.</title><p>Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> Blackman: male, labeled “[Capitan Mountains, N. Mex. 25 April, 1907], Hopk. US 3959, W.F. Fiske Collector, Type No. 43840 USNM” (USNM).</p></sec><sec sec-type="treatment-Non-type specimens"><title>Non-type specimens.</title><p><bold>CANADA:</bold><pmc-comment>PageBreak</pmc-comment><bold><italic>BRITISH COLUMBIA</italic>:</bold> Aspen Grove, 12.VII.1931, H. Richmond (CNCI-11, FMNH-2). Clinton, 6 mi N., 7.VII.1972, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (CNCI-7). Creston, 8.VI.1958, H. & A. Howden (EMEC-1). Fort Steele, 14.VI.[19]26, R. Hopping (CASC-1). Indian Meadows, Midday Creek, 13.VII.1920, R. Hopping, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CASC-2, CNCI-2). Lumby, Creighton Valley, 3.VI.[19]22, R. Hopping (CASC-1). Merritt, Midday Valley, 27.VI.1926, W. Mathers (CASC-2). Trinity Valley, 24.VI.1928, J.R. Howell, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content><named-content content-type="species">monticola</named-content></named-content></italic> (CASC-1), 10.VII.1928 (CASC-1). <bold>UNITED STATES:</bold><bold><italic>ARIZONA</italic>:</bold><italic>Pima Co.</italic>: Tucson, Mount Lemmon, 11.VI.1969, S.L. Wood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-2). <bold><italic>COLORADO</italic>:</bold><italic>Boulder Co.</italic>: Nederland, 5–7 km N., 5.VIII.2009, D.E. Bright, B.A. Barr, ex. branches of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (CNCI-4). [<italic>La Plata Co.</italic>]: Durango, Junction Creek Rd, 10000 ft, 12-17.VII.1968, E.C. Becker (CNCI-1). <italic>Larimer Co.</italic>: Roosevelt National Forest, Big Thompson Canyon, <named-content content-type="dwc:verbatimCoordinates">N40°24.456', W105°24.565'</named-content>, 7080 ft, 5.V.2010, S.M. Smith, D.E. Bright, B.A. Barr, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-36). <bold><italic>IDAHO</italic>:</bold> [<italic>Adams Co.</italic>]: Tamarack, 10 mi S., 3.VII.1967 (WFBM-1). <italic>Boise Co.</italic>: Idaho City, 2.VI.1970 (WFBM-9), 1.I.1971 (WFBM-4). Lowman, Edna Creek, 6.VII.[19]72, A. Vaccares, G. Starr, ex. Douglas fir stump [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (WFBM-1). <italic>Bonner Co.</italic>: 6.VI.1986, M.M. Furniss, J.B. Johnson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-7). Priest Lake, Indian Creek, 6.VI.1986, M.M. Furniss, J.B. Johnson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-25). Priest Lake, 6.VI.1986, M.M. Furniss, J.B. Johnson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-1). <italic>Boundary Co.</italic>: Parker Creek, 8.VI.1986, M.M. Furniss, J.B. Johnson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-5). <italic>Clearwater Co.</italic>: Elk River, 12 mi S.E., Hopk. U.S. 58536, 9.VII–16.VII.1973, R.D. Oakes (WFBM-1), 30.VII-6.VIII.1973 (WFBM-1); Hopk. U.S. 58771, 25.VI-1.VII.1974, J.M. Wells, ex. in flight (WFBM-1). <italic>Custer Co.</italic>: Herd Lake, Hopk. U.S. 60796-A, 3.IX.1978, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-9). MacKay, 9 mi N.E., 19.VII.1985, M.M. Furniss, J.B. Johnson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-3). Summit, W. of Pass Creek, 19.VII.1985, M.M. Furniss, J.B. Johnson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-12). <italic>Latah Co.</italic>: Big Sand Creek, Hopk. U.S. 53545, 11.VI.1969, M.M. Furniss (WFBM-1), 20.VI.1969 (WFBM-1). Moscow, 20.XI.1961, R.E. Stecker, ex. reared from Doug fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (WFBM-33). Moscow Mountain, Hopk. U.S. 48869, 18.VII.1967, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-3). Viola, 21.VI.1985, S.J. Gast, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-2). <italic>Nez Perce Co.</italic>: Forest, III.1985, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-6). <italic>Shoshone Co.</italic>: Red Ives Ranger Station, V.1983, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-1). <italic>Valley Co.</italic>: Cascade, 7 mi E., 14.VI.1966, R.L. Furniss (WFBM-1). <bold><italic>MONTANA</italic>:</bold><italic>Lake Co.</italic>: Swan Lake, 28.VI.1963, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-1). <italic>Madison Co.</italic>: Alder, 18 km S.W., 8.IX.1978, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-6), Alder, 12 mi S.W. (WFBM-16). <italic>Park Co.</italic>: Livingston, 10 mi S.E., 23.VII.1988, M.M. Furniss, J.B. Johnson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-2). <bold><italic>NEW MEXICO</italic>:</bold><italic>Otero Co.</italic>: Cloudcroft, 4.VI.1969, tree 53, S.L. Wood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-2). <italic>Sierra Co.</italic>: Emory Pass, 24.VII.1974, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (CNCI-1). <bold><italic>WASHINGTON</italic>:</bold> [<italic>Whitman Co.</italic>]: Pullman, 23.VI.1951, N.M. Downie (FMNH-1). <bold><italic>WYOMING</italic>:</bold> [<italic>Teton Co.</italic>]: Jackson, A.D. Hopkins, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (USNM-1).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: British Columbia. UNITED STATES: Arizona, Colorado, Idaho, Montana, New Mexico, Washington, Wyoming (Fig. <xref ref-type="fig" rid="F32">32</xref>).</p><fig id="F32" orientation="portrait" position="float"><label>Figure 32.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33385.jpg" xlink:href="zookeys-450-001-g032"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (Mirb.) Franco (Douglas fir).</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> colonizes suppressed limbs and branches of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> as well as fresh slash (<xref rid="B42" ref-type="bibr">Edson 1967</xref>; Smith, pers. obs.). The adult gallery is bayonet shaped, parallel to the grain of wood and consists of two egg galleries, one below and one above the nuptial chamber and a turning niche. The female constructs the adult gallery mostly in the cambium and lightly scores the sapwood. Larval mines are perpendicular to the egg gallery and turn to form a fan shaped pattern (<xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>; Smith, pers. obs.) (Fig. <xref ref-type="fig" rid="F24">24</xref>). The adult gallery measures 3.8–9.0 cm in length (<xref rid="B42" ref-type="bibr">Edson 1967</xref>). The following year adults emerge and excavate tunnels within twigs of Douglas fir for maturation feeding causing twig flagging (reported as <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic><xref rid="B96" ref-type="bibr">McMullen and Atkins 1962</xref>).</p><p>Like most <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species, there is a very limited amount of information regarding <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic>. For many years it was considered a synonym of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> and was thus referred to as <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> in publications. Very little was written about this species in the US (see <xref rid="B151" ref-type="bibr">Wood and Bright 1992</xref>) other than host preference and gallery descriptions (<xref rid="B77" ref-type="bibr">Keen 1938</xref>; <xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B17" ref-type="bibr">Bright 1976</xref>). <xref rid="B96" ref-type="bibr">McMullen and Atkins (1962)</xref> reported some notes on the biology of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> in British Columbia and appear to have reported a combined account of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic>. The authors noted the gallery of the <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> studied in their investigations as “in about 15 per cent of the galleries over 20 days of age were the type described by Chamberlin and Keen [<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic>, which has only a single egg gallery]; the majority were of the forked type and were similar to those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> [referencing a forked or bayonet shaped gallery that is made by <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic>]”. Their description matches the gallery description of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic>, which like has two egg galleries as part of a bayonet shaped adult gallery. Because most of the specimens are of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic>, this article does offer some information regarding the biology of the species be used with caution because not all of the findings may apply to <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic>.</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>The holotype of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> does not bear a locality label. <xref rid="B10" ref-type="bibr">Blackman’s (1934)</xref> description states the holotype was collected at the Capitan Mountains, New Mexico.</p><p><xref rid="B52" ref-type="bibr">Furniss and Johnson (2002)</xref> report <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> from Alberta, Canada. These specimens are likely <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> based on the geographic distribution of the species in the Rocky Mountains.</p><p><xref rid="B146" ref-type="bibr">Wood (1977</xref>: 388) placed <pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> in synonymy with <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> after examining both holotypes and 164 specimens from Arizona to British Columbia and concluded that there was too much intraspecific variation to recognize them as separate species. We assessed the intraspecific and interspecific variation (Tables <xref ref-type="table" rid="T6">6</xref> and <xref ref-type="table" rid="T7">7</xref>) for each of the four genes for each species. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> intraspecific variation for COI was low, 0.0–0.0231, and averaged 0.0183 among all sampled populations. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> intraspecific variation was also low, 0.0016–0.0282 and averaged 0.0121. Interspecific variation between <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> was much higher, 0.0331–0.0521 and averaged 0.043 among populations. Similar differences were also observed with CAD (Table <xref ref-type="table" rid="T7">7</xref>). The species have separate geographical ranges with <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> occurring primarily in the Rocky Mountains and British Columbia while <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> occurs in the Cascade and Sierra mountains from California to British Columbia. Both species are sympatric in the Interior Plateau of British Columbia near Merritt. The species are separated by the characters listed in the diagnosis and by the galleries. The galleries of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> contain two egg galleries; one above and one below the nuptial chamber. The gallery of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> only contains a single egg gallery. In addition, the gallery of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> lightly scores the sapwood while that of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> deeply scores the sapwood. After examining the types, 950 specimens of both species and testing the monophyly of each species using four genes, it is apparent that <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> is a distinct lineage and is here removed from synonymy with <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic>.</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></title><p><named-content content-type="taxon-authority">Wood, 1968</named-content></p><p><xref ref-type="fig" rid="F33">Fig. 33</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content><named-content content-type="comment"> Wood, 1968: 12.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Both sexes resemble the <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic> female. The female is distinguished by having a weakly developed epistomal process and the male is distinguished by having the apical margin of ventrite 1 produced, forming a carinate lip along the basal margin of ventrite 2 that is twice as produced as thick and by the host genus, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic>.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>3.0 mm long (n = 1); 1.5 times as long as wide. Head, antennae, pronotum, and abdominal venter dark red-brown, elytra and legs yellow-brown to light brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly emarginated; epistomal process weakly developed, low; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing convex when viewed laterally, slightly transversely impressed just above epistoma and between inner apices of eyes; moderately, finely, longitudinally aciculate-punctate; aciculations converging at epistoma; punctures small, coarse; moderately, uniformly covered by long, fine, yellow, erect hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, elongate, almost subquadrate, setose with partial septum, two arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, smooth apex; apex moderately emarginated at suture. Margin of apical edge bearing large, coarse punctures. Disc smooth, shining; interstriae not impressed, more than twice width of striae, punctures uniseriate, smaller than those of striae, punctures bearing sparse, long, erect, yellow hair-like setae (may be abraded); striae weakly impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 produced, forming carinate lip along basal margin of ventrite 2 that is twice as produced as thick, basal margin of ventrite 2 appearing impressed. Ventrite 2 nearly perpendicular to ventrite 1; surface shagreened, dull, finely punctate; punctures small, fine, shallow; setae moderately abundant, recumbent, short, about four times length of a puncture; surface weakly concave; apical margin armed with broad median denticle, occasionally absent; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 greater than combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F33" orientation="portrait" position="float"><label>Figure 33.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33386.jpg" xlink:href="zookeys-450-001-g033"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>2.5–3.5 mm long (mean = 3.27 mm; n = 6); 2.1–2.3 times as long as wide. Similar to male except epistoma feebly emarginated, epistomal process absent, frons more strongly convex when viewed laterally, weakly longitudinally aciculate, setae sparser, shorter, less than width of eye; weakly transversely impressed just above epistoma. Apical margin of ventrite 1 weakly elevated above base of ventrite 2. Second ventrite unarmed.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>27.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic> Wood: male, labeled “Tlaxco 11 mi N., 8900 ft, 9.VII.1967, S.L. W[ood], ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">religiosa</named-content></named-content></italic>” (USNM). Paratypes: <bold>MEXICO:</bold> [<bold><italic>Puebla</italic></bold>]: Tlaxco, 11 mi N., 8900 ft, 9.VII.1967, S.L. W[ood], ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">religiosa</named-content></named-content></italic> (USNM-21).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>MEXICO:</bold><bold><italic>CHIHUAHUA</italic>:</bold> La Magdalena, Hopk. U.S. 62081-B, 27.IV.1981, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">durangensis</named-content></named-content></italic> (USNM-1). <bold><italic>Nuevo León</italic>:</bold> Cerro Potosi, Hopk. U.S. 58615-C, 21.III.[19]74, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> sp. (USNM-2, WFBM-1).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>MEXICO: Chihuahua, Nuevo León, Puebla (Fig. <xref ref-type="fig" rid="F34">34</xref>).</p><fig id="F34" orientation="portrait" position="float"><label>Figure 34.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33387.jpg" xlink:href="zookeys-450-001-g034"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">religiosa</named-content></named-content></italic> (Kunth) Schltdl. & Cham (sacred fir), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">durangensis</named-content></named-content></italic> Martínez (Durango fir) and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (Mirb.) Franco (Douglas fir). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> species are likely the preferred hosts.</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p>There is a paucity of information regarding the biology of this uncommon species. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic> has been collected feeding in the cambium of large <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> slash greater than 10.0 cm in diameter (<xref rid="B142" ref-type="bibr">Wood 1967</xref>) and weakened and dying large <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (<xref rid="B32" ref-type="bibr">Cibrián Tovar et al. 1995</xref>). The adult galleries are transverse and perpendicular to the grain of the wood (<xref rid="B142" ref-type="bibr">Wood 1967</xref>). This species has been reported to colonize the same material as <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic> (<xref rid="B143" ref-type="bibr">Wood 1968</xref>).</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></title><p><named-content content-type="taxon-authority">Blackman, 1934</named-content></p><p><xref ref-type="fig" rid="F29">Figs 29</xref><xref ref-type="fig" rid="F35">, 35</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content><named-content content-type="comment"> Blackman, 1934: 24.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> is very morphologically similar to <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic>. Males are distinguished from those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> by the dull appearance of abdominal ventrite 2 and by the frons densely covered by long setae. Males can be distinguished from those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> by the following combination of characters: the base of the ventrite 2 spine extends from the apical margin to three-quarters the length of the segment and host genus. The female is distinguished from that of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> by the dull luster of ventrite 2 and is distinguished from that of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> by the frons moderately and coarsely aciculate-punctate; ventrite 1 rounded over onto surface of ventrite 2, not forming an obtuse angle, and the base of ventrite 2 finely impressed.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>2.3–4.0 mm long (mean = 3.1 mm; n = 15); 1.8–2.5 times as long as wide. Head, pronotum, and abdominal venter dark red-brown to black, elytra and legs light brown, antennae yellow-brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly emarginated; epistomal process present, moderately developed and elevated; median area above mandibles bearing dense patch of long yellow hair-like setae. Frons appearing flattened when viewed laterally from epistoma to vertex, moderately transversely impressed just above epistoma to vertex; strongly, coarsely, longitudinally aciculate-punctate; aciculations converging at epistoma; punctures sparse, large, coarse; moderately to densely uniformly covered by long, fine, yellow, erect, hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, irregularly ovoid, setose with partial septum, two arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, smooth apex; apex moderately emarginated at suture. Margin of apical edge bearing large, coarse punctures. Disc smooth, shining; interstriae not impressed, twice width of striae, punctures uniseriate, smaller than those of striae, bearing minute, recumbent setae length of interstrial punctures (may be abraded); striae weakly impressed. Declivity bearing sparse, short, erect, yellow setae. Metepimeron less than half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 rounded, marked by weak carina on ascendant part of venter. Ventrite 2 nearly perpendicular to ventrite 1; surface smooth, shining, finely punctate; punctures small, fine, shallow; covered with sparse setae less than length of segment 3; surface convex; apical margin armed with laterally compressed, median spine with base extending from apical margin to ¾ length of segment, apex rounded, rarely a slightly elevated vertical carina (Oregon: Dixie Pass and Frog Lake) instead of spine; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge equidistant between basal and apical margins of segment; length of ventrite 5 less than combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F35" orientation="portrait" position="float"><label>Figure 35.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33388.jpg" xlink:href="zookeys-450-001-g035"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>2.6–4.0 mm long (mean = 3.2 mm; n = 15); 2.1–2.9 times as long as wide. Similar to male except epistoma feebly emarginated, epistomal process weakly developed, frons convex when viewed laterally, weakly longitudinally aciculate, setae sparser, shorter, less than width of eye; weakly transversely impressed just above epistoma. Second ventrite apical margin armed with a blunted median denticle, with base extending from apical margin to a third length of segment. Ventrite 5 length greater than combined lengths of ventrites 3 and 4.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>294.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> Blackman: male, labeled “[Cedar Mountain, Moscow, ID] Hopk. US 225d, 8636 m [sic!], 6-20-[18]99, Type No. 43839” (USNM).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>CANADA:</bold><bold><italic>BRITISH COLUMBIA</italic>:</bold> Canoe, 12.VII.1933, A. Thrupp, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (CASC-3). Rossland, 21.IX.[19]61, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">laricina</named-content></named-content></italic> (CNCI-2). <bold>UNITED STATES:</bold><bold><italic>IDAHO</italic>:</bold><italic>Boundary Co.</italic>: [Idaho Panhandle National Forest], Robinson Creek campground, <named-content content-type="dwc:verbatimCoordinates">N48°58.197', W116°13.068'</named-content>, 2748 ft, 10.VIII.2010, S.M. Smith, [A.R. Gillogly], ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic>, emerged II.2011, M.M. Furniss (MSUC-94). <italic>Clearwater Co.</italic>: Elk River, V.1981, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (WFBM-17), VII.1981 (WFBM-11). Elk River, 13 mi S.W., Hopk. U.S. 60361-A, 23.IV.1975, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (WFBM-4). <italic>Idaho Co.</italic>: Salmon Mountain, 18.VIII.1985, M.M. Furniss, J.B. Johnson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">lyalli</named-content></named-content></italic> (WFBM-29). <italic>Kootenai Co.</italic>: Coeur d’Alene, Hopk. U.S. 16306-B, 2.VIII.1919, J.C. Evenden, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (WFBM-9), 8.VIII.1919 (MSUC-11). Deception Creek Experimental Forest, Hopk. U.S. 58889-A, 10.VII.1968, M.M. Furniss (OSAC-4), Hopk. U.S. 53376, 11.VII.1968 (WFBM-2), Hopk. U.S. 60320, 25.VII.1968, (OSAC-2), Hopk. U.S. 50421-B,C, 1.VII.1975 (OSAC-2), Hopk. U.S. 60356, 30.VII.[19]68, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (WFBM-6). <italic>Latah Co.</italic>: Moscow, 2.VIII.1930, 2560 ft, P. Rice, ex. trap (WFBM-1). Moscow Mountain, Hopk. U.S. 53632-F, 23.VI.1964, M.M. Furniss, ex. in flight (WFBM-1), Hopk. U.S. 60421-A, 1.VII.1975 (OSAC-1). <bold><italic>MONTANA</italic>:</bold><italic>Missoula Co</italic>: Missoula, 35 mi N.W., 3.XI.[19]65, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (WFBM-2). Nine Mile Creek, Hopk. U.S. 48830, 3.XI.1965, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (WFBM-2, OSAC-1). <italic>Ravalli Co.</italic>: [Bitterroot Mountains], Trapper Peak, 2.X.[19]88, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">lyalli</named-content></named-content></italic> (WFBM-3). [<italic>Unspecified County</italic>]: Libby, 60 mi S.E., 30.VI.[19]72, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">laricina</named-content></named-content></italic> (CNCI-7). <bold><italic>OREGON</italic>:</bold><italic>Crook Co.</italic>: Summit Prairie, 12.VI.1940, Schuh, Scott, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (CNCI-3, EMEC-2). [<italic>Grant Co.</italic>]: Dixie Pass, Malheur National Forest, 23.VI.1961, S.L. Wood, J.B. Karren, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (CNCI-6, USNM-6). <italic>Jefferson Co.</italic>: Camp Sherman, T.O. Thatcher, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">lyalli</named-content></named-content></italic> limbs (CSUC-10), 9.VII.[19]66, [L. Edson], ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">lyalli</named-content></named-content></italic> (CNCI-2). Suttle Lake, 4 mi N., 25.VII.1939, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">lyalli</named-content></named-content></italic> (CSUC-1); 28.VII.1939, F. Grey, J. Schuh, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (CASC-1, FMNH-6, MSUC-1, OSAC-14), W.J. Chamberlin, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">lyalli</named-content></named-content></italic> (EMEC-7, WFBM-3). Suttle Lake, 4 mi W., 15.VIII.1939, Schuh, Scott (MSUC-1). [<italic>Marion Co.</italic>]: Clear Lake, 17.VIII.[19]51, R. Kangur, ex. larch [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content></named-content></italic> sp.] (EMEC-4, WFBM-2). [<italic>Umatilla Co.</italic>]: Tollgate, 28.X.1948, C. Chastain, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (EMEC-1). [<italic>Wasco Co.</italic>]: [Mount Hood National Forest] Frog Lake [campground], 4S9 E17, 2.VIII.1951, R. Kangur (CNCI-3). <bold><italic>WASHINGTON</italic>:</bold><italic>Okanogan Co.</italic>: Disautel, 4.XI.1936, R.L. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (OSAC-3). [<italic>Pend Oreille Co.</italic>]: Metaline Falls, Hopk. U.S. 19905, 20.VII.1930, D. DeLeon, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (OSAC-2, WFBM-1). [<italic>Stevens Co.</italic>]: Northport, 18.VII.1929, R. Hopping (OSAC-1). [<italic>Unspecified County</italic>]: Mount Adams, 7000 ft, Hopk. U.S. 53359-C, 2.VIII.1968, M.M. Furniss, ex. on snowfield (WFBM-1).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: British Columbia. UNITED STATES: Idaho, Montana, Oregon, Washington (Fig. <xref ref-type="fig" rid="F36">36</xref>).</p><fig id="F36" orientation="portrait" position="float"><label>Figure 36.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33389.jpg" xlink:href="zookeys-450-001-g036"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> Nutt. (western larch) and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">lyalli</named-content></named-content></italic> Parl. (subalpine larch).</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> prefers to colonize suppressed lower limbs of its host and fresh slash.</p><p>The adult gallery is parallel to the grain of the wood and consists of two egg galleries and a central nuptial chamber (Fig. <xref ref-type="fig" rid="F24">24</xref>). The central nuptial chamber extends at a right angle to the rest of the gallery and opposite the nuptial chamber is a rounded turning niche. This gives the central part of the gallery an ovoid appearance that is different from all other Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic>. Egg galleries are extended parallel to the grain of the wood. The adult gallery scores the sapwood more deeply than the cambium, and ranges in length from 6.4–11.5 cm. The female constructs widely spaced egg niches on both sides of each egg gallery. Larval mines extend against the grain of wood and gradually diverge before terminating at pupation chambers that deeply score the sapwood (<xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>). There is one generation per year and broods overwinter as larvae (<xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>).</p></sec><sec sec-type="treatment-Collection notes"><title>Collection notes.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> was collected four times by the senior author and each time specimens were collected from fresh broken branches that were less than 5.0 cm in diameter with bright green needles. Specimens were also collected from a fallen <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> of 15.0 cm DBH.</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p><xref rid="B52" ref-type="bibr">Furniss and Johnson (2002)</xref> reported that specimens collected from <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">lyalli</named-content></named-content></italic> create a different gallery structure than those collected from <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic> (M.M. Furniss pers. comm.). The galleries on <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">lyalli</named-content></named-content></italic> are “shorter, less uniform, and with the entrance chamber often extended a short distance to the sides of the gallery” (<xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>). Specimens from <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">lyalli</named-content></named-content></italic> could not be collected during the coarse of our investigation. We noticed slight morphological variation among some populations but were unable to determine if the differences represent species limits. Further study is needed to address this question.</p><pmc-comment>PageBreak</pmc-comment>
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<p>The holotype of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> does not bear a locality label. <xref rid="B10" ref-type="bibr">Blackman’s (1934)</xref> description states A.D. Hopkins collected the holotype on Cedar Mountain near Moscow, Idaho. Numerous collectors listed in the material examined reported this species as collected from <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">lyalli</named-content></named-content></italic> in Oregon. This host species occurs east of the Cascade Range in the Wenatchee Mountains in Washington and is not known to occur in Oregon (<xref rid="B22" ref-type="bibr">Burns and Honkala 1990</xref>). It is probable that the host species for these specimens was <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content><named-content content-type="species">occidentalis</named-content></named-content></italic>.</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></title><p><named-content content-type="taxon-authority">(Swaine, 1917)</named-content></p><p><xref ref-type="fig" rid="F29">Figs 29</xref><xref ref-type="fig" rid="F37">, 37</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Eccoptogaster</named-content><named-content content-type="species">monticolae</named-content></named-content><named-content content-type="comment"> Swaine, 1917: 32.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content><named-content content-type="comment"> (Swaine, 1917): <xref rid="B76" ref-type="bibr">Keen 1929</xref>: 13.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> males are easily confused with <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> males, especially those exhibiting the wickhami phenotype. They are easily distinguished by the size of male ventrite 5. In <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic>, ventrite 5 is equal in length to width of ventrites 3 and 4 combined. In <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic>, ventrite 5 is equal in length to ventrite 4. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> lacks an epistomal process while <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> typically has a strongly developed epistomal process. Males are distinguished from those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> by the following combination of characters: surface of ventrite 2 shining but minutely reticulate; elytral striae not impressed; basal margin of ventrite 2 more pronounced and produced laterally; elytral strial punctures small, spaced 2–3 diameters of a puncture. Females of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> are distinguished from <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> females by having the apical margin of ventrite 1 weakly produced, never rounded and the surface of ventrite 2 smooth and flat, and are separated from those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> by elytral discal striae not impressed and ventrite 2 shining in luster.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>2.5–3.0 mm long (mean = 2.9 mm; n = 10); 2.0–2.5 times as long as wide. Head, pronotum, and abdominal venter dark red-brown, legs light brown, antennae yellow-brown, elytra usually dark red-brown but may be brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly emarginated; epistomal process weakly developed; median area above mandibles bearing dense patch of long yellow hair-like setae. Frons appearing flattened when viewed laterally, slightly transversely impressed just above epistoma; weakly aciculate-punctate, medial area appearing shagreened; aciculations converging at epistoma; punctures small, coarse; sparsely, uniformly covered by long, fine, yellow, erect, hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, irregularly ovoid, setose with partial septum, two arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on basal half, narrowing to subquadrate, smooth apex; apex moderately emarginated at suture. Margin of apical edge bearing large, coarse punctures. Disc glabrous, smooth, shining; interstriae not impressed, twice width of striae, punctures uniseriate, smaller than those of striae, bearing minute, recumbent setae length of interstrial punctures (may be abraded); striae not impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron less than half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 weakly elevated above base of ventrite 2, more pronounced and produced laterally forming two cups. Ventrite 2 nearly perpendicular to ventrite 1; surface smooth, shining, finely punctate; punctures small, fine, shallow; surface flattened, depressed above basal margin; apical margin unarmed; covered in recumbent setae twice width of a puncture; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 less than combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F37" orientation="portrait" position="float"><label>Figure 37.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33390.jpg" xlink:href="zookeys-450-001-g037"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>2.3–3.5 mm long (mean = 3.0 mm; n = 10); 2.3–2.7 times as long as wide. Similar to male except epistoma feebly emarginated, frons convex when viewed laterally, strigate, setae sparser, shorter, less than width of eye; weakly transversely impressed between inner apices of eyes. Second ventrite unarmed. Length of ventrite 5 greater than combined lengths of ventrites 3 and 4.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>123.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Lectotype: female, labeled “[Arrowhead, British Columbia], <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content><named-content content-type="species">monticola</named-content></named-content></italic>, 2423, J.M. Swaine Coll, female” (CNCI). Lectotype designated <xref rid="B15" ref-type="bibr">Bright 1967</xref>: 674.</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>CANADA:</bold><bold><italic>BRITISH COLUMBIA</italic>:</bold> Merritt, Midday Valley, 27.VI.1926, W. Mathers (USNM-2), 4.IX.1926 (USNM-2). Trinity Valley, 4.VII.1954, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> (USNM-4). <bold>UNITED STATES:</bold><bold><italic>COLORADO</italic>:</bold><italic>Custer Co.</italic>: Hillside, 6 mi W., Duckett Creek, FR331, Rita Alta Fuelwood area, 23.V.2000, D. Leatherman (CSUC-1). [<italic>Garfield Co.</italic>]: Glenwood Springs (CASC-1). <italic>Jefferson Co.</italic>: Buffalo Creek, 6.VIII.2004, D. Leatherman, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CASC-2). <bold><italic>IDAHO</italic>:</bold><italic>Benewah Co.</italic>: St. Maries, Hopk. U.S. 618074, 28.VIII.1978, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (USNM-32). <italic>Boise Co.</italic>: Boise National Forest, Bogus Basin, Bogus Basin Rd, <named-content content-type="dwc:verbatimCoordinates">N43°44.347', W116°07.099'</named-content>, 6047 ft, 8.VIII.2010, S.M. Smith, A.R. Gillogly, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-8). <italic>Bonner Co.</italic>: Priest River Experimental Forest, Hopk. U.S. 61809-A, 25.X.1978, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (USNM-26). <italic>Kootenai Co.</italic>: Coeur d’Alene, 30.VIII.1919, J.C. Evenden (MSUC-12); 7.VIII.1919, J.C. Evenden, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-4). <italic>Shoshone Co.</italic>: Coeur d’Alene National Forest, <named-content content-type="dwc:verbatimCoordinates">N47°25.708', W115°53.464'</named-content>, 3728 ft, 15.VIII.2010, S.M. Smith, A.R. Gillogly, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-19). Prichard, 23.VII.1920, J.C. Evenden, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (MSUC-2). <bold><italic>MONTANA</italic>:</bold> [<italic>Sanders Co.</italic>]: Trout Creek, 1.VIII.1981, J. Dunkel, ex. Douglas fir (USNM-2). <bold><italic>OREGON</italic>:</bold> [<italic>Unspecified County</italic>]: Santiam National [State] Forest, 22.VIII.[19]14, W.J. Chamberlin, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">amabalis</named-content></named-content></italic> (EMEC-1). <bold><italic>WASHINGTON</italic>:</bold> [<italic>Kittitas Co.</italic>]: Easton (USNM-1). [<italic>Yakima Co.</italic>]: Cliffdell, 7.VII.[19]35, R.H. Beaner (USNM-1). <bold><italic>WYOMING</italic>:</bold> [<italic>Park Co.</italic>]: Cody, Hopk. U.S. 34220-F, 10.V.[19]56, H.E. Ostmark, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CSUC-2).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: British Columbia. UNITED STATES: Colorado, Idaho, Montana, Oregon, Utah, Washington, Wyoming (Fig. <xref ref-type="fig" rid="F38">38</xref>).</p><fig id="F38" orientation="portrait" position="float"><label>Figure 38.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33391.jpg" xlink:href="zookeys-450-001-g038"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (Mirb.) Franco (Douglas fir).</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> commonly infests the bole, suppressed branches and fresh slash of Douglas fir. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> (as <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic>, see remarks below) has been reported to kill drought stressed sapling and pole-sized Douglas fir trees in British Columbia (<xref rid="B95" ref-type="bibr">McMullen and Atkins 1959</xref>) and Wyoming (<xref rid="B55" ref-type="bibr">Furniss and Carolin 1977</xref>).</p><p>Adult galleries strongly resemble those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic>, parallel to the grain of the wood, bayonet shaped and consist of two egg galleries and a central nuptial chamber (Fig. <xref ref-type="fig" rid="F24">24</xref>). From the central nuptial chamber, one egg gallery extends with the grain of the wood and the other egg gallery is slightly transversely extended and then is extended parallel to the grain. The nuptial chamber is oblique to the egg galleries. The adult gallery scores the sapwood more than the cambium and averages 5.0–9.0 cm in length. Egg niches are generally placed in pairs along the egg galleries and score the sapwood. Larvae extend their mines perpendicular to the egg gallery before diverging in a fan shaped pattern. Pupation may occur under the bark or in the sapwood. In Idaho, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> has one generation per year with flight occurring in July (<xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>; Smith, pers. obs.). Broods overwinter as larvae and emerge the following summer (<xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>).</p></sec><sec sec-type="treatment-Collection notes"><title>Collection notes.</title><p>The senior author found this species to be common in suppressed limbs and slash in Idaho.</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>The lectotype does not bear a locality label. <xref rid="B131" ref-type="bibr">Swaine’s (1917)</xref> description states the lectotype was collected at Arrowhead, British Columbia.</p><p>In their paper describing the biology of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic>, <xref rid="B95" ref-type="bibr">McMullen and Atkins (1959)</xref> considered <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> a synonym of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> based on correspondence with G.R. Hopping (page 417). <xref rid="B141" ref-type="bibr">Wood (1966</xref>: 30) formally synonymized <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> with <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic>, and later removed the species from synonymy (<xref rid="B148" ref-type="bibr">Wood 1982</xref>; M.M. Furniss, pers. comm.). However, <xref rid="B148" ref-type="bibr">Wood (1982)</xref> did not explicitly state that he was the author that recognized the species. These two species are closely related and have subtle morphological differences. This has led to a confusing account of both species in the literature. The majority of which seems to be applicable to <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> rather than <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic>. The above diagnostic characters and different biologies readily differentiate the species.</p><pmc-comment>PageBreak</pmc-comment>
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</sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></title><p><named-content content-type="taxon-authority">Wood, 1968</named-content></p><p><xref ref-type="fig" rid="F39">Fig. 39</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content><named-content content-type="comment"> Wood, 1968: 13.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>This species most strongly resembles <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic>. Both sexes are differentiated by elytral apices slightly emarginated only at interstria 3, by ventrite 3 covered with abundant erect setae that are greater than length of segment 3. The male can also be differentiated by the apical margin of ventrite 2 armed with a broad median denticle; larger size, the geographic distribution.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>4.0–4.5 mm long (mean = 4.35 mm; n = 4); 2.1–2.5 times as long as wide. Color dark brown to black, antennae red brown. Pronotum same color as elytra.</p><p><italic>Head.</italic> Epistoma weakly emarginated; epistomal process absent; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing convex when viewed laterally, moderately, transversely impressed just above epistoma; strongly, coarsely longitudinally, aciculate-punctate; aciculations converging at epistoma; punctures small, fine; densely, uniformly covered by long, fine, yellow, erect, hair-like setae, setae longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, nearly subquadrate, setose with partial septum, two arcuate sutures visible.</p><p><italic>Pronotum</italic> as long as wide; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical, anterolateral and lateral margins bearing abundant, erect, long, dark yellow-brown, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, smooth apex; elytral apices slightly emarginated at interstria 3; apex weakly emarginated at suture. Margin of apical edge bearing large, coarse punctures. Disc smooth, shining; interstriae weakly impressed, more than twice width of striae, punctures uniseriate, equal in size to those of striae, bearing moderately abundant long, erect, dark yellow-brown, hair-like setae; striae weakly impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 weakly elevated above base of ventrite 2. Ventrite 2 nearly perpendicular to ventrite 1; surface smooth, shining, finely punctate; punctures small, fine, shallow; covered with abundant erect setae greater than length of segment 3; surface flattened; apical margin armed with broad median denticle; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 greater than combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F39" orientation="portrait" position="float"><label>Figure 39.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33392.jpg" xlink:href="zookeys-450-001-g039"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>3.3–4.6 mm long (mean = 4.0 mm; n = 13); 1.8–2.6 times as long as wide. Similar to male except epistoma feebly emarginated, epistomal process absent, frons convex when viewed laterally, weakly longitudinally aciculate, setae sparser, shorter, less than width of eye; weakly transversely impressed between inner apices of eyes. Apical margin of second ventrite armed by small broad tumescence.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>28.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: male, labeled “11 mi, N Tlaxco, Puebla, Mexico, SL W[ood], <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">religiosa</named-content></named-content></italic>.” (USNM). Paratypes: <bold>MEXICO:</bold> [<bold><italic>Puebla</italic></bold>]: Tlaxco (Tlaxcala), 11 mi N., 8900 ft, 9.VII.1967, S.L. W[ood], ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">religiosa</named-content></named-content></italic> (USNM-21).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>MEXICO:</bold> [<bold><italic>Distrito Federal</italic></bold>]: Desierto de los Leones National Park, III.1951, J.M. Miller, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">religiosa</named-content></named-content></italic> (OSAC-2). <bold><italic>Hidalgo</italic>:</bold> El Chico, 31.IX.[19]77, E. Hernandez V., ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">religiosa</named-content></named-content></italic> (CNCI-2). <bold><italic>Oaxaca</italic>:</bold> Valle Nacional, 53 mi S., 10000 ft, 24.V.[19]71, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">religiosa</named-content></named-content></italic> (CNCI-2). <bold><italic>Tlaxcala</italic>:</bold> Villarreal Terrenate, 23.II.[19]78, E. Hernandez, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">religiosa</named-content></named-content></italic> (USNM-4, WFBM-5).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>MEXICO: Distrito Federal, Estado de México, Hidalgo, Michoacán, Morelos, Oaxaca, Puebla, Tlaxcala (Fig. <xref ref-type="fig" rid="F40">40</xref>).</p><fig id="F40" orientation="portrait" position="float"><label>Figure 40.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33393.jpg" xlink:href="zookeys-450-001-g040"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">religiosa</named-content></named-content></italic> (Kunth) Schltdl. & Cham. (sacred fir).</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic> is a serious pest of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">religiosa</named-content></named-content></italic> in Mexico and its life cycle and habits have been well-studied (see <xref rid="B32" ref-type="bibr">Cibrián Tovar et al. 1995</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic> attacks the tops of mature trees, trunks of recently cut trees and large logging slash greater than 10.0 cm in diameter (<xref rid="B143" ref-type="bibr">Wood 1968</xref>; <xref rid="B32" ref-type="bibr">Cibrián Tovar et al. 1995</xref>). Trees attacked by <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic> exhibit red foliage at the tree top. If the population density is high, subsequent generations will colonize the tree from the top down. However, attacks rarely occur at the base and so the host tree typically survives (<xref rid="B32" ref-type="bibr">Cibrián Tovar et al. 1995</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic> has been reported to colonize the same material as <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic> (<xref rid="B143" ref-type="bibr">Wood 1968</xref>).</p><p>The female initiates the attack by constructing an entrance in bark crevices at branch nodes. The adult galleries are transverse across the grain of the wood, similar in appearance to <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> and with a central nuptial chamber (see <xref rid="B32" ref-type="bibr">Cibrián Tovar et al. 1995</xref>, figs 130–131). Eggs are laid in niches excavated on each side of the central nuptial chamber. The larval mines extend parallel to the grain of wood and are first in the cambium and later penetrate into the sapwood. Pupation occurs in the sapwood and there are two generations per year with the first generation occurring from October to May and the second from June to October (<xref rid="B32" ref-type="bibr">Cibrián Tovar et al. 1995</xref>).</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></title><p><named-content content-type="taxon-authority">Wood, 1962</named-content></p><p><xref ref-type="fig" rid="F29">Figs 29</xref><xref ref-type="fig" rid="F41">, 41</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content><named-content content-type="comment"> Wood, 1962: 81.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Both sexes of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic> strongly resemble <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> and are distinguished by the presence of a small median denticle on the apical margin of ventrite 2.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>1.8–2.9 mm long (mean = 2.2 mm; n = 20); 2.1–2.9 times as long as wide. Color red brown to dark red-brown, antennae brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly emarginate; epistomal process weakly developed, low; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing convex when viewed laterally, slightly transversely impressed just above epistoma; moderately, coarsely aciculate-punctate; aciculations converging at epistoma; punctures small, coarse; moderately, uniformly, covered by long, fine, yellow, erect hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, ovoid, setose with partial septum, three arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, weakly serrate apex; apex weakly emarginated at suture. Margin of apical edge bearing large, coarse punctures. Disc glabrous, smooth, shining; interstriae not impressed, more than twice width of striae, punctures uniseriate, smaller than those of striae; striae not impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 strongly, acutely produced, forming lip along base of ventrite 2, basal margin of ventrite 2 appearing impressed. Apical margin of ventrite 1 weakly elevated above base of ventrite 2. Ventrite 2 nearly perpendicular to ventrite 1; surface shagreened, dull, finely punctate; punctures small, fine, shallow, punctures varying in size, larger and more abundant near basal margin; surface weakly concave; apical margin armed with acute median denticle; setae short, recumbent, about three times that of a puncture in length; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 equal to combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F41" orientation="portrait" position="float"><label>Figure 41.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33394.jpg" xlink:href="zookeys-450-001-g041"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>1.7–3.0 mm long (mean = 2.3 mm; n = 20); 2.0–2.7 times as long as wide. Similar to male except epistoma feebly emarginate, epistomal process absent, frons convex when viewed laterally, weakly aciculate, setae sparser, shorter, less than width of eye; weakly transversely impressed between inner apices of eyes. Apical margin of ventrite 1 moderately elevated above base of ventrite 2. Second ventrite armed with median tumescence or minute denticle.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>138.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: male, labeled “Payson Canyon, UT, S.L Wood, 14 May 1960, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic>, BLNO 001271” (USNM). Paratypes <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic><bold>UNITED STATES:</bold><bold><italic>UTAH</italic>:</bold> [<italic>Beaver Co.</italic>]: Beaver, 22.IV.1950, S.L. Wood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (USNM-4). <italic>Utah Co.</italic>: Payson Canyon, 14.V.1960, S.L. Wood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (USNM-6), 25.VI.1962 (USNM-2).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>United States:</bold><bold><italic>Arizona</italic>:</bold><italic>Coconino Co.</italic>: Coconino National Forest, Arizona Snow Bowl, <named-content content-type="dwc:verbatimCoordinates">N35°19.593', W111°42.681'</named-content>, 9230 ft, 27.V.2010, S.M. Smith, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic> [var. <italic>arizonica</italic>] (MSUC-62). Jacob Lake, 12 mi S., 31.V.1969, W. Harwood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> branch (USNM-4). <italic>Graham Co.</italic>: Pinaleno Mountains, 15.VII.1968, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (DEBC-3). [Coronado National Forest], [Pinaleno Mountains], Mount Graham, Hospital Flat, 9050 ft, 19.VIII.1952, H.B. Leech, J.W. Green (CASC-1), 8950 ft, 3.VIII.1965, H.B. Leech (CASC-5). <italic>Yavapai Co.</italic>: Prescott, 1 mi S., 3.VIII.1962, S.L. W[ood] (USNM-1). Prescott, 9 mi S., 5 mi E., 7800 ft, 5.VI.1969, W. Harwood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (USNM-6). [<italic>Unspecified County</italic>]: Santa Catalina Mountains, 8500 ft, 31.V.1969 (DEBC-1, FSCA-4). <bold><italic>COLORADO</italic>:</bold><italic>Costilla Co.</italic>: near Fort Garland, Forbes Trinchera Ranch, VII-VIII.1976, D. Leatherman, ex. white fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic>] (CSUC-4). <italic>Huerfano Co.</italic>: near Red Wing, 16.VII.1975, D. Leatherman, ex. white fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic>] (CSUC-2). <bold><italic>NEVADA</italic>:</bold><italic>Clark Co.</italic>: Mary Jane Falls, 7900 ft, 11.VI.1969, Harwood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (USNM-7). [<italic>White Pine Co.</italic>]: Baker, S9 T13N R69E, Mount Diablo Meridian, 17.V.1917, T.O. Thatcher, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic>, LCNM 39-12 (CSUC-8, USNM-1, WFBM-1). Mount Wheeler, 19.VIII.1974, S.L. Wood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (USNM-9). <bold><italic>NEW MEXICO</italic>:</bold> [<italic>Otero Co.</italic>]: Cloudcroft, 14-26.VII.[19]49, W.B.R. Stromberg, ex. fir tree [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> sp.] (USNM-1). [<italic>Unspecified County</italic>]: Sandia Mountains, 8090 ft, 29.V.1969, S.L. W[ood], ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (USNM-3), 30.V.1969 (USNM-9). <bold><italic>UTAH</italic>:</bold><italic>Utah Co.</italic>: Provo, Payson Canyon, VIII.1964, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (EMEC-4). [<italic>Unspecified County</italic>]: Bryce Canyon National Park, Hopk. U.S. 35-043, 11.VII.1952, R. Washburn, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (EMEC-2).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>UNITED STATES: Arizona, Colorado, Nevada, New Mexico, Utah (Fig. <xref ref-type="fig" rid="F27">27</xref>).</p></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (Gord. & Glend.) Lindl. ex Hildebr. (white fir) and <named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content><named-content content-type="infraspecific-rank">var.</named-content><named-content content-type="variety">arizonica</named-content></named-content> (Merriam) Lemmon (corkbark fir). Incidental host: <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>.</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic> is an uncommonly encountered species. The species prefers to colonize the limbs, tops and slash of its host. Infested material ranges from 3.0–10.0 cm in diameter (<xref rid="B148" ref-type="bibr">Wood 1982</xref>).</p><p>The adult gallery contains two egg galleries that branch from the central nuptial chamber. One egg gallery extends from the nuptial chamber perpendicular to the grain of wood and the second gallery extends at a 45° angle to the grain (Fig. <xref ref-type="fig" rid="F24">24</xref>). Galleries are frequently initiated near branch crotches and disguised under rough patches of bark. The adult gallery deeply scores the sapwood and lightly scores the cambium. The adult galleries range in size from 1.9–6.5 cm in length. Eggs are laid in niches on both sides of each egg gallery and larval mines radiate perpendicular to the egg gallery forming a fan shaped pattern. The larval mines gradually diverge and lightly etch the sapwood. Pupation occurs in the sapwood (<xref rid="B42" ref-type="bibr">Edson 1967</xref>).</p></sec><sec sec-type="treatment-Collection notes"><title>Collection notes.</title><p>The senior author collected specimens from suppressed 2.0–4.0 cm diameter branches of <named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content><named-content content-type="infraspecific-rank">var.</named-content><named-content content-type="variety">arizonica</named-content></named-content> that had been girdled at the base by porcupines. Needles of infested branches were pale green to yellow.</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>This species is sister to <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> and is very similar both in morphology and gallery architecture. Interspecific divergence in COI nucleotide difference between these two taxa is quite large and averages 10.36% (Table <xref ref-type="table" rid="T7">7</xref>).</p><pmc-comment>PageBreak</pmc-comment>
</sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></title><p><named-content content-type="taxon-authority">Blackman, 1934</named-content></p><p><xref ref-type="fig" rid="F29">Figs 29</xref><xref ref-type="fig" rid="F42">, 42</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content><named-content content-type="comment"> Blackman, 1934: 18.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic> is a rather distinctive species and both sexes are readily distinguished by having the apical margin of ventrite 1 thickened and on the surface of ventrite 2 and by the unarmed ventrite 2.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>2.6–3.6 mm long (mean = 3.3 mm; n = 20); 2.0–2.4 times as long as wide. Color dark red-brown, antennae brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly emarginate; epistomal process weakly developed; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing flattened when viewed laterally, slightly transversely impressed just above epistoma; moderately, coarsely, longitudinally aciculate-punctate; aciculations converging at epistoma; punctures sparse, small, coarse; moderately, uniformly covered by long, fine, yellow, erect, hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, irregularly ovoid, setose with partial septum, two strongly arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, smooth apex; apex weakly emarginated at suture. Margin of apical edge bearing large, coarse punctures. Disc glabrous, smooth, shining; interstriae not impressed, more than twice width of striae, punctures uniseriate, smaller than those of striae; striae weakly impressed. Declivity bearing sparse, short, erect, yellow setae. Metepimeron less than half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 not elevated above base of ventrite 2. Basal margin of ventrite 2 strongly thickened, lip-like; ventrite 2 nearly perpendicular to ventrite 1; surface shagreened, dull, finely punctate; punctures small, fine, shallow; surface weakly concave, weakly to strongly medially impressed just above base; apical margin unarmed; setae small, less than 1 diameter of a puncture; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 equal to combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F42" orientation="portrait" position="float"><label>Figure 42.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33395.jpg" xlink:href="zookeys-450-001-g042"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>2.8–4.0 mm long (mean = 3.23 mm; n = 20); 1.6–2.7 times as long as wide. Similar to male except epistoma entire, epistomal process absent, frons convex when viewed laterally, weakly longitudinally aciculate, setae sparser, shorter, less than width of eye; weakly transversely impressed between inner apices of eyes. Second ventrite unarmed, basal margin weakly thickened.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>96.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype, male, labeled “Ashland Ore., May 20/19, Glendinning, WE Colr, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic>, Hopk. US. 13399a, Type No. 43834 USNM” (USNM). Paratypes: <bold>UNITED STATES:</bold><bold><italic>Oregon</italic>:</bold> [<italic>Jackson Co.</italic>]: Ashland, Hopk. U.S. 14246-C, 10.III.[19]16, P.D. Sergent, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> (CNCI-2); II.[19]19, W.E Glendinning, Hopk. U.S. 1399-A (USNM-14).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>UNITED STATES:</bold><bold><italic>CALIFORNIA</italic>:</bold> [<italic>Del Norte Co.</italic>]: [Six Rivers National Forest], Gasquet R.S. [= Ranger Station], Hopk. U.S. 31722-H, 10.VII. R.L. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> (OSAC-23). <italic>Lake Co.</italic>: Middletown, 11.XI.1959, G.M. Thomas, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (OSAC-1). <italic>Los Angeles Co.</italic>: Angeles National Forest, Sawmill Mountain, <named-content content-type="dwc:verbatimCoordinates">34.6926°N, 118.5499°W</named-content>, V.28-VI.14.2007, Caterino, Chatzimanolis, ex. Lindgren trap (SBMN-4). <italic>Marin Co.</italic>: Alpine Lake, V.[19]57 (CASC-4). Mount Tamalpais 14.IX.[19]57, E.L. Smith, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (OSAC-1). Woodacre Creek, 9.VII.1951, P.S. Bartholomew (CASC-1). <italic>Napa Co.</italic>: Angwin, 2 mi N.N.E., N. side of Howell Mountain, 1300 ft, 16.VII.1974, H.B. Leech, ex. emerged from log of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (CASC-1), 21.VII.1974 (CASC-4, USNM-5), 22.VII.1974 (CASC-1), 25.VII.1974 (CASC-4, USNM-1), 7.VIII.1983 (CASC-1), 12.IX.1983 (CASC-2). Callistoga, 5.X.1947, S.L. Wood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> (USNM-2). Mount Saint Helena, Hopk. U.S. 15401-A, F.B. Herbert, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> (OSAC-1). <italic>Riverside Co.</italic>: Santa Barbara National Forest, Black Mountain Rd, <named-content content-type="dwc:verbatimCoordinates">33.8395°N, 116.7306°W</named-content>, 1.VII.2005, M. Caterino (SBMN-1). [<italic>Valyermo Co.</italic>]: Fenner Canyon, Hopk. U.S. 33853-A, 15.VIII.[19]51, A.D. Moore, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">macrocarpa</named-content></named-content></italic> (EMEC-3, OSAC-8, USNM-4). <bold><italic>OREGON</italic>:</bold><italic>Benton Co.</italic>: Marys Peak, Corvallis Watershed, 9.VIII.1963, ex. rotary traps (EMEC-1). <italic>Jackson Co.</italic>: Ashland, Hopk. U.S. 13363-B, 26.VI.1918, W.E.G, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic>, (OSAC-14). Mistletoe, Hopk. U.S. 15753-A, P.D. Sergent, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (OSAC-3). Rogue River National Forest, Rogue River Gorge viewpoint, <named-content content-type="dwc:verbatimCoordinates">N42°54.540', W122°26.733'</named-content>, 3489 ft, 21.VIII.2010, S.M. Smith, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-3). <italic>Linn/Lane Co.</italic>: Blue River, 11 mi N.E., H.J. Andrews experimental forest, 5.VIII.1986, Log Decomp Study, Site 2, SE1/4 S15 T15S RSE, Trap 2WA (OSAC-1). [<italic>Tillamook Co.</italic>]: Woods, 18.XI.[19]38 (OSAC-1). <italic>Wasco Co.</italic>: The Dalles, ODA Port/Mill survey, Trap #65-01a, 14.VIII.1997, ex. Lindgren funnel with α-pinene & ethanol lure (MSUC-1).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>UNITED STATES: California, Oregon, Washington (Fig. <xref ref-type="fig" rid="F30">30</xref>).</p></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (Mirb.) Franco (= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> Britton) (Douglas fir) and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">macrocarpa</named-content></named-content></italic> (Vasey) Mayr (bigcone Douglas fir).</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic> colonizes large limbs and tops of its host and also fresh slash (<xref rid="B42" ref-type="bibr">Edson 1967</xref>; Smith, pers. obs.).</p><p>The adult gallery is typically constructed parallel to the grain of the wood and has a central nuptial chamber (Fig. <xref ref-type="fig" rid="F24">24</xref>). The gallery structure is typically bayonet shaped but may also be longitudinal. Each egg gallery extends in opposite directions to the grain of the wood from the central nuptial chamber. The nuptial chamber is transverse to the egg galleries. The adult gallery deeply scores the sapwood and lightly scores the <pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment>cambium. The adult gallery averages 6.0–18.0 cm in length. Egg niches are closely spaced and deeply score the sapwood. Larvae extend their mines perpendicular to the egg gallery in a fan shaped pattern before terminating in pupation chambers, which are constructed in the sapwood (<xref rid="B42" ref-type="bibr">Edson 1967</xref>).</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>Specimens of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic> are very rarely collected and the species is perhaps the least common of the conifer-feeders in the United States. There are many gaps that occur in its distribution range, particularly between northern and southern California. Considerable variation is observed in the male ventrite 1 and 2 across the geographic range especially between northern California, Oregon and Washington and southern California. Southern California populations colonize <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">macrocarpa</named-content></named-content></italic> while individuals from the rest of the range colonize <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>. In addition, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic> males from California and particularly southern California have a greater impression of the second ventrite and thicker margin between ventrite 1 and 2 to specimens from Oregon and Washington.</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></title><p><named-content content-type="taxon-authority">(Swaine, 1910)</named-content></p><p><xref ref-type="fig" rid="F29">Figs 29</xref><xref ref-type="fig" rid="F43">, 43</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Eccoptogaster</named-content><named-content content-type="species">piceae</named-content></named-content><named-content content-type="comment"> Swaine, 1910: 34.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content><named-content content-type="comment"> (Swaine, 1910): <xref rid="B12" ref-type="bibr">Blatchley and Leng 1916</xref>: 589.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Both sexes are easily diagnosed by the large conical median spine on the surface of the second ventrite, the spine base never touches any margin, by the lack of lateral denticles on the apical margins of ventrites 2–4 and by the unicolorous elytra.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>2.5–3.0 mm long (mean = 2.4 mm; n = 12); 2.3–2.7 times as long as wide. Color red-brown to dark red brown, antennae yellow-brown, legs dark red-brown to yellow brown apically. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly emarginate; epistomal process present, moderately developed, low; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing flattened when viewed laterally from epistoma to vertex, slightly transversely impressed just above epistoma to inner apices of eyes; moderately, coarsely longitudinally aciculate-punctate; aciculations converging at epistoma; punctures large, dense, coarse; moderately, uniformly covered by long, fine, yellow, erect, hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, irregularly ovoid, setose with partial septum, two arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, smooth apex; apex weakly emarginated at suture. Margin of apical edge bearing large, coarse punctures. Disc glabrous, smooth, shining; interstriae weakly impressed, more than twice width of striae, punctures uniseriate, smaller than those of striae; striae weakly impressed. Declivity bearing sparse, short, erect, yellow setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 rounded, marked by weak carina on ascendant part of venter, more strongly marked laterally. Ventrite 2 nearly perpendicular to ventrite 1; surface reticulate, shagreened, dull, finely punctate; punctures small, fine, shallow punctures; covered with sparse setae that are about twice size of a puncture; surface convex; armed with large conical median spine, apex rounded; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 less than combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F43" orientation="portrait" position="float"><label>Figure 43.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33396.jpg" xlink:href="zookeys-450-001-g043"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>2.3–3.0 mm long (mean = 2.3 mm; n = 12); 2.3–2.7 times as long as wide. Similar to male except epistoma feebly emarginate, epistomal process weakly developed, frons convex when viewed laterally, weakly longitudinally aciculate, less coarsely punctate, setae sparser, shorter, less than width of eye; weakly transversely impressed just above epistoma. Second ventrite armed with smaller, rounded, median spine.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>467.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Lectotype: male, labeled “St. Anne’s, Que., July 21, 1907” (CUIC). Lectotype designated <xref rid="B15" ref-type="bibr">Bright 1967</xref>: 674. Paralectotype (<bold>here designated</bold>) <italic><named-content content-type="taxon-name"><named-content content-type="genus">Eccoptogaster</named-content><named-content content-type="species">piceae</named-content></named-content></italic>, female, <bold>CANADA:</bold><bold><italic>QUEBEC</italic>:</bold> St. Anne’s, 21.VII.1907 (CUIC-1). Paratypes: <bold>CANADA:</bold><bold><italic>QUEBEC</italic>:</bold> Hudson, 191[sic!] (CUIC-4).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>CANADA:</bold><bold><italic>ALBERTA</italic>:</bold> Banff (CNCI-2); IX.1916 (CNCI-1); 7.IX.1967, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content></named-content></italic> (CNCI-1). Cypress Hills, 16.VII.[19]32 (CNCI-3). Edmonton, 16.I.1916, Carr (CNCI-1), 27.VIII.1916 (CNCI-1), 28.VIII.1916 (CNCI-1), 5.VII.1924, (CUIC-1); 6.XII.1916 (CASC-2); 15.VII.[19]24 (CASC-2); 29.VI.[19]16, J.W. Green (CASC-3), 2.IX.[19]16 (CASC-1). [Jasper National Park], Athabasca Falls, 9.VIII.1919, J.M. Swaine (CNCI-5). Medicine Hat, 2.IX.[19]26 (CNCI-2). Seebe, 7.VII.1966, D.F.J. Hilton, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content></named-content></italic> (CNCI-8). <bold><italic>BRITISH COLUMBIA</italic>:</bold> Hixon, 9.VII.1972, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content></named-content></italic> sp. (CNCI-2). [Big Creek Provincial Park], Lorna [Lake], 6.VII.1924, G. Hopping, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">engelmannii</named-content></named-content></italic> (CASC-1), 10.VIII.1924 (CASC-1), 22.VII.1925 (CASC-1), 14.VII.1926, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content><named-content content-type="species">contorta</named-content></named-content></italic> (CASC-1), 9.VII.1926 (CASC-1); 10.VII.1926, H. Richmond ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">engelmannii</named-content></named-content></italic> (CNCI-1), 14.VII.1926 (CNCI-1). Pine Pass, 11.VII.1972, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content></named-content></italic> sp. (CNCI-6). Trinity Valley, 21.VII.[19]30 (CNCI-1), 29.VII.[19]30 (CNCI-1), 13.VII.1928, J.R. Howell, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> (CASC-1); 23.VII.[19]28, H. Richmond, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">engelmannii</named-content></named-content></italic> (CASC-1). <bold><italic>MANITOBA</italic>:</bold> Aweme, 20.IX.[19]06, E. Criddle (CNCI-1); 1.VII.1916, N. Criddle, (CNCI-2) ex. bred from spruce [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content></named-content></italic> sp.], 2.VII.1916 (CNCI-4); 10.VII.[19]16 (CNCI-10) ex. spruce [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content></named-content></italic> sp.], 27.VII.[19]10 (CNCI-20), 14.VI.1918 (CNCI-2), 15.VI.1918 (CNCI-3); 5.VII.1916 (CNCI-3); 8.VII.1916 (CNCI-6), 9.VII.1916 (CNCI-1); 31.VII.1916 (CNCI-1); 9.VIII.1916 (CNCI-1); 10.IX.1917 (CNCI-10); 11.IX.1917 (CNCI-6); 9.VI.19[18] (CNCI-1). Brandon, 28.V.1940, L. Peterson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">canadensis</named-content></named-content></italic> (CNCI-2). Gillam, 21.VII.950, W.J. Brown (CNCI-1). Glen Souris, 5.VI.[19]23, N. Criddle (CNCI-1). Grass River Provincial Park, 27.VII.1972, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content><named-content content-type="species">murrayana</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content><named-content content-type="species">contorta</named-content><named-content content-type="subspecies">murrayana</named-content></named-content></italic>] (CNCI-7). Onah, 11.VII.1924, R.M. White (CNCI-3); N. Criddle, 25.VIII.[19]72 (CNCI-1). Telford, 24.VII.1963, ex. life table plot (CNCI-1), 2.VIII.1963 (CNCI-1). Winnipeg, Hanham (USNM-1). <bold><italic>NEW BRUNSWICK</italic>:</bold> Kouchibouguac National Park, VIII.1977, D.E. Bright, Code-6224J (CNCI-1). McGraw Brook, 10 mi W., 7.VII.[19]70, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content></named-content></italic> (CNCI-1). <bold><italic>NEWFOUNDLAND</italic>:</bold> Deer Lake, 12 mi N.E., 23.VII.[19]70, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">mariana</named-content></named-content></italic> (CNCI-11). <bold><italic>NOVA SCOTIA</italic>:</bold> Cape Breton Highlands National Park, MacIntosh Brook, PG703866, 21.VII.1983, D.E. & J.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content></named-content></italic> (CNCI-7); 29.VII.1983, McKenzie, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content></named-content></italic> sp. (CNCI-10). Kejimkujik National Park, 16-17.VII.1967, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content></named-content></italic> (CNCI-14). <bold><italic>ONTARIO</italic>:</bold> Carp, 5.VIII.[19]66, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">mariana</named-content></named-content></italic> (CNCI-2). Lake of the Woods, 1.VIII.1972, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content></named-content></italic> sp. (CNCI-7). Ottawa, 24.VI.1913, J.M. Swaine, ex. crawling on dying <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content></named-content></italic> sp. (CNCI-1). Rainy River District, 3.VIII.[19]24, J.F. Brimley (CNCI-1). <bold><italic>QUEBEC</italic>:</bold> Aylmer, 12.VII.1924, B.B. Watson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content></named-content></italic> (CNCI-1); 30.VII.1924, A.R. Graham (CNCI-1). Hudson, 191[sic!] (CNCI-52). Limbor/Touranine, 27.VI.1974, R. Sexton (CNCI-1). [Gaspésie National Park], Mount Albert, 28.VII.1954, W.J. Brown, ex. north base 650 ft (CNCI-1). St. Anne’s [Sainte-Anne-de-Beaupré], 19.VI.[19]19 (CASC-2, CNCI-42). South March, 19.VI.1958, S.D. Hicks (CNCI-1). <bold><italic>SASKATCHEWAN</italic>:</bold> Big River, 23.VII.1972, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content></named-content></italic> (CNCI-1). Big River, 40 mi N.W., 22.VII.[19]72, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content></named-content></italic> (CNCI-1). Canoe Lake, 21.VII.1972, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content></named-content></italic> (CNCI-2). Christopher Lake, 5.VIII.1959, A. & J. Brooks (CNCI-1). Cypress Hills, 3.IX.1967, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content></named-content></italic> (CNCI-9); 24.IX.1964 (CNCI-2). <bold><italic>YUKON</italic>:</bold> Old Crow, 28.VI.[19]81, D.E. Bright (CNCI-5). <bold>UNITED STATES:</bold><bold><italic>ALASKA</italic>:</bold> [<italic>North Slope Borough</italic>]: Prudhoe Bay Rd, 8 mi N., South Fork Koyukuk River, <named-content content-type="dwc:verbatimCoordinates">67°13’N, 150°07’W</named-content>, 1000 ft [sic!], 8.VII.1978, Smetana, Campbell (CNCI-1). <italic>Fairbanks North Star Borough</italic>: Fairbanks, 2.VIII.[19]55, W.F. McCambridge, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content></named-content></italic> (DEBC-7). <bold><italic>CALIFORNIA</italic>:</bold><italic>Siskiyou Co</italic>: Callahan, 7 mi N.W., 16.VI.[19]63, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">engelmannii</named-content></named-content></italic> (DEBC-9, EMEC-6). Happy Camp, 18 mi N., 31.VII.[19]63, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">engelmannii</named-content></named-content></italic> (DEBC-4, EMEC-7). <bold><italic>COLORADO</italic>:</bold><italic>Denver Co.</italic>: Denver, Union Pacific, 14.IX.2000, USDA APHIS, ex. Lindgren funnel APEtOH Lure (CSUC-1). <italic>Larimer Co.</italic>: Livermore, 10.IX.2009, K. Smith, USDA APHIS, ex. Lindgren funnel APEtOH Lure, (CSUC-1). Roosevelt National Forest, Chambers Lake, 11.VIII.[19]68, 9200ft, L.A. Kelton (CNCI-1). <bold><italic>MAINE</italic>:</bold> [<italic>Oxford Co.</italic>]: Wilsons Mills, 9.VIII.[19]70, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content></named-content></italic> sp. (CNCI-1). <bold><italic>MASSACHUSETTS</italic>:</bold> [<italic>Norfolk Co.</italic>]: Dover, 8.VII.1933, C.W. Collins, ex. reared from blue spruce [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">pungens</named-content></named-content></italic>] (USNM-11). <bold><italic>MINNESOTA</italic>:</bold><italic>Cook Co.</italic>: Superior National Forest, Hwy 12 nr. Seagull Guard Station, <named-content content-type="dwc:verbatimCoordinates">48°6'29"N, 90°50'12"W</named-content>, 23.VI–9.VII.2003, K.J.K. Gandhi, ex. Lindgren funnel trap in <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content><named-content content-type="species">banksiana</named-content></named-content></italic>, wind-disturbed-salvaged-logged, site D (CASC-1). <italic>Mille Lacs Co.</italic>: 2.VII.[19]36, H.R. Dodge, ex. under bark of tamarack tree [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content></named-content></italic> sp.] (CASC-2). <bold><italic>MONTANA</italic>:</bold> Glacier National Park, 15.VII.[19]29 (CUIC-1). <bold><italic>NEW MEXICO</italic>:</bold> [<italic>Taos Co.</italic>]: Red River, 3 mi W., C.C. Hoff (AMNH-1). <bold><italic>NORTH DAKOTA</italic>:</bold><italic>Bottineau Co.</italic>: S30 T162 R75, Hopk. U.S. 56561-A, 27.V.1973, A.D. Tagestad, ex. collected from <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content><named-content content-type="subspecies">densata</named-content></named-content></italic> (USNM-3). <italic>Rolette Co.</italic>: S15 T162 R69, Hopk. U.S. 56446, 19.V.[19]72, A.D. Tagestad, ex. collected from <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">pungens</named-content></named-content></italic> (USNM-2). <bold><italic>OREGON</italic>:</bold> [<italic>Jefferson Co.</italic>]: Suttle Lake, 4 mi W., 3.IX.[19]39, Schuh, Gray, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">engelmannii</named-content></named-content></italic> (AMNH-12, FMNH-3). [<italic>Unspecified County</italic>]: Blue Mountains, 13.VII.[19]14, W.J. Chamberlain, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">engelmannii</named-content></named-content></italic> (CNCI-2). <bold><italic>SOUTH DAKOTA</italic>:</bold><italic>Lawrence Co.</italic>: near Leads, Brownsville Rd, <named-content content-type="dwc:verbatimCoordinates">N44.2922°, W103.7828°</named-content>, 5650 ft, 27.VII.2004, K.P. Dole, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content></named-content></italic> (MSUC-14). [<italic>Unspecified County</italic>]: Black Hills, 7.VII.[19]75, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content></named-content></italic> (CNCI-5). <bold><italic>UTAH</italic>:</bold> [<italic>Daggett Co.</italic>]: Mckee Draw, Ashley National Forest, 16.VI.1960, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">pungens</named-content></named-content></italic> (USNM-1). [<italic>Unspecified County</italic>]: Logan Canyon, S27 T13 NR4, 6.VII.1948, S.L. Wood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">engelmannii</named-content></named-content></italic> (USNM-8). <bold><italic>WYOMING</italic>:</bold><italic>Carbon Co.</italic>: [Medicine Bow National Forest], Mirror Lake, 4 mi W., 6.IX.2010, D.E. Bright, B.A. Barr, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">engelmannii</named-content></named-content></italic> branches (DEBC-1); [Medicine Bow National Forest] Snowy Mountains, WY130, Lake Marie, <named-content content-type="dwc:verbatimCoordinates">N41°19.965', W106°19.516'</named-content>, 3208 m, 26.VII.2011, S.M. Smith, D.E. Bright, B.A. Barr, ex. emerged 1-5.IX.2011, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">engelmannii</named-content></named-content></italic> (MSUC-36). <italic>Johnson Co.</italic>: Buffalo, 16 km SW, 20.VI.1968, S.L. W[ood], ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">engelmannii</named-content></named-content></italic> (USNM-1).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: Alberta, British Columbia, Manitoba, New Brunswick, Newfoundland, Northwest Territories, Nova Scotia, Ontario, Prince Edward island, Quebec, Saskatchewan, Yukon. UNITED STATES: Alaska, California, Colorado, Idaho, Maine, Massachusetts, Michigan, Minnesota, Montana, New Mexico, New York, North Dakota, Oregon, South Dakota, Utah, Washington, Wisconsin, Wyoming (Fig. <xref ref-type="fig" rid="F44">44</xref>).</p><fig id="F44" orientation="portrait" position="float"><label>Figure 44.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33397.jpg" xlink:href="zookeys-450-001-g044"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p>Most spruce species including <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">breweriana</named-content></named-content></italic> S. Watson (Brewer spruce), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">engelmannii</named-content></named-content></italic> Parry ex Engelm. (Engelmann spruce), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">glauca</named-content></named-content></italic> (Moench) Voss (white spruce), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">mariana</named-content></named-content></italic> (Mill.) B.S.P. (black spruce), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">pungens</named-content></named-content></italic> Engelm. (blue spruce), and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">rubens</named-content></named-content></italic> Sarg. (red spruce). This species rarely attacks <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content></named-content></italic> spp. (larch) and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">balsamea</named-content></named-content></italic> (L.) Mill (balsam fir).</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic> infests dead and dying limbs (<xref rid="B24" ref-type="bibr">Chamberlin 1939</xref>; <xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>) and is commonly found in branches of fallen spruce and suppressed limbs of standing hosts (Smith, pers. obs.).</p><p>The adult gallery is parallel to the grain of the wood and bayonet shaped (Fig. <xref ref-type="fig" rid="F24">24</xref>). From the central nuptial chamber, one egg gallery extends to the grain of the wood and the other egg gallery is slightly transversely extended and then extends parallel to the grain. The nuptial chamber is oblique to the egg galleries. The adult gallery equally scores the sapwood and cambium but in some instances may only lightly score the sapwood. Adult galleries average 5.0–8.0 cm in length. Ten to 30 egg niches are widely spaced and deeply score the sapwood. Larvae extend their mines perpendicular to the egg gallery before diverging in a fan shaped pattern. Pupation may occur under the bark or in the sapwood (<xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>). There is likely one generation per year (<xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>; <xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>).</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>The female paralectotype has been returned to CUIC from the CNCI following the repositories listed in <xref rid="B130" ref-type="bibr">Swaine (1910)</xref>.</p><pmc-comment>PageBreak</pmc-comment>
<pmc-comment>PageBreak</pmc-comment>
</sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></title><p><named-content content-type="taxon-authority">LeConte, 1876</named-content></p><p><xref ref-type="fig" rid="F45">Figs 45</xref><xref ref-type="fig" rid="F46">, 46</xref><xref ref-type="fig" rid="F47">, 47</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content><named-content content-type="comment"> LeConte, 1876: 373.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content><named-content content-type="comment"> Blackman, 1934: 21. <bold>syn. n.</bold></named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content><named-content content-type="comment"> Blackman, 1934: 20. <bold>syn. n.</bold></named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> is a morphologically variable species across its range. Both sexes most closely resemble <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic> and are differentiated by the absence of a median tubercle on the apical margin of ventrite 2 in both sexes. The female is differentiated from that of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic> by the presence of a weakly developed and almost indistinct epistomal process and smaller size.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>1.8–3.2 mm long (mean = 2.5 mm; n = 20); 2.0–2.9 times as long as wide. Color dark red-brown to black, antennae yellow-brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly emarginate; epistomal process absent; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing convex when viewed laterally, slightly transversely impressed just above epistoma and along median line to upper level of eyes (may not be impressed in some specimens); moderately, coarsely, aciculate-punctate; aciculations converging at epistoma; punctures small, coarse; moderately, uniformly covered by long, fine, yellow, erect, hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, ovoid, setose with partial septum, two arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, smooth apex; apex weakly emarginated at suture. Margin of apical edge bearing large, coarse punctures. Disc glabrous, smooth, shining; interstriae not impressed, more than twice width of striae, punctures uniseriate, smaller than those of striae; striae not impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 strongly, acutely produced forming lip along base of ventrite 2, basal margin of ventrite 2 appearing impressed. Ventrite 2 nearly perpendicular to ventrite 1; surface glabrous, shagreened, finely punctate (obscurely punctate in some specimens); punctures small, fine, shallow; surface weakly concave; apical margin unarmed or armed with a longitudinal carina and blunt tubercle, appearing keel-shaped or a low median longitudinal carina; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 equal to combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F45" orientation="portrait" position="float"><label>Figure 45.</label><caption><p>Male venters. <bold>A</bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic><bold>B</bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> abietis phenotype <bold>C</bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> opacus phenotype <bold>D</bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic><bold>E</bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content><named-content content-type="subspecies">wickhami</named-content></named-content></italic> phenotype (from <xref rid="B42" ref-type="bibr">Edson 1967</xref>).</p></caption><graphic id="oo_33398.jpg" xlink:href="zookeys-450-001-g045"></graphic></fig><fig id="F46" orientation="portrait" position="float"><label>Figure 46.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33399.jpg" xlink:href="zookeys-450-001-g046"></graphic></fig><fig id="F47" orientation="portrait" position="float"><label>Figure 47.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> abietis phenotype <bold>A</bold> lateral male habitus <bold>B</bold> male venter <bold>C</bold> male venter oblique <bold>D</bold> lateral female habitus <bold>E</bold> female venter <bold>F</bold> female venter oblique <bold>G</bold> lateral opacus phenotype male habitus <bold>H</bold> opacus phenotype male venter <bold>I</bold> opacus phenotype male venter oblique; opacus phenotype female <bold>J</bold> opacus phenotype lateral female habitus <bold>K</bold> opacus phenotype female venter <bold>L</bold> opacus phenotype female venter oblique.</p></caption><graphic id="oo_33400.jpg" xlink:href="zookeys-450-001-g047"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>1.7–3.5 mm long (mean = 2.69 mm; n = 20); 2.0–3.0 times as long as wide. Similar to male except epistoma feebly emarginate, epistomal process absent, frons more strongly convex when viewed laterally, weakly aciculate, setae sparser, shorter, less than width of eye; entirely convex to weakly transversely impressed just above epistoma. Apical margin of ventrite 1 weakly elevated above base of ventrite 2. Apical margin of ventrite 2 unarmed or weakly longitudinally tumescent.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>627.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Lectotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> LeConte: female, labeled “Cali. [California, Calaveras, Crotch Coll], Type 967” (MCZC). Lectotype designated <xref rid="B148" ref-type="bibr">Wood 1982</xref>: 439. Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic> Blackman: male, labeled “8637iss [Sandpoint, Idaho], 6-20-[18]99, [A.D. Hopkins], Hopk. US 224, Type No. 43837” (USNM). Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content></italic> Blackman: male, labeled “Ouray, Colo[rado], 7,500-8,000 ft., July 1-15, [18]97, HF Wickham, Type No. 43836” (USNM).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>CANADA:</bold><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><bold><italic>BRITISH COLUMBIA</italic>:</bold> [Big Creek Provincial Park], Lorna [Lake], 5.VIII.1926, H. Richmond, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic> (CASC-2). Duncan, Genoa Bay, 16.VIII.1928, W.G. Mathers, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (CNCI-10), 17.VIII.1928 (CASC-3). <bold>UNITED STATES:</bold><bold><italic>CALIFORNIA</italic>:</bold><italic>Alpine Co.</italic>: Humboldt-Toiyabe National Forest, Hwy 88, <named-content content-type="dwc:verbatimCoordinates">N38°45.824', W119°51.498'</named-content>, 6262 ft, 24.VII.2010, S.M. Smith, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (MSUC-3). <italic>El Dorado Co.</italic>: Georgetown, 10 mi E., University of California Blodgett [Experimental] Forest, VI.[19]62, R.W. Stark, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (DEBC-3, EMEC-8); 30.V.1986, K. Hobson (EMEC-2), 1-2.VI.1986 (EMEC-2), 21-24.VI.1986 (EMEC-2), 1-7.VII.1986 (EMEC-9), 9-16.VII.1986 (EMEC-3); 2.VI.2003, K. Apigian (EMEC-2), 5.VI.2003 (EMEC-1). Ice House Reservoir, 25.V.2007, A.I. Cognato (MSUC-5). [Lake Tahoe], Fallen Leaf Lake, 6300 ft, 5.VII.1940, H.T. Reynolds (EMEC-2). Pacific House, 0.7 mi E., 2.VII.1989, F.G. Andrews, ex. flume (CSCA-1). Pacific House, 1 mi E., 1.V.1994, C.B. Barr, ex. seining El Dorado ditch (EMEC-1). South Lake Tahoe, 16.VI.1988, D. Adams, ex. reared from <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (CSCA-1). [<italic>Fresno Co.</italic>]: Huntington Lake, 23.VII.[19]19, F.C. Clark (CASC-52). Shaver [Lake], 14.IX.[19]08, Miller, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (CNCI-3). [<italic>Madera Co.</italic>]: North Fork, 22.VI.1935, R.P. Allen (EMEC-1). <italic>Marin Co.</italic>: Inverness, 8.VIII.[19]62, C.A. Toschi (EMEC-1). <italic>Mariposa Co.</italic>: Yosemite Valley, 7.VII.1921 (CASC-21). <italic>Mendocino Co.</italic>: Noyo River, VI.1896 (CASC-2). <italic>Modoc Co.</italic>: Alturas, Knox Mountain, 8.VII.1964, D.L. Dahlsten (EMEC-1). Warner Mountains, 4.VII.1919, G. Hopping, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (CASC-4, USNM-2). <italic>Monterey Co.</italic>: 30.VII.1964, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">bracteata</named-content></named-content></italic> (CASC-8). Carmel Valley, 15 mi S., 22.VI.1963, C.J. Wray, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">bracteata</named-content></named-content></italic> (CNCI-7, DEBC-1, EMEC-5). Williams Canyon, Los Padres National Forest, 24.IV.1992, D.E. Bright, G. Ferrell, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">bracteata</named-content></named-content></italic> limbs (CNCI-22). <italic>Nevada Co.</italic>: Donner Memorial State Park, 10.IX.[19]87, S. Seybold, ex. collected on <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (CNCI-2, EMEC-1), 11.IX.[19]87 (CNCI-1, EMEC-1), 29.X.1987 (CNCI-2, EMEC-2). Nevada City, 28.V.1939, R.P. Allen (CNCI-5). <italic>Riverside Co.</italic>: Mount San Jacinto State Park, <named-content content-type="dwc:verbatimCoordinates">33.807°N, 116.654°W</named-content>, 15.VII.2003, M. Caterino (SBMN-1). Santa Rosa Mountain, 15.IX.[19]56, D.E. Bright, D.N. King, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (EMEC-4). <italic>San Bernardino Co.</italic>: San Bernardino National Forest, E. of Arrowbear Lake, <named-content content-type="dwc:verbatimCoordinates">34.2076°N, 117.0584°W</named-content>, 29.V.2004, M. Caterino (SBMN-2); S. Fork Trail, <named-content content-type="dwc:verbatimCoordinates">34.1297°N, 117.8426°W</named-content>, 28.V.2004, M. Caterino (SBMN-1). San Bernardino Mountains, Dollar Lake trail, 10.VII.1956, R.W. Bushing (EMEC-2), 11.VII.1956 (DEBC-1, EMEC-10). <italic>Sierra Co.</italic>: Calpine, 2.3 mi N.W., 5200ft, 27.VIII.1961, H.B. Leech, ex. under bark of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> sp. (CASC-9). <italic>Siskiyou Co.</italic>: McCloud, 14.VI.1961, S.L. Wood, J.B. Karren, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (DEBC-5). Mount Shasta, 28.VII.1980, A.J. Gilbert (CASC-1). <italic>Trinity Co.</italic>: Klamath National Forest, FR 41N16, 0.2 mi E. FR 93, <named-content content-type="dwc:verbatimCoordinates">N41°14.822', W122°53.562'</named-content>, 5081ft, 28.VII.2010, S.M. Smith, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (MSUC-2). <italic>Tuolumne Co.</italic>: Cow Creek, 5 mi N. Strawberry, 18.VII.1964, C.W. O’Brien, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> sp. (CASC-102, DEBC-11). Pinecrest, Hopk. U.S. 19192-A, 29.VII.1930, G.R. Struble, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (DEBC-3); IX.1966, emerged 28.III.1967, G.T. Ferrell, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (EMEC-4). Sierra Village, 10.VII.1966, B.A. Tolden (EMEC-1). [<italic>Unspecified County</italic>]: Shasta [-Trinity] National Forest, Hopk. U.S. 21078, 10.V.1934, K.A. Salmon (DEBC-4). <bold><italic>COLORADO</italic>:</bold><italic>Clear Creek Co.</italic>: Idaho Springs, 12 mi S., 12.VII.2007, D.E. Bright, B.A. Barr, ex. bole <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic> (CSUC-2, DEBC-4). <italic>Eagle Co.</italic>: Basalt Mountain, 20.VI.1996, D. Leatherman (CSUC-1). <bold><italic>IDAHO</italic>:</bold> [<italic>Bonner Co.</italic>]: Sandpoint, 2.VII.1964, N.M. Downie (FMNH-1). <italic>Clearwater Co.</italic>: Angel Butte thinning, T37N R3E, sec 3, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> #86 (USNM-10). <italic>Kootenai Co.</italic>: Coeur d’Alene, 28.IV.1922, H.J. Rust, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (MSUC-3); 15.V.1951, 2157 ft, W.F. Barr, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (EMEC-3). <italic>Latah Co.</italic>: Moscow Mountain, 46.8042, -116.830183, 2713ft, 10.VIII.2010, S.M. Smith, [A.R. Gillogly], M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic>, emerged II.2011, M.M. Furniss (MSUC-72). <italic>Lewis Co.</italic>: Nezperce, 3 mi N., 3.VII.[19]69, R.C. Biggum (EMEC-1). <bold><italic>OREGON</italic>:</bold><italic>Deschutes Co.</italic>: Deschutes National Forest, Black Butte Rd, Black Butte, <named-content content-type="dwc:verbatimCoordinates">N44°24.924', W121°38.323'</named-content>, 4212 ft, 31.VII.2010, S.M. Smith, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (MSUC-91). Paulina Lake, 2 mi W., 12.VI.1940, Schuh, Scott, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (EMEC-2). <italic>Douglas Co.</italic>: Diamond Lake, 9.VII.[19]64, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> sp. (EMEC-7). <italic>Jackson Co.</italic>: Copper, 2 mi N., 3.IX.1970, W.G. Harwood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic>, emerged 21.X.1970 (MSUC-7). [<italic>Klamath Co.</italic>]: Keno, 16.VII.[19]29, J.A. Beal, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (CSUC-2). <italic>Linn/Lane Co.</italic>: Blue River, 11 mi N.E., H.J. Andrews Experimental Forest, 5.VIII.1988, log decomp study, site 2, SE1/4 S15 T15S RSE, trap 2WA (OSAC-1). [<italic>Umatilla Co.</italic>]: Tollgate, 6.VI.[19]49, C. Chastain, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (CSUC-3, EMEC-2, FMNH-1). <bold><italic>UTAH</italic>:</bold><italic>Utah Co.</italic>: Hobble Creek Canyon, 14.VI.[19]60, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (DEBC-2). [<italic>Unspecified County</italic>]: (CNCI-1). 1952, T.O. Thatcher (CSUC-10). <bold><italic>WYOMING</italic>:</bold><italic>Albany Co.</italic>: Snowy Mountains, [Medicine Bow National Forest], Spruce campground, 6.IX.2010, D.E. Bright, B.A. Barr coll., ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> branches (DEBC-2). [<italic>Carbon Co.</italic>]: Saratoga, Hopk. U.S. 31518-E, 1.IX.[19]38, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic> (CSUC-1).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: Alberta, British Columbia. UNITED STATES: California, Colorado, Idaho, Montana, Oregon, Utah, Washington, Wyoming (Fig. <xref ref-type="fig" rid="F48">48</xref>).</p><fig id="F48" orientation="portrait" position="float"><label>Figure 48.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33401.jpg" xlink:href="zookeys-450-001-g048"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (Gord. & Glend.) Lindl. ex Hildebr. (white fir), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (Douglas ex D. Don) Lindl. (grand fir), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic> (Hook.) Nutt. (subalpine fir) and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">bracteata</named-content></named-content></italic> (D. Don) Poit. (bristlecone fir).</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> prefers to colonize fresh slash but is also found in small limbs and tops of small standing trees (<xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>).</p><p>The adult gallery contains two egg galleries that branch from the central nuptial chamber. Typically, one egg gallery extends from the nuptial chamber perpendicular to the grain of wood and the second egg gallery extends at a 45° angle to the grain (Fig. <xref ref-type="fig" rid="F24">24</xref>). Occasionally both egg galleries are extended obliquely to the grain, at a 45° angle to the grain or perpendicular to the grain (<xref rid="B42" ref-type="bibr">Edson 1967</xref>). Galleries are frequently initiated near branch crotches and disguised under rough patches of bark. The adult gallery equally scores the sapwood and cambium. The adult galleries range in size from 2.5–6.4 cm in length. Eggs are laid in niches on both sides of each egg gallery and larval mines radiate perpendicular to the egg gallery. The larval mines gradually diverge forming a fan shaped pattern and lightly etch the sapwood. Pupation occurs in the sapwood (<xref rid="B42" ref-type="bibr">Edson 1967</xref>). There is one generation per year and broods overwinter as larvae (<xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>).</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>There has been a great deal of historical uncertainty regarding the placement of <pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content></italic> since they were described. The species are extremely morphologically similar and thus difficult to differentiate. This similarity led several authors to treat <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic> as either a synonym of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content></italic> (<xref rid="B148" ref-type="bibr">Wood 1982</xref>) or as a subspecies (<xref rid="B17" ref-type="bibr">Bright 1976</xref>). <xref rid="B46" ref-type="bibr">Equihua-Martínez and Furniss (2009)</xref> removed <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic> from synonymy with <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content></italic> based on differences in host use, submentum shape, and striations on the underside of the head and characters of the second ventrite including setae, punctures and spine shape. The species <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> were not recovered as monophyletic in any of our analyses (Figs <xref ref-type="fig" rid="F2">2</xref>, <xref ref-type="fig" rid="F8">8</xref>, <xref ref-type="fig" rid="F9">9</xref>). We observed very low intraspecific COI and ArgK nucleotide differences among populations (COI: 0–0.0049; mean = 0.0032), (ArgK: 0–0.0044; mean = 0.0024) and no differences in 28S or CAD (Table <xref ref-type="table" rid="T6">6</xref>). Considerable variation was also observed in the shape of the spine on the male second ventrite, especially within <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic>. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">abietis</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">opacus</named-content></named-content></italic> are here placed in synonymy with <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> because they were recovered as polyphyletic (Figs <xref ref-type="fig" rid="F2">2</xref>, <xref ref-type="fig" rid="F8">8</xref>, <xref ref-type="fig" rid="F9">9</xref>), there are minute molecular difference among populations sampled from California, Oregon, Idaho and Wyoming (Table <xref ref-type="table" rid="T6">6</xref>), the species have identical galleries (<xref rid="B42" ref-type="bibr">Edson 1967</xref>, <xref rid="B148" ref-type="bibr">Wood 1982</xref>) and variability of the spine on male ventrite 2.</p><p>The lectotype of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> bears a partial locality label. <xref rid="B82" ref-type="bibr">LeConte’s (1868)</xref> description states that the lectotype was collected at Calaveras, California by Mr. Crotch.</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></title><p><named-content content-type="taxon-authority">Blackman, 1934</named-content></p><p><xref ref-type="fig" rid="F45">Figs 45</xref><xref ref-type="fig" rid="F49">, 49</xref><xref ref-type="fig" rid="F50">–50</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content><named-content content-type="comment"> Blackman, 1934: 13.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">virgatus</named-content></named-content><named-content content-type="comment"> Bright, 1972: 1490. <bold>syn. n.</bold></named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content><named-content content-type="comment"> Blackman, 1934: 13. <bold>syn. n.</bold></named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Males exhibiting the reflexus phenotype are readily distinguished by the presence of a strong recurved subapical carinate ridge on ventrite 5 that is medially produced to form a slightly recurved and subrostriform ridge that appears “reflexed”. Males of this species exhibiting the wickhami phenotype are easily confused with males of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic>. They are easily distinguished by the size of male ventrite 5. In <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic>, ventrite 5 is equal in length to ventrite 4. In <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic>, ventrite 5 is equal in length to ventrites 3 and 4 combined. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> lacks an epistomal process while <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> typically has a strongly developed epistomal process. Females are distinguished from those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> by having the apical margin of ventrite 1 rounded and by the ventrite 2 surface rugose, shining, coarsely punctate and convex.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>2.4–3.8 mm long (mean = 3.2 mm; n = 20); 1.8–2.5 times as long as wide. Color dark red-brown to black, antenna brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly to acutely, deeply emarginate; epistomal process weakly to strongly developed and elevated; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing flattened when viewed laterally, slightly transversely impressed just above epistoma; moderately, longitudinally aciculate, deeply, coarsely punctate; aciculations converging at epistoma; punctures small, coarse; moderately, uniformly covered by long, fine, yellow, erect, hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, irregularly ovoid, setose with partial septum, three sharply arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical, anterolateral and lateral margins bearing sparse, erect, yellow, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, smooth apex; apex moderately emarginated at suture. Margin of apical edge bearing small, fine punctures. Disc smooth, shining; interstriae weakly impressed, more than twice width of striae, punctures uniseriate, smaller than those of striae, bearing minute, recumbent setae less than length of a puncture; striae weakly impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 rounded, marked by weak carina on vertical surface of segment. Ventrite 2 nearly perpendicular to ventrite 1; surface rugose, shagreened, finely punctate; punctures small, coarse, shallow; surface convex; setae moderately abundant, long, erect and longer than length of ventrite 3; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 typically armed with strong, recurved, subapical, carinate ridge, occasionally modified and medially produced to form a slightly recurved and subrostriform ridge appearing “reflexed” (Chiricahua Mountains, Arizona and Chihuahua, Mexico populations). Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 less than combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F49" orientation="portrait" position="float"><label>Figure 49.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33402.jpg" xlink:href="zookeys-450-001-g049"></graphic></fig><fig id="F50" orientation="portrait" position="float"><label>Figure 50.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> wickhami phenotype <bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique.</p></caption><graphic id="oo_33403.jpg" xlink:href="zookeys-450-001-g050"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>2.5–4.0 mm long (mean = 3.2 mm; n = 20); 1.87–2.5 times as long as wide. Similar to male except epistoma entire, epistomal process absent, frons convex when viewed laterally, weakly longitudinally aciculate, setae sparser, shorter, less than width of eye; weakly transversely impressed just above epistoma and between inner apices of eyes. Second ventrite unarmed, setae sparse, erect, measuring length of three diameters of a puncture. Length of ventrite 5 greater than combined lengths of ventrites 3 and 4, armed with apical strongly recurved subapical carinate ridge.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>358.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> Blackman: male, labeled “Sta Catalina Mts, Ariz., Chrisman, M. Coll, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic>, Hopk. U.S. 12210, Reared June 1-14 H.B. Kirk, Type No. 43831 USNM” (USNM). Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">virgatus</named-content></named-content></italic> Bright: male, labeled “MEX., N.L., Cerro Potosi, V.2.71, 8500', D.E. Bright, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>, CNC No. 12604” (CNCI). Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic> Blackman: male, labeled “Buena Vista, Col. H.F. Wickham July 1-6 ’96, 7,900-8,000 ft, Type No. 43832 USNM” (USNM). Paratypes <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic><bold>UNITED STATES:</bold><bold><italic>ARIZONA</italic>:</bold> [<italic>Cochise Co.</italic>]: Chiricahua Mountains, 18.VI.[?], Hubbard, Schwarz (EMEC-1). <italic>Pima/ Pinal Co.</italic>: Santa Catalina Mountains, M. Chrisman, reared 25.VI.[19]14, H.B. Kirk, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-1). Paratypes <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic><bold>UNITED STATES:</bold><bold><italic>ARIZONA</italic>:</bold> [<italic>Cochise Co.</italic>]: Chiricahua Mountains, Hopk. U.S. 5558-A, 5.VIII.[19]07, J.L. Webb, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CNCI-1). <bold><italic>NEW MEXICO</italic>:</bold> [<italic>Lincoln Co.</italic>]: Capitan Mountains, Hopk. U.S. 5674, J.L. Webb, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CNCI-1). Paratypes <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">virgatus</named-content></named-content></italic><bold>MEXICO:</bold><bold><italic>Nuevo León</italic>:</bold> Cerro Potosi, 8500 ft, 2.V.[19]71, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (CNCI-4).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>MEXICO:</bold><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><bold><italic>CHIHUAHUA</italic>:</bold> San Juanito, 16 km N.E., 50 m, 19.VII.1960, S.L. Wood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-3, USNM-7). San Juanito, Hopk. U.S. 58592, 16.III.1974, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content></named-content></italic> sp. (USNM-1, WFBM-5). <bold><italic>DURANGO</italic>:</bold> Durango, Hopk. U.S. 58685, 24.III.1974, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (USNM-14). <bold>UNITED STATES:</bold><bold><italic>ARIZONA</italic>:</bold><italic>Cochise Co.</italic>: Chiricahua Mountains, 21.VII.1916, C.R. Bruck, (DEBC-2), 29.IX.[19]47, D.J. & J.N. Knull (DEBC-4). Coronado National Forest, Chiricahua Mountains, <named-content content-type="dwc:verbatimCoordinates">N31°54.915', W109°16.040'</named-content>, 8196 ft, 20.V.2010, S.M. Smith, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-7). Huachuca Mountains, Upper Carr Canyon, 7500 ft, 6–10.VIII.[19]52, H.B. Leech, J.W. Green (CASC-1). [<italic>Coconino Co.</italic>]: Jacob Lake, Kaibab National Forest, 19.VI.[19]66, [L. Edson], ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (EMEC-3). <italic>Greenlee Co.</italic>: Hannagan [Meadows] campground, 12.VII.1968, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (CNCI-5). <italic>Pima Co.</italic>: Tucson, Mount Lemmon, 11.VI.1969, S.L. Wood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-1). <italic>Pima/Pinal Co.</italic>: Santa Catalina Mountains, 9000 ft, 6.VI.1926, R.B. Streets (CASC-1); 5.VIII.1968, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (CNCI-1); Bear Wallow, 7800 ft, 11.VI.1969, S.L. Wood (USNM-5). <italic>Santa Cruz Co.</italic>: Carr Canyon, 8.VIII.1962, S.L. Wood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (USNM-12). <italic>Yavapai Co.</italic>: Prescott National Forest, Mount Union, Lake. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (EMEC-1). <bold><italic>COLORADO</italic>:</bold> [<italic>Boulder Co.</italic>]: Boulder, Hopk. U.S. 17700-Y, 21.IV.[19]37, J.A. Beal, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CSUC-3, USNM-5). <italic>Chaffee Co.</italic>: N. of Poncha Pass, 25.VII.1997, D. Leatherman, ex. ponderosa pine [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content><named-content content-type="species">ponderosa</named-content></named-content></italic>] (CSUC-1). <italic>Clear Creek Co.</italic>: Idaho Springs, 7.5 mi S., 12.VII.2007, D. Bright, B.A. Barr, ex. bole <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (DEBC-3). Lawson, 26.VII.1994, D. Leatherman, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CSUC-4). Near Lawson, 26.VII.1994, S. Kelley, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (CNCI-2). [<italic>Garfield Co.</italic>]: Glenwood Springs, VII.[?] (CASC-1). <italic>Jefferson Co.</italic>: 1–15.VII.2010, Colorado Dept of Agriculture, ex. Lindgren funnel UHR EtOH & Ω-pinene conifer (CSUC-3). Buffalo Creek, 6.VIII.2004, D. Leatherman, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CSUC-1). <italic>Larimer Co.</italic>: Estes Park, 19.VI.[19]35, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (USNM-3); 28-VI-13.VII.2010, Colorado Dept of Agriculture, ex. Lindgren funnel UHR EtOH & Ω-pinene conifer (CSUC-3). Fort Collins, 29.VI.[19]35 (USNM-2). Mount Margaret trailhead, 4.VIII.1994, D. Leatherman, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CSUC-1). Pingree Park, 30.VIII.1995, D. Leatherman (CSUC-1). Poudre Canyon, 27.V.1975, D. Leatherman, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CSUC-1). Rist Canyon, 28.X.[19]56, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (DEBC-1); S28 T8 NR70W, 3.III.1957, T.O. Thatcher, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (DEBC-7). Rist Canyon Picnic Area, 26.VI.2008, D. Leatherman, ex. fallen Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CSUC-3). Roosevelt National Forest, Big Thompson Canyon, <named-content content-type="dwc:verbatimCoordinates">N40°24.456', W105°24.565'</named-content>, 7080 ft, 5.V.2010, S.M. Smith, D.E. Bright, B.A Barr, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-2). Red Feather Lakes, 12.III.2003, D. Leatherman, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CSUC-1), VI.2000 (CSUC-5). CR 63E, 2 mi S. off Hwy 14, 4.VII.2007, D. Leatherman, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CSUC-11). <italic>Mineral Co.</italic>: Wolf Creek Pass off US 160, Sheep Mountain, 25.VII.2005, D. Leatherman, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CSUC-1). <italic>Pueblo Co.</italic>: SR 165, 5 mi S.E. San Isabel Millset trailhead, 23.VI.2000, D. Leatherman, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CSUC-3). <italic>Teller Co.</italic>: Ridgewood Subdivision, 5.VIII.2004, D. Leatherman, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CSUC-1). <bold><italic>NEVADA</italic>:</bold> [<italic>Lander Co.</italic>]: Austin, 12.VIII.[19]40, D.E. Hardy (USNM-1). <bold><italic>NEW MEXICO</italic>:</bold><italic>Otero Co.</italic>: Cloudcroft, 4.VI.1969, tree 53, S.L. Wood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-14); 11-13.VII.[19]74, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (CNCI-4). Lincoln National Forest, Apache Point Observatory, <named-content content-type="dwc:verbatimCoordinates">N32°47.046', W105°48.841'</named-content>, 9116 ft, 15.V.2010, S.M. Smith, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-145). [<italic>Sandoval Co.</italic>]: Jemez Springs, Hopk. U.S. 37218-F, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (USNM-4); Hopk. U.S. 37214-D, 1.IX.[19]57, F.M. Yasinski, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (USNM-15). <bold><italic>TEXAS</italic>:</bold><italic>Culberson Co.</italic>: Guadalupe Mountains National Park, The Bowl, 17.VII.1974, Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (CNCI-6). <bold><italic>UTAH</italic>:</bold> [<italic>Utah Co.</italic>]: Wasatch National Forest, Mount Timpanogos, 13.VII.1957, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (DEBC-1). [<italic>Unspecified County</italic>]: Logan Canyon, 5000 ft, 31.XII.1945, S.L. Wood, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (USNM-3), 28.IV.1946 (USNM-2), 16.VI.1946 (USNM-4), 3.VII.1946 (USNM-4).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>MEXICO: Chihuahua, Durango, Nuevo León. UNITED STATES: Arizona, Colorado, Nevada, New Mexico, Texas, Utah (Fig. <xref ref-type="fig" rid="F51">51</xref>).</p><fig id="F51" orientation="portrait" position="float"><label>Figure 51.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33404.jpg" xlink:href="zookeys-450-001-g051"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (Mirb.) Franco (= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> Britton) (Douglas fir).</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> is relatively common throughout its range and is found in fresh slash and branches of Douglas fir (<xref rid="B42" ref-type="bibr">Edson 1967</xref>).</p><p>The adult gallery consists of two egg galleries and a central nuptial chamber. The adult gallery is parallel to the grain of the wood and bayonet shaped (Fig. <xref ref-type="fig" rid="F24">24</xref>). From the central nuptial chamber, one egg gallery extends parallel to the grain of the wood and the other egg gallery is slightly transversely extended and then extends parallel to the grain. The nuptial chamber is oblique to the egg galleries. The adult gallery equally scores the sapwood and cambium and averages 3.9–7.6 cm in length. Egg niches are closely spaced and score the sapwood. Larvae extend their mines perpendicular to the egg gallery before diverging in a fan shaped pattern. Pupation may occur under the bark or in the sapwood (<xref rid="B42" ref-type="bibr">Edson 1967</xref>).</p></sec><sec sec-type="treatment-Collection notes"><title>Collection notes.</title><p>The senior author has collected <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> in the Chiricahua Mountains, Arizona and Apache Point Observatory, New Mexico killing small (less than 10.0 cm DBH) Douglas fir trees. This species was also reported killing Douglas fir in the Sacramento Mountains and Santa Fe National Forest of New Mexico, and was tentatively and incorrectly identified as <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> (<xref rid="B136" ref-type="bibr">USFS 2004</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> strongly resembles <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> (see diagnosis) but does not occur in New Mexico.</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p><xref rid="B146" ref-type="bibr">Wood (1977</xref>: 388) placed <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic> in synonymy with <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic>. <xref rid="B148" ref-type="bibr">Wood (1982)</xref> removed <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> from synonymy with <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic>. In this publication he also listed <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic> as a synonym of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> and cited <xref rid="B146" ref-type="bibr">Wood (1977)</xref> for the synonymy. <xref rid="B151" ref-type="bibr">Wood and Bright (1992</xref>: 364) also cite <xref rid="B146" ref-type="bibr">Wood (1977)</xref> for the synonymy. However, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic> was never removed from synonymy with <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> and designated as a synonym of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic>.</p><p>In his description of <pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">virgatus</named-content></named-content></italic>, <xref rid="B16" ref-type="bibr">Bright (1972)</xref> posited that the species was closely related to <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic> or either a subspecies or variety. <xref rid="B148" ref-type="bibr">Wood (1982)</xref> considered <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic> a synonym of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic>, but as discussed above, never formally placed it in synonymy. All three species are here treated as one slightly morphologically variable species. We assessed intraspecific variation within these three species for four genes (Table <xref ref-type="table" rid="T6">6</xref>). It was small for each gene with the average divergence of 1.3% for COI and less than 0.12%, 0.03% and 0.29% for 28S, CAD and ArgK respectively among sampled populations (Table <xref ref-type="table" rid="T6">6</xref>). These ranges are consistent with variation observed within other species. There are two main characteristics found that vary among the populations: the male fifth ventrite carina and the male epistomal process. There are two main phenotypes observed in populations: reflexus and wickhami. The reflexus phenotype includes individuals that were previously considered to be <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> with the male ventrite 5 armed with a strong recurved subapical carinate ridge that at its crest is closer to the basal than the apical margin of the segment. The length and height of the subapical carinate ridge is also variable within populations. This character is only found in the Chiricahua Mountains, Arizona and in Mexico. The wickhami phenotype includes individuals that were formerly considered <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">virgatus</named-content></named-content></italic>. In the wickhami phenotype the process on the male fifth ventrite is reduced in the wickhami phenotype and but still forms strong recurved subapical carinate ridge. However it is important to note that both the reflexus and wickhami phenotypes are sympatric in the Chiricahua Mountains. The male epistomal process also varies from a strongly developed and elevated ridge above the epistoma to weakly developed and elevated ridge. In general, the ridge is more strongly developed in the reflexus phenotype but considerable variation is observed especially in the wickhami phenotype. Additional variation is observed in the density of punctures on the male second ventrite and the coarseness of the male frons aciculations. In addition, the gallery structure of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic> are identical; the gallery of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">virgatus</named-content></named-content></italic> is has not been noted. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">wickhami</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">virgatus</named-content></named-content></italic> are here designated as synonyms of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic>.</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></title><p><named-content content-type="taxon-authority">Blackman, 1934</named-content></p><p><xref ref-type="fig" rid="F52">Figs 52</xref><xref ref-type="fig" rid="F53">–53</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content><named-content content-type="comment"> Blackman, 1934: 19.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>The <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic> male is quite distinctive with its strongly flattened frons, apical margin of ventrite 1 strongly, acutely produced forming a lip along the base of ventrite 2, basal margin of ventrite 2 appearing impressed and by the unarmed apical margin of ventrite 2. The female is morphologically similar to that of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic>. The <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic> female is distinguished by having the apical margin of ventrite 1 forming a carinate lip along the basal margin of ventrite 2.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>2.5–4.0 mm long (mean = 3.2 mm; n = 20); 1.8–2.7 times as long as wide. Head, pronotum and abdominal venter dark red-brown, antennae light brown, legs dark red-brown to light brown, elytra red-brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma moderately emarginate; epistomal process strongly developed and elevated; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing flattened when viewed laterally, slightly transversely impressed just above epistoma; moderately, coarsely, longitudinally aciculate-punctate; aciculations converging at epistoma; punctures small, coarse; moderately and uniformly covered by long, fine, yellow erect hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, irregularly ovoid, setose with partial septum, two sharply arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, weakly serrate apex; apex moderately emarginated at suture. Margin of apical edge bearing large, coarse punctures. Disc smooth, shining; interstriae not impressed, more than twice width of striae, interstrial punctures uniseriate, smaller than those of striae, bearing minute, recumbent setae less than length of a puncture; striae weakly impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 strongly, acutely produced forming lip along base of ventrite 2, basal margin of ventrite 2 appearing impressed. Ventrite 2 nearly perpendicular to ventrite 1; surface glabrous, shining, rugose, finely punctate; punctures small, fine, shallow; surface flattened, unarmed or with weak median tumescence on apical margin; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 equal to combined lengths of ventrites 3 and 4; setal patch or median depression is absent.</p><fig id="F52" orientation="portrait" position="float"><label>Figure 52.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> male venters (from <xref rid="B42" ref-type="bibr">Edson 1967</xref>).</p></caption><graphic id="oo_33405.jpg" xlink:href="zookeys-450-001-g052"></graphic></fig><fig id="F53" orientation="portrait" position="float"><label>Figure 53.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33406.jpg" xlink:href="zookeys-450-001-g053"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>2.2–4.1 mm long (mean = 3.2 mm; n = 20); 1.9–2.5 times as long as wide. Similar to male except epistoma feebly emarginate, epistomal process less strong<pmc-comment>PageBreak</pmc-comment>ly developed and elevated, frons convex when viewed laterally, weakly longitudinally aciculate, setae sparser, shorter, less than width of eye; weakly transversely impressed between inner apices of eyes. Second ventrite unarmed.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>143.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic> Blackman: male, labeled “Prescott N.F. Ariz., VII-24-[19]30, M.W. Blackman Collector, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic>, Hopk. US 20410E, Type No. 43835 USNM” (USNM).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>UNITED STATES:</bold><bold><italic>ARIZONA</italic>:</bold> [<italic>Coconino Co.</italic>]: Grand Canyon North Rim, Kaibab National Forest, VI.[19]66, [L. Edson] (EMEC-4). [<italic>Graham Co.</italic>]: Pinaleno Mountains, Swift Trail, 3 mi, Lady Bug Saddle, 11.IX.1964, C.W. O’Brien, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> sp. (CASC-1, CNCI-4, EMEC-2); Coronado National Forest, SR 366, <named-content content-type="dwc:verbatimCoordinates">N32°37.702', W109°49.472'</named-content>, 7896 ft, 24.V.2010, S.M. Smith, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (MSUC-2). <bold><italic>COLORADO</italic>:</bold><italic>Costilla Co.</italic>: near Fort Garland, Forbes Trinchera Ranch, VII-VIII.1976, D. Leatherman ex. white fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic>] (CSUC-1). <italic>Huerfano Co.</italic>: near Red Wing, 16.VII.1975, D. Leatherman, ex. white fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic>] (CSUC-4). Pass Creek, 20.VIII.1975, D. Leatherman, ex. white fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic>] (CSUC-2). <italic>La Plata Co.</italic>: Columbine Lake, 11.VIII.2004, D. Leatherman, ex. subalpine fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic>] (CSUC-1). <bold><italic>NEVADA</italic>:</bold><italic>Clark Co.</italic>: 19.VII.[19]29, C.C. Searl (DEBC-1, EMEC-4). Cathedral Rock, emerged 7-9.V.2002, R. Turnbow, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> sp. (WFBM-2). <italic>White Pine Co.</italic>: Baker, 17.V.1939, T.O. Thatcher (CNCI-2). <bold><italic>NEW MEXICO</italic>:</bold><italic>Bernalillo Co.</italic>: Cibola National Forest, Sandia Peak, 9.VII.[19]68, D.E. Bright (CNCI-1); NM536, <named-content content-type="dwc:verbatimCoordinates">N35°12.853', W106°24.743'</named-content>, 8753 ft, 10.V.2010, S.M. Smith, A.I. Cognato, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (MSUC-15), <named-content content-type="dwc:verbatimCoordinates">N35°11.655', W106°24.075'</named-content>, 8317 ft (MSUC-30). [<italic>Santa Fe Co.</italic>]: Santa Fe, Little Teseque Canyon, 14.VI.[19]35, Van Dyke (CASC-3). [<italic>Unspecified County</italic>]: Sandia Mountains, tree 9, 29.V.1969, S.L. Wood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (MSUC-13), tree 18, 30.V.1969 (MSUC-19). <bold><italic>TEXAS</italic>:</bold> [<italic>Unspecified county</italic>]: San Antonio, Hopk. U.S. 3938, W.F. Fiske, ex. bred 15.VI.[19]07 (USNM-3). <bold><italic>UTAH</italic>:</bold><italic>Cache Co.</italic>: Logan (CNCI-1). [<italic>San Juan Co.</italic>]: Mount Navajo, 8500 ft, 10.VI.[19]36, McAbee (CASC-4). <italic>Utah Co.</italic>: Payson Canyon, 14.V.1960, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (CNCI-3); 20.V.1961, S.L. Wood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (USNM-9). [<italic>Unspecified County</italic>]: Bryce Canyon National Park, 10.V.1981, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (WFBM-10). <bold><italic>UNSPECIFIED STATE</italic>:</bold> (CASC-1).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>UNITED STATES: Arizona, Colorado, Nevada, New Mexico, Texas, Utah (Fig. <xref ref-type="fig" rid="F34">34</xref>).</p></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p>Principal host: <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (Gord. & Glend.) Lindl. ex Hildebr (white fir). Incidental host: <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic> (Hook.) Nutt (subalpine fir).</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic> is found in the limbs and tops of its host and fresh slash. Galleries are often initiated near limb bases (<xref rid="B42" ref-type="bibr">Edson 1967</xref>).</p><p>The adult gallery contains two egg galleries that branch from the central nuptial chamber. Gallery shape is quite variable and ranges from ‘S’ shaped at an oblique angle to the grain to nearly perpendicular against the grain (Fig. <xref ref-type="fig" rid="F25">25</xref>). Typically one egg gallery obliquely extends from the central nuptial chamber against the grain of wood for a short distance before becoming apically recurved. The second egg gallery is identical to the first but runs in the opposite direction. However galleries may extend perpendicular <pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment>to the grain of the wood or may not be apically recurved. The adult gallery deeply scores the sapwood and ranges in size from 2.5–7.0 cm in length. Egg niches are distinct, score the sapwood and are closely spaced along the egg galleries. Larval mines rapidly diverge perpendicular to the adult gallery and are parallel to the grain of the wood. Larval mines terminate in pupation chambers that score the sapwood (<xref rid="B42" ref-type="bibr">Edson 1967</xref>).</p></sec><sec sec-type="treatment-Collection notes"><title>Collection notes.</title><p>The senior author collected this species from 10.0 cm diameter slash with thick (5.0 mm) bark.</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>A relatively large amount of intraspecific variation (0.0537) was observed in mitochondrial COI sequences from specimens collected from Arizona and New Mexico and low variation observed between the two Arizona populations (Table <xref ref-type="table" rid="T6">6</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic> only occurs on <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic>. In southeastern Arizona, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> spp. and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic> are confined to high elevation sites on the sky islands. These Arizona populations are isolated from the New Mexico populations by large areas that are unsuitable for <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic>. No specific geographic morphological differences were observed among the examined specimens.</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></title><p><named-content content-type="taxon-authority">Bright, 1972</named-content><named-content content-type="taxon-status">valid sp.</named-content></p><p><xref ref-type="fig" rid="F54">Fig. 54</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content><named-content content-type="comment"> Bright, 1972: 1489.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>The male is distinguished from other species by having the apical margin of ventrite 4 thickened, forming a broad carina with a blunt median tubercle. The female is distinguished from the morphologically similar male of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic> by having the apical margin of ventrite 1 produced, forming a carinate lip along the basal margin of ventrite 2 that is about half as produced as thick and by the host and is distinguished from the <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic> female by having a strongly developed epistomal process.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>3.0 mm long (mean = 3.0 mm; n = 1); 2.1 times as long as wide. Color dark red-brown to black. Pronotum same color as elytra.</p><p><italic>Head.</italic> Epistoma moderately, broadly emarginate; epistomal process weakly developed; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing flattened when viewed laterally, slightly transversely impressed just above epistoma; moderately, coarsely, aciculate-punctate; aciculations converging at epistoma; punctures small, coarse; moderately, uniformly covered by long, fine, erect, yellow-brown, hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, almost subquadrate, setose with partial septum, two broadly arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, dark yellow-brown setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, smooth apex; apex moderately emarginated at suture. Margin of apical edge bearing small, fine punctures. Disc smooth, shining; interstriae not impressed, more than twice width of striae, punctures uniseriate, smaller than those of striae; bearing sparse, recumbent, long, dark yellow-brown setae; striae weakly impressed. Declivity bearing abundant, long, erect dark yellow-brown hair-like setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 moderately elevated above base of ventrite 2, ventrite 2 appearing impressed. Ventrite 2 nearly perpendicular to ventrite 1; surface smooth, shining, finely punctate; punctures small, fine, shallow; setae small, about two diameters of a puncture in length; surface flattened; lateral margins of ventrites 2–3 unarmed. Apical margin of ventrite 4 thickened forming a broad carina with blunt median tubercle. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 greater than combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F54" orientation="portrait" position="float"><label>Figure 54.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33407.jpg" xlink:href="zookeys-450-001-g054"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>3.3–3.6 mm long (mean = 3.4 mm; n = 3); 2.2–2.8 times as long as wide. Similar to male except epistomal process more strongly developed, frons convex when viewed laterally, weakly aciculate, setae sparse, shorter, less than width of eye; weakly transversely impressed between inner apices of eye. Second ventrite unarmed.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>4.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic> Bright: male, labeled “MEX., N.L., Cerro Potosi, V.3.[19]71, 1100', D.E. Bright, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>, CNC No. 12603” (CNCI). Allotype, female, identical data as holotype. Paratypes, identical data as holotype (CNCI-2).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p>None examined.</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>MEXICO: Nuevo León (Fig. <xref ref-type="fig" rid="F32">32</xref>).</p></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (Mirb.) Franco (Douglas fir).</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic> is only known from a single collecting event from broken branches of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (<xref rid="B16" ref-type="bibr">Bright 1972</xref>). The gallery and biology of this species are unknown but the gallery structure is presumably similar to that of other <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content></named-content></italic> feeding <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic>, parallel to the grain of the wood.</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p><xref rid="B145" ref-type="bibr">Wood (1975</xref>: 22) placed <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic> in synonymy with <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic> because it occured on the same mountain as <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic>. Wood reasoned that only one <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species could occur in the same location and that the observed morphological variation was due to intraspecific differences. In addition, he stated that Bright’s image of the <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic> male was a normal male of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic>. We here remove <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic> from synonymy with <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic> because of the many distinct morphological and host differences originally noted by <xref rid="B16" ref-type="bibr">Bright (1972)</xref>. In the male of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic>, the apical margin of ventrite 1 is moderately thickened and weakly produced, the posterior margin of ventrite 3 is slightly medially thickened and the posterior margin of ventrite 4 is in strongly produced and thickened medially, forming a broad carina with a blunt median tubercle. In the male of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic> ventrite 1 apical margin is thickened and strongly produced and ventrites 3–4 are flat and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic> colonizes <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> species rather than <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content></named-content></italic>. It is also not uncommon for multiple <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species to have overlapping distributions.</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></title><p><named-content content-type="taxon-authority">LeConte, 1876</named-content></p><p><xref ref-type="fig" rid="F52">Figs 52</xref><xref ref-type="fig" rid="F55">, 55</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content><named-content content-type="comment"> LeConte, 1876: 373.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>The male most closely resembles those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic>. It is distinguished from that of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic> by the sparse, obscure, fine and shallow punctures of ventrite 2, by the dull luster of ventrite 2 and by the geographic distribution. The male can be distinguished from that of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> by the presence of a median denticle on the apical margin of ventrite 2. Females closely resemble those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> and are distinguished by the distinctly, moderately longitudinally aciculate and weakly punctate frons, and by the weakly produced apical margin of ventrite 1 that forms a weak carinate lip along the basal margin of ventrite 2.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>2.0–4.3 mm long (mean = 3.5 mm; n = 20); 1.8–2.3 times as long as wide. Head, pronotum and abdominal venter dark red-brown, antennae light brown, legs dark red-brown to light brown, elytra red brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma moderately, very broadly emarginate; epistomal process moderately developed and elevated; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing convex when viewed laterally, slightly transversely impressed just above epistoma; moderately, coarsely, longitudinally aciculate-punctate; aciculations converging at epistoma; punctures minute, fine; moderately and uniformly covered by long, fine, yellow, erect hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, ovoid, setose with partial septum, three arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, smooth apex; apex moderately emarginated at suture. Margin of apical edge bearing small, fine punctures. Disc smooth, shining; interstriae not impressed, more than twice width of striae, punctures uniseriate, smaller than those of striae; punctures bearing short, sparse, recumbent yellow setae slightly longer than size of a puncture (may be abraded); striae weakly impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 strongly, acutely produced forming lip along base of ventrite 2, basal margin of ventrite 2 appearing impressed. Ventrite 2 nearly perpendicular to ventrite 1; surface glabrous, shagreened, dull, finely and obscurely punctate; punctures small, fine, shallow; surface weakly concave; apical margin armed with broad median denticle; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 greater than combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F55" orientation="portrait" position="float"><label>Figure 55.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33408.jpg" xlink:href="zookeys-450-001-g055"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>3.1–5.0 mm long (mean = 3.85 mm; n = 20); 1.9–2.6 times as long as wide. Similar to male except epistoma feebly emarginate, epistomal process absent, frons more strongly convex when viewed laterally, weakly longitudinally aciculate, setae sparser, shorter, less than width of eye. Apical margin of ventrite 1 weakly elevated above base of ventrite 2. Second ventrite unarmed, surface weakly rugose, punctures larger, deeper.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>111.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Lectotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic> LeConte: female, Vanc. [Vancouver, B.C.], Type 968” (MCZC). Lectotype designated <xref rid="B148" ref-type="bibr">Wood 1982</xref>: 443.</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>UNITED STATES:</bold><bold><italic>CALIFORNIA</italic>:</bold><italic>Alpine Co.</italic>: Stanislaus National Forest, Hermit Valley, Hopk. U.S. 19193-A, J.M. Miller, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (EMEC-1, OSAC-4). Humboldt-Toiyabe National Forest, Hwy 89, 6.6 mi E. NF 4188, <named-content content-type="dwc:verbatimCoordinates">N38°39.906', W119°38.540'</named-content>, 8011 ft, 24.VII.2010, S.M. Smith, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">magnifica</named-content></named-content></italic> (MSUC-3). <italic>El Dorado Co.</italic>: Echo Lake, Hopk. U.S. 18381-A, 27.V.[19]31, J.M. Miller, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">magnifica</named-content></named-content></italic> (CSUC-1, EMEC-17, OSAC-20, USNM-3). Lake Tahoe, Fallen Leaf Lake, 22.VII.1930, A.C. Browne (CASC-1). Uncle Tom’s, 0.1 road mile W., 28.VII-12.VIII.1978, J.A. Benedictis, ex. from pheromone trap baited with E-11 tetradecenyl acetate (EMEC-1). [<italic>Madera Co.</italic>]: Bass Lake, Hopk. U.S. 19376-A, 2.VII.[19]32, G.R. Struble, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (EMEC-2, OSAC-3). Sugar Pine [community], 12.VIII.1920, E. Schiffel (CASC-1). <italic>Placer Co.</italic>: Lake Tahoe, 1 mi N., 3.VIII.[19]67, G.T. Ferrell (EMEC-3). <italic>Plumas Co.</italic>: LaPort, Hopk. U.S. 17933-A, J.M. Miller, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">magnifica</named-content></named-content></italic> (EMEC-2, OSAC-2). [<italic>Unspecified County</italic>]: Stanislaus National Forest, Hopk. U.S. 19818, J.M. Miller, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (OSAC-1). <bold><italic>IDAHO</italic>:</bold><italic>Clearwater Co.</italic>: Pierce, 4 mi W.N.W., 18.VII.1973, H.L. Osborne, ex. flight trap (USNM-1). <italic>Latah Co.</italic>: 1992, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (WFBM-1). Flat Creek, 11.II.1995, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (WFBM-4). Harvard, 3.5 mi N.N.E., 6.VIII.1973, H.L. Osborne, ex. flight trap (USNM-1). Moscow, 20.VII.[19]73, LC-1 (USNM-1), 24.VII.[19]73, LC-1 (USNM-2). Potlatch, 4 mi N.E., 3.VIII.1973, H.L. Osbourne, ex. flight trap (USNM-1). <bold><italic>OREGON</italic>:</bold> [<italic>Benton Co.</italic>]: [Corvallis], Kiger Island, IX.1922, W.J. Chamberlain (USNM-1). [<italic>Klamath Co.</italic>]: Crater Lake [National Park], 11.VI.[19]33, Hopk. U.S. 18966-A, W.J. Buckhorn, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (EMEC-4), 14.XII.[19]33 (EMEC-3, OSAC-20, WFBM-1). <bold><italic>WASHINGTON</italic>:</bold> [<italic>Chelan Co.</italic>]: Lake Wenatchee [State Park], VII.[19]69 (EMEC-1). [<italic>King Co.</italic>]: Seattle, 9.VI.[19]12 (OSAC-2). [<italic>Thurston Co.</italic>]: Olympia, 25.IV.[18]94 (OSAC-2).</p><pmc-comment>PageBreak</pmc-comment>
<pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: British Columbia. UNITED STATES: California, Idaho, Montana, Oregon, Washington (Fig. <xref ref-type="fig" rid="F56">56</xref>).</p><fig id="F56" orientation="portrait" position="float"><label>Figure 56.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic> distribution map.</p></caption><graphic id="oo_33409.jpg" xlink:href="zookeys-450-001-g056"></graphic></fig></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p>Principal host: <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (Dougl. Ex D. Don) Lindl. (grand fir). Incidental hosts: <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic> (Hook.) Nutt. (subalpine fir), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">magnifica</named-content></named-content></italic> A. Murr. (red fir).</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><xref rid="B53" ref-type="bibr">Furniss and Kegley (2011)</xref> provide a detailed and thorough account of the biology of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic>. Adults infest suppressed branches in the crowns of mature trees and are rarely found in slash (<xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B53" ref-type="bibr">Furniss and Kegley 2011</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic> is also associated with a staining fungus, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Spicaria</named-content><named-content content-type="species">anomala</named-content></named-content></italic> (Corda) Harz. that kills the host cambium (<xref rid="B152" ref-type="bibr">Wright 1938</xref>).</p><p>Adult galleries are distinct from those of other <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species. They are epsilon (“<bold>ε</bold>”) shaped and deeply excavated in the sapwood (Fig. <xref ref-type="fig" rid="F25">25</xref>). The gallery consists of a central nuptial chamber and two egg galleries that are recurved around the nuptial chamber (<xref rid="B42" ref-type="bibr">Edson 1967</xref>). The adult gallery is 2.0–5.0 cm in length and varies by branch size, with larger branches having larger galleries (<xref rid="B53" ref-type="bibr">Furniss and Kegley 2011</xref>). Egg niches lightly score the sapwood. In Oregon 12–30 eggs are laid per gallery and in Idaho the upper limit appears to be 12 (<xref rid="B53" ref-type="bibr">Furniss and Kegley 2011</xref>). Larval mines radiate from the egg gallery in all directions, often crossing each other. The larval mines are located in the phloem and cambium for about the first centimeter of their length. After the first centimeter, larval mines lightly score the sapwood. Pupation chambers are formed in the cambium or outer bark. There is one generation per year and the brood overwinters as larvae. In Idaho, flight occurs in July and adults leave the gallery once eggs have been laid (<xref rid="B53" ref-type="bibr">Furniss and Kegley 2011</xref>).</p></sec><sec sec-type="treatment-Collection notes"><title>Collection notes.</title><p>Old galleries of this species were observed by the senior author while hiking along the Tuolumne Grove Trail in Yosemite National Park on 23.VII.2010.</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>The lectotype bears a partial locality label. <xref rid="B82" ref-type="bibr">LeConte’s (1868)</xref> description states the lectotype was collected at Vancouver Island.</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></title><p><named-content content-type="taxon-authority">(Swaine, 1917)</named-content></p><p><xref ref-type="fig" rid="F52">Figs 52</xref><xref ref-type="fig" rid="F57">, 57</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Eccoptogaster</named-content><named-content content-type="species">tsugae</named-content></named-content><named-content content-type="comment"> Swaine, 1917: 32.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content><named-content content-type="comment"> (Swaine, 1917): <xref rid="B76" ref-type="bibr">Keen 1929</xref>: 12.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> most strongly resembles <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> and the two species are easily and often confused. Both sexes are distinguished from those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> by the impressed elytral discal striae, giving the elytra a corrugated appearance, by the dull luster of ventrite 2 and the host genus <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content></named-content></italic>.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>2.8–3.4 mm long (mean = 3.1 mm; n = 16); 2.1–2.5 times as long as wide. Body dark red-brown and antennae light brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly emarginate; epistomal process absent; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing flattened when viewed laterally, slightly transversely impressed just above epistoma; weakly longitudinally aciculate, moderately punctate; aciculations converging at epistoma; punctures small, coarse; moderately, uniformly covered by long, fine, erect, yellow, hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, irregularly ovoid, setose with partial septum, two arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, dark yellow-brown, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on basal half, narrowing to subquadrate, smooth apex; apex moderately emarginated at suture. Margin of apical edge bearing large, coarse punctures. Disc glabrous, smooth, shining; interstriae weakly impressed, more than twice width of striae, interstrial punctures uniseriate, smaller than those of striae; striae weakly impressed, elytra with a corrugated appearance. Declivity bearing sparse, short, erect yellow setae. Metepimeron less than half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 weakly, continuously elevated above base of ventrite 2. Ventrite 2 nearly perpendicular to ventrite 1; surface glabrous, shagreened, dull, finely punctate; punctures small, fine, shallow; surface flattened; apical margin unarmed; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 greater than combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F57" orientation="portrait" position="float"><label>Figure 57.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33410.jpg" xlink:href="zookeys-450-001-g057"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>2.3–3.5 mm long (mean = 3.0 mm; n = 16); 1.9–2.8 times as long as wide. Similar to male except epistoma feebly emarginate, epistomal process present, moderately developed, low, frons convex when viewed laterally, weakly strigate, setae sparser, shorter, less than width of eye; weakly transversely impressed between inner apices of eyes.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>369.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Lectotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Eccoptogaster</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> Swaine: female, labeled “Entomological Branch, Ottawa, Canada No. 2251, female, J.M. Swaine Coll., Lectotype CNCNo. 9239” (CNCI). Lectotype designated <xref rid="B15" ref-type="bibr">Bright 1967</xref>: 674. Paralectotypes <italic><named-content content-type="taxon-name"><named-content content-type="genus">Eccoptogaster</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> (CNCI), [<italic>Unspecified locality</italic>]: 2251 (CNCI-12, CUIC-2, EMEC-1), 2327 (CNCI-2).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>CANADA:</bold><bold><italic>BRITISH COLUMBIA</italic>:</bold> Adams Lake, 8.V.[19]22, R. Hopping, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (CASC-1). Bowman Creek, 10.VIII.1928, R. Hopping, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic> (CASC-2). Garibaldi, 7.VII.1988, R.J. Rabaglia (RJRC-1). Merritt, Midday Valley, 5.VI.1926, W. Mathers, 17134, lot 94, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content><named-content content-type="species">ponderosa</named-content></named-content></italic> (CASC-1), 11.VI.1926, 17190, lot 23 (CNCI-1). North Vancouver, Lynn Canyon, 1.VI.[19]23, 17003, N.L. Cutler, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">amabalis</named-content></named-content></italic> (CASC-12, CNCI-2). Pender Harbour, 17189, lot 1, 11.V.[19]26, G.R. Hopping, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic> (CNCI-9); lot 2, 12.V.[19]26 (CASC-1, CNCI-8), lot 4, 29.V.[19]26 (CASC-1, CNCI-6), lot 5, 1926 (CNCI-1), 13.VI.1928 (CASC-4). Terrace, Mrs. W.W. Hippisley (CNCI-1). Trinity Valley, 16.VIII.1927, J.R. Howell (CASC-2), 1722, lot 46, 10.VII.1928, (CASC-1), lot 50, 24.VII.1928 (CASC-1), 17339, lot 12, 19.VI.[19]30, (CASC-2); 16.VI.1927, 17213, lot 27, E.A. Rendell (CASC-1). Vancouver, 11.VI.1935, A. Graham, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (CASC-2); 27.V.1939, W.G. Mathers, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic> (CASC-2), 31.V.1939 (CASC-5), 5.VI.1939 (CASC-3), 12.VI.1939 (CASC-3), 27.VI.1939 (CASC-5), 4.VII.1939 (CASC-1), 10.VII.1939 (CASC-2). <bold>UNITED STATES:</bold><bold><italic>CALIFORNIA</italic>:</bold><italic>Alpine Co.</italic>: Ebbetts Pass, 8730 ft, 13.VIII.[19]63, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (CNCI-8, EMEC-9). <italic>El Dorado Co.</italic>: [Georgetown, 10 mi E.], Blodgett [Experimental] Forest, 2.VI.2003, K. Apigian (EMEC-1), 30.V.1986, K. Hobson (EMEC-1). Pollock Pines, 22.VI.[19]48, A. Bartel (EMEC-1). <italic>Lassen Co.</italic>: Grassy Lake, 27.IX.[19]14, lot 142, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (CASC-2). <italic>Nevada Co.</italic>: Tahoe National Forest, Sagehen Basin, Carpenter Ridge, <named-content content-type="dwc:verbatimCoordinates">39.4149°N, 120.3109°W</named-content>, 15.VII.2003, M. Caterino (SBMN-1). [<italic>Unspecified County</italic>]: Yosemite National Park, Hopk. U.S. 15727-A, 2.VII.1918, J.E. Patterson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (OSAC-4). <bold><italic>IDAHO</italic>:</bold><italic>Bonner Co.</italic>: Priest River Experimental Forest, Hopk. U.S. 61810, 28.X.1978, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic> (USNM-8). Priest Lake, Reader Bay, 6.VIII.1985, M.M. Furniss, J.B. Johnson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic> (WFBM-54). <italic>Boundary Co.</italic>: Idaho Panhandle National Forest, Roman Nose, <named-content content-type="dwc:verbatimCoordinates">N48°40.911', W116°34.345'</named-content>, 4353 ft, 12.VIII.2010, S.M. Smith, A.R. Gillogly, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic> (MSUC-7). <italic>Kootenai Co.</italic>: Deception Creek Experimental Forest, Hopk. U.S. 58885-B, 10.VII.1968, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic> (OSAC-1), 16.VII.1968 (OSAC-1), 24.VII.1968 (OSAC-4), 30.VII.1968 (OSAC-2), 8.VIII.1968 (OSAC-1). Magee, VII.[19]29, R.L. Furniss, ex. hemlock [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content></named-content></italic> sp.] (OSAC-4). <bold><italic>MONTANA</italic>:</bold> [<italic>Unspecified County</italic>]: Glacier [National] Park, 8.VII.[19]49, D. Giuliani (CASC-1). <bold><italic>OREGON</italic>:</bold> [<italic>Douglas Co.</italic>]: Diamond Lake, Hopk. U.S. 20959-A, VII.[19]31, R.L. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (OSAC-18). <italic>Hood River Co.</italic>: Mount Hood National Forest, Hwy 35, Sherwood Forest campground, <named-content content-type="dwc:verbatimCoordinates">N45°19.278', W121°37.104'</named-content>, 4293 ft, 2.VIII.2010, S.M. Smith, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic> (MSUC-5). <italic>Klamath Co.</italic>: Crescent Lake, 6.VII.[19]60, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (CASC-4). Crater Lake [National Park], Hopk. U.S. 18.916-A, 22.V.[19]30, W.J. Buckhorn, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (OSAC-30), Hopk. U.S. 18,950-A, 16.VI.[19]31 (OSAC-44); Hopk. U.S. 18851-A, 14.VI.[19]31, J.A. Beal, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (OSAC-6, USNM-2); Hopk. U.S. 20537-A, 17.VI.[19]33, F.P. Keen, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (OSAC-19); 24.VIII.[19]62, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (CNCI-1); 12.VIII.1984, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (WFBM-16); Hopk. U.S. 20807-A, R.L. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic> (OSAC-2). <italic>Linn Co.</italic>: Santiam Pass 7.VII.[19]64 (EMEC-1). [<italic>Yamhill Co.</italic>]: McMinnville, 27.XI.1937, K.M. & D.M. Fender (OSAC-1). <bold><italic>Washington</italic>:</bold><italic>King Co.</italic>: Seattle, 27.V.[19]07 (OSAC-1), 10.IV.[19]12 (OSAC-2), 12.IV.[19]12 (OSAC-1). [<italic>Snohomish Co.</italic>]: [labeled <italic>King Co.</italic>] Mountlake Terrace, 20.VIII.[19]62, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic> (CNCI-4). <bold><italic>UNSPECIFIED LOCALITY</italic>:</bold> Hopk. U.S. 13247-A, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (OSAC-2). Summit Viola Trail, 9.IX.1910, J.M. Miller, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (OSAC-1).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: Alberta, British Columbia. UNITED STATES: California, Idaho, Montana, Oregon, Washington (Fig. <xref ref-type="fig" rid="F36">36</xref>).</p></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic> Sarg. (western hemlock) and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (Bong.) Carrière (mountain hemlock).</p></sec><sec sec-type="treatment-Common name"><title>Common name.</title><p>Hemlock engraver.</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> attacks fresh slash, the main bole and large branches of hemlock (<xref rid="B42" ref-type="bibr">Edson 1967</xref>).</p><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> is an uncommon and poorly studied species. The adult galleries are typically perpendicular to the grain of the wood and 4.0–10.0 cm in length (Fig. <xref ref-type="fig" rid="F25">25</xref>) (<xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>). Adult galleries score the cambium slightly more than the sapwood. However, <xref rid="B42" ref-type="bibr">Edson (1967)</xref> reported that specimens from a series in northern California produced an adult gallery that was oriented obliquely to the grain of the wood. Galleries consist of a central nuptial chamber and two egg galleries. Each egg gallery is extended perpendicular to the grain of the wood from the central nuptial chamber. Egg niches are irregularly spaced and faintly score the sapwood. Larval tunnels are extended parallel to the grain of the wood, etching the sapwood lightly at first and deeply near the pupation chamber. There is one generation per year and broods overwinter as larvae (<xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>).</p></sec><sec sec-type="treatment-Collection notes"><title>Collection notes.</title><p>The senior author collected this species from fresh logging slash limbs that were 6.0–10.0 cm in diameter in Idaho and Oregon.</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>The lectotype does not bear a locality label. <xref rid="B131" ref-type="bibr">Swaine’s (1917)</xref> description states the type series was collected at “Cherry Creek valley, Vernon District, British Columbia, Glacier, B.C., Jasper Park, Alta.” from both <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">mucronata</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>]. In <xref rid="B15" ref-type="bibr">Bright’s (1967)</xref> lectotype designation he lists the locality of the lectotype as “Glacier, BC, XI-26-15, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic>”.</p><p>For many years <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> was considered a synonym of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> (see <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> remarks). In their paper describing the biology of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic>, <xref rid="B95" ref-type="bibr">McMullen and Atkins (1959)</xref> actually described the biology of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic>. In their investigation the species studied was from <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> rather than <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content></named-content></italic> spp. and created vertical instead of transverse galleries.</p></sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></title><p><named-content content-type="taxon-authority">LeConte, 1876</named-content></p><p><xref ref-type="fig" rid="F52">Figs 52</xref><xref ref-type="fig" rid="F58">, 58</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content><named-content content-type="comment"> LeConte, 1876: 372.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sobrinus</named-content></named-content><named-content content-type="comment"> Blackman, 1934: 23.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> is very morphologically similar to <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic>. Males are distinguished from those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> by the frons flattened when viewed laterally, never deeply impressed, less moderately abundant frontal setae (compared to dense) and by the host genus, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content></named-content></italic>. Males are distinguished from those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> by the following combination of characters: abdominal venter dull in luster, the base of the ventrite 2 spine extending from the apical margin to half the length of the segment and by the geographical distribution. The female is distinguished from that of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> by the dull luster of ventrite 2 and is distinguished from that of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> by the finely aciculate-punctate frons; ventrite 1 joining base of ventrite 2 more obtusely, base of ventrite 2 not finely impressed, flush with ventrite 1; epistomal process weakly developed, almost indistinct.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>2.2–3.2 mm long (mean = 2.7 mm; n = 15); 2.3–2.7 times as long as wide. Head, antennae, pronotum, and abdominal venter dark red-brown. Elytra and legs yellow-brown to light brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly emarginate; epistomal process present, weakly developed, low; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing flattened when viewed laterally from epistoma to vertex, slightly transversely impressed just above epistoma to inner apices of eyes; moderately, coarsely, longitudinally aciculate-punctate; aciculations converging at epistoma; punctures sparse, small, fine; sparsely, uniformly covered by long, fine, erect, yellow, hair-like setae, thesee longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, irregularly ovoid, setose with partial septum, two arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, smooth apex; apex moderately emarginated at suture. Margin of apical edge bearing small, fine punctures. Disc smooth, shining; interstriae not impressed, twice width of striae, punctures uniseriate, smaller than those of striae; bearing short, sparse, recumbent yellow setae slightly longer than size of a puncture (may be abraded); striae weakly impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron less than half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 rounded, marked by weak carina. Ventrite 2 nearly perpendicular to ventrite 1; surface shagreened, dull, finely punctate; punctures small, fine, shallow, nearly obscure; covered with sparse setae that are less than length of segment 3; surface convex; apical margin armed with laterally compressed, median spine with base extending from apical margin to half length of segment, apex rounded; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 less than combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F58" orientation="portrait" position="float"><label>Figure 58.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33411.jpg" xlink:href="zookeys-450-001-g058"></graphic></fig></sec><sec sec-type="treatment-Female"><title>Female.</title><p>2.2–3.0 mm long (mean = 2.7 mm; n = 15); 2.3–2.7 times as long as wide. Similar to male except epistoma feebly emarginate, epistomal process feebly <pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment>developed, frons convex when viewed laterally, weakly longitudinally aciculate, setae sparser, shorter, less than width of eye; weakly transversely impressed between epistoma and inner apices of eyes. Second ventrite apical margin armed with acute median denticle, with base extending from apical margin to half-length of segment.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>654.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Lectotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> LeConte: male, labeled “Or. [Oregon], Type 965” (MCZC). Lectotype designated <xref rid="B148" ref-type="bibr">Wood 1982</xref>: 432. Holotype <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sobrinus</named-content></named-content></italic> Blackman: male, labeled “Burke Colr, Kent, WA, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>], Hopk. US 41900, Type No. 43838” (USNM). Synonymy: <xref rid="B141" ref-type="bibr">Wood 1966</xref>: 30. Paratypes <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sobrinus</named-content></named-content></italic> Blackman <bold>UNITED STATES:</bold><bold><italic>OREGON</italic>:</bold> [<italic>Jackson Co.</italic>]: Little Applegate River, Hopk. U.S. 14238-B, J.E. Patterson (EMEC-6). Ashland, Hopk. U.S. 14264-T, F.P. Keen (EMEC-3). <bold><italic>WASHINGTON</italic>:</bold> [<italic>King Co.</italic>]: Kent, Hopk. U.S. 4190-A, [H.E.] Burke, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-5).</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>CANADA:</bold><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><bold><italic>BRITISH COLUMBIA</italic>:</bold> Cloverdale, 28.XII.[19]22, N.L. Cutler, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic> (CASC-32, CNCI-11). Duncan, Genoa Bay, 10.VII.1928, W.G. Mathers, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CNCI-1). Indian Meadows, Midday Creek, 13.VII.1920, R. Hopping, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content><named-content content-type="species">ponderosa</named-content></named-content></italic> (CNCI-1). Merritt, Midday Valley, 31.V.1926, W. Mathers (CNCI-1), 2.VII.1926 (CNCI-2); R. Hopping 3.VIII.1923 (CNCI-1), 15.VIII.19[23] (CNCI-1), 7.VII.1926 (CNCI-1). Nanaimo, [Pacific] Biological Station, 23.VI.1920 (CASC-1). Nelson Island, West Lake, 1701, 23.VI.1922, R. Hopping, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CNCI-12). Oliver, 25 mi N.W., 15.VI.1958, H. & A. Howden (EMEC-3). Pender Harbour, 30.VI.1928, G. Hopping, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CASC-2). Squamish, Diamond Head trail, 3200 ft, 9.VIII.1953, G.J. Spencer (CNCI-1). Shuswap, 24.IV.[19]13, T. Wilson, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CNCI-1). Vancouver, 7.VI.1934, W. Mathers ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CASC-1), 11.VI.1934 (CASC-2). Vanguard, 23.VI.[19]32, R. Hopping, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">mucronata</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CASC-2, CNCI-2). Vernon, 27.V.1932, R. Hopping (CASC-1). <bold>UNITED STATES:</bold><bold><italic>CALIFORNIA</italic>:</bold><italic>Del Norte Co.</italic>: 3.VII.[19]24, E.B. Leach (EMEC-1). <italic>El Dorado Co.</italic>: [Georgetown, 10mi E.], Blodgett Experimental Forest, 4000 ft, 26.V.1986, Hobson, Atkinson, ex. Lindgren trap, ponderosa pine resin, H&L B.P. fractions, burned over area (EMEC-3); 27.V.1986, Hobson, Irving, ex. Lindgren trap, ponderosa pine resin, untreated Oleo, burned over area (EMEC-2); 30.V.1986, Hobson, Atkinson, ex. Lindgren trap, unbaited, logged area (EMEC-1); 30.V.1986, Hobson, Atkinson, ex. Lindgren trap, unbaited, burned over area (EMEC-2). Placerville, Hopk. U.S. 33961-A, 26.III.[19]54, M.M. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-16). Pollack Pines, 22.VI.[19]48, R.C. Bynum (EMEC-6). <italic>Humboldt Co.</italic>: Blocksburg, 13.V.[19]34, B.P. Biven (CASC-2). Orleans, 15 mi N.W., 13.VI.[19]64, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-7). Orick, 11 mi S., 11.VI.[19]62, D.E. Bright, B.A. Barr, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (DEBC-4, EMEC-3). <italic>Lake Co.</italic>: Whispering Pines, 14.IV.1964, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CASC-1). <italic>Los Angeles Co.</italic>: Coquillett (EMEC-2). <italic>Madera Co.</italic>: 1.VII.[19]38 (EMEC-1). <italic>Marin Co.</italic>: Inverness, 11.X.[19]61, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (DEBC-3, EMEC-2). [<italic>Mariposa Co.</italic>]: [Yosemite National Park], Yosemite Valley, Hopk. U.S. 2810, Hopkins (EMEC-4). <italic>Mendocino Co.</italic>: (CASC-7). 14.VII.[19]22, E.R. Leach (CASC-4). Bransomb, CDFA#870697, 11.VII.[20]01, M. Garvin, ex. Lindgren funnel trap with ISP (CSCA-1), 25.VII.[20]01 (CSCA-3). Flynn Creek between Comptche and Navarro, 17.VIII.1953, P.S. Bartholomew (CASC-1). Mill Creek, 20.III.[19]59, R.E. Stevens, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-13). Noyo River, VI.1896 (CASC-14). <italic>Napa Co.</italic>: Angwin, 2 mi N.N.E., North side of Howell Mountain, 1300 ft, 24.V.1974, H.B. Leech, ex. emerged from log of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (CASC-3, USNM-6), 7.VII.1974 (CASC-3), 20.VII.1974 (CASC-16), 21.VII.1974 (CASC-11), 22.VII.1974 (CASC-4), 23.VII.1974 (CASC-8), 25.VII.1974 (CASC-15), 26.VII.1974 (CASC-2, USNM-3), 27.VII.1974 (CASC-1), 28.VII.1974 (CASC-6, USNM-3), 1.VIII.1974 (CASC-3), 3.VIII.1974 (CASC-3). Callistoga, 26.V.[19]57 (CASC-4); 4.X.1947, T.O. Thatcher, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-3); 1.V.[19]63, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (EMEC-3). <italic>Nevada Co.</italic>: Grass Valley, 17.I.1961, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CASC-2). Middleton [labeled as <italic>Lake Co.</italic>], Hopk. U.S. 37588-A, R.W. Bushing, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (EMEC-3), G.M. Thomas (EMEC-3). [<italic>Placer Co.</italic>]: Colfax, 1 mi E., Hopk. U.S. 34068-A, 31.I.[19]37, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (EMEC-4). Towle, 11.XI.1932 (EMEC-1). <italic>Plumas Co.</italic>: Walker Mine, 15.VII.[19]30 (EMEC-3). [<italic>San Francisco Co.</italic>]: San Francisco, Hopk. U.S. 8557, 11.V.[18]99, ex. on pine [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content></named-content></italic> sp.] (EMEC-1), 15.V.[18]99 (EMEC-1). <italic>Santa Cruz Co.</italic>: 16.IV.[19]59, J.E. Henry (WFBM-16). Aptos, New Brighton Beach State Park, 13.IX.1986, D. Adams, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content><named-content content-type="species">radiata</named-content></named-content></italic> (EMEC-54). <italic>Shasta Co.</italic>: Hat Creek, 17.VI.[19]62, D.E. Bright, B.A. Barr, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (EMEC-2). <italic>Siskiyou Co.</italic>: Grass Valley, 16.X.[19]60, R.W. Bushing, emerged XI.1960, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (EMEC-6). <italic>Sonoma Co.</italic>: (CNCI-1). Fort Ross, 2 mi E., 2.XI.1947, T.O. Thatcher, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-3). Mount Saint Helena, 3.VI.[19]31, E.C. Zimmermann, ex. digger pine [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content><named-content content-type="species">sabiniana</named-content></named-content></italic>] (EMEC-1); Hopk. U.S. 21,125-F, 3.IV.1934, R.L. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-4). Sebastopol, Hopk. U.S. 32638-A, 20.I.1940, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (DEBC-4, EMEC-5). Stillwater Cove, 12.V.1951, H.R. Moffitt (EMEC-1). <italic>Trinity Co.</italic>: 14.V.[19]23, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CASC-4). [<italic>Unspecified County</italic>]: (CUIC-7). Yosemite National Park, Hopk. U.S. 20953-B, 19.VI.1932, J.M. Miller, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-8). <bold><italic>IDAHO</italic>:</bold><italic>Bonner Co.</italic>: Sandpoint, 13.VI.931, N.M. Downie (FMNH-1), 9.VII.1977 (FMNH-1); 9.VII.1977 (FMNH-1). <bold><italic>OREGON</italic>:</bold> [<italic>Benton Co.</italic>]: Corvallis, 17.IV.[19]16, W.J. Chamberlin, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-3); 8.VII.1946, K.R. Hobbs (EMEC-1). <italic>Coos Co.</italic>: Myrtle Point, 18.VI.[19]64, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (EMEC-10). <italic>Curry Co.</italic>: Agness, 6 mi S.W., 10.VII.1990, M.M. Furniss, J.B. Johnson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-6). Brookings, 12 mi E.N.E., 12.VIII.1990, M.M. Furniss, J.B. Johnson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-4). Illahe, 4 mi N., 9.VIII.[19]90, M.M. Furniss, J.B. Johnson, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (WFBM-2). <italic>Deschutes Co.</italic>: Deschutes National Forest, Black Butte Rd, Black Butte, <named-content content-type="dwc:verbatimCoordinates">N44°24.924', W121°38.323'</named-content>, 4212 ft, 31.VIII.2010, S.M. Smith, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-44). <italic>Douglas Co.</italic>: Roseburg, W.J. Chamberlin, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (DEBC-2, EMEC-6, WFBM-2). [<italic>Grant Co.</italic>]: Dixie Pass, Malheur National Forest, 23.VI.1961, S.L. Wood, J.B. Karren, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (DEBC-5). <italic>Jackson Co.</italic>: Dead Indian Spring, 17.V.1962, J. Schuh (CNCI-1). Mistletoe, Hopk. U.S. 15753-A, P.D. Sergent, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-3), Hopk. U.S. 15753-B, 21.VIII.1918, P.D. Sargent (DEBC-5, EMEC-10). Pinehurst, 2 mi E., 18.VII.[19]64, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (CNCI-5, EMEC-6). Prospect, 10.VII.[19]64, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (CNCI-11, EMEC-14). [<italic>Klamath Co.</italic>]: Klamath Falls, 17.VI.[19]64, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (EMEC-2). [<italic>Lane Co.</italic>]: McCredie Springs, 19.VI.1961, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">heterophylla</named-content></named-content></italic> (CNCI-2). Vaughn, J. Pierce, ex. Douglas fir limbs [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (WFBM-4); 21.VI.1955, J.A. Rudinsky, J.R. Pierce, ex. Doug fir branches [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (WFBM-3). <italic>Linn Co.</italic>: Santiam Pass, 19.VI.1951, S.L. Wood, J.B. Karren, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (DEBC-1). [<italic>Malheur Co.</italic>]: Monument Peak, 25.IX.[19]53, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-3). [<italic>Marion Co.</italic>]: Clear Lake, 17.VIII.[19]51, R. Kangur, ex. Larch [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Larix</named-content></named-content></italic> sp.] ((EMEC-4, WFBM-2). [<italic>Multnomah Co.</italic>]: Portland, Hubbard, Schwarz (EMEC-2). [<italic>Umatilla Co.</italic>]: Tollgate, 30.VI.1950, E.S. McClurskey, ex. on aluminum roof (CNCI-1), 1.IX.1950 (CNCI-1). [<italic>Unspecified County</italic>]: Detroit, 25 mi E., 17.VII.1939, Schuh, Scott, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content><named-content content-type="species">mertensiana</named-content></named-content></italic> (FMNH-6). Middle Sister Mountain, 8000 ft, Hopk. U.S. 53349-G, 4.VIII.1968, M.M. Furniss, ex. on snowfield (WFBM-2). Portland, Wickham (CNCI-1). Tillamook burn, 18.VII.1941, R. Kangur (EMEC-1). Warm Springs Indian Reservation, 15.VIII.[19]51, R. Kangur, ex. Douglas fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-4). <bold><italic>WASHINGTON</italic>:</bold> [<italic>Clallam Co.</italic>]: Port Angeles, Hopk. U.S. 130, A.D. Hopkins, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-1). Port Williams [Marlyn Nelson County Park at Port Williams], Hopk. U.S. 168-F, A.D. Hopkins, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-1). <italic>Cowlitz Co.</italic>: Castle Rock, 10 mi E., 27.VI.[19]64, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (DEBC-18, EMEC-9). [<italic>Grays Harbor Co.</italic>]: Humptulips, 28.V.1914, E.C. VanDyke (CASC-3). [<italic>Kittitas Co.</italic>]: Easton (CASC-6, EMEC-2). <italic>Okanogan Co.</italic>: Disautel, 4.XI.1936, R.L. Furniss, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (OSAC-17, WFBM-7). [<italic>Pend Oreille Co.</italic>]: Metaline Falls, Hopk. U.S. 21340, 17.VII.1931, W.D. Bedard, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (WFBM-15). <italic>Skamania Co.</italic>: Mineral Springs, 27.VI.[19]64, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (EMEC-9). [<italic>Snohomish Co.</italic>]: [labeled <italic>King Co.</italic>] Mountlake Terrace, 20.VIII.[19]62, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (CNCI-3). <italic>Thurston Co.</italic>: Olympia, 7-30.V.1996, E. LaGasa, ex. Washington Department of Agriculture port trapping survey (WFBM-1). <italic>Yakima Co.</italic>: Naches, 14 mi W., Dry Creek Ridge, Snoqualamie National Forest, 3.VII.1965, R.B. Hutt (DEBC-2). [<italic>Unspecified County</italic>]: Mount Adams, Bird Creek, 6000-7000 ft, 2.VII.1925 (CASC-1). Mount Rainier National Park, 21.VIII.[19]62, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (CNCI-6).</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: British Columbia. UNITED STATES: California, Idaho, Oregon, Washington (Fig. <xref ref-type="fig" rid="F51">51</xref>).</p></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (Mirb.) Franco (Douglas fir) but also likely occurs in <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">macrocarpa</named-content></named-content></italic> (Vasey) Mayr (bigcone Douglas fir) in Southern California. Incidental in <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pinus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Tsuga</named-content></named-content></italic>.</p></sec><sec sec-type="treatment-Common name"><title>Common name.</title><p>Douglas-fir engraver.</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> is very common (<xref rid="B24" ref-type="bibr">Chamberlin 1939</xref>; Smith pers. obs.) and attacks the boles and branches of weakened, injured, dying and recently killed Douglas fir. Populations of this species can build up in windfalls, slash and during drought when the species becomes capable of killing young trees (<xref rid="B77" ref-type="bibr">Keen 1938</xref>; <xref rid="B24" ref-type="bibr">Chamberlin 1939</xref>; <xref rid="B139" ref-type="bibr">Wood et al. 2003</xref>). Outbreaks are sporadic and are of short duration (<xref rid="B77" ref-type="bibr">Keen 1938</xref>; <xref rid="B139" ref-type="bibr">Wood et al. 2003</xref>).</p><p>The adult gallery of this species consists of a single egg gallery that extends with the grain of the wood (Fig. <xref ref-type="fig" rid="F25">25</xref>). The nuptial chamber includes a short extension at a 45° angle from the egg gallery and is located at one end on the egg gallery. The gallery deeply scores the sapwood and lightly scores the cambium (<xref rid="B25" ref-type="bibr">Chamberlin 1958</xref>). The female deposits 40–100 eggs in egg niches on both sides of the egg gallery 0.5–1.0 mm apart (<xref rid="B40" ref-type="bibr">Doane et al. 1936</xref>). Larvae extend their mines at a right angle to the egg gallery forming a fan shaped pattern (<xref rid="B24" ref-type="bibr">Chamberlin 1939</xref>). There are two generations per year in California and the broods overwinter as either eggs or larvae (<xref rid="B77" ref-type="bibr">Keen 1938</xref>; <xref rid="B139" ref-type="bibr">Wood et al. 2003</xref>). Adults emerge from late April through July (Keen 1958). In Oregon one generation per year has been observed at high elevations in and two at lower elevations (<xref rid="B24" ref-type="bibr">Chamberlin 1939</xref>).</p><p><xref rid="B96" ref-type="bibr">McMullen and Atkins (1962)</xref> reported some notes on the biology of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> in British Columbia and appear to have reported a combined account of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> (see <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> biology for more information). This paper was the main source of information regarding the biology of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> and served as the basis for describing the biology of the species in numerous publications (including <xref rid="B17" ref-type="bibr">Bright 1976</xref>; <xref rid="B55" ref-type="bibr">Furniss and Carolin 1977</xref>; <xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>; <xref rid="B139" ref-type="bibr">Wood et al. 2003</xref>). <xref rid="B24" ref-type="bibr">Chamberlin (1939</xref>, <xref rid="B25" ref-type="bibr">1958</xref>) provides the most reliable source of information regarding the biology of the species.</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>There are 25 <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">sobrinus</named-content></named-content></italic> specimens bearing paratype labels found in the EMEC. These specimens are not designated as such by <xref rid="B10" ref-type="bibr">Blackman (1934)</xref> and have been labeled “not Paratype” by the authors. These specimens bear the following Hopkins numbers: 4205-A (EMEC-2), 4272-A (EMEC-4), 1968-D1 (EMEC-1), 1968-D3 (EMEC-1), 4273-B (EMEC-1), 4204 (EMEC-2), 4220-A (EMEC-3), 4226-D (EMEC-1), 4201-A (EMEC-2), 13232-C (EMEC-1), 17 (EMEC-1), 18 (EMEC-1), 19 (EMEC-2) and Sonoma, California (EMEC-3).</p><p>This species occurs along the western coastal states of the United States and British Columbia, Canada. The range of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> does not overlap with that of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> except in south central British Columbia where species are sympatric in the Interior Plateau near Merritt.</p><pmc-comment>PageBreak</pmc-comment>
</sec></sec><sec sec-type="taxon-treatment"><sec-meta><kwd-group><label>Taxon classification</label><kwd><named-content content-type="kingdom">Animalia</named-content></kwd><kwd><named-content content-type="order">Coleoptera</named-content></kwd><kwd><named-content content-type="family">Curculionidae</named-content></kwd></kwd-group></sec-meta><title><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></title><p><named-content content-type="taxon-authority">LeConte, 1868</named-content></p><p><xref ref-type="fig" rid="F52">Figs 52</xref><xref ref-type="fig" rid="F59">, 59</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content><named-content content-type="comment"> LeConte, 1868: 167.</named-content></p></list-item></list><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Males are distinguished from those of other species by having the base of ventrite 2 elevated, the surface of ventrite 2 flat, the apical margin of ventrite 2 often bearing a median denticle, and by the glabrous ventrite 2. Females most closely resemble those of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic>. Females are distinguished from those of both species by the indistinctly and weakly aciculate and strongly punctate frons and by the apical margin of ventrite 1 flush with basal margin of ventrite 2, appearing rounded.</p></sec><sec sec-type="treatment-Description (male)"><title>Description (male).</title><p>3.0–4.0 mm long (mean = 3.65 mm; n = 20); 2.1–2.7 times as long as wide. Head, pronotum and abdominal venter dark red-brown, antennae light brown, elytra and legs yellow-brown to light brown. Pronotum typically darker than elytra.</p><p><italic>Head.</italic> Epistoma weakly emarginate; epistomal process weakly elevated; median area above mandibles bearing dense patch of long, yellow, hair-like setae. Frons appearing convex when viewed laterally, slightly transversely impressed just above epistoma; moderately, coarsely, longitudinally aciculate-punctate; aciculations converging at epistoma; punctures small, coarse; moderately, uniformly covered by long, fine, yellow, erect, hair-like setae, these longer than width of midpoint of eye. Antennal scape short, elongate; club flattened, irregularly ovoid, setose with partial septum, three arcuate sutures visible.</p><p><italic>Pronotum</italic> wider than long; apical margin broadly rounded, median area between eyes lined with scales; sides distinctly arcuate, strongly constricted near apex, forming a weak transverse impression near apical margin; surface smooth, shining, punctures on disc fine, shallow, moderately abundant, larger and more abundant laterally and on apical constriction; apical and anterolateral margins bearing sparse, erect, yellow, hair-like setae; base weakly bisinuate.</p><p><italic>Elytra</italic> with sides sub-parallel on apical half, narrowing to subquadrate, smooth apex; apex moderately emarginated at suture. Margin of apical edge bearing large, coarse punctures. Disc glabrous, smooth, shining; interstriae not impressed, more than twice width of striae, interstrial punctures uniseriate, smaller than those of striae; striae weakly impressed. Declivity bearing sparse, short, erect yellow setae. Metepimeron half-length of metanepisternum.</p><p><italic>Venter.</italic> Apical margin of ventrite 1 weakly elevated above base of ventrite 2. Ventrite 2 nearly perpendicular to ventrite 1; surface glabrous, shagreened, dull, finely punctate; punctures small, fine, shallow; surface flattened; apical margin armed with broad median denticle, occasionally absent; lateral margins of ventrites 2–3 and ventrite 4 unarmed. Ventrite 5 carinate ridge closer to apical margin of segment; length of ventrite 5 less than combined lengths of ventrites 3 and 4; setal patch and median depression absent.</p><fig id="F59" orientation="portrait" position="float"><label>Figure 59.</label><caption><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic><bold>A</bold> dorsal male habitus <bold>B</bold> lateral male habitus <bold>C</bold> male frons <bold>D</bold> male frons oblique <bold>E</bold> male venter <bold>F</bold> male venter oblique <bold>G</bold> dorsal female habitus <bold>H</bold> lateral female habitus <bold>I</bold> female frons <bold>J</bold> female frons oblique <bold>K</bold> female venter <bold>L</bold> female venter oblique.</p></caption><graphic id="oo_33412.jpg" xlink:href="zookeys-450-001-g059"></graphic></fig><pmc-comment>PageBreak</pmc-comment>
<pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="treatment-Female"><title>Female.</title><p>2.2–4.6 mm long (mean = 3.56 mm; n = 20); 2.1–2.7 times as long as wide. Similar to male except epistoma feebly emarginate, epistomal process absent, frons convex when viewed laterally, weakly longitudinally aciculate, setae sparser, shorter, less than width of eye; weakly transversely impressed between epistoma and inner apices of eyes. Second ventrite unarmed.</p></sec><sec sec-type="treatment-Specimens examined"><title>Specimens examined.</title><p>324.</p></sec><sec sec-type="treatment-Type material"><title>Type material.</title><p>Lectotype: male, labeled “[Washington Territory] Type 970” (MCZC). Lectotype designated <xref rid="B148" ref-type="bibr">Wood 1982</xref>: 441.</p></sec><sec sec-type="treatment-Non-type material"><title>Non-type material.</title><p><bold>CANADA:</bold><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><bold><italic>BRITISH COLUMBIA</italic>:</bold> Duncan, Genoa Bay, 30.VI.1928, W.G. Mathers, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (CNCI-4), 21.VI.1928 (CNCI-1), 23.VI.1928 (CASC-1), 24.VI.1928 (CASC-1), 16.VII.1928 (CASC-1), 13.VIII.1936 (CASC-1). Steelhead, 6.VII.1933, H.B. Leech, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">amabalis</named-content></named-content></italic> (CNCI-1), 27.VII.1933 (CASC-1), 24.VII.1933 (CNCI-1), 26.VII.1933 (CNCI-1). Trail, 22 km S.E., 24.V.1991, J.E. Macias, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (CNCI-1). Vancouver, 10.VI.1935, K. Graham, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (CASC-12). <bold>MEXICO:</bold><bold><italic>Baja California</italic>:</bold> San Pedro Mártir, Hopk. U.S. 32716-A, 5.VI.1944, F.P. Keen, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (EMEC-1). <bold>UNITED STATES:</bold><bold><italic>ARIZONA</italic>:</bold><italic>Cochise Co.</italic>: Coronado National Forest, Chiricahua Mountains, 14.VII.2009, J. Hulcr, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic> (MSUC-3). <bold><italic>CALIFORNIA</italic>:</bold> [<italic>Amador Co.</italic>]: Jackson, 31.III.[19]55, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (EMEC-1). <italic>Calaveras Co.</italic>: Big Trees [State Park], VII.1930, F.E. Blaisdell (CASC-1). <italic>El Dorado Co.</italic>: Bijou at south end of Lake Tahoe, H.B. Leech, ex. Douglas fir log [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>], emerged San Francisco, 18-20.V.1965, W.E. Kuhl (CASC-8). [Georgetown, 10 mi E.], Blodgett [Experimental] Forest, 30.V.1986, K. Hobson (EMEC-1), 11.VI.1986 (EMEC-1), 21–24.VI.1986 (EMEC-6), 25-27.VI.1986 (EMEC-1), 1–7.VII.1986 (EMEC-2), 9–16.VII.1986 (EMEC-4); 30.V.1986, Hobson, Atkinson, ex. Lindgren trap, unbaited over burned area (EMEC-3), ex. Lindgren trap, turpentine bait, logged area (EMEC-1); 27.V.1986, Hobson, Irving, ex. Lindgren trap, ponderosa pine resin H&L B.P. fractions, logged area (EMEC-1), ex. Lindgren trap, ponderosa pine resin, untreated oleo, burned over area (EMEC-1). Ice House Reservoir, 23.VI.[19]90, J.B. Johnson (EMEC-1). Pacific, 8.VII.[19]62, R.L. McDonald (CASC-1). <italic>Fresno Co.</italic>: Huntington Lake, 16.VII.[19]19, F.E. Blaisdell (CASC-5). <italic>Lassen Co.</italic>: Lassen National Forest, 20–25.VII.1994, S1/2 S35 T34N R7E, A.I. Cognato (MSUC-20, SBMN-4); Black Mountain Experimental Forest, VII.1995, A.I. Cognato (MSUC-1). Near Loon Lake campground, 6300 ft, 28.VII–12.VIII.1978, J.A. De Benedictis (EMEC-1). Norval Flats, 5500 ft, 18.VII.[19]20, J.O. Martin, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (CASC-47). Uncle Tom’s, 0.1 road mile W., 28.VII-12.VIII.1978, J.A. De Benedictis (EMEC-1). <italic>Marin Co.</italic>: VI-VIII.[19]56, D. Giuliani (CASC-1). <italic>Mariposa Co.</italic>: Miami Ranger Station, 2mi S., 19.VII.1946, H.P. Chandler (DEBC-3). Miami Lodge, 17.VII.1946, G.R. Struble, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (CSUC-1). <italic>Mendocino Co.</italic>: Noyo River, VI.1896 (CASC-22). <italic>Modoc Co.</italic>: Modoc National Forest, Hwy 299, 5.3 mi W. Cedarville at creek near exit from ski area, <named-content content-type="dwc:verbatimCoordinates">41°32.9’N 120°14’W</named-content>, 9.IX.1995, J. Schweikert, ex. swept creek side plants (CASC-1). Davis Creek, 4 mi N.E., Sugar Hill, 6300 ft, 9.VI.1970, W. Middlekauff (EMEC-1). <italic>Nevada Co.</italic>: Sagehen Creek Field Station, <named-content content-type="dwc:verbatimCoordinates">39.4298°N, 120.2429°W</named-content>, 12-18.VIII.2003, M. Caterino, ex. FIT (SBNM-1). Near Sagehen campground, <named-content content-type="dwc:verbatimCoordinates">39.4344°N, 120.2808°W</named-content>, 15.VIII.2003, M. Caterino (SBNM-1). Sagehen Creek, 20.VII.1966, W.J. Turner (EMEC-1). [<italic>Placer Co.</italic>]: Cisco (USNM-1). <italic>Plumas Co.</italic>: Bartle, 9 mi N., 12-15.VI.1974, L. Green (EMEC-1). Chester, 8mi N.W., Warner Creek, 5000 ft, 21.VI.1989, E.E. Lundquist (CNCI-2). Janesville, 15-21.VIII.[19]50, M. Wasbauer, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (EMEC-1). <italic>Riverside Co.</italic>: Mount San Jacinto State Park, <named-content content-type="dwc:verbatimCoordinates">33.800°N, 116.673°W</named-content>, 15.VII.2003, M. Caterino (SBMN-1). Santa Rosa Mountains, 10.IV.[19]63, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (EMEC-1). [<italic>San Bernardino Co.</italic>]: San Bernardino Mountains, Dollar Lake trail, 10.VII.1956, H.W. Michalk (CSCA-1). <italic>Siskiyou Co.</italic>: Happy Camp, 18 mi N., 31.VII.[19]63, C.J. Wray, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Picea</named-content><named-content content-type="species">breweriana</named-content></named-content></italic> (DEBC-2, EMEC-1). Klamath National Forest, goosenest adaptive management area, <named-content content-type="dwc:verbatimCoordinates">41.5°N, 121.9°W</named-content>, 26.VII.2000, ex. pitfall 3-840 (SBNM-1). McCloud, 22.VI.1914 (CASC-23), 14.VI.1962, D.E. Bright, B.A. Barr, C. Hector (CASC-1). <italic>Tehama Co.</italic>: Mineral, 11.VIII.[19]35 (CASC-1). <italic>Trinity Co.</italic>: Carville, 30.VI.1913, ex. dug out of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (CASC-1), 2400-2500 ft, 23.V.1934 (CASC-1). Nash Mine, 12.VI.1913 (CASC-1). <italic>Tulare Co.</italic>: Giant Forest, 6500 ft, VII.1908, Hopping (CASC-3). Kaweah, 100 ft, Hopping (CASC-1). Sequoia National Park, Redwood Canyon, 20.IX.1980, S.F. Muzzio (CASC-2). [<italic>Unspecified county</italic>]: <bold><italic>COLORADO</italic>:</bold><italic>Costilla Co.</italic>: Pass Creek, 17.VI.1976, D. Leatherman, ex. white fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic>] (CSUC-4). <italic>Huerfano Co.</italic>: near Red Wing, 16.VI.1975, D. Leatherman, ex. white fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic>] (CSUC-1). <italic>Las Animas Co.</italic>: Monument Lake, 6.VII.1994, S. Kelley, ex. white fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic>] (DEBC-2). <italic>Pueblo Co.</italic>: SR 165, 5 mi S.E. San Isabel Millset trailhead, D. Leatherman, ex. burned white fir [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic>] (CSUC-6). <bold><italic>IDAHO</italic>:</bold><italic>Benewah Co.</italic>: Plummer, 4 mi S., 14.VIII.1956, W.F. Barr, E.C. Clark, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (FMNH-1). <italic>Boundary Co.</italic>: Idaho Panhandle National Forest, Hwy 95, Robinson Lake campground, <named-content content-type="dwc:verbatimCoordinates">N48°58.200', W116°13.067'</named-content>, 2696 ft, 13.VIII.2010, S.M. Smith, A.R. Gillogly, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (MSUC-6). <italic>Kootenai Co.</italic>: Coeur d’Alene, 5.IX.1919, J.C. Evenden, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (MSUC-3). <bold><italic>MONTANA</italic>:</bold> [<italic>Unspecified county</italic>]: Glacier [National] Park, 15.VII.[19]31 (CUIC-1). <bold><italic>NEW MEXICO</italic>:</bold><italic>Bernalillo Co.</italic>: Cibola National Forest, Sandia Peak, NM536, <named-content content-type="dwc:verbatimCoordinates">N35°11.655', W106°24.075'</named-content>, 8317 ft, 10.V.2010, S.M. Smith, A.I. Cognato, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (MSUC-1). [<italic>Otero Co.</italic>]: Cloudcroft, 900 ft, W. Knaus (CNCI-1). <italic>Santa Fe Co.</italic>: Little Tesuque Canyon, 14.VI.[19]35 (CASC-1). <italic>Socorro Co.</italic>: VII.[?], Wickham (CASC-1). <bold><italic>OREGON</italic>:</bold><italic>Benton Co.</italic>: Corvallis, VIII.1919, W.J. Chamberlin, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">taxifolia</named-content></named-content></italic> [= <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic>] (EMEC-1), Kiger Island, VII.[19]22, W.J. Chamberlin (EMEC-2). Mary’s Peak, 14 mi W. Corvallis, XII.1958 (EMEC-2). [<italic>Deschutes Co.</italic>]: Bend, Hopk. U.S. 33,531-B, 8.VII.[19]58, W.J. Buckhorn, P.W. Orr, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (OSAC-8). [<italic>Klamath Co.</italic>]: Klamath Falls, Geary Ranch, 4.X.1962, J. Schuh, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (CNCI-2, FMNH-4, MSUC-7). [<italic>Linn Co.</italic>]: McMinnville, 7.VIII.1951, R. Kangur (WFBM-2). [<italic>Wallowa Co.</italic>]: Wallowa Lake, 19.VII.[19]51, Quintus, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (WFBM-1). <bold><italic>UTAH</italic>:</bold><italic>Juab Co.</italic>: Mount Nebo, 20.VII.1958, D.E. Bright, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (CNCI-2). <italic>Utah Co.</italic>: Payson Canyon, 20.V.1961, S.L. Wood, ex. <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (USNM-18). <bold><italic>WASHINGTON</italic>:</bold> [<italic>Jefferson Co.</italic>]: Quilcene, Hopk. U.S. 65564, 30.IX.1946 (OSAC-6). [<italic>Pend Oreille Co.</italic>]: Metaline Falls, Hopk. U.S. 21405, 17.VII.[19]31, W.D. Bedard, ex. flying (USNM-4). [<italic>Stevens Co.</italic>]: Northport, 1.IX.1930, R. Hopping (CASC-3). [<italic>Walla Walla Co.</italic>]: Walla Walla, VIII.1933, M.C. Lume (USNM-11). [<italic>Yakima Co.</italic>]: [Mount Baker], Snoqualmie National Forest, Dry Creek Ridge, 14 mi W. of Naches, 3.VII.1965, R.B. Hutt (DEBC-1).</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>CANADA: British Columbia. MEXICO: Baja California. UNITED STATES: Arizona, California, Colorado, Idaho, Montana, Nevada, New Mexico, Oregon, Utah, Washington, Wyoming (Fig. <xref ref-type="fig" rid="F40">40</xref>).</p></sec><sec sec-type="treatment-Hosts"><title>Hosts.</title><p>Principle hosts: <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">concolor</named-content></named-content></italic> (Gord. & Glend.) Lindl. ex Hildebr. (white fir), <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">grandis</named-content></named-content></italic> (Douglas ex D. Don) Lindl. (grand fir), and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">magnifica</named-content></named-content></italic> A. Murray (red fir). Incidental hosts: <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">lasiocarpa</named-content></named-content></italic> (Hook.) Nutt. (subalpine fir).</p></sec><sec sec-type="treatment-Common name"><title>Common name.</title><p>Fir engraver.</p></sec><sec sec-type="treatment-Biology"><title>Biology.</title><p><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> can cause significant fir mortality and is the most destructive conifer-feeding <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species in the Nearctic (<xref rid="B77" ref-type="bibr">Keen 1938</xref>; <xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>). During a period from 1924–1936, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> killed 15% and damaged an additional 25% of the merchantable fir in California. It has also been reported to be quite destructive in Oregon (<xref rid="B77" ref-type="bibr">Keen 1938</xref>). <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> is associated with a symbiotic stain fungus, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Trichasporium</named-content><named-content content-type="species">symbioticum</named-content></named-content></italic> Wright, which the adult beetle introduces when it excavates the adult gallery. The fungus spreads in all directions around the gallery system (<xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>). Due to the potential of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> to kill fir trees, this species is the most well studied native <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> in North America. Attacks usually occur on the boles of weakened and stressed standing trees from a few feet above the base to the top of the tree, but can also occur in large slash and fresh and fallen trees (<xref rid="B25" ref-type="bibr">Chamberlin 1958</xref>; <xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>). Attacks at the top of the tree are more common on overmature standing trees during drought and healthy, vigorous trees are not preferred (<xref rid="B25" ref-type="bibr">Chamberlin 1958</xref>; <xref rid="B106" ref-type="bibr">Raffa and Berryman 1987</xref>). Trees can also become successively attacked over a period of years and slowly die. Healthy trees may survive the attacks but can develop rots and defects that reduce timber value (<xref rid="B127" ref-type="bibr">Struble 1937</xref>). Unlike most tree-killing or primary bark beetles, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> does not have pheromone to aggregate conspecifics to host trees. The beetles locate suitable hosts via primary attraction to host volatiles (<xref rid="B89" ref-type="bibr">Macías-Sámano et al. 1998a</xref>,<xref rid="B90" ref-type="bibr">b</xref>).</p><p>Adult galleries are perpendicular to the grain of the wood (Fig. <xref ref-type="fig" rid="F25">25</xref>), deeply score the sapwood, lightly score the cambium and consist of two egg galleries with a central nuptial chamber (<xref rid="B42" ref-type="bibr">Edson 1967</xref>). The nuptial chamber is typically short and at a right angle to the egg galleries but may also extend at an oblique angle. When this occurs, one of the egg galleries is briefly extended at an oblique angle against the grain before becoming perpendicular to the grain (<xref rid="B42" ref-type="bibr">Edson 1967</xref>). Eggs are deposited singly in triangular niches spaced 1.0–1.5 mm apart on each side of the egg gallery with 80–300 niches per gallery. Galleries range in size from 8.0–30.0 cm in length (<xref rid="B25" ref-type="bibr">Chamberlin 1958</xref>; <xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>; <xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>). Larval mines are perpendicular to the egg gallery and parallel to the grain of the wood. Larval mines are also parallel to each other both above and below the egg gallery, giving the gallery a diamond shaped appearance (<xref rid="B77" ref-type="bibr">Keen 1938</xref>; <xref rid="B42" ref-type="bibr">Edson 1967</xref>). Larval mines lightly score the <pmc-comment>PageBreak</pmc-comment>sapwood and deeply score the cambium. Larvae construct pupation chambers in the phloem or outer bark (<xref rid="B42" ref-type="bibr">Edson 1967</xref>). Broods overwinter as larvae or adults (<xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>). The number of generations per year varies both geographically and with elevation. Development time can range from as little as 41 days at low latitudes and elevations to as many as 380 days at high latitudes and elevations (<xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>). There is typically one generation per year (<xref rid="B20" ref-type="bibr">Bright and Stark 1973</xref>). In Idaho pupation occurs from June to July and peak flight occurs in July (<xref rid="B52" ref-type="bibr">Furniss and Johnson 2002</xref>).</p></sec><sec sec-type="treatment-Remarks"><title>Remarks.</title><p>The lectotype does not bear a locality label. <xref rid="B82" ref-type="bibr">LeConte’s (1868)</xref> description states that George Gibbs collected the lectotype in the Washington Territory.</p></sec></sec></sec><sec sec-type="Key to the Nearctic adults of Scolytus"><title>Key to the Nearctic adults of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic></title><p>This is the first key to both sexes of Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> adults. Unlike previous keys (e.g. <xref rid="B42" ref-type="bibr">Edson 1967</xref>; <xref rid="B148" ref-type="bibr">Wood 1982</xref>) apriori knowledge of the gender is not required. Only one sex is necessary to identify the specimen and host species or gallery types are not needed. Because the sexual dimorphic characters are not consistent for the Nearctic species (see sexual dimorphism discussion above), the sexes could not be evenly split. In general, males and females of native hardwood-feeding species will key out beginning at couplet 3 and both sexes of some <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content></named-content></italic> feeding species beginning at couplet 23. The key lists the gender of the identified specimen. If no gender is specified, the identification applies to both sexes. The length of frons setae is measured relative to the width of the midpoint of the eye.</p><table-wrap content-type="key" orientation="portrait" id="d36e29322" position="anchor"><table frame="hsides" rules="all"><tbody><tr><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">Elytral apices narrowly rounded (Fig. <xref ref-type="fig" rid="F60">60A</xref>)</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic></bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Elytral apices subquadrate (Fig. <xref ref-type="fig" rid="F60">60B</xref>) or broadly rounded</td><td rowspan="1" colspan="1"><bold>2</bold></td></tr><tr><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">Lateral profile of frons clearly flattened and/or impressed (Fig. <xref ref-type="fig" rid="F1">1A</xref>)</td><td rowspan="1" colspan="1"><bold>3</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Lateral profile of frons convex (Fig. <xref ref-type="fig" rid="F1">1C</xref>)</td><td rowspan="1" colspan="1"><bold>23</bold></td></tr><tr><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">Apical margin of ventrite 3 and/or 4 armed with teeth or spines laterally and/ or medially</td><td rowspan="1" colspan="1"><bold>4</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Apical margin of ventrite 3 and 4 unarmed</td><td rowspan="1" colspan="1"><bold>7</bold></td></tr><tr><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">Ventrite 3 and 4 armed with lateral teeth or spines</td><td rowspan="1" colspan="1"><bold>5</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Ventrites 3 and 4 not armed with lateral teeth or spines</td><td rowspan="1" colspan="1"><bold>6</bold></td></tr><tr><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">Apical margin of ventrite 3 armed by three acute spines (two lateral and one medial); apical margin of ventrite 4 armed by one median tooth; ventrite 1 apically descending; ventrite 2 deeply concave, basal margin produced and bearing a median tubercle</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Apical margin of ventrite 3 and 4 each armed by two lateral teeth; ventrite 1 horizontal; ventrite 2 convex and bearing a blunt median tubercle on the basal margin</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">6</td><td rowspan="1" colspan="1">Apical margin of ventrite 4 armed by an acute median denticle; apical margins of ventrites 2 and 3 may also each bear a smaller median denticle (variable)</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1"><pmc-comment>PageBreak</pmc-comment>–</td><td rowspan="1" colspan="1">Apical margin of ventrite 4 thickened forming a broad carina with a blunt median tubercle; apical margins of ventrites 2 and 3 always unarmed</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">7</td><td rowspan="1" colspan="1">Ventrites 2–4 covered with abundant, erect, long, hair-like setae</td><td rowspan="1" colspan="1"><bold>8</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Ventrites 2–4 glabrous or covered with minute ground vestiture or short, sparse, recumbent, hair-like or scale-like setae</td><td rowspan="1" colspan="1"><bold>10</bold></td></tr><tr><td rowspan="1" colspan="1">8</td><td rowspan="1" colspan="1">Frons with hair-like setae on lateral and dorsal margins thicker, longer, incurved, remaining areas of frons largely devoid of setae; frons strongly longitudinally aciculate</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic></bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Frons with equally distributed hair-like setae of uniform length; frons weakly to moderately longitudinally aciculate or weakly aciculate-punctate</td><td rowspan="1" colspan="1"><bold>9</bold></td></tr><tr><td rowspan="1" colspan="1">9</td><td rowspan="1" colspan="1">Apical margin of elytra produced between interstriae 1 and 2, deeply emarginate at interstria 3, produced on interstria 4 and deeply emarginate at stria 4; ventrite 2 armed with a laterally compressed tubercle that extends from the apical margin of ventrite 2 to approximately ¾ the length of ventrite; frons weakly longitudinally aciculate, strongly punctate; epistoma strongly emarginate</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Apical margin of elytra slightly emarginate at interstriae 3; apical margin of ventrite 2 armed with a median denticle; frons moderately longitudinally aciculate, almost impunctate; epistoma entire to faintly emarginate</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">10</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 rounded; basal margin of ventrite 2 flat or marked by a weak carina; surface of ventrite 2 convex, often armed with a spine or tubercle (Fig. <xref ref-type="fig" rid="F61">61A</xref>)</td><td rowspan="1" colspan="1"><bold>11</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 thickened or produced on the ventral and/or lateral margins, forming a carinate lip along the basal margin of ventrite 2, often weakly produced in <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic>; surface of ventrite 2 flat to impressed (Fig. <xref ref-type="fig" rid="F61">61B</xref>)</td><td rowspan="1" colspan="1"><bold>19</bold></td></tr><tr><td rowspan="1" colspan="1">11</td><td rowspan="1" colspan="1">Frons impressed just above epistoma or medially impressed, strongly longitudinally aciculate</td><td rowspan="1" colspan="1"><bold>12</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Frons flat, weakly to moderately longitudinally aciculate or coarsely punctate</td><td rowspan="1" colspan="1"><bold>15</bold></td></tr><tr><td rowspan="1" colspan="1">12</td><td rowspan="1" colspan="1">Ventrite 2 unarmed; elytral surface shining and glabrous</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Ventrite 2 armed with a spine; elytral surface dull, sparse setae present (rarely the spine on ventrite 2 is absent in <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic>)</td><td rowspan="1" colspan="1"><bold>13</bold></td></tr><tr><td rowspan="1" colspan="1">13</td><td rowspan="1" colspan="1">Base of spine touching apical margin of ventrite 2; frons strongly longitudinally aciculate</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Base of spine not touching apical margin of ventrite 2; frons strigose-punctate</td><td rowspan="1" colspan="1"><bold>14</bold></td></tr><tr><td rowspan="1" colspan="1">14</td><td rowspan="1" colspan="1">Spine conical, narrow; elytra always unicolorous; elytral disc shining and glabrous; subapical carina on ventrite 5 located 1/2 length of segment from apex</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1"><pmc-comment>PageBreak</pmc-comment>–</td><td rowspan="1" colspan="1">Spine broadly conical with blunted apex; elytra often with a banded appearance; subapical carina on ventrite 5 located just before end of segment</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">15</td><td rowspan="1" colspan="1">Ventrite 2 armed with a rounded spine</td><td rowspan="1" colspan="1"><bold>16</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Ventrite 2 unarmed</td><td rowspan="1" colspan="1"><bold>17</bold></td></tr><tr><td rowspan="1" colspan="1">16</td><td rowspan="1" colspan="1">Spine on ventrite 2 extending from apical margin to three-fourths length of ventrite; ventrite 2 surface shining (Fig. <xref ref-type="fig" rid="F62">62A</xref>)</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Spine on ventrite 2 extending from apical margin to one half length of ventrite; ventrite 2 surface dull (Fig. <xref ref-type="fig" rid="F62">62B</xref>)</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">17</td><td rowspan="1" colspan="1">Ventrite 5 length at middle longer than or equal to that of ventrites 3 and 4 combined (Fig. <xref ref-type="fig" rid="F63">63A</xref>); ventrite 5 without a transverse carina near base</td><td rowspan="1" colspan="1"><bold>18</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Ventrite 5 length at middle shorter than that of ventrites 3 and 4 combined (Fig. <xref ref-type="fig" rid="F63">63B</xref>); ventrite 5 with a transverse carinate ridge near base that may or may not be reflexed (variable)</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1"><pmc-comment>PageBreak</pmc-comment>18</td><td rowspan="1" colspan="1">Frons either granulate or faintly longitudinally aciculate; frons with setae uniformly distributed, fewer setae on lateral and dorsal margins, shorter, finer; elytra glabrous (except on declivity); ventrite 5 unarmed</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Frons moderately longitudinally aciculate, with long, fine, incurved setae predominately on lateral and dorsal margins, fewer, shorter and finer setae medially; elytra with minute ground vestiture</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">19</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 moderately produced and not forming a distinct lip along base of ventrite 2</td><td rowspan="1" colspan="1">2<bold>0</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 distinctly thickened or strongly produced, forming a lip along base of ventrite 2</td><td rowspan="1" colspan="1"><bold>21</bold></td></tr><tr><td rowspan="1" colspan="1">20</td><td rowspan="1" colspan="1">Surface of ventrite 2 shining but minutely reticulate; elytral striae not impressed; basal margin of ventrite 2 more pronounced and produced laterally; elytral strial punctures small, spaced 2–3 diameters of a puncture (Fig. <xref ref-type="fig" rid="F64">64A</xref>)</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Surface of ventrite 2 opaque; elytral discal striae impressed; basal margin of ventrite 2 continuously and evenly elevated; elytral strial punctures large, spaced 1–2 diameters of a puncture (Fig. <xref ref-type="fig" rid="F64">64B</xref>)</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">21</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 distinctly thickened, at most slightly produced over base of ventrite 2</td><td rowspan="1" colspan="1"><bold>22</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 strongly acutely produced forming a lip along base of ventrite 2, surface of ventrite 2 appearing concave</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">22</td><td rowspan="1" colspan="1">Base of ventrite 2 strongly thickened; surface of ventrite 2 often weakly medially impressed just above the base; apical margin of ventrite 2 unarmed (Fig. <xref ref-type="fig" rid="F65">65A</xref>)</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Base of ventrite 2 faintly elevated; surface of ventrite 2 flat; apical margin of ventrite 2 slightly elevated often with a median denticle (Fig. <xref ref-type="fig" rid="F65">65B</xref> )</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1"><pmc-comment>PageBreak</pmc-comment>23</td><td rowspan="1" colspan="1">Apical margin of elytra produced between interstriae 1 and 2, deeply emarginate at interstria 3, produced on interstria 4 and deeply emarginate at stria 4</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Apical margin of elytra entire or slightly emarginate at interstriae 3</td><td rowspan="1" colspan="1"><bold>24</bold></td></tr><tr><td rowspan="1" colspan="1">24</td><td rowspan="1" colspan="1">Ventrite 2 armed either on the surface or on apical margin by either a spine or a low median tumescence</td><td rowspan="1" colspan="1"><bold>25</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Ventrite 2 unarmed</td><td rowspan="1" colspan="1"><bold>35</bold></td></tr><tr><td rowspan="1" colspan="1">25</td><td rowspan="1" colspan="1">Surface of ventrite 2 armed with a rounded, keel shaped or broadly acute spine that is at least as long as its basal width</td><td rowspan="1" colspan="1"><bold>26</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Surface of ventrite 2 apical margin armed by a low median tumescence or small denticle</td><td rowspan="1" colspan="1"><bold>30</bold></td></tr><tr><td rowspan="1" colspan="1">26</td><td rowspan="1" colspan="1">Base of spine on ventrite 2 touches the basal margin of segment; apical margins of ventrites 3 and 4 each armed by two lateral teeth</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Base of spine on ventrite 2 never touches basal margin of segment; apical margins of ventrites 3 and 4 unarmed</td><td rowspan="1" colspan="1"><bold>27</bold></td></tr><tr><td rowspan="1" colspan="1">27</td><td rowspan="1" colspan="1">Base of spine not touching apical margin of ventrite 2</td><td rowspan="1" colspan="1"><bold>28</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Base of spine touching apical margin of ventrite 2</td><td rowspan="1" colspan="1"><bold>29</bold></td></tr><tr><td rowspan="1" colspan="1">28</td><td rowspan="1" colspan="1">Spine conical, narrow; elytra always unicolorous; elytral disc shining and glabrous; subapical carina on ventrite 5 located 1/5 length of segment from apex</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Spine broadly conical with blunted apex; elytra often with a banded appearance; subapical carina on ventrite 5 located just before end of segment</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1"><pmc-comment>PageBreak</pmc-comment>29</td><td rowspan="1" colspan="1">Ventrite 2 armed with a stout and broadly acute spine extending from the apical margin to one half-length of the ventrite, never with a longitudinal carina</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Ventrite 2 armed with a longitudinal carina and a blunt tubercle, appearing keel-shaped, or a low median longitudinal carina</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic>, in part</bold></td></tr><tr><td rowspan="1" colspan="1">30</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 produced, forming a carinate lip along the basal margin of ventrite 2; ventrite 2 appearing impressed (Fig. <xref ref-type="fig" rid="F61">61B</xref>)</td><td rowspan="1" colspan="1"><bold>31</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 rounded; basal margin of ventrite 2 flat or marked by a weak carina; surface of ventrite 2 convex (Fig. <xref ref-type="fig" rid="F61">61A</xref>)</td><td rowspan="1" colspan="1"><bold>34</bold></td></tr><tr><td rowspan="1" colspan="1">31</td><td rowspan="1" colspan="1">Apical margin of ventrite 2 weakly longitudinally tumescent, never pointed</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Apical margin of ventrite 2 with a small median denticle</td><td rowspan="1" colspan="1"><bold>32</bold></td></tr><tr><td rowspan="1" colspan="1"><pmc-comment>PageBreak</pmc-comment>32</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 slightly elevated; ventrite 2 appearing convex; median denticle on apical margin of ventrite 2 broad, broadly pointed</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 strongly, acutely produced, forming a lip along the base of ventrite 2, surface of ventrite 2 appearing impressed, median denticle on apical margin of ventrite 2 narrow, acutely pointed</td><td rowspan="1" colspan="1"><bold>33</bold></td></tr><tr><td rowspan="1" colspan="1">33</td><td rowspan="1" colspan="1">Second ventrite punctures abundant, fine and moderately impressed, appearing distinct; second ventrite subopaque</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></named-content></italic></bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Second ventrite punctures sparse, fine and shallow, almost appearing indistinct; second ventrite appearing strongly opaque</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1"><pmc-comment>PageBreak</pmc-comment>34</td><td rowspan="1" colspan="1">Frons finely longitudinally aciculate-punctate (Fig. <xref ref-type="fig" rid="F66">66A</xref>); ventrite 1 joining base of ventrite 2 obtusely, base of ventrite 2 not finely impressed, flush with ventrite 1</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Frons moderately and coarsely longitudinally aciculate-punctate (Fig. <xref ref-type="fig" rid="F66">66B</xref>); ventrite 1 rounded over onto surface of ventrite 2, not forming an obtuse angle, base of ventrite 2 finely impressed</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">35</td><td rowspan="1" colspan="1">Epistomal process absent (Fig. <xref ref-type="fig" rid="F67">67A</xref>)</td><td rowspan="1" colspan="1"><bold>36</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Epistomal process present, may be indistinct (Fig. <xref ref-type="fig" rid="F67">67B</xref>)</td><td rowspan="1" colspan="1"><bold>37</bold></td></tr><tr><td rowspan="1" colspan="1">36</td><td rowspan="1" colspan="1">Frons weakly longitudinally aciculate, strongly punctate, with uniformly distributed long setae; elytral striae weakly impressed</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Frons finely longitudinally aciculate and glabrous or minutely setose; elytral striae not impressed</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">37</td><td rowspan="1" colspan="1">Setae on frons at least 1.5 times the width of the eye at the middle</td><td rowspan="1" colspan="1"><bold>38</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Setae on frons equal in length to the width of the eye at the middle</td><td rowspan="1" colspan="1"><bold>42</bold></td></tr><tr><td rowspan="1" colspan="1">38</td><td rowspan="1" colspan="1">Ventrites 2–5 shining; thorax and apical and lateral margins of elytra covered with long hair-like setae as long as the length of ventrites 3 and 4 combined, with pointed apices</td><td rowspan="1" colspan="1"><bold>39</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Ventrites 2–5 opaque; thorax covered with setae as long as the length of ventrite 3, with quadrate apices</td><td rowspan="1" colspan="1"><bold>41</bold></td></tr><tr><td rowspan="1" colspan="1">39</td><td rowspan="1" colspan="1">Epistomal process weakly developed, and faintly emarginate medially</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Epistomal process strongly developed, and strongly emarginate medially</td><td rowspan="1" colspan="1"><bold>40</bold></td></tr><tr><td rowspan="1" colspan="1">40</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 produced, forming a carinate lip along the basal margin of ventrite 2 that is about half as produced as thick; <italic><named-content content-type="taxon-name"><named-content content-type="genus">Pseudotsuga</named-content><named-content content-type="species">menziesii</named-content></named-content></italic></td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1"><pmc-comment>PageBreak</pmc-comment>–</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 produced, forming a carinate lip along the basal margin of ventrite 2 that is twice as produced as thick; <italic><named-content content-type="taxon-name"><named-content content-type="genus">Abies</named-content><named-content content-type="species">religiosa</named-content></named-content></italic></td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></named-content></italic> (male)</bold></td></tr><tr><td rowspan="1" colspan="1">41</td><td rowspan="1" colspan="1">Frons distinctly, moderately longitudinally aciculate, weakly punctate (Fig. <xref ref-type="fig" rid="F68">68A</xref>); apical margin of ventrite 1 weakly produced, forming a weak carinate lip along the basal margin of ventrite 2</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Frons indistinctly, weakly longitudinally aciculate, strongly punctate (Fig. <xref ref-type="fig" rid="F68">68B</xref>); apical margin of ventrite 1 flush with basal margin of ventrite 2, appearing rounded</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1"><pmc-comment>PageBreak</pmc-comment>42</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 rounded (Fig. <xref ref-type="fig" rid="F61">61A</xref>); surface of ventrite 2 rugose, shining, coarsely punctate and convex</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 with thickened lip or produced (Fig. <xref ref-type="fig" rid="F61">61B</xref>), never rounded; surface of ventrite 2 smooth and flat</td><td rowspan="1" colspan="1"><bold>43</bold></td></tr><tr><td rowspan="1" colspan="1">43</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 thickened, on surface of ventrite 2 and nearly flush with the surface of ventrite 2</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 not thickened or on the surface of ventrite 2, often slightly apically produced (unequally carinate in <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic>)</td><td rowspan="1" colspan="1"><bold>44</bold></td></tr><tr><td rowspan="1" colspan="1">44</td><td rowspan="1" colspan="1">Ventrite 2 punctulate with small or minute, shallow punctures</td><td rowspan="1" colspan="1"><bold>45</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Ventrite 2 distinctly punctate with large and deep punctures</td><td rowspan="1" colspan="1"><bold>47</bold></td></tr><tr><td rowspan="1" colspan="1">45</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 equally and continuously carinate (Fig. <xref ref-type="fig" rid="F69">69A</xref>)</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic>, in part (female)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Apical margin of ventrite 1 unequally carinate, more laterally produced forming cusps on each side of ventrite 2 (Fig. <xref ref-type="fig" rid="F69">69B</xref>)</td><td rowspan="1" colspan="1"><bold>46</bold></td></tr><tr><td rowspan="1" colspan="1">46</td><td rowspan="1" colspan="1">Elytral discal striae not impressed; ventrite 2 shining in luster (Fig. <xref ref-type="fig" rid="F70">70A</xref>)</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Elytral discal striae impressed, giving the elytra a corrugated appearance; ventrite 2 opaque, dull in luster (Fig. <xref ref-type="fig" rid="F70">70B</xref>)</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">47</td><td rowspan="1" colspan="1">Ventrite 2 shining</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Ventrite 2 dull, opaque</td><td rowspan="1" colspan="1"><bold>48</bold></td></tr><tr><td rowspan="1" colspan="1">48</td><td rowspan="1" colspan="1">Epistomal process strongly developed, distinct; frons evenly convex</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></named-content></italic> (female)</bold></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Epistomal process weakly developed, almost indistinct; frons flattened or convex with slight medial depression</td><td rowspan="1" colspan="1"><bold><italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic>, in part</bold></td></tr><pmc-comment>PageBreak</pmc-comment>
</tbody></table></table-wrap><fig id="F60" orientation="portrait" position="float"><label>Figure 60.</label><caption><p>Elytral apices consist of two general shapes: <bold>A</bold> narrowly rounded (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic>) <bold>B</bold> subquadrate (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic>).</p></caption><graphic id="oo_33413.jpg" xlink:href="zookeys-450-001-g060"></graphic></fig><fig id="F61" orientation="portrait" position="float"><label>Figure 61.</label><caption><p>Apical margin of ventrite 1: <bold>A</bold> rounded (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic>) <bold>B</bold> thickened and produced forming a carinate lip (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></named-content></italic>).</p></caption><graphic id="oo_33414.jpg" xlink:href="zookeys-450-001-g061"></graphic></fig><fig id="F62" orientation="portrait" position="float"><label>Figure 62.</label><caption><p>Spine on ventrite 2: <bold>A</bold> extending from apical margin to three-fourths length of ventrite <bold>B</bold> extending from apical margin to one half length of ventrite (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic>)</p></caption><graphic id="oo_33415.jpg" xlink:href="zookeys-450-001-g062"></graphic></fig><fig id="F63" orientation="portrait" position="float"><label>Figure 63.</label><caption><p>Length of ventrite 5 at middle: <bold>A</bold> longer than or equal to that of ventrites 3 and 4 combined (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic>) <bold>B</bold> shorter than that of ventrites 3 and 4 combined (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic>).</p></caption><graphic id="oo_33416.jpg" xlink:href="zookeys-450-001-g063"></graphic></fig><fig id="F64" orientation="portrait" position="float"><label>Figure 64.</label><caption><p>Surface of ventrite 2: <bold>A</bold> shining but minutely reticulate (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic>) <bold>B</bold> opaque (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic>).</p></caption><graphic id="oo_33417.jpg" xlink:href="zookeys-450-001-g064"></graphic></fig><fig id="F65" orientation="portrait" position="float"><label>Figure 65.</label><caption><p>Base of ventrite 2: <bold>A</bold> distinctly thickened (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></named-content></italic>) <bold>B</bold> faintly elevated (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic>).</p></caption><graphic id="oo_33418.jpg" xlink:href="zookeys-450-001-g065"></graphic></fig><fig id="F66" orientation="portrait" position="float"><label>Figure 66.</label><caption><p>Frons: <bold>A</bold> finely longitudinally aciculate-punctate (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></named-content></italic>) <bold>B</bold> moderately and coarsely longitudinally aciculate-punctate (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></named-content></italic>).</p></caption><graphic id="oo_33419.jpg" xlink:href="zookeys-450-001-g066"></graphic></fig><fig id="F67" orientation="portrait" position="float"><label>Figure 67.</label><caption><p>Epistomal process: <bold>A</bold> absent (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic>) <bold>B</bold> present (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></named-content></italic>).</p></caption><graphic id="oo_33420.jpg" xlink:href="zookeys-450-001-g067"></graphic></fig><fig id="F68" orientation="portrait" position="float"><label>Figure 68.</label><caption><p>Frons: <bold>A</bold> distinctly, moderately longitudinally aciculate, weakly punctate (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></named-content></italic>) <bold>B</bold> indistinctly, weakly longitudinally aciculate, strongly punctate (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic>).</p></caption><graphic id="oo_33421.jpg" xlink:href="zookeys-450-001-g068"></graphic></fig><fig id="F69" orientation="portrait" position="float"><label>Figure 69.</label><caption><p>Apical margin of ventrite 1: <bold>A</bold> equally and continuously carinate (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></named-content></italic>) <bold>B</bold> unequally carinate, more laterally produced forming cusps on each side of ventrite 2 (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic>).</p></caption><graphic id="oo_33422.jpg" xlink:href="zookeys-450-001-g069"></graphic></fig><fig id="F70" orientation="portrait" position="float"><label>Figure 70.</label><caption><p>Ventrite 2: <bold>A</bold> shining in luster (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></named-content></italic>) <bold>B</bold> opaque, dull in luster (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></named-content></italic>).</p></caption><graphic id="oo_33423.jpg" xlink:href="zookeys-450-001-g070"></graphic></fig></sec></sec><sec sec-type="Discussion"><title>Discussion</title><p>Morphology failed to resolve the <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> phylogeny and very few synapomorphic characters were found (Fig. <xref ref-type="fig" rid="F2">2</xref>). Unique combinations of homoplastic characters define species limits. Characters that are among the most informative are sexually dimorphic and male including: male fifth ventrite carina (character 36), size and coarseness of frons punctures (character 7), first ventrite apical margin (character 27), second ventrite surface (character 29) and presence of an epistomal process (character 2). Other non-dimorphic informative characters included elytral apex emargination (character 20), host (character 43), and strial impression (character 18). The lack of phylogenetically informative characters necessitated the need for further investigation using molecular characters to illuminate species boundaries.</p><p>As in other phylogenetic studies, COI data failed to recover many species as monophyletic (Fig. <xref ref-type="fig" rid="F3">3</xref>) (<xref rid="B35" ref-type="bibr">Cognato et al. 2005</xref>; <xref rid="B70" ref-type="bibr">Jordal and Hewitt 2004</xref>). Several relationships found in this phylogeny did not agree with the topologies obtained from different analyses. For example a hardwood-feeder, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></named-content></italic>, was found to be sister to the conifer clade in the Bayesian analysis and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></named-content></italic> was found to be sister to the Nearctic conifer clade. In addition, the hardwood clade was paraphyletic with <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic> forming one lineage and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic> as sister to <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></named-content></italic>. Saturation of nucleotide substitutions and a lack of lineage sorting are potential reasons for the discordance among these mitochondrial and nuclear phylogenies (<xref rid="B124" ref-type="bibr">Sota and Vogler 2001</xref>; <xref rid="B50" ref-type="bibr">Funk and Ohmland 2003</xref>; <xref rid="B88" ref-type="bibr">Lin and Danforth 2004</xref>). The mitochondrial phylogeny may suffer from both phenomena because of the observed poor resolution and support for deeper nodes and short-branch lengths among the conifer-feeding species (Fig. <xref ref-type="fig" rid="F3">3</xref>). As with other studies, concatenating the COI dataset with multiple genes and morphology remedied its deficiency and elucidated the relationships among the <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> species (<xref rid="B70" ref-type="bibr">Jordal and Hewitt 2004</xref>).</p><pmc-comment>PageBreak</pmc-comment>
<p>When analyzed together, the genes used in this study (COI, 28S, CAD, ArgK) were useful in resolving the <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> phylogeny and species limits. These genes have demonstrated similar phylogenetic utility in other scolytine studies (<xref rid="B33" ref-type="bibr">Cognato and Sperling 2000</xref>; <xref rid="B49" ref-type="bibr">Farrell et al. 2001</xref>; <xref rid="B72" ref-type="bibr">Jordal et al. 2002</xref>; <xref rid="B70" ref-type="bibr">Jordal and Hewitt 2004</xref>; <xref rid="B35" ref-type="bibr">Cognato et al. 2005</xref>; <xref rid="B34" ref-type="bibr">Cognato and Sun 2007</xref>; <xref rid="B73" ref-type="bibr">Jordal et al. 2008</xref>; <xref rid="B41" ref-type="bibr">Dole et al. 2010</xref>; <xref rid="B36" ref-type="bibr">Cognato et al. 2011</xref>; <xref rid="B74" ref-type="bibr">Jordal et al. 2011</xref>; <xref rid="B71" ref-type="bibr">Jordal and Cognato 2012</xref>). Overall, the model-based rates of nucleotide evolution utilized in Bayesian analysis combined with all four genes and morphology allowed us to recover a well-supported and sufficiently resolved phylogeny which enabled the revision of <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> (Fig. <xref ref-type="fig" rid="F4">4</xref>).</p><p>Nearctic <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content></named-content></italic> were recovered as paraphyletic in two clades, native hardwood and conifer (Fig. <xref ref-type="fig" rid="F4">4</xref>). A group of introduced Palearctic species was also found. Members of the native hardwood clade are sister to the <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">scolytus</named-content></named-content></italic> subgenus <italic>sensu</italic><xref rid="B23" ref-type="bibr">Butovitsch (1929)</xref> rather than the native conifer clade. This is supported by morphological similarities between the two groups. The clade containing the <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">scolytus</named-content></named-content></italic> subgenus <italic>sensu</italic> Butovitsch and the native hardwood-feeders share a median impression on male ventrite 5 (except <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laevis</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic>) and a setal patch on ventrite 5 (except <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laevis</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></named-content></italic>). Several species in these groups also share the absence of a carina on male ventrite 5 (<italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laevis</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ratzeburgii</named-content></named-content></italic>, <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></named-content></italic> and <italic><named-content content-type="taxon-name"><named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></named-content></italic>).</p></sec><sec sec-type="supplementary-material"><title>Supplementary Material</title><supplementary-material id="zookeys.450.7452-treatment1" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment1.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment2" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mali</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment2.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment3" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">multistriatus</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment3.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment4" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">rugulosus</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment4.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment5" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">schevyrewi</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment5.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment6" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fagi</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment6.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment7" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">muticus</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment7.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment8" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">quadrispinosus</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment8.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment9" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">aztecus</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment9.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment10" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">dentatus</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment10.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment11" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">fiskei</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment11.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment12" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">hermosus</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment12.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment13" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">laricis</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment13.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment14" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">monticolae</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment14.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment15" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">mundus</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment15.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment16" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">obelus</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment16.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment17" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">oregoni</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment17.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment18" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">piceae</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment18.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment19" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">praeceps</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment19.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment20" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">reflexus</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment20.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment21" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">robustus</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment21.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment22" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">silvaticus</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment22.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment23" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">subscaber</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment23.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment24" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">tsugae</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment24.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment25" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">unispinosus</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment25.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.450.7452-treatment26" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Scolytus</named-content><named-content content-type="species">ventralis</named-content></title></caption><media xlink:href="zookeys.450.7452-treatment26.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material></sec></body><back><ack><title>Acknowledgements</title><p>This study was enhanced by thoughtful discussions with Donald Bright, Malcolm Furniss, Gary Parsons, Deborah McCullough, Therese Poland, Aaron Smith and Bernice DeMarco. Special thanks to Thomas Atkinson (University of Texas) for assistance with creating distribution maps. The manuscript was greatly improved through the thoughtful comments of the two reviewers Bjarte Jordal (University of Bergen), and Don Bright. Lewis Edson graciously allowed us to use his drawings from his <xref rid="B42" ref-type="bibr">1967</xref> publication.</p><p>Fieldwork would not have been as productive without the assistance of Debra Allen-Reid (USDA-FS), Barbara Barr, Donald Bright, Ken Bush, Malcolm Furniss, and Alan and Pat Gillogly. Specimens were generously donated to this research by the following individuals: Thomas Atkinson, Jiri Hulcr (University of Florida), Miloš Knížek (Forestry and Game Management Research Institute), Bjarte Jordal, Åke Lindelöw (Swedish University of Agricultural Sciences), Alexander Petrov and Robert Rabaglia (USDA-FS). David Furth (USNM) facilitated the senior author’s studies at the Smithsonian. Thomas Smith provided logistical and photographic assistance. We also wish to thank the curators listed herein for providing access to the specimens used in this investigation.</p><p>This research was supported by USDA-FS Early Detection and Rapid Response of Exotic Bark Beetles (07-DG-11420004-182), Michigan State University Competitive Discretionary Funding Program awarded to AIC. Support was also provided by the Norwegian Research Council (214232) and the University of Bergen, Norway (F20) awarded to Bjarte H. Jordal (University of Bergen). Additional funding from <pmc-comment>PageBreak</pmc-comment>the American Museum of Natural History Theodore Roosevelt Memorial Grant, Entomological Society of America Systematics and Evolutionary Biology student travel grant, Museum of Comparative Zoology Ernst Mayr Travel Grant in Animal Systematics, Michigan State University (Department of Entomology Hutson Funds and the Hagen Bashian Trust support scholarship from the MSU college of Agriculture and Natural Resources awarded to SMS.</p></ack><ref-list><title>References</title><ref id="B1"><mixed-citation publication-type="journal"><person-group><name><surname>Alonso-Zarazaga</surname><given-names>MA</given-names></name><name><surname>Lyal</surname><given-names>CHC</given-names></name></person-group> (<year>2009</year>) <article-title>A catalogue of family and genus group names in <named-content content-type="taxon-name"><named-content content-type="subfamily">Scolytinae</named-content></named-content> and <named-content content-type="taxon-name"><named-content content-type="subfamily">Platypodinae</named-content></named-content> with nomenclatural remarks (<named-content content-type="taxon-name"><named-content content-type="order">Coleoptera</named-content></named-content>: <named-content content-type="taxon-name"><named-content content-type="family">Curculionidae</named-content></named-content>).</article-title><source>Zootaxa</source><volume>2258</volume>: <fpage>1</fpage>–<lpage>134</lpage>.</mixed-citation></ref><ref id="B2"><mixed-citation publication-type="book"><person-group><name><surname>Arnett</surname><given-names>RH</given-names><suffix>Jr</suffix></name><name><surname>Samuelson</surname><given-names>GA</given-names></name><name><surname>Nishida</surname><given-names>GM</given-names></name></person-group> (<year>1993</year>) <source>The insect and spider collections of the world. 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HfdIndexSelect -h $EXPLOR_AREA/Data/Pmc/Corpus/RBID.i -Sk "pubmed:25408617" \
| HfdSelect -Kh $EXPLOR_AREA/Data/Pmc/Corpus/biblio.hfd \
| NlmPubMed2Wicri -a CheneBelgiqueV1
| This area was generated with Dilib version V0.6.27. |  |