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<mods:title id="43288B559985BB7AF8CFA10F7E8DA97F">A review of the genus Dryocoetiops Schedl, with new species, new synonymy and a key to species (Coleoptera: Curculionidae: Scolytinae)</mods:title>
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<subSubSection id="C3496577FFF4FF9F5988001EFDFD7318" box="[151,606,1593,1619]" pageId="1" pageNumber="237" type="nomenclature">
<paragraph id="8BEC36FCFFF4FF9F5988001EFDFD7318" blockId="1.[151,606,1593,1619]" box="[151,606,1593,1619]" pageId="1" pageNumber="237">
<heading id="D0A48190FFF4FF9F5988001EFDFD7318" bold="true" box="[151,606,1593,1619]" fontSize="11" level="1" pageId="1" pageNumber="237" reason="1">
<treatmentCitationGroup id="AB4311D2FFF4FF9F5988001EFDFD7318" box="[151,606,1593,1619]" pageId="1" pageNumber="237">
<emphasis id="B927EAEEFFF4FF9F5988001EFDFD7318" bold="true" box="[151,606,1593,1619]" pageId="1" pageNumber="237">
Genus
<taxonomicName id="4C534D7FFFF4FF9F59F1001EFDFD7318" ID-CoL="8HHSH" authority="Schedl, 1957: 13" authorityName="Schedl" authorityPageNumber="13" authorityYear="1957" box="[238,606,1593,1619]" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF4FF9F59F1001EFE2F7318" bold="true" box="[238,396,1593,1619]" italics="true" pageId="1" pageNumber="237">Dryocoetiops</emphasis>
<treatmentCitation id="0AF210EDFFF4FF9F588B001EFDFD7318" author="Schedl, K. E." box="[404,606,1593,1619]" page="13" pageId="1" pageNumber="237" year="1957">
<bibRefCitation id="EFC24B0DFFF4FF9F588B001EFDFD7318" author="Schedl, K. E." box="[404,606,1593,1619]" pageId="1" pageNumber="237" pagination="1 - 163" refId="ref12403" refString="Schedl, K. E. (1957) Scolytoidea nouveaux du Congo Belge. II. Mission R. Mayne-K. E. Schedl 1952. Annales du Musee Royal du Congo Belge, Tervuren, serie 8 o, Sciences Zoologiques, 56, 1 - 163." type="journal article" year="1957">Schedl, 1957: 13</bibRefCitation>
</treatmentCitation>
</taxonomicName>
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</heading>
</paragraph>
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<subSubSection id="C3496577FFF4FF9F59880059FD0C73DE" box="[151,687,1661,1686]" pageId="1" pageNumber="237" type="materials_examined">
<paragraph id="8BEC36FCFFF4FF9F59880059FD0C73DE" blockId="1.[151,687,1661,1686]" box="[151,687,1661,1686]" pageId="1" pageNumber="237">
<typeStatus id="54E8885EFFF4FF9F59880059FF6A73DD" box="[151,201,1662,1686]" pageId="1" pageNumber="237">Type</typeStatus>
species:
<taxonomicName id="4C534D7FFFF4FF9F58390059FD0C73DE" authority="Strohmeyer 1911: 22" authorityName="Strohmeyer" authorityPageNumber="22" authorityYear="1911" box="[294,687,1661,1686]" class="Insecta" family="Curculionidae" genus="Ozopemon" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="237" phylum="Arthropoda" rank="species" species="laevis">
<emphasis id="B927EAEEFFF4FF9F58390059FE7773DE" box="[294,468,1661,1685]" italics="true" pageId="1" pageNumber="237">Ozopemon laevis</emphasis>
<bibRefCitation id="EFC24B0DFFF4FF9F58C5005AFD0C73DE" author="Strohmeyer, H." box="[474,687,1661,1686]" pageId="1" pageNumber="237" pagination="17 - 27" refId="ref12742" refString="Strohmeyer, H. (1911) Borkenkafer der Philippinen. Philippine Journal of Science, D. General Biology, 6, 17 - 27." type="journal article" year="1911">Strohmeyer 1911: 22</bibRefCitation>
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="C3496577FFF4FF9C598800E4FD3D742C" lastPageId="2" lastPageNumber="238" pageId="1" pageNumber="237" type="diagnosis">
<paragraph id="8BEC36FCFFF4FF9C598800E4FD3D742C" blockId="1.[151,1436,1731,2010]" lastBlockId="2.[151,1437,153,2015]" lastPageId="2" lastPageNumber="238" pageId="1" pageNumber="237">
<emphasis id="B927EAEEFFF4FF9F598800E4FEB17395" bold="true" box="[151,274,1731,1758]" pageId="1" pageNumber="237">Diagnosis.</emphasis>
The genus
<taxonomicName id="4C534D7FFFF4FF9F588C00E3FDDF7395" authority="Schedl" authorityName="Schedl" authorityYear="1957" box="[403,636,1732,1758]" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF4FF9F588C00E3FD897396" box="[403,554,1732,1757]" italics="true" pageId="1" pageNumber="237">Dryocoetiops</emphasis>
Schedl
</taxonomicName>
belongs in the tribe
<taxonomicName id="4C534D7FFFF4FF9F5A7D00E3FC4B7395" box="[866,1000,1732,1758]" class="Insecta" family="Curculionidae" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="237" phylum="Arthropoda" rank="tribe" tribe="Dryocoetini">Dryocoetini</taxonomicName>
(
<bibRefCitation id="EFC24B0DFFF4FF9F5AE900E3FBDF7396" author="Wood, S. L." box="[1014,1148,1731,1758]" pageId="1" pageNumber="237" pagination="1 - 126" refId="ref12814" refString="Wood, S. L. (1986) A reclassification of the genera of Scolytidae (Coleoptera). Great Basin Naturalist Memoirs, 10, 1 - 126." type="journal article" year="1986">Wood 1986</bibRefCitation>
; Alonso-Zarazaga &amp; Lyal 2009). Generic boundaries in the tribe are not always clear (
<bibRefCitation id="EFC24B0DFFF4FF9F5A2600CFFC627249" author="Wood, S. L." box="[825,961,1767,1794]" pageId="1" pageNumber="237" pagination="1 - 126" refId="ref12814" refString="Wood, S. L. (1986) A reclassification of the genera of Scolytidae (Coleoptera). Great Basin Naturalist Memoirs, 10, 1 - 126." type="journal article" year="1986">Wood 1986</bibRefCitation>
), and
<taxonomicName id="4C534D7FFFF4FF9F5D1B00CFFB38724A" authorityName="Schedl" authorityYear="1957" box="[1028,1179,1768,1793]" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF4FF9F5D1B00CFFB38724A" box="[1028,1179,1768,1793]" italics="true" pageId="1" pageNumber="237">Dryocoetiops</emphasis>
</taxonomicName>
is no exception. Many members of the tribe appear to have undergone mosaic evolution, and genera can be distinguished only by combinations of characters.
<taxonomicName id="4C534D7FFFF4FF9F58680117FDAD7202" authorityName="Schedl" authorityYear="1957" box="[375,526,1840,1865]" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF4FF9F58680117FDAD7202" box="[375,526,1840,1865]" italics="true" pageId="1" pageNumber="237">Dryocoetiops</emphasis>
</taxonomicName>
is distinguished morphologically from other genera included in the tribe by the following combination of characters: 1) antennal funicle four-segmented (excluding the pedicel), club flattened or weakly obliquely truncate, sutures strongly displaced apically on posterior face, anterior face with basal segment confined to basal fifth or quarter, its apical margin convex to straight;
<emphasis id="B927EAEEFFF4FF9F5A8101BBFC6872FD" bold="true" box="[926,971,1948,1974]" pageId="1" pageNumber="237">and</emphasis>
2) eye emarginate, in a few species rather large, coarsely facetted and extending onto the frons;
<emphasis id="B927EAEEFFF4FF9F5BF001E7FCBF7291" bold="true" box="[751,796,1984,2010]" pageId="1" pageNumber="237">and</emphasis>
3) female frons without a brush of setae, sparsely setose, not aciculate;
<emphasis id="B927EAEEFFF7FF9C582C06BEFEC375F8" bold="true" box="[307,352,153,179]" pageId="2" pageNumber="238">and</emphasis>
4) pronotum about as long as wide or wider than long, strongly declivous anteriorly, usually with rather coarse, rasp-like asperities, unequal in size and larger antero-laterally, the summit obtuse, humped, at or a little behind the middle, a weak to distinct transverse impression behind it, disc flat in lateral view, usually granulate-punctate;
<emphasis id="B927EAEEFFF7FF9C58260722FEC57454" bold="true" box="[313,358,261,287]" pageId="2" pageNumber="238">and</emphasis>
5) elytra with distinct striae and interstriae;
<emphasis id="B927EAEEFFF7FF9C5A730722FC3A7454" bold="true" box="[876,921,261,287]" pageId="2" pageNumber="238">and</emphasis>
6) elytral declivity steep, convex, interstriae granulate except in one species;
<emphasis id="B927EAEEFFF7FF9C58E1070EFD887408" bold="true" box="[510,555,297,323]" pageId="2" pageNumber="238">and</emphasis>
7) protibiae with four socketed teeth (rarely five) in apical half, without further smaller teeth on external margin more basally.
</paragraph>
</subSubSection>
<subSubSection id="C3496577FFF7FF9C59D80757FB6B76C8" pageId="2" pageNumber="238" type="description">
<paragraph id="8BEC36FCFFF7FF9C59D80757FD2F7614" blockId="2.[151,1437,153,2015]" pageId="2" pageNumber="238">
<emphasis id="B927EAEEFFF7FF9C59D80757FEFA74C0" bold="true" box="[199,345,368,395]" pageId="2" pageNumber="238">Description.</emphasis>
Female. Antenna with 4-segmented funicle (excluding the pedicel), club flattened or weakly obliquely truncate, without septum, sutures strongly displaced apically on posterior face, basal segment not extending beyond basal quarter on anterior face, its apical margin convex, sinuate or approximately straight. Eyes distinctly but not deeply emarginate anteriorly, either of normal size, finely faceted, or in a few species enlarged, more coarsely faceted, and extending onto frons. Frons punctate or granulate-punctate, usually with a median raised line, sometimes with a median tubercle. Pronotum strongly declivous anteriorly, rising to an obtuse hump-like elevation, approximately in the middle, with a weak to distinct transverse impression behind it. Anterior slope of pronotum with rasp-like asperities, often extending to anterior margin, usually increasing in size antero-laterally, the disc flat in lateral view, granulate-punctate. Elytra convex, striae and interstriae distinct, strial punctures with short, fine, recumbent setae, interstrial punctures with stouter, longer, erect setae, occasionally flattened on declivity. Elytral declivity steep, convex, striae 1, and less often striae 23 impressed, interstriae usually granulate, 13 raised and costate in two species. Procoxae usually narrowly separated, and prosternal process pointed, but contiguous and prosternal process blunt in a few species. Protibiae with four socketed teeth (rarely five), meso- and metatibiae usually with six teeth. Body length 1.94.0 mm.
</paragraph>
<paragraph id="8BEC36FCFFF7FF9C59D8054EFB6B76C8" blockId="2.[151,1437,153,2015]" box="[199,1224,873,899]" pageId="2" pageNumber="238">
Male. Unknown. Presumed to be dwarfed and flightless as in
<taxonomicName id="4C534D7FFFF7FF9C5A65054EFB6776C8" authority="Eichhoff, 1878 a" authorityName="Eichhoff" authorityYear="1878" box="[890,1220,873,899]" class="Insecta" family="Curculionidae" genus="Coccotrypes" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C5A65054EFBA576C8" box="[890,1030,873,899]" italics="true" pageId="2" pageNumber="238">Coccotrypes</emphasis>
<bibRefCitation id="EFC24B0DFFF7FF9C5D12054EFB6776C8" author="Eichhoff, W. J." box="[1037,1220,873,899]" pageId="2" pageNumber="238" pagination="383 - 392" refId="ref11503" refString="Eichhoff, W. J. (1878 a) Neue oder noch unbeschreibene Tomicinen. Entomologische Zeitung, Stettin, 39, 383 - 392." type="journal article" year="1878">Eichhoff, 1878a</bibRefCitation>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C3496577FFF7FF9C59D805ABFAE37238" pageId="2" pageNumber="238" type="distribution">
<paragraph id="8BEC36FCFFF7FF9C59D805ABFF6476A4" blockId="2.[151,1437,153,2015]" pageId="2" pageNumber="238">
<emphasis id="B927EAEEFFF7FF9C59D805ABFEC276EC" bold="true" box="[199,353,908,935]" pageId="2" pageNumber="238">Distribution.</emphasis>
The genus is primarily palaeotropical in its distribution from
<collectingCountry id="F344766CFFF7FF9C5D0605AAFBF076EC" box="[1049,1107,909,935]" name="India" pageId="2" pageNumber="238">India</collectingCountry>
through South-East Asia and
<collectingCountry id="F344766CFFF7FF9C59880596FEA77680" box="[151,260,945,971]" name="Indonesia" pageId="2" pageNumber="238">Indonesia</collectingCountry>
to New
<collectingCountry id="F344766CFFF7FF9C587A0596FE157680" box="[357,438,945,971]" name="Guinea-Bissau" pageId="2" pageNumber="238">Guinea</collectingCountry>
and northern
<collectingCountry id="F344766CFFF7FF9C5B4C0596FD1E7680" box="[595,701,945,971]" name="Australia" pageId="2" pageNumber="238">Australia</collectingCountry>
. Two species (
<emphasis id="B927EAEEFFF7FF9C5A7D0595FBDE7680" box="[866,1149,945,971]" italics="true" pageId="2" pageNumber="238">
<taxonomicName id="4C534D7FFFF7FF9C5A7D0595FC567680" authorityName="Eichhoff" authorityYear="1875" box="[866,1013,945,971]" class="Insecta" family="Curculionidae" genus="Dryocoetes" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="species" species="apatoides">D. apatoides</taxonomicName>
,
<taxonomicName id="4C534D7FFFF7FF9C5D1E0595FBDE7680" baseAuthorityName="Blandford" baseAuthorityYear="1894" box="[1025,1149,946,971]" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="species" species="moestus">D. moestus</taxonomicName>
</emphasis>
) occur as far north as
<collectingCountry id="F344766CFFF7FF9C5C630596FF6076A4" name="Japan" pageId="2" pageNumber="238">Japan</collectingCountry>
.
</paragraph>
<paragraph id="8BEC36FCFFF7FF9C59D805DFFAFE7340" blockId="2.[151,1437,153,2015]" pageId="2" pageNumber="238">
<emphasis id="B927EAEEFFF7FF9C59D805DFFED37158" bold="true" box="[199,368,1016,1043]" pageId="2" pageNumber="238">Relationships.</emphasis>
Early phylogenetic studies (
<bibRefCitation id="EFC24B0DFFF7FF9C5BA605DEFC2B7158" author="Jordal, B. H. &amp; Normark, B. B. &amp; Farrell, B. D." box="[697,904,1017,1043]" pageId="2" pageNumber="238" pagination="483 - 499" refId="ref11778" refString="Jordal, B. H., Normark, B. B. &amp; Farrell, B. D. (2000) Evolutionary radiation of an inbreeding haplodiploid beetle lineage (Curculionidae, Scolytinae). Biological Journal of the Linnaean Society, 71, 483 - 499. https: // doi. org / 10.1111 / j. 1095 - 8312.2000. tb 01270. x" type="journal article" year="2000">
Jordal
<emphasis id="B927EAEEFFF7FF9C5A1605DDFCE77158" box="[777,836,1017,1043]" italics="true" pageId="2" pageNumber="238">et al.</emphasis>
2000
</bibRefCitation>
,
<bibRefCitation id="EFC24B0DFFF7FF9C5A8905DEFC737158" author="Jordal, B. H. &amp; Normark, B. B. &amp; Farrell, B. D. &amp; Kirkendall, L. R." box="[918,976,1017,1043]" pageId="2" pageNumber="238" pagination="171 - 188" refId="ref11847" refString="Jordal, B. H., Normark, B. B., Farrell, B. D. &amp; Kirkendall, L. R. (2002) Extraordinary haplotype diversity in haplodiploid inbreeders: phylogenetics and evolution of the bark beetle genus Coccotrypes. Molecular Phylogenetics and Evolution, 23, 171 - 188. https: // doi. org / 10.1016 / S 1055 - 7903 (02) 00013 - 1" type="journal article" year="2002">2002</bibRefCitation>
) suggested that
<taxonomicName id="4C534D7FFFF7FF9C5D9205DDFA877158" authorityName="Schedl" authorityYear="1957" box="[1165,1316,1018,1043]" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C5D9205DDFA877158" box="[1165,1316,1018,1043]" italics="true" pageId="2" pageNumber="238">Dryocoetiops</emphasis>
</taxonomicName>
should be included within the genus
<taxonomicName id="4C534D7FFFF7FF9C58D9023AFDF1717C" authorityName="Eichhoff" authorityYear="1878" box="[454,594,1053,1079]" class="Insecta" family="Curculionidae" genus="Coccotrypes" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C58D9023AFDF1717C" box="[454,594,1053,1079]" italics="true" pageId="2" pageNumber="238">Coccotrypes</emphasis>
</taxonomicName>
. The studies were based on
<taxonomicName id="4C534D7FFFF7FF9C5A860239FBAD717C" authorityName="Eggers" authorityYear="1923" box="[921,1038,1053,1079]" class="Insecta" family="Curculionidae" genus="Dryocoetes" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="species" species="coffeae">
<emphasis id="B927EAEEFFF7FF9C5A860239FBAD717C" box="[921,1038,1053,1079]" italics="true" pageId="2" pageNumber="238">D. coffeae</emphasis>
</taxonomicName>
and
<taxonomicName id="4C534D7FFFF7FF9C5D570239FB54717C" authorityName="Schedl" authorityYear="1942" box="[1096,1271,1053,1079]" class="Insecta" family="Curculionidae" genus="Dryocoetes" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="species" species="eugeniae">
<emphasis id="B927EAEEFFF7FF9C5D570239FBC7717C" box="[1096,1124,1054,1079]" italics="true" pageId="2" pageNumber="238">D.</emphasis>
cf.
<emphasis id="B927EAEEFFF7FF9C5D8D0239FB54717C" box="[1170,1271,1054,1079]" italics="true" pageId="2" pageNumber="238">eugeniae</emphasis>
</taxonomicName>
[
<emphasis id="B927EAEEFFF7FF9C5C160239FA8B717C" box="[1289,1320,1054,1079]" italics="true" pageId="2" pageNumber="238">sic</emphasis>
]. Both
<taxonomicName id="4C534D7FFFF7FF9C5C9E0239FF4B7110" authorityName="Eggers" authorityYear="1923" class="Insecta" family="Curculionidae" genus="Dryocoetes" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="species" species="coffeae">
<emphasis id="B927EAEEFFF7FF9C5C9E0239FF4B7110" italics="true" pageId="2" pageNumber="238">D. coffeae</emphasis>
</taxonomicName>
and
<taxonomicName id="4C534D7FFFF7FF9C58010265FE067110" authorityName="Schedl" authorityYear="1942" box="[286,421,1090,1115]" class="Insecta" family="Curculionidae" genus="Dryocoetes" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="species" species="eugeniae">
<emphasis id="B927EAEEFFF7FF9C58010265FE067110" box="[286,421,1090,1115]" italics="true" pageId="2" pageNumber="238">D. eugeniae</emphasis>
</taxonomicName>
are now considered to be synonyms of
<taxonomicName id="4C534D7FFFF7FF9C5A7D0265FC7D7110" baseAuthorityName="Blandford" baseAuthorityYear="1894" box="[866,990,1090,1115]" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="species" species="moestus">
<emphasis id="B927EAEEFFF7FF9C5A7D0265FC7D7110" box="[866,990,1090,1115]" italics="true" pageId="2" pageNumber="238">D. moestus</emphasis>
</taxonomicName>
(see below). Later phylogenetic studies (
<bibRefCitation id="EFC24B0DFFF7FF9C59800242FE097134" author="Jordal, B. H. &amp; Cognato, A. I." box="[159,426,1125,1151]" pageId="2" pageNumber="238" pagination="133" refId="ref11719" refString="Jordal, B. H. &amp; Cognato, A. I. (2012) Molecular phylogeny of bark and ambrosia beetles reveals multiple origins of fungus farming during periods of global warming. BMC Evolutionary Biology, 12, 133. https: // doi. org / 10.1186 / 1471 - 2148 - 12 - 133" type="journal article" year="2012">Jordal &amp; Cognato 2012</bibRefCitation>
;
<bibRefCitation id="EFC24B0DFFF7FF9C58AA0242FDD77134" author="Gohli, J. &amp; Kirkendall, L. R. &amp; Smith, S. M. &amp; Cognato, A. I. &amp; Hulcr, J. &amp; Jordal, B. H." box="[437,628,1125,1151]" pageId="2" pageNumber="238" pagination="1258 - 1272" refId="ref11589" refString="Gohli, J., Kirkendall, L. R., Smith, S. M., Cognato, A. I., Hulcr, J. &amp; Jordal, B. H. (2017) Biological factors contributing to bark and ambrosia beetle species diversification. Evolution, 71, 1258 - 1272. https: // doi. org / 10.1111 / evo. 13219" type="journal article" year="2017">
Gohli
<emphasis id="B927EAEEFFF7FF9C58E30241FD977134" box="[508,564,1125,1151]" italics="true" pageId="2" pageNumber="238">et al.</emphasis>
2017
</bibRefCitation>
) suggest that
<taxonomicName id="4C534D7FFFF7FF9C5A110241FC067134" authorityName="Schedl" authorityYear="1957" box="[782,933,1126,1151]" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C5A110241FC067134" box="[782,933,1126,1151]" italics="true" pageId="2" pageNumber="238">Dryocoetiops</emphasis>
</taxonomicName>
should rather be considered as a sister genus to
<taxonomicName id="4C534D7FFFF7FF9C59AB02AEFEE371E8" authorityName="Eichhoff" authorityYear="1878" box="[180,320,1161,1187]" class="Insecta" family="Curculionidae" genus="Coccotrypes" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C59AB02AEFEE371E8" box="[180,320,1161,1187]" italics="true" pageId="2" pageNumber="238">Coccotrypes</emphasis>
</taxonomicName>
. It appears to have the same haplo-diploid breeding system as
<taxonomicName id="4C534D7FFFF7FF9C5D1E02AEFACE71E8" authority="(Jordal et al. 2002)" baseAuthorityName="Jordal et al." baseAuthorityYear="2002" box="[1025,1389,1161,1187]" class="Insecta" family="Curculionidae" genus="Coccotrypes" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C5D1E02AEFB2E71E8" box="[1025,1165,1161,1187]" italics="true" pageId="2" pageNumber="238">Coccotrypes</emphasis>
(
<bibRefCitation id="EFC24B0DFFF7FF9C5D8302AEFAC671E8" author="Jordal, B. H. &amp; Normark, B. B. &amp; Farrell, B. D. &amp; Kirkendall, L. R." box="[1180,1381,1161,1187]" pageId="2" pageNumber="238" pagination="171 - 188" refId="ref11847" refString="Jordal, B. H., Normark, B. B., Farrell, B. D. &amp; Kirkendall, L. R. (2002) Extraordinary haplotype diversity in haplodiploid inbreeders: phylogenetics and evolution of the bark beetle genus Coccotrypes. Molecular Phylogenetics and Evolution, 23, 171 - 188. https: // doi. org / 10.1016 / S 1055 - 7903 (02) 00013 - 1" type="journal article" year="2002">
Jordal
<emphasis id="B927EAEEFFF7FF9C5DF602ADFA8171E8" box="[1257,1314,1161,1187]" italics="true" pageId="2" pageNumber="238">et al.</emphasis>
2002
</bibRefCitation>
)
</taxonomicName>
, but is morphologically distinct. Species of
<taxonomicName id="4C534D7FFFF7FF9C5B550289FD42718C" authorityName="Schedl" authorityYear="1957" box="[586,737,1198,1223]" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C5B550289FD42718C" box="[586,737,1198,1223]" italics="true" pageId="2" pageNumber="238">Dryocoetiops</emphasis>
</taxonomicName>
can be distinguished from
<taxonomicName id="4C534D7FFFF7FF9C5D0F028AFB3F718C" box="[1040,1180,1197,1223]" class="Insecta" family="Curculionidae" genus="Coccotrypes" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C5D0F028AFB3F718C" box="[1040,1180,1197,1223]" italics="true" pageId="2" pageNumber="238">Coccotrypes</emphasis>
</taxonomicName>
most easily by characters of the pronotum. In
<taxonomicName id="4C534D7FFFF7FF9C58B302F5FDE071A0" authorityName="Schedl" authorityYear="1957" box="[428,579,1234,1259]" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C58B302F5FDE071A0" box="[428,579,1234,1259]" italics="true" pageId="2" pageNumber="238">Dryocoetiops</emphasis>
</taxonomicName>
, 1) pronotum dome-shaped with a distinct summit near the middle, as wide as or wider than long, steeply sloping anteriorly, asperities often increasing in size antero-laterally, anterior margin broadly rounded, without a distinct row of asperities on the margin, slightly to distinctly impressed behind summit, disc flat in lateral view;
<emphasis id="B927EAEEFFF7FF9C58BC031AFE73701C" bold="true" box="[419,464,1341,1367]" pageId="2" pageNumber="238">and</emphasis>
2) frons without aciculation. In
<taxonomicName id="4C534D7FFFF7FF9C5A2A031AFC62701C" box="[821,961,1341,1367]" class="Insecta" family="Curculionidae" genus="Coccotrypes" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C5A2A031AFC62701C" box="[821,961,1341,1367]" italics="true" pageId="2" pageNumber="238">Coccotrypes</emphasis>
</taxonomicName>
, the pronotum is usually longer than wide, and if there is a distinct summit, it is often behind the middle. The anterior slope is less steep, and the asperities usually smaller, and not increasing in size antero-laterally. In some species, there is a row of larger asperities on the anterior margin only. The species of
<taxonomicName id="4C534D7FFFF7FF9C5B28038DFD6D7088" authorityName="Schedl" authorityYear="1957" box="[567,718,1450,1475]" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C5B28038DFD6D7088" box="[567,718,1450,1475]" italics="true" pageId="2" pageNumber="238">Dryocoetiops</emphasis>
</taxonomicName>
are myelophagous (Kirkendall
<emphasis id="B927EAEEFFF7FF9C5D2B038DFBCD7088" box="[1076,1134,1449,1475]" italics="true" pageId="2" pageNumber="238">et al.</emphasis>
2015) breeding in the pith of small branches and twigs; those of
<taxonomicName id="4C534D7FFFF7FF9C5B2403EAFD6470AC" box="[571,711,1485,1511]" class="Insecta" family="Curculionidae" genus="Coccotrypes" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C5B2403EAFD6470AC" box="[571,711,1485,1511]" italics="true" pageId="2" pageNumber="238">Coccotrypes</emphasis>
</taxonomicName>
breed in a variety of habitats, including fruits and seeds, phloem, leafstalks, and mangrove propagules (
<bibRefCitation id="EFC24B0DFFF7FF9C5B2003D6FD787340" author="Browne, F. G." box="[575,731,1521,1547]" pageId="2" pageNumber="238" pagination="1 - 255" refId="ref11306" refString="Browne, F. G. (1961) The biology of Malayan Scolytidae and Platypodidae. Malayan Forest Records, 22, 1 - 255." type="journal article" year="1961">Browne 1961</bibRefCitation>
;
<bibRefCitation id="EFC24B0DFFF7FF9C5BF803D6FC0E7340" author="Jordal, B. H. &amp; Normark, B. B. &amp; Farrell, B. D. &amp; Kirkendall, L. R." box="[743,941,1521,1547]" pageId="2" pageNumber="238" pagination="171 - 188" refId="ref11847" refString="Jordal, B. H., Normark, B. B., Farrell, B. D. &amp; Kirkendall, L. R. (2002) Extraordinary haplotype diversity in haplodiploid inbreeders: phylogenetics and evolution of the bark beetle genus Coccotrypes. Molecular Phylogenetics and Evolution, 23, 171 - 188. https: // doi. org / 10.1016 / S 1055 - 7903 (02) 00013 - 1" type="journal article" year="2002">
Jordal
<emphasis id="B927EAEEFFF7FF9C5A2D03D5FCC87340" box="[818,875,1521,1547]" italics="true" pageId="2" pageNumber="238">et al.</emphasis>
2002
</bibRefCitation>
), but almost never in the pith of twigs.
</paragraph>
<paragraph id="8BEC36FCFFF7FF9C59D80032FAE37238" blockId="2.[151,1437,153,2015]" pageId="2" pageNumber="238">
Besides
<emphasis id="B927EAEEFFF7FF9C58390032FE1B7364" box="[294,440,1557,1583]" italics="true" pageId="2" pageNumber="238">
<taxonomicName id="4C534D7FFFF7FF9C58390032FE167364" box="[294,437,1557,1583]" class="Insecta" family="Curculionidae" genus="Coccotrypes" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">Coccotrypes</taxonomicName>
,
</emphasis>
the genus is most similar morphologically to
<taxonomicName id="4C534D7FFFF7FF9C5AA30032FB5E7364" authority="(Schedl 1957)" baseAuthorityName="Schedl" baseAuthorityYear="1957" box="[956,1277,1556,1583]" class="Insecta" family="Curculionidae" genus="Taphrorychus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C5AA30032FBF67364" box="[956,1109,1557,1583]" italics="true" pageId="2" pageNumber="238">Taphrorychus</emphasis>
(
<bibRefCitation id="EFC24B0DFFF7FF9C5D7A0032FB557364" author="Schedl, K. E." box="[1125,1270,1556,1583]" pageId="2" pageNumber="238" pagination="1 - 163" refId="ref12403" refString="Schedl, K. E. (1957) Scolytoidea nouveaux du Congo Belge. II. Mission R. Mayne-K. E. Schedl 1952. Annales du Musee Royal du Congo Belge, Tervuren, serie 8 o, Sciences Zoologiques, 56, 1 - 163." type="journal article" year="1957">Schedl 1957</bibRefCitation>
)
</taxonomicName>
, although it is not very closely related (
<bibRefCitation id="EFC24B0DFFF7FF9C58B2001EFD187318" author="Jordal, B. H. &amp; Cognato, A. I." box="[429,699,1593,1619]" pageId="2" pageNumber="238" pagination="133" refId="ref11719" refString="Jordal, B. H. &amp; Cognato, A. I. (2012) Molecular phylogeny of bark and ambrosia beetles reveals multiple origins of fungus farming during periods of global warming. BMC Evolutionary Biology, 12, 133. https: // doi. org / 10.1186 / 1471 - 2148 - 12 - 133" type="journal article" year="2012">Jordal &amp; Cognato 2012</bibRefCitation>
;
<bibRefCitation id="EFC24B0DFFF7FF9C5BD9001EFC2A7318" author="Gohli, J. &amp; Kirkendall, L. R. &amp; Smith, S. M. &amp; Cognato, A. I. &amp; Hulcr, J. &amp; Jordal, B. H." box="[710,905,1593,1619]" pageId="2" pageNumber="238" pagination="1258 - 1272" refId="ref11589" refString="Gohli, J., Kirkendall, L. R., Smith, S. M., Cognato, A. I., Hulcr, J. &amp; Jordal, B. H. (2017) Biological factors contributing to bark and ambrosia beetle species diversification. Evolution, 71, 1258 - 1272. https: // doi. org / 10.1111 / evo. 13219" type="journal article" year="2017">
Gohli
<emphasis id="B927EAEEFFF7FF9C5A12001DFCE57318" box="[781,838,1593,1619]" italics="true" pageId="2" pageNumber="238">et al.</emphasis>
2017
</bibRefCitation>
), and has a very different biology and breeding system (see below). However, the two genera have been confused in the past, and some species now placed in
<taxonomicName id="4C534D7FFFF7FF9C5C460079FF5773D0" authorityName="Schedl" authorityYear="1957" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C5C460079FF5773D0" italics="true" pageId="2" pageNumber="238">Dryocoetiops</emphasis>
</taxonomicName>
have previously been placed in
<taxonomicName id="4C534D7FFFF7FF9C5B7400A6FCA773D0" baseAuthorityName="Schedl" baseAuthorityYear="1957" box="[619,772,1665,1691]" class="Insecta" family="Curculionidae" genus="Taphrorychus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C5B7400A6FCA773D0" box="[619,772,1665,1691]" italics="true" pageId="2" pageNumber="238">Taphrorychus</emphasis>
</taxonomicName>
(see below). Hence, we provide distinguishing characters here.
<taxonomicName id="4C534D7FFFF7FF9C59C80081FECD73F4" authorityName="Schedl" authorityYear="1957" box="[215,366,1702,1727]" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C59C80081FECD73F4" box="[215,366,1702,1727]" italics="true" pageId="2" pageNumber="238">Dryocoetiops</emphasis>
</taxonomicName>
can be most easily distinguished morphologically as follows: 1) basal segment of labial palp enlarged, barrel-shaped; 2) frons without a brush of setae;
<emphasis id="B927EAEEFFF7FF9C5A3C00EEFCF373A8" bold="true" box="[803,848,1737,1763]" pageId="2" pageNumber="238">and</emphasis>
3) pronotum with distinct summit and steep anterior slope;
<emphasis id="B927EAEEFFF7FF9C59FC00CAFEB3724C" bold="true" box="[227,272,1773,1799]" pageId="2" pageNumber="238">and</emphasis>
4) pronotal asperities coarse, increasing in size antero-laterally;
<emphasis id="B927EAEEFFF7FF9C5AE800CAFB87724C" bold="true" box="[1015,1060,1773,1799]" pageId="2" pageNumber="238">and</emphasis>
5) elytra with distinct striae and interstriae;
<emphasis id="B927EAEEFFF7FF9C58080136FEE77260" bold="true" box="[279,324,1809,1835]" pageId="2" pageNumber="238">and</emphasis>
6) without a circular impression on the elytral declivity;
<emphasis id="B927EAEEFFF7FF9C5AD40136FC5B7260" bold="true" box="[971,1016,1809,1835]" pageId="2" pageNumber="238">and</emphasis>
7) protibiae without additional small denticles. In
<emphasis id="B927EAEEFFF7FF9C58330112FE6F7204" box="[300,460,1845,1871]" italics="true" pageId="2" pageNumber="238">
<taxonomicName id="4C534D7FFFF7FF9C58330112FE6B7204" baseAuthorityName="Schedl" baseAuthorityYear="1957" box="[300,456,1845,1871]" class="Insecta" family="Curculionidae" genus="Taphrorychus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">Taphrorychus</taxonomicName>
,
</emphasis>
the basal segment of the labial palp is of normal size. The other characters can vary depending on the species and the sex of the individual, but the combination of all of them is never present.
</paragraph>
</subSubSection>
<subSubSection id="C3496577FFF7FF9D59D8015BFC877774" lastPageId="3" lastPageNumber="239" pageId="2" pageNumber="238" type="biology_ecology">
<paragraph id="8BEC36FCFFF7FF9D59D8015BFB4774C0" blockId="2.[151,1437,153,2015]" lastBlockId="3.[151,1437,153,575]" lastPageId="3" lastPageNumber="239" pageId="2" pageNumber="238">
<emphasis id="B927EAEEFFF7FF9C59D8015BFE8472DC" bold="true" box="[199,295,1916,1943]" pageId="2" pageNumber="238">Biology.</emphasis>
The biology of
<taxonomicName id="4C534D7FFFF7FF9C58F70159FDDC72DC" authorityName="Schedl" authorityYear="1957" box="[488,639,1918,1943]" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF7FF9C58F70159FDDC72DC" box="[488,639,1918,1943]" italics="true" pageId="2" pageNumber="238">Dryocoetiops</emphasis>
</taxonomicName>
has been investigated only in
<taxonomicName id="4C534D7FFFF7FF9C5AFA0159FBC772DC" baseAuthorityName="Blandford" baseAuthorityYear="1894" box="[997,1124,1918,1943]" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="species" species="moestus">
<emphasis id="B927EAEEFFF7FF9C5AFA0159FBC772DC" box="[997,1124,1918,1943]" italics="true" pageId="2" pageNumber="238">D. moestus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFC24B0DFFF7FF9C5D69015AFAB572DC" author="Browne, F. G." box="[1142,1302,1917,1943]" pageId="2" pageNumber="238" pagination="1 - 255" refId="ref11306" refString="Browne, F. G. (1961) The biology of Malayan Scolytidae and Platypodidae. Malayan Forest Records, 22, 1 - 255." type="journal article" year="1961">Browne 1961</bibRefCitation>
;
<bibRefCitation id="EFC24B0DFFF7FF9C5C3B015AFF7172F0" author="Kalshoven, L. G. E." pageId="2" pageNumber="238" pagination="157 - 180" refId="ref11924" refString="Kalshoven, L. G. E. (1958) Studies on the biology of Indonesian Scolytoidea. 4. Data on the habits of Scolytidae. First part. Tijdschrift voor Entomologie, 101, 157 - 180." type="journal article" year="1958">Kalshoven 1958</bibRefCitation>
, both as
<taxonomicName id="4C534D7FFFF7FF9C582D0185FDAB72F0" authorityName="Eggers" authorityYear="1923" box="[306,520,1953,1979]" class="Insecta" family="Curculionidae" genus="Dryocoetes" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="238" phylum="Arthropoda" rank="species" species="coffeae">
<emphasis id="B927EAEEFFF7FF9C582D0185FDAB72F0" box="[306,520,1953,1979]" italics="true" pageId="2" pageNumber="238">Dryocoetes coffeae</emphasis>
</taxonomicName>
), but is probably similar in other species. The species are inbreeding, polyphagous twig-borers. Only the female disperses and constructs the nest. A longitudinal tunnel is constructed in the centre of the twig, usually extending both upwards and downwards from the entrance.
<bibRefCitation id="EFC24B0DFFF6FF9D5D0506BEFB6775FF" author="Browne, F. G." box="[1050,1220,153,180]" pageId="3" pageNumber="239" pagination="1 - 255" refId="ref11306" refString="Browne, F. G. (1961) The biology of Malayan Scolytidae and Platypodidae. Malayan Forest Records, 22, 1 - 255." type="journal article" year="1961">Browne (1961)</bibRefCitation>
notes that the eggs are laid in clusters and are very variable in size. The larvae feed on the tissues surrounding the gallery, and eventually pupate in single file (
<bibRefCitation id="EFC24B0DFFF6FF9D58CB06C6FDD775B7" author="Browne, F. G." box="[468,628,225,252]" pageId="3" pageNumber="239" pagination="1 - 255" refId="ref11306" refString="Browne, F. G. (1961) The biology of Malayan Scolytidae and Platypodidae. Malayan Forest Records, 22, 1 - 255." type="journal article" year="1961">Browne 1961</bibRefCitation>
).
<bibRefCitation id="EFC24B0DFFF6FF9D5B9606C6FC9675B7" author="Browne, F. G." box="[649,821,225,252]" pageId="3" pageNumber="239" pagination="1 - 255" refId="ref11306" refString="Browne, F. G. (1961) The biology of Malayan Scolytidae and Platypodidae. Malayan Forest Records, 22, 1 - 255." type="journal article" year="1961">Browne (1961)</bibRefCitation>
also states that the species is monogamous with both parents playing a part in the care of the nest and the young brood. However, this part of his account of the biology is mistaken. As noted above, all known adults are female, and the male remains unknown. It is likely that Browne observed two females occupying the same gallery system. Dissections of females of
<taxonomicName id="4C534D7FFFF6FF9D5D4A0769FB71742C" baseAuthorityName="Blandford" baseAuthorityYear="1894" box="[1109,1234,334,359]" class="Insecta" family="Curculionidae" genus="Dryocoetiops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="239" phylum="Arthropoda" rank="species" species="moestus">
<emphasis id="B927EAEEFFF6FF9D5D4A0769FB71742C" box="[1109,1234,334,359]" italics="true" pageId="3" pageNumber="239">D. moestus</emphasis>
</taxonomicName>
caught in ethanol traps show that the spermathecae contain sperm, indicating that the species is not parthenogenetic.
</paragraph>
<paragraph id="8BEC36FCFFF6FF9D59D807B2FC877774" blockId="3.[151,1437,153,575]" pageId="3" pageNumber="239">
By contrast, the morphologically similar species of
<taxonomicName id="4C534D7FFFF6FF9D5A1607B2FC0174E4" baseAuthorityName="Schedl" baseAuthorityYear="1957" box="[777,930,405,431]" class="Insecta" family="Curculionidae" genus="Taphrorychus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="239" phylum="Arthropoda" rank="genus">
<emphasis id="B927EAEEFFF6FF9D5A1607B2FC0174E4" box="[777,930,405,431]" italics="true" pageId="3" pageNumber="239">Taphrorychus</emphasis>
</taxonomicName>
are outbreeding bark beetles, making gallery systems below the bark of angiosperm trees, particularly of the family
<taxonomicName id="4C534D7FFFF6FF9D5AA6079EFB867498" authorityName="Dumortier" authorityYear="1829" box="[953,1061,441,467]" class="Magnoliopsida" family="Fagaceae" kingdom="Plantae" order="Fagales" pageId="3" pageNumber="239" phylum="Tracheophyta" rank="family">Fagaceae</taxonomicName>
. Both male and female are of the same size, and the sex ratio at emergence is approximately 1: 1. Both sexes are capable of flight, disperse, and are involved in gallery construction. The males are harem polygynous (Kirkendall
<emphasis id="B927EAEEFFF6FF9D5D390425FBC27750" box="[1062,1121,513,539]" italics="true" pageId="3" pageNumber="239">et al.</emphasis>
2015), with more than one female associated with each male in most gallery systems.
</paragraph>
</subSubSection>
</treatment>
</document>