288 lines
25 KiB
XML
288 lines
25 KiB
XML
<document ID-DOI="http://dx.doi.org/10.3897/zookeys.416.7706" ID-GBIF-Dataset="47852319-33aa-4966-89af-68e6352e05ae" ID-PMC="PMC4109512" ID-Pensoft-Pub="1313-2970-416-113" ID-PubMed="25061349" ID-ZBK="CE0561FB5EE4498BA2C2EDF9B14F241D" ModsDocAuthor="" ModsDocDate="2014" ModsDocID="1313-2970-416-113" ModsDocOrigin="ZooKeys 416" ModsDocTitle="An unexpected clade of South American ground beetles (Coleoptera, Carabidae, Bembidion)" checkinTime="1451245749490" checkinUser="pensoft" docAuthor="Maddison, David R." docDate="2014" docId="A6C88BA1561BC8EC698F397DA5D9B7DB" docLanguage="en" docName="ZooKeys 416: 113-155" docOrigin="ZooKeys 416" docSource="http://dx.doi.org/10.3897/zookeys.416.7706" docTitle="Bembidion (Nothonepha) tetrapholeon Maddison, 2014, sp. n." docType="treatment" docUuid="90188564-6B1E-4F0E-B411-5AD99047F716" docUuidSource="ZooBank" docVersion="4" lastPageNumber="136" masterDocId="7643F6149609155E6212FF95A85FFFBF" masterDocTitle="An unexpected clade of South American ground beetles (Coleoptera, Carabidae, Bembidion)" masterLastPageNumber="155" masterPageNumber="113" pageNumber="132" updateTime="1668158654224" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>An unexpected clade of South American ground beetles (Coleoptera, Carabidae, Bembidion)</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Maddison, David R.</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>ZooKeys</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2014</mods:date>
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<mods:detail type="volume">
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<mods:number>416</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>113</mods:start>
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<mods:end>155</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:location>
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<mods:url>http://dx.doi.org/10.3897/zookeys.416.7706</mods:url>
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</mods:location>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.416.7706</mods:identifier>
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<mods:identifier type="Pensoft-Pub">1313-2970-416-113</mods:identifier>
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<mods:identifier type="ZBK">CE0561FB5EE4498BA2C2EDF9B14F241D</mods:identifier>
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<mods:identifier type="ZooBank">CE0561FB5EE4498BA2C2EDF9B14F241D</mods:identifier>
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</mods:mods>
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<treatment ID-GBIF-Taxon="152053570" LSID="urn:lsid:zoobank.org:act:90188564-6B1E-4F0E-B411-5AD99047F716" httpUri="http://treatment.plazi.org/id/A6C88BA1561BC8EC698F397DA5D9B7DB" lastPageId="23" lastPageNumber="136" pageId="19" pageNumber="132">
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<subSubSection pageId="19" pageNumber="132" type="multiple">
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<paragraph pageId="19" pageNumber="132">Taxon classification Animalia Coleoptera Carabidae</paragraph>
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</subSubSection>
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<subSubSection pageId="19" pageNumber="132" type="nomenclature">
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<paragraph pageId="19" pageNumber="132">
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<taxonomicName LSID="http://zoobank.org/90188564-6B1E-4F0E-B411-5AD99047F716" class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion (Nothonepha) tetrapholeon" order="Coleoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="species" species="tetrapholeon" subGenus="Nothonepha">Bembidion (Nothonepha) tetrapholeon</taxonomicName>
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<taxonomicNameLabel pageId="19" pageNumber="132">sp. n.</taxonomicNameLabel>
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Figs 5A, B, 9A, 15, 16, 17A, 18
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</paragraph>
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</subSubSection>
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<subSubSection pageId="19" pageNumber="132" type="holotype">
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<paragraph pageId="19" pageNumber="132">Holotype</paragraph>
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<paragraph pageId="19" pageNumber="132">
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male (IADIZA), with 3 labels: "Argentina:
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<normalizedToken originalValue="Neuquén">Neuquen</normalizedToken>
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: Arroyo /
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<normalizedToken originalValue="Queñi">Queni</normalizedToken>
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at Lago
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<normalizedToken originalValue="Queñi">Queni</normalizedToken>
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, 830m, /
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<geoCoordinate direction="south" orientation="latitude" precision="5" value="-40.1575">40.1575°S</geoCoordinate>
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,
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<geoCoordinate direction="west" orientation="longitude" precision="55" value="-71.721">71.721°W</geoCoordinate>
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, / 10-11.ii.2007. DRM 07.035. / D.R. Maddison, S.A.Roig", "David R. Maddison / DNA2356 / DNA Voucher" [printed on pale green paper], and "HOLOTYPE / Bembidion / tetrapholeon / David R. Maddison" [printed on red paper]. Genitalia in glycerine in small plastic vial beneath specimen; extracted DNA stored separately. GenBank accession numbers for DNA sequences of the holotype are KJ653049 (28S), KJ653145 (COI), KJ653112 (CAD), KJ653181 (Topo), KJ653215 (wg), and KJ653082 (ArgK).
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="20" lastPageNumber="133" pageId="19" pageNumber="132" type="paratypes">
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<paragraph pageId="19" pageNumber="132">Paratypes.</paragraph>
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<paragraph lastPageId="20" lastPageNumber="133" pageId="19" pageNumber="132">
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Total of 244, in IADIZA, MACN, MNNC, OSAC, MNHN, BMNH, EMEC, CTVR, and CMNH, from "Argentina:
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<normalizedToken originalValue="Neuquén">Neuquen</normalizedToken>
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: Arroyo /
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<normalizedToken originalValue="Queñi">Queni</normalizedToken>
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at Lago
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<normalizedToken originalValue="Queñi">Queni</normalizedToken>
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, 830m, /
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<geoCoordinate direction="south" orientation="latitude" precision="5" value="-40.1575">40.1575°S</geoCoordinate>
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,
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<geoCoordinate direction="west" orientation="longitude" precision="55" value="-71.721">71.721°W</geoCoordinate>
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, / 10-11.ii.2007. DRM 07.035. / D.R. Maddison, S.A.Roig" [135 exx.], Argentina:
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<normalizedToken originalValue="Neuquén">Neuquen</normalizedToken>
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: Rio Pichi / Traful nr Lago Traful, 810m, /
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<geoCoordinate direction="south" orientation="latitude" precision="5" value="-40.4867">40.4867°S</geoCoordinate>
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,
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<geoCoordinate direction="west" orientation="longitude" precision="5" value="-71.5958">71.5958°W</geoCoordinate>
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, / 12.ii.2007. DRM 07.039. / D.R. Maddison, S.A.Roig" [95 exx], "Argentina: Chubut: Rio / Azul at Lago Puelo, 200m /
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<geoCoordinate direction="south" orientation="latitude" precision="5" value="-42.0929">42.0929°S</geoCoordinate>
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,
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<geoCoordinate direction="west" orientation="longitude" precision="5" value="-71.6244">71.6244°W</geoCoordinate>
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/ 13.ii.2007. DRM 07.044. / D.R. Maddison" [12 exx],
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<pageBreakToken pageId="20" pageNumber="133" start="start">"</pageBreakToken>
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Argentina: Chubut: Rio / Azul at Lago Puelo, 200m /
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<geoCoordinate direction="south" orientation="latitude" precision="5" value="-42.0929">42.0929°S</geoCoordinate>
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,
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<geoCoordinate direction="west" orientation="longitude" precision="5" value="-71.6244">71.6244°W</geoCoordinate>
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, / 13.ii.2007. DRM 07.045. / S.A.Roig, D.R. Maddison" [1 exx].
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</paragraph>
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</subSubSection>
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<subSubSection pageId="20" pageNumber="133" type="additional material examined">
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<paragraph pageId="20" pageNumber="133">Additional material examined.</paragraph>
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<paragraph pageId="20" pageNumber="133">
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CHILE: Reg. X,
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<normalizedToken originalValue="Chiloé">Chiloe</normalizedToken>
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: Rio Puntra at rt 5, 55m,
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<geoCoordinate direction="south" orientation="latitude" precision="5" value="-42.1661">42.1661°S</geoCoordinate>
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,
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<geoCoordinate direction="west" orientation="longitude" precision="5" value="-73.7256">73.7256°W</geoCoordinate>
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, 19.i.2006. DRM 06.075. D.R. Maddison [5 exx, OSAC, MNNC]. CHILE: Region X, Rio Pullinque at Puente Huanehue, 8 km NE Panguipulli. 16 Jan 2002,
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<geoCoordinate direction="south" orientation="latitude" precision="5" value="-39.6162">39.6162°S</geoCoordinate>
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,
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<geoCoordinate direction="west" orientation="longitude" precision="5" value="-72.2286">72.2286°W</geoCoordinate>
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, 1590 ft. W. D. Shepard [2 exx, OSAC].
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</paragraph>
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</subSubSection>
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<subSubSection pageId="20" pageNumber="133" type="additional identified material">
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<paragraph pageId="20" pageNumber="133">Additional identified material.</paragraph>
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<paragraph pageId="20" pageNumber="133">
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The following specimens have been examined by Luca Toledano and confirmed to belong to this species (based upon photographs we have shared). CHILE: Reg. X, Los Lagos, P.N. Vicente
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<normalizedToken originalValue="Péres">Peres</normalizedToken>
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Rosales,
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<normalizedToken originalValue="Petrohué">Petrohue</normalizedToken>
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, Lago Todos los Santos, 190m, mouth Rio El Caulle,
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<geoCoordinate direction="south" orientation="latitude" precision="5" value="-41.0924">41.0924°S</geoCoordinate>
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,
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<geoCoordinate direction="west" orientation="longitude" precision="5" value="-72.395">72.3950°W</geoCoordinate>
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. 5.i.2014. L. Toledano, R. Olivieri, J.P. Morales. [1 ex, CTVR]; CHILE: Region XI, Parque Nat. Rio Simpson, H. Franz [4 exx, NHMW]; CHILE: Reg. X, I.
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<normalizedToken originalValue="Chiloé">Chiloe</normalizedToken>
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, R. Punta, 31.i.1986. M. Spies. [1 ex, USNM].
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</paragraph>
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</subSubSection>
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<subSubSection pageId="20" pageNumber="133" type="type locality">
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<paragraph pageId="20" pageNumber="133">Type locality.</paragraph>
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<paragraph pageId="20" pageNumber="133">
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Argentina:
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<normalizedToken originalValue="Neuquén">Neuquen</normalizedToken>
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: Arroyo
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<normalizedToken originalValue="Queñi">Queni</normalizedToken>
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at Lago
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<normalizedToken originalValue="Queñi">Queni</normalizedToken>
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, 830m,
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<geoCoordinate direction="south" orientation="latitude" precision="5" value="-40.1575">40.1575°S</geoCoordinate>
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,
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<geoCoordinate direction="west" orientation="longitude" precision="5" value="-71.721">71.7210°W</geoCoordinate>
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. The habitat at the type locality is a cobble, gravel, and coarse sand river shore (Fig. 6D); the river is cold and has crystal-clear water. In the same habitat members of the genus
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidarenas" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidarenas" order="Coleoptera" pageId="20" pageNumber="133" phylum="Arthropoda" rank="genus">Bembidarenas</taxonomicName>
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Erwin are abundant, as is
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion (Antiperyphanes) rufoplagiatum" order="Coleoptera" pageId="20" pageNumber="133" phylum="Arthropoda" rank="species" species="rufoplagiatum" subGenus="Antiperyphanes">Bembidion (Antiperyphanes) rufoplagiatum</taxonomicName>
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.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="20" pageNumber="133" type="derivation of specific epithet">
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<paragraph pageId="20" pageNumber="133">Derivation of specific epithet.</paragraph>
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<paragraph pageId="20" pageNumber="133">
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From the Greek
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<normalizedToken originalValue="“tetra”">"tetra"</normalizedToken>
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, meaning
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<normalizedToken originalValue="“four”">"four"</normalizedToken>
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, and
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<normalizedToken originalValue="“pholeon”">"pholeon"</normalizedToken>
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, meaning
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<normalizedToken originalValue="“pit”">"pit"</normalizedToken>
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, referring to the four prominent pits visible on the underside of adults.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="21" pageNumber="134" type="diagnosis">
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<paragraph pageId="21" pageNumber="134">
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<pageBreakToken pageId="21" pageNumber="134" start="start">Diagnosis</pageBreakToken>
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.
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</paragraph>
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<paragraph pageId="21" pageNumber="134">
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A large, sleek, shiny
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="21" pageNumber="134" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
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, with an unusual body form (Figs 5A, B) of narrow forebody and large elytra. With its shape, color, and smoothness it is one of the most distinctive
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="21" pageNumber="134" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
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species in South America, and no other known species is likely to be confused with it; it is more reminiscent of some species in New Zealand, e.g.,
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion (Zeplataphus) dehiscens" order="Coleoptera" pageId="21" pageNumber="134" phylum="Arthropoda" rank="species" species="dehiscens" subGenus="Zeplataphus">Bembidion (Zeplataphus) dehiscens</taxonomicName>
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Broun (
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<bibRefCitation author="Lindroth, CH" journalOrPublisher="New Zealand Journal of Zoology" pageId="26" pageNumber="139" pagination="161 - 198" title="Genus Bembidion Latreille (Coleoptera: Carabidae) in New Zealand: a revision." url="10.1080/03014223.1976.9517912" volume="3" year="1976">Lindroth 1976</bibRefCitation>
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).
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</paragraph>
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<paragraph pageId="21" pageNumber="134">Length (4.7-5.7 mm, with most specimens above 5.0 mm). Color piceous (Fig. 5A), with legs and antennae in some specimens slightly paler, and with a few specimens having a large orange spot just in front of the elytral apices (Fig. 5B).</paragraph>
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<paragraph pageId="21" pageNumber="134">Head with shallow and parallel frontal furrows.</paragraph>
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<paragraph pageId="21" pageNumber="134">Pronotum narrow, cordate, with hind angles flaring outward (Fig. 15). Very smooth, without punctures, and with a linear basolateral foveae; without distinct carina at hind angle. Lateral bead of pronotum not complete, not reaching front angle of prothorax and only in some specimens reaching the hind angle. One midlateral and one basolateral seta on each side.</paragraph>
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<caption pageId="21" pageNumber="134">
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<paragraph pageId="21" pageNumber="134">
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Figure 15. Pronotum of
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion tetrapholeon" order="Coleoptera" pageId="21" pageNumber="134" phylum="Arthropoda" rank="species" species="tetrapholeon">Bembidion tetrapholeon</taxonomicName>
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, DRM voucher V100781. Scale bar 0.1 mm.
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</paragraph>
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</caption>
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<paragraph pageId="21" pageNumber="134">
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Each elytron with two discal setae (ed3 and ed5); ed3 in third stria. Elytral striae with prominent punctures in their basal half, but with striae 2-7 absent in about the hind 40% or more of the elytra, such that the posterior discal seta, ed5, is in a region without striae. Striae 7 absent in many specimens. In many specimens the striae are effaced anteriorly, especially striae 2 and 3. Lateral bead of elytron effaced anteriorly, not extended onto shoulder (Fig. 16A), similar to that of
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion (Nothonepha) lonae" order="Coleoptera" pageId="21" pageNumber="134" phylum="Arthropoda" rank="species" species="lonae" subGenus="Nothonepha">Bembidion (Nothonepha) lonae</taxonomicName>
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(Fig. 16B), but unlike most other
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="21" pageNumber="134" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
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(e.g.,
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion (Nothonepha) germainianum" order="Coleoptera" pageId="21" pageNumber="134" phylum="Arthropoda" rank="species" species="germainianum" subGenus="Nothonepha">Bembidion (Nothonepha) germainianum</taxonomicName>
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, Fig. 16C).
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</paragraph>
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<caption pageId="21" pageNumber="134">
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<paragraph pageId="21" pageNumber="134">
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Figure 16. Humeral region of left elytron and posterior corner of pronotum of three
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion (Nothonepha)" order="Coleoptera" pageId="21" pageNumber="134" phylum="Arthropoda" rank="subGenus" subGenus="Nothonepha">Bembidion (Nothonepha)</taxonomicName>
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species. A
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion tetrapholeon" order="Coleoptera" pageId="21" pageNumber="134" phylum="Arthropoda" rank="species" species="tetrapholeon">Bembidion tetrapholeon</taxonomicName>
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, DRM voucher V100781 B
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion lonae" order="Coleoptera" pageId="21" pageNumber="134" phylum="Arthropoda" rank="species" species="lonae">Bembidion lonae</taxonomicName>
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, DRM voucher V100786 C
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion germainianum" order="Coleoptera" pageId="21" pageNumber="134" phylum="Arthropoda" rank="species" species="germainianum">Bembidion germainianum</taxonomicName>
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, DRM voucher V100782. Arrows show the anterior end of the lateral elytral groove and bead. Scale bar 0.1 mm.
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</paragraph>
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</caption>
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<paragraph pageId="21" pageNumber="134">Mesothorax with prominent pits in the mesepisternum (Fig. 9A), which appear internally as large intrusions that touch in the middle (Figs 10A, C, E). Smaller pits are present ventrally at the junction of abdominal segments II and III (Fig. 13A), which are evident internally as knob-like intrusions (Fig. 14A).</paragraph>
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<paragraph pageId="21" pageNumber="134">Hind wings full.</paragraph>
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<paragraph pageId="21" pageNumber="134">Microsculpture absent from entire dorsal surface of the body except for the cervical region of the head, labrum, and faintly on the clypeus; the beetles are thus brilliantly shiny. Microsculpture absent from most of the ventral surface as well, with the most notable microsculpture being on the undersurface of the head.</paragraph>
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<paragraph pageId="21" pageNumber="134">Aedeagus with nearly straight ventral margin, tip of variable width (Fig. 17). Prominent brush sclerite, and with flagellum not clearly evident from the left side. There is no evident correlation between aedeagal structure and presence or absence of orange spots on the elytra (compare Figs 17A, B to Figs 17C, D).</paragraph>
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<caption pageId="21" pageNumber="134">
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<paragraph pageId="21" pageNumber="134">
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Figure 17. Male aedeagus of
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion tetrapholeon" order="Coleoptera" pageId="21" pageNumber="134" phylum="Arthropoda" rank="species" species="tetrapholeon">Bembidion tetrapholeon</taxonomicName>
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. A Black form, Chile: Region X, Rio Pullinque at Puente Huanehue, 8 km NE Panguipulli, DRM voucher DNA1752 B Black form, Chile: Reg. X,
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<normalizedToken originalValue="Chiloé">Chiloe</normalizedToken>
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: Rio Puntra at route 5, DRM voucher DNA2236 C Orange-spotted form, Argentina:
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<normalizedToken originalValue="Neuquén">Neuquen</normalizedToken>
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: Rio Pichi Traful nr Lago Traful, DRM voucher DNA2564 D Orange-spotted form, Argentina:
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<normalizedToken originalValue="Neuquén">Neuquen</normalizedToken>
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: Rio Pichi Traful nr Lago Traful, DRM voucher DNA2555. Scale bar 0.1 mm.
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</paragraph>
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</caption>
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</subSubSection>
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<subSubSection lastPageId="22" lastPageNumber="135" pageId="21" pageNumber="134" type="morphological variation">
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<paragraph pageId="21" pageNumber="134">Morphological variation.</paragraph>
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<paragraph lastPageId="22" lastPageNumber="135" pageId="21" pageNumber="134">
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The most noted variation is in color of the elytra. Of the 257 specimens examined (including the six specimens identified by Luca Toledano), 245 have uniformly piceous elytra (Fig. 5A); the remaining 12 have a large, diffuse orange spot occupying most of the posterior third of the elytra (Fig. 5B). Ten of these orange-spotted specimens are from the three localities in Argentina, with at least one orange-spotted specimen from each locality, and two of the orange-spotted specimens are from Chile. In most of orange-spotted specimens, the posterior discal seta (ed5) is in the orange region, but immediately around the seta is a small dark spot. No other
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<pageBreakToken pageId="22" pageNumber="135" start="start">aspect</pageBreakToken>
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of morphological or molecular variation was observed to be correlated with presence or absence of the orange spot.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="22" pageNumber="135" type="dna sequence variation">
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<paragraph pageId="22" pageNumber="135">DNA sequence variation.</paragraph>
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<paragraph pageId="22" pageNumber="135">As noted above under Results, there was minor variation present in COI and the wingless gene, and no variation in the other genes studied.</paragraph>
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</subSubSection>
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<subSubSection pageId="22" pageNumber="135" type="habitat and seasonality">
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<paragraph pageId="22" pageNumber="135">Habitat and seasonality.</paragraph>
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<paragraph pageId="22" pageNumber="135">
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At all four localities where habitat data were recorded,
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion tetrapholeon" order="Coleoptera" pageId="22" pageNumber="135" phylum="Arthropoda" rank="species" species="tetrapholeon">Bembidion tetrapholeon</taxonomicName>
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specimens were found on cobble, gravel, and coarse sand shores of clear, fast-flowing rivers (Fig. 6D), from 55 m elevation (Rio Puntra, Isla Grande de
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<normalizedToken originalValue="Chiloé">Chiloe</normalizedToken>
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, Chile) to 830 m elevation (Arroyo
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<normalizedToken originalValue="Queñi">Queni</normalizedToken>
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,
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<normalizedToken originalValue="Neuquén">Neuquen</normalizedToken>
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|
, Argentina). These shorelines lack vascular plants. The beetles occur close to the water, most within 1 m. Specimens have been found in January and February.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="22" pageNumber="135" type="geographic distribution">
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<paragraph pageId="22" pageNumber="135">Geographic distribution.</paragraph>
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<paragraph pageId="22" pageNumber="135">
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In southern Argentina and Chile (Fig. 18). In Argentina this species has been found in
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<normalizedToken originalValue="Neuquén">Neuquen</normalizedToken>
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and Chubut, and in Chile from Regions X and XI.
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</paragraph>
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<caption pageId="22" pageNumber="135">
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<paragraph pageId="22" pageNumber="135">
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Figure 18. Geographic distribution of
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<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion tetrapholeon" order="Coleoptera" pageId="22" pageNumber="135" phylum="Arthropoda" rank="species" species="tetrapholeon">Bembidion tetrapholeon</taxonomicName>
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. Circles filled in black are based upon specimens I have examined and that have been sequenced; circles filled in gray are based upon specimens identified by Luca Toledano. Five cities are included as landmarks.
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</paragraph>
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</caption>
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</subSubSection>
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<subSubSection pageId="23" pageNumber="136" type="relationship to other bembidion">
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<paragraph pageId="23" pageNumber="136">
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<pageBreakToken pageId="23" pageNumber="136" start="start">Relationship</pageBreakToken>
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|
to other
|
|
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="23" pageNumber="136" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
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|
.
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</paragraph>
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|
<paragraph pageId="23" pageNumber="136">
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|
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion tetrapholeon" order="Coleoptera" pageId="23" pageNumber="136" phylum="Arthropoda" rank="species" species="tetrapholeon">Bembidion tetrapholeon</taxonomicName>
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|
is a member of subgenus
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<taxonomicName class="Insecta" family="Carabidae" genus="Nothonepha" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nothonepha" order="Coleoptera" pageId="23" pageNumber="136" phylum="Arthropoda" rank="genus">Nothonepha</taxonomicName>
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|
, as strongly supported by DNA sequences (Table 3) and the presence of shared, derived mesepisternal pits.
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|
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion tetrapholeon" order="Coleoptera" pageId="23" pageNumber="136" phylum="Arthropoda" rank="species" species="tetrapholeon">Bembidion tetrapholeon</taxonomicName>
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|
appears to be the sister of remaining
|
|
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion (Nothonepha)" order="Coleoptera" pageId="23" pageNumber="136" phylum="Arthropoda" rank="subGenus" subGenus="Nothonepha">Bembidion (Nothonepha)</taxonomicName>
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|
(Figs 7, 8). Four genes support this placement (CAD, wg, ArgK, and 18S; Table 3), although the support is weak or moderate in single-gene analyses.
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</paragraph>
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</subSubSection>
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</treatment>
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</document> |