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<document id="E499DCA63F9669F437D085494DDB12F6" ID-DOI="10.5281/zenodo.189922" ID-GBIF-Dataset="921683d4-ada3-4068-aee4-0765c6cc91e4" ID-ISSN="1175-5326" ID-Zenodo-Dep="189922" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1461106188744" checkinUser="plazi" docAuthor="Maddison, David R. &amp; Arnold, Elizabeth" docDate="2009" docId="03E2D11B83489136F0FF94A1FD4EFBDB" docLanguage="en" docName="zt02214p061.pdf" docOrigin="Zootaxa 2214" docStyle="DocumentStyle:890A69B780ED73D6DB8551B71C8AC79E.4:Zootaxa.2009-2012.journal_article" docStyleId="890A69B780ED73D6DB8551B71C8AC79E" docStyleName="Zootaxa.2009-2012.journal_article" docStyleVersion="4" docTitle="Bembidion paraenulum Maddison, n.sp." docType="treatment" docVersion="10" lastPageNumber="60" masterDocId="FFDBA96383439139F068911EFF9AFFF0" masterDocTitle="A review of the Bembidion (Odontium) aenulum subgroup (Coleoptera: Carabidae), with description of a new species" masterLastPageNumber="61" masterPageNumber="45" pageNumber="56" updateTime="1698593221765" updateUser="plazi">
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<mods:title id="B3EC97EA3FC2A38651B5410EA5A26732">A review of the Bembidion (Odontium) aenulum subgroup (Coleoptera: Carabidae), with description of a new species</mods:title>
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<mods:namePart id="EF45EEDCBA4A24DA685200E11078EF76">Maddison, David R.</mods:namePart>
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<mods:date id="26D654693F9EC80038296DE23AB4A5C2">2009</mods:date>
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<treatment id="03E2D11B83489136F0FF94A1FD4EFBDB" ID-DOI="http://doi.org/10.5281/zenodo.5615781" ID-GBIF-Taxon="119623097" ID-Zenodo-Dep="5615781" LSID="urn:lsid:plazi:treatment:03E2D11B83489136F0FF94A1FD4EFBDB" httpUri="http://treatment.plazi.org/id/03E2D11B83489136F0FF94A1FD4EFBDB" lastPageId="15" lastPageNumber="60" pageId="11" pageNumber="56">
<subSubSection id="C351338683489132F0FF94A1FDF5FA0C" pageId="11" pageNumber="56" type="nomenclature">
<paragraph id="8BF4600D83489132F0FF94A1FD1EFA29" blockId="11.[151,644,1471,1532]" box="[151,644,1471,1497]" pageId="11" pageNumber="56">
<heading id="D0BCD76183489132F0FF94A1FD1EFA29" bold="true" box="[151,644,1471,1497]" fontSize="11" level="1" pageId="11" pageNumber="56" reason="1">
<emphasis id="B93FBC1F83489132F0FF94A1FD1EFA29" bold="true" box="[151,644,1471,1497]" pageId="11" pageNumber="56">
<taxonomicName id="4C4B1B8E83489132F0FF94A1FDA6FA29" ID-CoL="68HP7" authority="Maddison" authorityName="Maddison" box="[151,572,1471,1497]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="56" phylum="Arthropoda" rank="species" species="paraenulum" status="sp. nov.">
<emphasis id="B93FBC1F83489132F0FF94A1FE2CFA29" bold="true" box="[151,438,1471,1497]" italics="true" pageId="11" pageNumber="56">Bembidion paraenulum</emphasis>
Maddison
</taxonomicName>
,
<taxonomicNameLabel id="A20C016483489132F22294A1FD1EFA29" box="[586,644,1471,1497]" pageId="11" pageNumber="56" rank="species">n.sp.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BF4600D83489132F0FF94FCFDF5FA0C" blockId="11.[151,644,1471,1532]" box="[151,623,1506,1532]" pageId="11" pageNumber="56">
(
<figureCitation id="13707C8883489132F0F794FCFF7EFA0C" box="[159,228,1506,1532]" captionStart="FIGURE 1" captionStartId="4.[151,256,1352,1376]" captionTargetBox="[151,1435,370,1315]" captionTargetId="figure@4.[151,1436,349,1329]" captionTargetPageId="4" captionText="FIGURE 1. Habitat of Bembidion aenulum (A) and Bembidion paraenulum (B D). A: USA: Iowa: Webster Co., Des Moines River near Stratford. B. aenulum occurs in the coarse, well-drained sand; the river bank also houses Bembidion integrum Casey, B. chalceum, B. cordatum LeConte, B. grandiceps Hayward, B. confusum, and B. rapidum LeConte. B: USA: New Hampshire: Hillsborough Co., Souhegan River, Merrimack. B. paraenulum is found on the sand island on the right along with B. carinula, B. chalceum, B. inaequale Say, and B. nigrum Say. C: Type locality of Bembidion paraenulum, USA: Virginia: Bedford Co., Big Otter River at route 24. Bembidion (Pseudoperyphus) arenobilis Maddison is also abundant in this habitat. The substrate here consists of coarse sand with some gravel. D: USA: Mississippi: Walthall Co., 5.9 mi S Tylertown, Magees Creek. B. paraenulum occurs on the pure sand banks." httpUri="https://zenodo.org/record/189923/files/figure.png" pageId="11" pageNumber="56">Figs 1</figureCitation>
BD, 4B, 5C, 5D, 6B, 7DF, 8B)
</paragraph>
</subSubSection>
<subSubSection id="C351338683489134F0FF9731FACBFA3C" lastPageId="13" lastPageNumber="58" pageId="11" pageNumber="56" type="materials_examined">
<paragraph id="8BF4600D83489132F0FF9731FEE3F8A8" blockId="11.[151,1438,1583,1880]" pageId="11" pageNumber="56">
<emphasis id="B93FBC1F83489132F0FF9731FEC1F9B9" bold="true" box="[151,347,1583,1609]" pageId="11" pageNumber="56">
<typeStatus id="54F0DEAF83489132F0FF9731FF4CF9B9" box="[151,214,1583,1609]" pageId="11" pageNumber="56">Type</typeStatus>
Material.
</emphasis>
<typeStatus id="54F0DEAF83489132F10E9731FE40F9B9" box="[358,474,1583,1609]" pageId="11" pageNumber="56" type="holotype">Holotype</typeStatus>
male (in
<collectionCode id="ED5AF8C883489132F23D9731FD2BF9B9" LSID="urn:lsid:biocol.org:col:34871" box="[597,689,1583,1609]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="11" pageNumber="56">USNM</collectionCode>
, specimen number ADP115779), labelled: &quot;
<collectingCountry id="F35C209D83489132F4B39731FABBF9B9" box="[1243,1313,1583,1609]" name="United States of America" pageId="11" pageNumber="56">USA</collectingCountry>
: Virginia: Bedford Co., Big Otter River at route
<quantity id="4CB3CDE883489132F2FF9748FC8DF980" box="[663,791,1622,1648]" metricMagnitude="4" metricUnit="m" metricValue="2.4175" pageId="11" pageNumber="56" unit="m" value="24175.0">24, 175m</quantity>
,
<geoCoordinate id="EE7F06CA83489132F34E9748FC3FF980" box="[806,933,1622,1648]" direction="north" orientation="latitude" pageId="11" pageNumber="56" precision="55" value="37.246">37.246°N</geoCoordinate>
<geoCoordinate id="EE7F06CA83489132F3C79748FBDFF980" box="[943,1093,1622,1648]" direction="west" orientation="longitude" pageId="11" pageNumber="56" precision="5" value="-79.3495">79.3495°W</geoCoordinate>
,
<date id="FFF546CD83489132F43C9748FB7EF980" box="[1108,1252,1622,1648]" pageId="11" pageNumber="56" value="2004-04-20">20.iv.2004</date>
.
<collectionCode id="ED5AF8C883489132F49A9748FAA4F980" box="[1266,1342,1622,1648]" pageId="11" pageNumber="56">DRM</collectionCode>
04.026. D.R.Maddison, A.E.Arnold&quot; / &quot;David R. Maddison
<collectionCode id="ED5AF8C883489132F29F9763FCADF967" LSID="urn:lsid:biocol.org:col:15536" box="[759,823,1661,1687]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15536" name="Department of Natural Resources, Environment, The Arts and Sport" pageId="11" pageNumber="56">DNA</collectionCode>
1856
<collectionCode id="ED5AF8C883489132F3E89763FC5AF967" LSID="urn:lsid:biocol.org:col:15536" box="[896,960,1661,1687]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15536" name="Department of Natural Resources, Environment, The Arts and Sport" pageId="11" pageNumber="56">DNA</collectionCode>
Voucher&quot; [pale green paper] / &quot;David R. Maddison V
<date id="FFF546CD83489132F15897BDFE16F94D" box="[304,396,1699,1725]" pageId="11" pageNumber="56" value="83-10-01">100183</date>
Voucher Specimen&quot; / &quot;Chromosomes 2n=22+XY David R. Maddison&quot; [pale purple paper] / &quot;
<typeStatus id="54F0DEAF83489132F16297D4FE35F914" box="[266,431,1738,1764]" pageId="11" pageNumber="56" type="holotype">HOLOTYPE</typeStatus>
<taxonomicName id="4C4B1B8E83489132F1D097D4FCFFF914" authority="Maddison" authorityName="Maddison" box="[440,869,1738,1764]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="56" phylum="Arthropoda" rank="species" species="paraenulum">Bembidion paraenulum Maddison</taxonomicName>
&quot; [red paper]. Genitalia in glycerine vial with specimen, with chromosome slide nearby; extracted
<collectionCode id="ED5AF8C883489132F36A97EFFCD8F8FB" LSID="urn:lsid:biocol.org:col:15536" box="[770,834,1777,1803]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15536" name="Department of Natural Resources, Environment, The Arts and Sport" pageId="11" pageNumber="56">DNA</collectionCode>
stored separately. GenBank accession numbers for
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sequences from the
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are
<accessionNumber id="9418FDEE83489132F2359609FD45F8C1" box="[605,735,1815,1841]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/GQ424282" pageId="11" pageNumber="56">GQ424282</accessionNumber>
(28S rDNA),
<accessionNumber id="9418FDEE83489132F3ED9609FB9DF8C1" box="[901,1031,1815,1841]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/GQ424254" pageId="11" pageNumber="56">GQ424254</accessionNumber>
(
<collectionCode id="ED5AF8C883489132F47F9609FBD5F8C1" LSID="urn:lsid:biocol.org:col:14498" box="[1047,1103,1815,1841]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:14498" name="University of Coimbra Botany Department" pageId="11" pageNumber="56">COI</collectionCode>
),
<accessionNumber id="9418FDEE83489132F40A9609FB7EF8C1" box="[1122,1252,1815,1841]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/GQ424288" pageId="11" pageNumber="56">GQ424288</accessionNumber>
(
<emphasis id="B93FBC1F83489132F49D9606FAC0F8C1" box="[1269,1370,1816,1841]" italics="true" pageId="11" pageNumber="56">wingless</emphasis>
), and
<accessionNumber id="9418FDEE83489132F0FF9620FE83F8A8" box="[151,281,1854,1880]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/GQ424226" pageId="11" pageNumber="56">GQ424226</accessionNumber>
(
<collectingCountry id="F35C209D83489132F1419620FEF7F8A8" box="[297,365,1854,1880]" name="Chad" pageId="11" pageNumber="56">
<collectionCode id="ED5AF8C883489132F1419620FEF7F8A8" box="[297,365,1854,1880]" pageId="11" pageNumber="56">CAD</collectionCode>
</collectingCountry>
).
</paragraph>
<caption id="DF343085834F9135F0FF9614FDA8F802" httpUri="https://zenodo.org/record/189929/files/figure.png" pageId="12" pageNumber="57" targetBox="[252,1317,214,1773]" targetPageId="12">
<paragraph id="8BF4600D834F9135F0FF9614FDA8F802" blockId="12.[151,1437,1802,2034]" pageId="12" pageNumber="57">
<emphasis id="B93FBC1F834F9135F0FF9614FE83F8D2" bold="true" box="[151,281,1802,1826]" pageId="12" pageNumber="57">FIGURE 7.</emphasis>
Aedeagus, left side, of
<taxonomicName id="4C4B1B8E834F9135F27C9615FD70F8D2" box="[532,746,1803,1826]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834F9135F27C9615FD70F8D2" box="[532,746,1803,1826]" italics="true" pageId="12" pageNumber="57">Bembidion aenulum</emphasis>
</taxonomicName>
(AC) and
<taxonomicName id="4C4B1B8E834F9135F3009615FB9FF8D2" box="[872,1029,1803,1826]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F834F9135F3009615FB9FF8D2" box="[872,1029,1803,1826]" italics="true" pageId="12" pageNumber="57">B. paraenulum</emphasis>
</taxonomicName>
(DF). A:
<taxonomicName id="4C4B1B8E834F9135F4149615FB6CF8D2" box="[1148,1270,1803,1826]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834F9135F4149615FB6CF8D2" box="[1148,1270,1803,1826]" italics="true" pageId="12" pageNumber="57">B. aenulum</emphasis>
</taxonomicName>
, DRM voucher V100290, USA: Iowa: Webster Co., Des Moines River near Stratford. B:
<taxonomicName id="4C4B1B8E834F9135F3D59633FBA3F8B4" box="[957,1081,1837,1860]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834F9135F3D59633FBA3F8B4" box="[957,1081,1837,1860]" italics="true" pageId="12" pageNumber="57">B. aenulum</emphasis>
</taxonomicName>
, DRM voucher V100291, USA: Iowa: Jones Co., Wapsipinicon River W of Oxford Junction. C:
<taxonomicName id="4C4B1B8E834F9135F328964EFC22F897" box="[832,952,1872,1895]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834F9135F328964EFC22F897" box="[832,952,1872,1895]" italics="true" pageId="12" pageNumber="57">B. aenulum</emphasis>
</taxonomicName>
, DRM voucher V100292, USA: Iowa: Cedar Co., Cedar River at route X40. D:
<taxonomicName id="4C4B1B8E834F9135F26C966DFD3BF87A" box="[516,673,1907,1930]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F834F9135F26C966DFD3BF87A" box="[516,673,1907,1930]" italics="true" pageId="12" pageNumber="57">B. paraenulum</emphasis>
</taxonomicName>
, DRM voucher V100293, USA: Virginia: Bedford Co., Big Otter River at route 24. E:
<taxonomicName id="4C4B1B8E834F9135F156968BFE78F85C" box="[318,482,1941,1964]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F834F9135F156968BFE78F85C" box="[318,482,1941,1964]" italics="true" pageId="12" pageNumber="57">B. paraenulum</emphasis>
</taxonomicName>
, DRM voucher V100295, USA: New Hampshire: Hillsborough Co., Souhegan River, Merrimack. F:
<taxonomicName id="4C4B1B8E834F9135F15E96A6FE4EF83F" box="[310,468,1976,1999]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="57" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F834F9135F15E96A6FE4EF83F" box="[310,468,1976,1999]" italics="true" pageId="12" pageNumber="57">B. paraenulum</emphasis>
</taxonomicName>
, DRM voucher V100514, USA: Mississippi: Stone Co., Red Creek W.M.A., near Wiggins. All to same scale; scale bar is 0.1 mm.
</paragraph>
</caption>
<caption id="DF343085834E9134F0FF9245FB8AFC2A" httpUri="https://zenodo.org/record/189930/files/figure.png" pageId="13" pageNumber="58" targetBox="[259,1327,194,832]" targetPageId="13">
<paragraph id="8BF4600D834E9134F0FF9245FB8AFC2A" blockId="13.[151,1436,859,986]" pageId="13" pageNumber="58">
<emphasis id="B93FBC1F834E9134F0FF9245FE87FC83" bold="true" box="[151,285,859,883]" pageId="13" pageNumber="58">FIGURE 8.</emphasis>
Details of internal sac of male genitalia, showing central sclerites, from left side. Consistent differences between two species indicated with arrows. A:
<taxonomicName id="4C4B1B8E834E9134F2F69260FCE0FC65" box="[670,890,894,917]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834E9134F2F69260FCE0FC65" box="[670,890,894,917]" italics="true" pageId="13" pageNumber="58">Bembidion aenulum</emphasis>
</taxonomicName>
, DRM voucher V100291, USA: Iowa: Jones Co., Wapsipinicon River W of Oxford Junction. B:
<taxonomicName id="4C4B1B8E834E9134F2ED92BFFCB8FC48" box="[645,802,929,952]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F834E9134F2ED92BFFCB8FC48" box="[645,802,929,952]" italics="true" pageId="13" pageNumber="58">B. paraenulum</emphasis>
</taxonomicName>
, DRM voucher V100293, USA: Virginia: Bedford Co., Big Otter River at route 24. VSP: Ventral sclerite patch; CSC: Central sclerite complex.
</paragraph>
</caption>
<paragraph id="8BF4600D834E9134F0AE9517FC08FAFB" blockId="13.[151,1437,1033,1987]" pageId="13" pageNumber="58">
<typeStatus id="54F0DEAF834E9134F0AE9517FEA7FBD3" box="[198,317,1033,1059]" pageId="13" pageNumber="58" type="paratype">Paratypes</typeStatus>
:
<specimenCount id="9D4DAB84834E9134F1209517FE6FFBD3" box="[328,501,1033,1059]" pageId="13" pageNumber="58" type="generic">103 specimens</specimenCount>
from the following localities, deposited in
<collectionCode id="ED5AF8C8834E9134F3869517FBD3FBD3" box="[1006,1097,1033,1059]" pageId="13" pageNumber="58">BMNH</collectionCode>
,
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,
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,
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,
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,
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<collectionCode id="ED5AF8C8834E9134F2D69531FC92FBB9" box="[702,776,1071,1097]" httpUri="http://grbio.org/cool/tizb-nd41" name="University of Arizona" pageId="13" pageNumber="58">UAIC</collectionCode>
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, and
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: Same data as
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(25);
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<collectingCountry id="F35C209D834E9134F53F9531FA0CFBB9" box="[1367,1430,1071,1097]" name="United States of America" pageId="13" pageNumber="58">USA</collectingCountry>
: VIRGINIA
</emphasis>
:
<emphasis id="B93FBC1F834E9134F1539548FD8DFB80" bold="true" box="[315,535,1110,1136]" pageId="13" pageNumber="58">Danville City Co</emphasis>
., Danville, Dan River at Robertson Bridge,
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(33);
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(6);
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(6);
<emphasis id="B93FBC1F834E9134F3359562FB95FB66" bold="true" box="[861,1039,1148,1174]" pageId="13" pageNumber="58">Campbell Co</emphasis>
., Roanoke River at Brookneal,
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(2); Altavista, Roanoke River,
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<geoCoordinate id="EE7F06CA834E9134F3DE95BDFBA5FB4D" box="[950,1087,1187,1213]" direction="west" orientation="longitude" pageId="13" pageNumber="58" precision="5" value="-79.2863">79.2863°W</geoCoordinate>
(8).
<emphasis id="B93FBC1F834E9134F41395BDFE8EFB13" bold="true" pageId="13" pageNumber="58">
<collectionCode id="ED5AF8C8834E9134F41395BDFB7DFB4D" box="[1147,1255,1187,1213]" pageId="13" pageNumber="58">NORTH</collectionCode>
CAROLINA: Stokes Co
</emphasis>
., Town Fork Creek at Walnut Cove,
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<geoCoordinate id="EE7F06CA834E9134F33795D4FC72FB14" box="[863,1000,1226,1252]" direction="west" orientation="longitude" pageId="13" pageNumber="58" precision="5" value="-80.1403">80.1403°W</geoCoordinate>
(14);
<emphasis id="B93FBC1F834E9134F45B95D7FA9BFB13" bold="true" box="[1075,1281,1225,1251]" pageId="13" pageNumber="58">Rockingham Co</emphasis>
., Dan River, Madison,
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<geoCoordinate id="EE7F06CA834E9134F1FB95EFFD80FAFB" box="[403,538,1265,1291]" direction="west" orientation="longitude" pageId="13" pageNumber="58" precision="5" value="-79.993">79.9930°W</geoCoordinate>
(3);
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<geoCoordinate id="EE7F06CA834E9134F2B295EFFCFBFAFB" box="[730,865,1265,1291]" direction="west" orientation="longitude" pageId="13" pageNumber="58" precision="5" value="-79.9918">79.9918°W</geoCoordinate>
(6).
</paragraph>
<paragraph id="8BF4600D834E9134F0AD9409FD15FA55" blockId="13.[151,1437,1033,1987]" pageId="13" pageNumber="58">
<emphasis id="B93FBC1F834E9134F0AD9409FEF5FAC1" bold="true" box="[197,367,1303,1329]" pageId="13" pageNumber="58">
<typeStatus id="54F0DEAF834E9134F0AD9409FE99FAC1" box="[197,259,1303,1329]" pageId="13" pageNumber="58">Type</typeStatus>
locality.
</emphasis>
<collectingCountry id="F35C209D834E9134F11F9409FE2DFAC1" box="[375,439,1303,1329]" name="United States of America" pageId="13" pageNumber="58">USA</collectingCountry>
: Virginia: Bedford Co., Big Otter River at route
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,
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<geoCoordinate id="EE7F06CA834E9134F5669409FA02FAC1" box="[1294,1432,1303,1329]" direction="west" orientation="longitude" pageId="13" pageNumber="58" precision="5" value="-79.3495">79.3495°W</geoCoordinate>
. This is about
<quantity id="4CB3CDE8834E9134F15F9420FE13FAA8" box="[311,393,1342,1368]" metricMagnitude="3" metricUnit="m" metricValue="5.5" pageId="13" pageNumber="58" unit="km" value="5.5">5.5 km</quantity>
WNW of Evington, at the mouth of Lick Run where it empties into the Big Otter River, at the NW corner of the bridge of route 24. The species is abundant there, and shares the habitat with the similarly common
<taxonomicName id="4C4B1B8E834E9134F11C9492FD15FA55" authority="Maddison." authorityName="Maddison." box="[372,655,1419,1445]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="arenobilis">
<emphasis id="B93FBC1F834E9134F11C9492FD91FA55" box="[372,523,1420,1445]" italics="true" pageId="13" pageNumber="58">B. arenobilis</emphasis>
Maddison.
</taxonomicName>
</paragraph>
<paragraph id="8BF4600D834E9134F0AD94AFFACBFA3C" blockId="13.[151,1437,1033,1987]" box="[197,1361,1457,1484]" pageId="13" pageNumber="58">
<emphasis id="B93FBC1F834E9134F0AD94AFFE55FA3B" bold="true" box="[197,463,1457,1483]" pageId="13" pageNumber="58">Nomenclatorial notes</emphasis>
. This is the species called &quot;
<taxonomicName id="4C4B1B8E834E9134F37C94ACFC09FA3B" box="[788,915,1458,1483]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="genus">
<emphasis id="B93FBC1F834E9134F37C94ACFC09FA3B" box="[788,915,1458,1483]" italics="true" pageId="13" pageNumber="58">Bembidion</emphasis>
</taxonomicName>
sp. nr.
<taxonomicName id="4C4B1B8E834E9134F38094ACFBD7FA3B" box="[1000,1101,1458,1483]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834E9134F38094ACFBD7FA3B" box="[1000,1101,1458,1483]" italics="true" pageId="13" pageNumber="58">aenulum</emphasis>
</taxonomicName>
&quot; in
<bibRefCitation id="EFDA1DFC834E9134F41794ACFAD7FA3C" author="Maddison" box="[1151,1357,1458,1484]" lastPageId="13" lastPageNumber="61" pageId="13" pageNumber="60" refString="Maddison, D. R. (1993) Systematics of the Holarctic beetle subgenus Bracteon and related Bembidion (Coleoptera: Carabidae). Bulletin of the Museum of Comparative Zoology, 153, 143 - 299." type="journal article" year="1993">Maddison (1993)</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C3513386834E9134F0AD94C6FB6BF833" pageId="13" pageNumber="58" type="diagnosis">
<paragraph id="8BF4600D834E9134F0AD94C6FDAAF8D8" blockId="13.[151,1437,1033,1987]" pageId="13" pageNumber="58">
<emphasis id="B93FBC1F834E9134F0AD94C6FEA7FA02" bold="true" box="[197,317,1496,1522]" pageId="13" pageNumber="58">Diagnosis</emphasis>
. Most specimens larger than specimens of
<taxonomicName id="4C4B1B8E834E9134F35094C7FC24FA02" box="[824,958,1497,1522]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834E9134F35094C7FC24FA02" box="[824,958,1497,1522]" italics="true" pageId="13" pageNumber="58">B. aenulum</emphasis>
</taxonomicName>
(
<figureCitation id="13707C88834E9134F3A694C7FB8FFA03" box="[974,1045,1497,1523]" captionStart="FIGURE 4" captionStartId="9.[151,255,1907,1931]" captionTargetBox="[151,1436,294,1898]" captionTargetId="figure@9.[151,1436,290,1900]" captionTargetPageId="9" captionText="FIGURE 4. Male habitus. A: Bembidion aenulum, DRM voucher V 100128, Iowa: Des Moines River near Stratford. B: Bembidion paraenulum, DRM voucher V 100078, New Hampshire: Merrimack. C: Specimens of each species showing typical size difference: Bembidion aenulum (left), DRM voucher V 100129, Iowa: Des Moines River near Stratford and B. paraenulum (right), DRM voucher V 100130, USA: North Carolina: Rockingham Co., Dan River, Madison." httpUri="https://zenodo.org/record/189926/files/figure.png" pageId="13" pageNumber="58">Fig. 4</figureCitation>
C). Prothorax more parallel sided than
<taxonomicName id="4C4B1B8E834E9134F0B8971EFECCF9E9" box="[208,342,1536,1561]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834E9134F0B8971EFECCF9E9" box="[208,342,1536,1561]" italics="true" pageId="13" pageNumber="58">B. aenulum</emphasis>
</taxonomicName>
, and more rounded sides (
<figureCitation id="13707C88834E9134F2E294E1FD4EF9E9" box="[650,724,1535,1561]" captionStart="FIGURE 5" captionStartId="10.[151,257,1332,1356]" captionTargetBox="[151,1435,194,1294]" captionTargetId="figure@10.[151,1436,194,1308]" captionTargetPageId="10" captionText="FIGURE 5. Pronota and elytral shoulder. A: Bembidion aenulum, DRM voucher V 100515, Iowa: Des Moines River near Stratford. B: Comparison of shoulder area and hind pronotal angle of B. aenulum and B. paraenulum. Note how the humeral sulcus is nearly parallel to the hind margin of the pronotum in B. aenulum, whereas in B. paraenulum the sulcus diverges from the pronotal hind margin toward the middle of the beetle. This also means that mesal end of the sulcus points more posteriad in B. paraenulum than B. aenulum. C: B. paraenulum holotype, Virginia: Big Otter River. D: B. paraenulum, DRM voucher V 100126, Mississippi: Perry Co., DeSoto N. F., Black Creek at Fairley Bridge Landing. All photographs to same scale; scale bar is 0.5 mm." httpUri="https://zenodo.org/record/189927/files/figure.png" pageId="13" pageNumber="58">Figs 5</figureCitation>
C, D); base of pronotum more evidently narrower than elytra (
<figureCitation id="13707C88834E9134F0F79738FF7DF9B0" box="[159,231,1574,1600]" captionStart="FIGURE 4" captionStartId="9.[151,255,1907,1931]" captionTargetBox="[151,1436,294,1898]" captionTargetId="figure@9.[151,1436,290,1900]" captionTargetPageId="9" captionText="FIGURE 4. Male habitus. A: Bembidion aenulum, DRM voucher V 100128, Iowa: Des Moines River near Stratford. B: Bembidion paraenulum, DRM voucher V 100078, New Hampshire: Merrimack. C: Specimens of each species showing typical size difference: Bembidion aenulum (left), DRM voucher V 100129, Iowa: Des Moines River near Stratford and B. paraenulum (right), DRM voucher V 100130, USA: North Carolina: Rockingham Co., Dan River, Madison." httpUri="https://zenodo.org/record/189926/files/figure.png" pageId="13" pageNumber="58">Fig. 4</figureCitation>
B); pronotum more convex, with front angles more depressed; laterobasal depressions deeper, and more evident (
<figureCitation id="13707C88834E9134F0949753FEDCF997" box="[252,326,1613,1639]" captionStart="FIGURE 5" captionStartId="10.[151,257,1332,1356]" captionTargetBox="[151,1435,194,1294]" captionTargetId="figure@10.[151,1436,194,1308]" captionTargetPageId="10" captionText="FIGURE 5. Pronota and elytral shoulder. A: Bembidion aenulum, DRM voucher V 100515, Iowa: Des Moines River near Stratford. B: Comparison of shoulder area and hind pronotal angle of B. aenulum and B. paraenulum. Note how the humeral sulcus is nearly parallel to the hind margin of the pronotum in B. aenulum, whereas in B. paraenulum the sulcus diverges from the pronotal hind margin toward the middle of the beetle. This also means that mesal end of the sulcus points more posteriad in B. paraenulum than B. aenulum. C: B. paraenulum holotype, Virginia: Big Otter River. D: B. paraenulum, DRM voucher V 100126, Mississippi: Perry Co., DeSoto N. F., Black Creek at Fairley Bridge Landing. All photographs to same scale; scale bar is 0.5 mm." httpUri="https://zenodo.org/record/189927/files/figure.png" pageId="13" pageNumber="58">Figs 5</figureCitation>
C, D). Humeral sulcus of elytron medially diverging more from basal margin of pronotum than in
<taxonomicName id="4C4B1B8E834E9134F0DF976AFED8F97D" box="[183,322,1652,1677]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834E9134F0DF976AFED8F97D" box="[183,322,1652,1677]" italics="true" pageId="13" pageNumber="58">B. aenulum</emphasis>
</taxonomicName>
(
<figureCitation id="13707C88834E9134F13B976DFE3AF97D" box="[339,416,1651,1677]" captionStart="FIGURE 5" captionStartId="10.[151,257,1332,1356]" captionTargetBox="[151,1435,194,1294]" captionTargetId="figure@10.[151,1436,194,1308]" captionTargetPageId="10" captionText="FIGURE 5. Pronota and elytral shoulder. A: Bembidion aenulum, DRM voucher V 100515, Iowa: Des Moines River near Stratford. B: Comparison of shoulder area and hind pronotal angle of B. aenulum and B. paraenulum. Note how the humeral sulcus is nearly parallel to the hind margin of the pronotum in B. aenulum, whereas in B. paraenulum the sulcus diverges from the pronotal hind margin toward the middle of the beetle. This also means that mesal end of the sulcus points more posteriad in B. paraenulum than B. aenulum. C: B. paraenulum holotype, Virginia: Big Otter River. D: B. paraenulum, DRM voucher V 100126, Mississippi: Perry Co., DeSoto N. F., Black Creek at Fairley Bridge Landing. All photographs to same scale; scale bar is 0.5 mm." httpUri="https://zenodo.org/record/189927/files/figure.png" pageId="13" pageNumber="58">Figs 5</figureCitation>
C,D). Shallower microsculpture than
<taxonomicName id="4C4B1B8E834E9134F303976AFC6FF97D" box="[875,1013,1652,1677]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834E9134F303976AFC6FF97D" box="[875,1013,1652,1677]" italics="true" pageId="13" pageNumber="58">B. aenulum</emphasis>
</taxonomicName>
, and thus shinier, most notably on pronotum. Mentum tooth narrower, more parallel-sided (
<figureCitation id="13707C88834E9134F3529784FC19F944" box="[826,899,1690,1716]" captionStart="FIGURE 6" captionStartId="11.[151,255,1293,1317]" captionTargetBox="[318,1244,515,1245]" captionTargetId="figure@11.[317,1265,472,1259]" captionTargetPageId="11" captionText="FIGURE 6. Ventral surface of mentum. A: Bembidion aenulum, DRM voucher V 100290, Iowa: Des Moines River near Stratford. B: B. paraenulum, DRM voucher V 100293, Virginia: Big Otter River at route 24. Drawings to same scale; scale bar is 0.1 mm." httpUri="https://zenodo.org/record/189928/files/figure.png" pageId="13" pageNumber="58">Fig. 6</figureCitation>
B). Male aedeagus longer, with narrower and less blunt apex than
<taxonomicName id="4C4B1B8E834E9134F1E397DFFD8BF92A" box="[395,529,1729,1754]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834E9134F1E397DFFD8BF92A" box="[395,529,1729,1754]" italics="true" pageId="13" pageNumber="58">B. aenulum</emphasis>
</taxonomicName>
(
<figureCitation id="13707C88834E9134F24A97DFFDF6F92B" box="[546,620,1729,1755]" captionStart="FIGURE 7" captionStartId="12.[151,255,1802,1826]" captionTargetBox="[252,1317,214,1773]" captionTargetId="figure@12.[248,1332,194,1782]" captionTargetPageId="12" captionText="FIGURE 7. Aedeagus, left side, of Bembidion aenulum (A C) and B. paraenulum (D F). A: B. aenulum, DRM voucher V 100290, USA: Iowa: Webster Co., Des Moines River near Stratford. B: B. aenulum, DRM voucher V 100291, USA: Iowa: Jones Co., Wapsipinicon River W of Oxford Junction. C: B. aenulum, DRM voucher V 100292, USA: Iowa: Cedar Co., Cedar River at route X 40. D: B. paraenulum, DRM voucher V 100293, USA: Virginia: Bedford Co., Big Otter River at route 24. E: B. paraenulum, DRM voucher V 100295, USA: New Hampshire: Hillsborough Co., Souhegan River, Merrimack. F: B. paraenulum, DRM voucher V 100514, USA: Mississippi: Stone Co., Red Creek W. M. A., near Wiggins. All to same scale; scale bar is 0.1 mm." httpUri="https://zenodo.org/record/189929/files/figure.png" pageId="13" pageNumber="58">Figs 7</figureCitation>
DF); ventral sclerite patch with dorsal ridge straighter, and meeting the ventral margin of the anterior basin at an angle (
<figureCitation id="13707C88834E9134F29497F9FCD8F8F1" box="[764,834,1767,1793]" captionStart="FIGURE 8" captionStartId="13.[151,257,859,883]" captionTargetBox="[259,1327,194,832]" captionTargetId="figure@13.[259,1328,194,835]" captionTargetPageId="13" captionText="FIGURE 8. Details of internal sac of male genitalia, showing central sclerites, from left side. Consistent differences between two species indicated with arrows. A: Bembidion aenulum, DRM voucher V 100291, USA: Iowa: Jones Co., Wapsipinicon River W of Oxford Junction. B: B. paraenulum, DRM voucher V 100293, USA: Virginia: Bedford Co., Big Otter River at route 24. VSP: Ventral sclerite patch; CSC: Central sclerite complex." httpUri="https://zenodo.org/record/189930/files/figure.png" pageId="13" pageNumber="58">Fig. 8</figureCitation>
B, VSP); right lobe of central sclerite complex less rounded (
<figureCitation id="13707C88834E9134F16F9610FED7F8D8" box="[263,333,1806,1832]" captionStart="FIGURE 8" captionStartId="13.[151,257,859,883]" captionTargetBox="[259,1327,194,832]" captionTargetId="figure@13.[259,1328,194,835]" captionTargetPageId="13" captionText="FIGURE 8. Details of internal sac of male genitalia, showing central sclerites, from left side. Consistent differences between two species indicated with arrows. A: Bembidion aenulum, DRM voucher V 100291, USA: Iowa: Jones Co., Wapsipinicon River W of Oxford Junction. B: B. paraenulum, DRM voucher V 100293, USA: Virginia: Bedford Co., Big Otter River at route 24. VSP: Ventral sclerite patch; CSC: Central sclerite complex." httpUri="https://zenodo.org/record/189930/files/figure.png" pageId="13" pageNumber="58">Fig. 8</figureCitation>
B, CSC right lobe).
</paragraph>
<paragraph id="8BF4600D834E9134F0AD962AFB6BF833" blockId="13.[151,1437,1033,1987]" pageId="13" pageNumber="58">
<emphasis id="B93FBC1F834E9134F0AD962AFD92F8BE" bold="true" box="[197,520,1844,1870]" pageId="13" pageNumber="58">Additional characteristics</emphasis>
. ABL 3.8 to 5.0 mm, with most specimens longer than
<quantity id="4CB3CDE8834E9134F4FB962BFB6BF8BF" box="[1171,1265,1845,1871]" metricMagnitude="-3" metricUnit="m" metricValue="4.3" pageId="13" pageNumber="58" unit="mm" value="4.3">4.3 mm</quantity>
. Color as in
<emphasis id="B93FBC1F834E9134F5EB962BFE99F885" italics="true" pageId="13" pageNumber="58">
<taxonomicName id="4C4B1B8E834E9134F5EB962BFE9AF885" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="aenulum">B. aenulum</taxonomicName>
,
</emphasis>
except often with a less-intense metallic reflection, which is also rarely as gold or reddish as in
<taxonomicName id="4C4B1B8E834E9134F5EB9642FF64F86B" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="58" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834E9134F5EB9642FF64F86B" italics="true" pageId="13" pageNumber="58">B. aenulum</emphasis>
</taxonomicName>
, and more frequently greenish. Other characteristics as in the subgroup description. Chromosome number is 2n = 22+XY (three males examined, two from Virginia: Danville, and the
<typeStatus id="54F0DEAF834E9134F41196B7FB7FF833" box="[1145,1253,1961,1987]" pageId="13" pageNumber="58" type="holotype">holotype</typeStatus>
).
</paragraph>
</subSubSection>
<subSubSection id="C3513386834D9136F0AD9189FBC1FE6A" lastPageId="15" lastPageNumber="60" pageId="14" pageNumber="59" type="description">
<paragraph id="8BF4600D834D9137F0AD9189FD54FF0F" blockId="14.[151,1437,151,255]" pageId="14" pageNumber="59">
<emphasis id="B93FBC1F834D9137F0AD9189FE42FF41" bold="true" box="[197,472,151,177]" pageId="14" pageNumber="59">Geographic variation</emphasis>
. The smallest specimens (
<quantity id="4CB3CDE8834D9137F3709186FCEDFF42" box="[792,887,152,178]" metricMagnitude="-3" metricUnit="m" metricValue="3.8" pageId="14" pageNumber="59" unit="mm" value="3.8">3.8 mm</quantity>
) are males from New Hampshire. Specimens from the other regions are all larger than
<quantity id="4CB3CDE8834D9137F21391A0FD43FF28" box="[635,729,190,216]" metricMagnitude="-3" metricUnit="m" metricValue="4.1" pageId="14" pageNumber="59" unit="mm" value="4.1">4.1 mm</quantity>
. Specimens from Mississippi have larger, more convex, and wider prothoraces (compare
<figureCitation id="13707C88834D9137F18E91FBFDB0FF0F" box="[486,554,229,255]" captionStart="FIGURE 5" captionStartId="10.[151,257,1332,1356]" captionTargetBox="[151,1435,194,1294]" captionTargetId="figure@10.[151,1436,194,1308]" captionTargetPageId="10" captionText="FIGURE 5. Pronota and elytral shoulder. A: Bembidion aenulum, DRM voucher V 100515, Iowa: Des Moines River near Stratford. B: Comparison of shoulder area and hind pronotal angle of B. aenulum and B. paraenulum. Note how the humeral sulcus is nearly parallel to the hind margin of the pronotum in B. aenulum, whereas in B. paraenulum the sulcus diverges from the pronotal hind margin toward the middle of the beetle. This also means that mesal end of the sulcus points more posteriad in B. paraenulum than B. aenulum. C: B. paraenulum holotype, Virginia: Big Otter River. D: B. paraenulum, DRM voucher V 100126, Mississippi: Perry Co., DeSoto N. F., Black Creek at Fairley Bridge Landing. All photographs to same scale; scale bar is 0.5 mm." httpUri="https://zenodo.org/record/189927/files/figure.png" pageId="14" pageNumber="59">Fig. 5</figureCitation>
D to
<figureCitation id="13707C88834D9137F20D91FBFD30FF0F" box="[613,682,229,255]" captionStart="FIGURE 5" captionStartId="10.[151,257,1332,1356]" captionTargetBox="[151,1435,194,1294]" captionTargetId="figure@10.[151,1436,194,1308]" captionTargetPageId="10" captionText="FIGURE 5. Pronota and elytral shoulder. A: Bembidion aenulum, DRM voucher V 100515, Iowa: Des Moines River near Stratford. B: Comparison of shoulder area and hind pronotal angle of B. aenulum and B. paraenulum. Note how the humeral sulcus is nearly parallel to the hind margin of the pronotum in B. aenulum, whereas in B. paraenulum the sulcus diverges from the pronotal hind margin toward the middle of the beetle. This also means that mesal end of the sulcus points more posteriad in B. paraenulum than B. aenulum. C: B. paraenulum holotype, Virginia: Big Otter River. D: B. paraenulum, DRM voucher V 100126, Mississippi: Perry Co., DeSoto N. F., Black Creek at Fairley Bridge Landing. All photographs to same scale; scale bar is 0.5 mm." httpUri="https://zenodo.org/record/189927/files/figure.png" pageId="14" pageNumber="59">Fig. 5</figureCitation>
C).
</paragraph>
<caption id="DF343085834D9137F0FF9492FADBF9FB" httpUri="https://zenodo.org/record/189931/files/figure.png" pageId="14" pageNumber="59" targetBox="[168,1420,332,1379]" targetPageId="14">
<paragraph id="8BF4600D834D9137F0FF9492FADBF9FB" blockId="14.[151,1436,1420,1547]" pageId="14" pageNumber="59">
<emphasis id="B93FBC1F834D9137F0FF9492FE83FA54" bold="true" box="[151,281,1420,1444]" pageId="14" pageNumber="59">FIGURE 9.</emphasis>
Geographic distribution of
<taxonomicName id="4C4B1B8E834D9137F2569493FC8EFA54" box="[574,788,1421,1444]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="59" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834D9137F2569493FC8EFA54" box="[574,788,1421,1444]" italics="true" pageId="14" pageNumber="59">Bembidion aenulum</emphasis>
</taxonomicName>
(stars) and
<taxonomicName id="4C4B1B8E834D9137F3E79493FBB6FA54" box="[911,1068,1421,1444]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="59" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F834D9137F3E79493FBB6FA54" box="[911,1068,1421,1444]" italics="true" pageId="14" pageNumber="59">B. paraenulum</emphasis>
</taxonomicName>
(circles). Black symbols represent specimens identified using DNA sequences and morphological data; gray symbols are those identified only with morphological data. The open circles represent specimens of
<taxonomicName id="4C4B1B8E834D9137F35D94CCFC4DFA19" box="[821,983,1490,1513]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="59" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F834D9137F35D94CCFC4DFA19" box="[821,983,1490,1513]" italics="true" pageId="14" pageNumber="59">B. paraenulum</emphasis>
</taxonomicName>
, but with labels only indicating the state; these localities are considered questionable as they are so far outside the well-documented range of this species.
</paragraph>
</caption>
<paragraph id="8BF4600D834D9137F0AD9720FC05F8E9" blockId="14.[151,1438,1598,2011]" pageId="14" pageNumber="59">
Specimens from the three geographic regions also differ in DNA sequences. The three New Hampshire specimens have three unique bases in COI (1 of which is a unique within the other
<taxonomicName id="4C4B1B8E834D9137F41F977BFB6DF98E" box="[1143,1271,1637,1662]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="59" phylum="Arthropoda" rank="genus">
<emphasis id="B93FBC1F834D9137F41F977BFB6DF98E" box="[1143,1271,1637,1662]" italics="true" pageId="14" pageNumber="59">Bembidion</emphasis>
</taxonomicName>
sampled, and thus possibly a synapomorphy), the Mississippi population has 2 unique bases (both of which are unique within
<taxonomicName id="4C4B1B8E834D9137F08397ACFEF1F93B" box="[235,363,1714,1739]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="59" phylum="Arthropoda" rank="genus">
<emphasis id="B93FBC1F834D9137F08397ACFEF1F93B" box="[235,363,1714,1739]" italics="true" pageId="14" pageNumber="59">Bembidion</emphasis>
</taxonomicName>
), and the Virginia/North Carolina populations have three unique bases, none of which are unique within
<taxonomicName id="4C4B1B8E834D9137F12997C7FE25F902" box="[321,447,1753,1778]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="59" phylum="Arthropoda" rank="genus">
<emphasis id="B93FBC1F834D9137F12997C7FE25F902" box="[321,447,1753,1778]" italics="true" pageId="14" pageNumber="59">Bembidion</emphasis>
</taxonomicName>
. Thus, sequences from the three regions differ consistently, one from each other, by five or six bases out of the 762 bases sequenced for all specimens.
</paragraph>
<paragraph id="8BF4600D834D9136F0AD9638FDC5FED6" blockId="14.[151,1438,1598,2011]" lastBlockId="15.[151,1437,152,1067]" lastPageId="15" lastPageNumber="60" pageId="14" pageNumber="59">
In 28S, site 496 of the analyzed sequence varies between geographic regions. This site is the tenth in the sequence CACTTCTCCCCTTGTAGGA. All specimens of
<taxonomicName id="4C4B1B8E834D9137F33E9653FB99F896" box="[854,1027,1869,1894]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="59" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F834D9137F33E9653FB99F896" box="[854,1027,1869,1894]" italics="true" pageId="14" pageNumber="59">B. paraenulum</emphasis>
</taxonomicName>
examined contain 28S copies with this sequence within their genomes, as all specimens have a dominant peak of a C at that site in chromatograms. In addition, most specimens have a secondary peak of T at that site, suggesting they also contain some 28S copies with the sequence CACTTCTCCTCTTGTAGGA. In the Virginia, North Carolina, and Mississippi specimens, the T peak is less than 7% of the C peak, suggesting that almost all of the copies of ribosomal DNA in the genome have C at that site, with only a few copies having T. In contrast, the three New Hampshire specimens have the T peak 1950% of the C peak, indicating a higher fraction of the copies are likely T at that site in New Hampshire.
</paragraph>
<paragraph id="8BF4600D834C9136F0AD902CFBC1FE6A" blockId="15.[151,1437,152,1067]" pageId="15" pageNumber="60">
<emphasis id="B93FBC1F834C9136F0AD902CFEBDFEBC" bold="true" box="[197,295,306,332]" pageId="15" pageNumber="60">Habitat</emphasis>
. We have collected this species at nine localities in New Hampshire, Virginia, North Carolina, and Mississippi along rivers and creeks on pure sand shores, or sand with some gravel (
<figureCitation id="13707C88834C9136F4C99047FB74FE83" box="[1185,1262,345,371]" captionStart="FIGURE 1" captionStartId="4.[151,256,1352,1376]" captionTargetBox="[151,1435,370,1315]" captionTargetId="figure@4.[151,1436,349,1329]" captionTargetPageId="4" captionText="FIGURE 1. Habitat of Bembidion aenulum (A) and Bembidion paraenulum (B D). A: USA: Iowa: Webster Co., Des Moines River near Stratford. B. aenulum occurs in the coarse, well-drained sand; the river bank also houses Bembidion integrum Casey, B. chalceum, B. cordatum LeConte, B. grandiceps Hayward, B. confusum, and B. rapidum LeConte. B: USA: New Hampshire: Hillsborough Co., Souhegan River, Merrimack. B. paraenulum is found on the sand island on the right along with B. carinula, B. chalceum, B. inaequale Say, and B. nigrum Say. C: Type locality of Bembidion paraenulum, USA: Virginia: Bedford Co., Big Otter River at route 24. Bembidion (Pseudoperyphus) arenobilis Maddison is also abundant in this habitat. The substrate here consists of coarse sand with some gravel. D: USA: Mississippi: Walthall Co., 5.9 mi S Tylertown, Magees Creek. B. paraenulum occurs on the pure sand banks." httpUri="https://zenodo.org/record/189923/files/figure.png" pageId="15" pageNumber="60">Figs. 1</figureCitation>
BD). In most localities in Virginia and North Carolina it occurs in the company of
<taxonomicName id="4C4B1B8E834C9136F3D5909EFBCEFE69" box="[957,1108,384,409]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="60" phylum="Arthropoda" rank="species" species="arenobilis">
<emphasis id="B93FBC1F834C9136F3D5909EFBCEFE69" box="[957,1108,384,409]" italics="true" pageId="15" pageNumber="60">B. arenobilis</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C3513386834C9136F0AD90B8FD4EFBDB" pageId="15" pageNumber="60" type="materials_examined">
<paragraph id="8BF4600D834C9136F0AD90B8FD4EFBDB" blockId="15.[151,1437,152,1067]" pageId="15" pageNumber="60">
<emphasis id="B93FBC1F834C9136F0AD90B8FE5AFE30" bold="true" box="[197,448,422,448]" pageId="15" pageNumber="60">Material examined</emphasis>
(
<figureCitation id="13707C88834C9136F1BA90B8FDBAFE30" box="[466,544,422,448]" captionStart="FIGURE 9" captionStartId="14.[151,255,1420,1444]" captionTargetBox="[168,1420,332,1379]" captionTargetId="figure@14.[151,1436,313,1397]" captionTargetPageId="14" captionText="FIGURE 9. Geographic distribution of Bembidion aenulum (stars) and B. paraenulum (circles). Black symbols represent specimens identified using DNA sequences and morphological data; gray symbols are those identified only with morphological data. The open circles represent specimens of B. paraenulum, but with labels only indicating the state; these localities are considered questionable as they are so far outside the well-documented range of this species." httpUri="https://zenodo.org/record/189931/files/figure.png" pageId="15" pageNumber="60">Fig. 9</figureCitation>
, circles).
<quantity id="4CB3CDE8834C9136F2F490B8FD76FE30" box="[668,748,422,448]" metricMagnitude="0" metricUnit="m" metricValue="5.6388" pageId="15" pageNumber="60" unit="in" value="222.0">222 in</quantity>
total. In addition to the
<typeStatus id="54F0DEAF834C9136F44890B8FBCCFE30" box="[1056,1110,422,448]" pageId="15" pageNumber="60">type</typeStatus>
material, specimens were examined from the following localities:
<emphasis id="B93FBC1F834C9136F21790D3FB5EFE17" bold="true" box="[639,1220,461,487]" pageId="15" pageNumber="60">
<collectingCountry id="F35C209D834C9136F21790D3FD25FE17" box="[639,703,461,487]" name="United States of America" pageId="15" pageNumber="60">USA</collectingCountry>
:
<collectionCode id="ED5AF8C8834C9136F2A690D3FC8FFE17" box="[718,789,461,487]" httpUri="http://grbio.org/cool/93c9-w2q7" name="University of Newcastle" pageId="15" pageNumber="60">NEW</collectionCode>
<collectionCode id="ED5AF8C8834C9136F37590D3FC40FE17" box="[797,986,461,487]" pageId="15" pageNumber="60">HAMPSHIRE</collectionCode>
: Hillsborough Co.
</emphasis>
, Souhegan River, Merrimack,
<geoCoordinate id="EE7F06CA834C9136F14590EAFE2BFDFE" box="[301,433,500,526]" direction="north" orientation="latitude" pageId="15" pageNumber="60" precision="5" value="42.861">42.8610°N</geoCoordinate>
<geoCoordinate id="EE7F06CA834C9136F1D290EAFDDEFDFE" box="[442,580,500,526]" direction="west" orientation="longitude" pageId="15" pageNumber="60" precision="5" value="-71.4924">71.4924°W</geoCoordinate>
(13).
<emphasis id="B93FBC1F834C9136F2F890EDFBD2FDFD" bold="true" box="[656,1096,499,525]" pageId="15" pageNumber="60">
<collectionCode id="ED5AF8C8834C9136F2F890EDFD67FDFD" box="[656,765,499,525]" pageId="15" pageNumber="60">NORTH</collectionCode>
CAROLINA: Wayne Co.
</emphasis>
, The Cliffs State Park (21).
<emphasis id="B93FBC1F834C9136F0FF9304FE2DFDC4" bold="true" box="[151,439,538,564]" pageId="15" pageNumber="60">
<collectingCountry id="F35C209D834C9136F0FF9304FEB3FDC4" box="[151,297,538,564]" name="Georgia" pageId="15" pageNumber="60">GEORGIA</collectingCountry>
: Clarke Co
</emphasis>
., Whitehall Forest (8);
<emphasis id="B93FBC1F834C9136F2A09304FCD9FDC4" bold="true" box="[712,835,538,564]" pageId="15" pageNumber="60">Ware Co.,</emphasis>
<quantity id="4CB3CDE8834C9136F3249304FC1BFDC4" box="[844,897,538,564]" metricMagnitude="4" metricUnit="m" metricValue="1.2874752" pageId="15" pageNumber="60" unit="mi" value="8.0">8 mi</quantity>
NW Waycross,
<collectingCountry id="F35C209D834C9136F4289304FBFCFDC4" box="[1088,1126,538,564]" name="United States of America" pageId="15" pageNumber="60">US</collectingCountry>
23 &amp;
<collectingCountry id="F35C209D834C9136F4C09304FB48FDC4" box="[1192,1234,538,564]" name="United States of America" pageId="15" pageNumber="60">US</collectingCountry>
1 at Satilla R. (7).
<emphasis id="B93FBC1F834C9136F0FF935EFE20FDAA" bold="true" box="[151,442,576,602]" pageId="15" pageNumber="60">FLORIDA: Liberty Co</emphasis>
.,
<quantity id="4CB3CDE8834C9136F1B8935FFD9CFDAB" box="[464,518,577,603]" metricMagnitude="3" metricUnit="m" metricValue="1.609344" pageId="15" pageNumber="60" unit="mi" value="1.0">1 mi</quantity>
W of Bristol, Apalachicola River (1); Torreya State Park Primitive Campsite, Apalachicola River (3);
<emphasis id="B93FBC1F834C9136F1D09379FDE9FD71" bold="true" box="[440,627,615,641]" pageId="15" pageNumber="60">Santa Rosa Co</emphasis>
., Blackwater River State Park (1);
<emphasis id="B93FBC1F834C9136F47F9379FB29FD71" bold="true" box="[1047,1203,615,641]" pageId="15" pageNumber="60">Suwanee Co</emphasis>
., Suwanee River at route 249 (5); Suwanee River State Park (1).
<emphasis id="B93FBC1F834C9136F2DD9390FC6FFD58" bold="true" box="[693,1013,654,680]" pageId="15" pageNumber="60">ALABAMA: Baldwin Co</emphasis>
., Dixie Landing, Alabama River,
<quantity id="4CB3CDE8834C9136F5E59390FF24FD3F" metricMagnitude="3" metricUnit="m" metricValue="8.0" pageId="15" pageNumber="60" unit="km" value="8.0">8 km</quantity>
W Chrysler (1);
<emphasis id="B93FBC1F834C9136F1FE93ABFDB1FD3F" bold="true" box="[406,555,693,719]" pageId="15" pageNumber="60">Monroe Co</emphasis>
.,
<quantity id="4CB3CDE8834C9136F22D93ABFD0FFD3F" box="[581,661,693,719]" metricMagnitude="3" metricUnit="m" metricValue="5.632704" pageId="15" pageNumber="60" unit="mi" value="3.5">3.5 mi</quantity>
S Uriah, Little River (2).
<emphasis id="B93FBC1F834C9136F3B793ABFABAFD3F" bold="true" box="[991,1312,693,719]" pageId="15" pageNumber="60">MISSISSIPPI: Stone Co</emphasis>
.,
<quantity id="4CB3CDE8834C9136F55193ABFA18FD3F" box="[1337,1410,693,719]" metricMagnitude="4" metricUnit="m" metricValue="1.9312128" pageId="15" pageNumber="60" unit="mi" value="12.0">12 mi</quantity>
E Wiggins on Hwy 26, Black Creek (6); Red Creek W.M.A., near Wiggins,
<geoCoordinate id="EE7F06CA834C9136F44C93C2FB03FD06" box="[1060,1177,732,758]" direction="north" orientation="latitude" pageId="15" pageNumber="60" precision="55" value="30.774">30.774°N</geoCoordinate>
<geoCoordinate id="EE7F06CA834C9136F4CA93C2FA85FD06" box="[1186,1311,732,758]" direction="west" orientation="longitude" pageId="15" pageNumber="60" precision="55" value="-88.914">88.914°W</geoCoordinate>
(21); Red Creek X hwy 15 (16);
<emphasis id="B93FBC1F834C9136F1F5921CFDD1FCEC" bold="true" box="[413,587,770,796]" pageId="15" pageNumber="60">Covington Co</emphasis>
., Okatoma Creek, south of Seminary (5); Okatoma Ck at Lux (2/
<quantity id="4CB3CDE8834C9136F52D921CFAE2FCEC" box="[1349,1400,770,796]" metricMagnitude="3" metricUnit="m" metricValue="4.828032" pageId="15" pageNumber="60" unit="mi" value="3.0">3 mi</quantity>
N. Co. line) (2); Okatoma R. near Lux (1);
<emphasis id="B93FBC1F834C9136F2EA9236FC86FCB2" bold="true" box="[642,796,808,834]" pageId="15" pageNumber="60">Walthall Co</emphasis>
.,
<quantity id="4CB3CDE8834C9136F35D9237FCF7FCB3" box="[821,877,809,835]" metricMagnitude="3" metricUnit="m" metricValue="8.04672" pageId="15" pageNumber="60" unit="mi" value="5.0">5 mi</quantity>
SE Lexie (2);
<emphasis id="B93FBC1F834C9136F44A9236FB02FCB2" bold="true" box="[1058,1176,808,834]" pageId="15" pageNumber="60">Perry Co</emphasis>
., DeSoto N.F., Black Creek at Fairley Bridge Landing,
<geoCoordinate id="EE7F06CA834C9136F249924EFD3BFC9A" box="[545,673,848,874]" direction="north" orientation="latitude" pageId="15" pageNumber="60" precision="5" value="30.9197">30.9197°N</geoCoordinate>
<geoCoordinate id="EE7F06CA834C9136F2C0924EFCB5FC9A" box="[680,815,848,874]" direction="west" orientation="longitude" pageId="15" pageNumber="60" precision="5" value="-88.9661">88.9661°W</geoCoordinate>
(10);
<quantity id="4CB3CDE8834C9136F31E924EFC5AFC9A" box="[886,960,848,874]" metricMagnitude="3" metricUnit="m" metricValue="9.4951296" pageId="15" pageNumber="60" unit="mi" value="5.9">5.9 mi</quantity>
S Tylertown, Magees Creek,
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<geoCoordinate id="EE7F06CA834C9136F0FF9268FE85FC60" box="[151,287,886,912]" direction="west" orientation="longitude" pageId="15" pageNumber="60" precision="5" value="-90.1922">90.1922°W</geoCoordinate>
(2). In addition, there is a specimen in the
<collectionCode id="ED5AF8C8834C9136F34E9268FCFCFC60" LSID="urn:lsid:biocol.org:col:33791" box="[806,870,886,912]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:33791" name="Museum of Comparative Zoology" pageId="15" pageNumber="60">MCZ</collectionCode>
labelled &quot;Mo&quot;, presumably an abbreviation for Missouri, and another in the H.C. Fall Collection at the
<collectionCode id="ED5AF8C8834C9136F34D9283FCFFFC47" LSID="urn:lsid:biocol.org:col:33791" box="[805,869,925,951]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:33791" name="Museum of Comparative Zoology" pageId="15" pageNumber="60">MCZ</collectionCode>
labelled &quot;Wis&quot;, presumably for Wisconsin, both of which belong to this species. Missouri and Wisconsin are both notably outside the otherwise-known range (
<figureCitation id="13707C88834C9136F0F792F4FF72FBF4" box="[159,232,1002,1028]" captionStart="FIGURE 9" captionStartId="14.[151,255,1420,1444]" captionTargetBox="[168,1420,332,1379]" captionTargetId="figure@14.[151,1436,313,1397]" captionTargetPageId="14" captionText="FIGURE 9. Geographic distribution of Bembidion aenulum (stars) and B. paraenulum (circles). Black symbols represent specimens identified using DNA sequences and morphological data; gray symbols are those identified only with morphological data. The open circles represent specimens of B. paraenulum, but with labels only indicating the state; these localities are considered questionable as they are so far outside the well-documented range of this species." httpUri="https://zenodo.org/record/189931/files/figure.png" pageId="15" pageNumber="60">Fig. 9</figureCitation>
); we suspect those specimens are mislabelled. If valid, however, these localities would suggest that
<taxonomicName id="4C4B1B8E834C9136F5EB92F4FF66FBDA" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="60" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834C9136F5EB92F4FF66FBDA" italics="true" pageId="15" pageNumber="60">B. aenulum</emphasis>
</taxonomicName>
and
<taxonomicName id="4C4B1B8E834C9136F15C950FFE79FBDA" box="[308,483,1041,1066]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="60" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F834C9136F15C950FFE79FBDA" box="[308,483,1041,1066]" italics="true" pageId="15" pageNumber="60">B. paraenulum</emphasis>
</taxonomicName>
might be sympatric.
</paragraph>
</subSubSection>
</treatment>
</document>