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<document id="9ED29FE6A827A06C3C8E29456A103FC2" 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="03E2D11B834B9132F0FF9448FD29FE30" 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 aenulum Hayward 1901" docType="treatment" docVersion="10" lastPageNumber="55" masterDocId="FFDBA96383439139F068911EFF9AFFF0" masterDocTitle="A review of the Bembidion (Odontium) aenulum subgroup (Coleoptera: Carabidae), with description of a new species" masterLastPageNumber="61" masterPageNumber="45" pageNumber="53" updateTime="1698593221765" updateUser="plazi">
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<mods:title id="574692B4C46883424A4C4167CDBCC66A">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="046853C804064A59D10AF90C276AA8B9">Maddison, David R.</mods:namePart>
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<treatment id="03E2D11B834B9132F0FF9448FD29FE30" ID-DOI="http://doi.org/10.5281/zenodo.5615779" ID-GBIF-Taxon="119623096" ID-Zenodo-Dep="5615779" LSID="urn:lsid:plazi:treatment:03E2D11B834B9132F0FF9448FD29FE30" httpUri="http://treatment.plazi.org/id/03E2D11B834B9132F0FF9448FD29FE30" lastPageId="11" lastPageNumber="55" pageId="8" pageNumber="53">
<subSubSection id="C3513386834B9131F0FF9448FDAFFA67" pageId="8" pageNumber="53" type="nomenclature">
<paragraph id="8BF4600D834B9131F0FF9448FDC9FA80" blockId="8.[151,595,1366,1431]" box="[151,595,1366,1392]" pageId="8" pageNumber="53">
<heading id="D0BCD761834B9131F0FF9448FDC9FA80" bold="true" box="[151,595,1366,1392]" fontSize="11" level="1" pageId="8" pageNumber="53" reason="1">
<taxonomicName id="4C4B1B8E834B9131F0FF9448FDC9FA80" ID-CoL="68HP5" authority="Hayward, 1901" authorityName="Hayward" authorityYear="1901" box="[151,595,1366,1392]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="53" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834B9131F0FF9448FDC9FA80" bold="true" box="[151,595,1366,1392]" pageId="8" pageNumber="53">
<emphasis id="B93FBC1F834B9131F0FF9448FE17FA80" bold="true" box="[151,397,1366,1392]" italics="true" pageId="8" pageNumber="53">Bembidion aenulum</emphasis>
<bibRefCitation id="EFDA1DFC834B9131F1FD9448FDC9FA80" author="Hayward" box="[405,595,1366,1392]" pageId="8" pageNumber="60" refString="Hayward, R. T. (1901) Synonymical notes on Bembidium and descriptions of new species. Transactions of the American Entomological Society, 27, 156 - 158." type="journal article" year="1901">Hayward, 1901</bibRefCitation>
</emphasis>
</taxonomicName>
</heading>
</paragraph>
<paragraph id="8BF4600D834B9131F0FF9463FDAFFA67" blockId="8.[151,595,1366,1431]" box="[151,565,1405,1431]" pageId="8" pageNumber="53">
(
<figureCitation id="13707C88834B9131F0F79463FF6EFA67" box="[159,244,1405,1431]" 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="8" pageNumber="53">Figs. 1</figureCitation>
A, 4A, 5A, 6A, 7AC, 8A)
</paragraph>
</subSubSection>
<subSubSection id="C3513386834B9131F0FF94D4FE05F87E" pageId="8" pageNumber="53" type="materials_examined">
<paragraph id="8BF4600D834B9131F0FF94D4FD0EF9C2" blockId="8.[151,1437,1482,2011]" pageId="8" pageNumber="53">
<emphasis id="B93FBC1F834B9131F0FF94D4FED1FA14" bold="true" box="[151,331,1482,1508]" pageId="8" pageNumber="53">
<typeStatus id="54F0DEAF834B9131F0FF94D4FF4EFA14" box="[151,212,1482,1508]" pageId="8" pageNumber="53">Type</typeStatus>
material
</emphasis>
.
<typeStatus id="54F0DEAF834B9131F13594D4FE54FA14" box="[349,462,1482,1508]" pageId="8" pageNumber="53" type="holotype">Holotype</typeStatus>
female (in
<collectionCode id="ED5AF8C8834B9131F23594D4FD39FA14" LSID="urn:lsid:biocol.org:col:33791" box="[605,675,1482,1508]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:33791" name="Museum of Comparative Zoology" pageId="8" pageNumber="53">MCZ</collectionCode>
), examined, labelled: &quot;Cedar Co. Iowa,
<date id="FFF546CD834B9131F4E794D4FB5DFA14" box="[1167,1223,1482,1508]" pageId="8" pageNumber="53" value="1931-05">V.31</date>
['Iowa' machineprinted, remainder handwritten] / Roland Hayward Coll. /
<typeStatus id="54F0DEAF834B9131F30A94EFFC04F9FB" box="[866,926,1521,1547]" pageId="8" pageNumber="53">Type</typeStatus>
2006 ['2006' handwritten, on red paper] / Bembidium
<taxonomicName id="4C4B1B8E834B9131F1419706FE7BF9C2" authority="Hayw." authorityName="Hayw." box="[297,481,1560,1586]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="53" phylum="Arthropoda" rank="species" species="aenulum">aenulum Hayw.</taxonomicName>
[handwritten]&quot;
</paragraph>
<paragraph id="8BF4600D834B9131F0AD9720FE05F87E" blockId="8.[151,1437,1482,2011]" pageId="8" pageNumber="53">
<emphasis id="B93FBC1F834B9131F0AD9720FE47F9A8" bold="true" box="[197,477,1598,1624]" pageId="8" pageNumber="53">Nomenclatorial notes</emphasis>
. The female
<typeStatus id="54F0DEAF834B9131F2EB9720FD75F9A8" box="[643,751,1598,1624]" pageId="8" pageNumber="53" type="holotype">holotype</typeStatus>
of
<taxonomicName id="4C4B1B8E834B9131F3729721FBBAF9A8" authority="Hayward" authorityName="Hayward" box="[794,1056,1598,1624]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="53" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834B9131F3729721FC3DF9A8" box="[794,935,1599,1624]" italics="true" pageId="8" pageNumber="53">B. aenulum</emphasis>
Hayward
</taxonomicName>
has all the external diagnostic character states of this western species, including the deeper microsculpture, less angulate humeral angle, distinctive prothorax shape, and wider mentum tooth. While identification of the
<typeStatus id="54F0DEAF834B9131F4269792FB2FF956" box="[1102,1205,1676,1702]" pageId="8" pageNumber="53" type="holotype">holotype</typeStatus>
as a member of the western species would be more certain if the specimen were a male, or was sequenceable, the seven specimens whose
<collectionCode id="ED5AF8C8834B9131F10797C7FE35F903" LSID="urn:lsid:biocol.org:col:15536" box="[367,431,1753,1779]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15536" name="Department of Natural Resources, Environment, The Arts and Sport" pageId="8" pageNumber="53">DNA</collectionCode>
were sequenced from the
<typeStatus id="54F0DEAF834B9131F28497C7FC85F903" box="[748,799,1753,1779]" pageId="8" pageNumber="53">type</typeStatus>
locality, Cedar County, Iowa, and neighboring Jones County all belong to this species. In addition, our 2003 collections from these two counties yielded 49 additional specimens, all of which show the morphological traits of the western species. Thus, it appears the eastern species does not occur at the
<typeStatus id="54F0DEAF834B9131F2249653FDE5F897" box="[588,639,1869,1895]" pageId="8" pageNumber="53">type</typeStatus>
locality, providing further evidence of membership of the
<typeStatus id="54F0DEAF834B9131F55D9653FA06F897" box="[1333,1436,1869,1895]" pageId="8" pageNumber="53" type="holotype">holotype</typeStatus>
in the western species.
</paragraph>
</subSubSection>
<subSubSection id="C3513386834B9133F0AD9684FCB7F918" lastPageId="10" lastPageNumber="55" pageId="8" pageNumber="53" type="diagnosis">
<paragraph id="8BF4600D834B9130F0AD9684FC14FF0F" blockId="8.[151,1437,1482,2011]" lastBlockId="9.[151,1436,152,255]" lastPageId="9" lastPageNumber="54" pageId="8" pageNumber="53">
<emphasis id="B93FBC1F834B9131F0AD9684FEDAF844" bold="true" box="[197,320,1946,1972]" pageId="8" pageNumber="53">Diagnosis</emphasis>
. The smallest species of
<taxonomicName id="4C4B1B8E834B9131F2189685FD7DF844" box="[624,743,1947,1972]" class="Insecta" family="Carabidae" genus="Odontium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="53" phylum="Arthropoda" rank="genus">
<emphasis id="B93FBC1F834B9131F2189685FD7DF844" box="[624,743,1947,1972]" italics="true" pageId="8" pageNumber="53">Odontium</emphasis>
</taxonomicName>
. Pronotum with hind angles wider than front angles, thus with a trapezoidal pronotum; sides straighter than
<taxonomicName id="4C4B1B8E834B9131F34D96DFFC7EF82A" box="[805,996,1985,2010]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="53" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F834B9131F34D96DFFC7EF82A" box="[805,996,1985,2010]" italics="true" pageId="8" pageNumber="53">B. paraenulum</emphasis>
</taxonomicName>
, and with less evident laterobasal depressions (
<figureCitation id="13707C88834A9130F1509186FE1BFF42" box="[312,385,152,178]" 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="9" pageNumber="54">Fig. 5</figureCitation>
A). Humeral sulcus of elytron nearly parallel to basal margin of pronotum (
<figureCitation id="13707C88834A9130F5439186FAEEFF42" box="[1323,1396,152,178]" 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="9" pageNumber="54">Fig. 5</figureCitation>
A). Microsculpture coarse, notably granulate in places, especially at the back of the pronotum. Mentum tooth wider at base, more trapezoidal than in
<taxonomicName id="4C4B1B8E834A9130F20D91FBFC8EFF0E" box="[613,788,229,254]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="54" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F834A9130F20D91FBFC8EFF0E" box="[613,788,229,254]" italics="true" pageId="9" pageNumber="54">B. paraenulum</emphasis>
</taxonomicName>
(
<figureCitation id="13707C88834A9130F34B91FBFCF3FF0F" box="[803,873,229,255]" 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="9" pageNumber="54">Fig. 6</figureCitation>
A).
</paragraph>
<caption id="DF343085834A9130F0FF966DFAABF803" httpUri="https://zenodo.org/record/189926/files/figure.png" pageId="9" pageNumber="54" targetBox="[151,1436,294,1898]" targetPageId="9">
<paragraph id="8BF4600D834A9130F0FF966DFAABF803" blockId="9.[151,1436,1907,2035]" pageId="9" pageNumber="54">
<emphasis id="B93FBC1F834A9130F0FF966DFE89F87B" bold="true" box="[151,275,1907,1931]" pageId="9" pageNumber="54">FIGURE 4</emphasis>
. Male habitus. A:
<taxonomicName id="4C4B1B8E834A9130F1B4966AFD29F87B" box="[476,691,1908,1931]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="54" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834A9130F1B4966AFD29F87B" box="[476,691,1908,1931]" italics="true" pageId="9" pageNumber="54">Bembidion aenulum</emphasis>
</taxonomicName>
, DRM voucher V100128, Iowa: Des Moines River near Stratford. B:
<taxonomicName id="4C4B1B8E834A9130F0FF9689FE0EF85E" box="[151,404,1943,1966]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="54" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F834A9130F0FF9689FE0EF85E" box="[151,404,1943,1966]" italics="true" pageId="9" pageNumber="54">Bembidion paraenulum</emphasis>
</taxonomicName>
, DRM voucher V100078, New Hampshire: Merrimack. C: Specimens of each species showing typical size difference:
<taxonomicName id="4C4B1B8E834A9130F1FB96A4FDF1F821" box="[403,619,1978,2001]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="54" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F834A9130F1FB96A4FDF1F821" box="[403,619,1978,2001]" italics="true" pageId="9" pageNumber="54">Bembidion aenulum</emphasis>
</taxonomicName>
(left), DRM voucher V100129, Iowa: Des Moines River near Stratford and
<taxonomicName id="4C4B1B8E834A9130F0FF96C2FEAEF803" box="[151,308,2012,2035]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="54" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F834A9130F0FF96C2FEAEF803" box="[151,308,2012,2035]" italics="true" pageId="9" pageNumber="54">B. paraenulum</emphasis>
</taxonomicName>
(right), DRM voucher V100130, USA: North Carolina: Rockingham Co., Dan River, Madison.
</paragraph>
</caption>
<caption id="DF34308583499133F0FF942AFD09F9EB" httpUri="https://zenodo.org/record/189927/files/figure.png" pageId="10" pageNumber="55" targetBox="[151,1435,194,1294]" targetPageId="10">
<paragraph id="8BF4600D83499133F0FF942AFD09F9EB" blockId="10.[151,1436,1332,1563]" pageId="10" pageNumber="55">
<emphasis id="B93FBC1F83499133F0FF942AFE8CFABC" bold="true" box="[151,278,1332,1356]" pageId="10" pageNumber="55">FIGURE 5</emphasis>
. Pronota and elytral shoulder. A:
<taxonomicName id="4C4B1B8E83499133F2E4942BFCFDFABC" box="[652,871,1333,1356]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="55" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F83499133F2E4942BFCFDFABC" box="[652,871,1333,1356]" italics="true" pageId="10" pageNumber="55">Bembidion aenulum</emphasis>
</taxonomicName>
, DRM voucher V100515, Iowa: Des Moines River near Stratford. B: Comparison of shoulder area and hind pronotal angle of
<taxonomicName id="4C4B1B8E83499133F3DB9449FBB6FA9E" box="[947,1068,1367,1390]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="55" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F83499133F3DB9449FBB6FA9E" box="[947,1068,1367,1390]" italics="true" pageId="10" pageNumber="55">B. aenulum</emphasis>
</taxonomicName>
and
<taxonomicName id="4C4B1B8E83499133F4089449FB64FA9E" box="[1120,1278,1367,1390]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="55" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F83499133F4089449FB64FA9E" box="[1120,1278,1367,1390]" italics="true" pageId="10" pageNumber="55">B. paraenulum</emphasis>
</taxonomicName>
. Note how the humeral sulcus is nearly parallel to the hind margin of the pronotum in
<taxonomicName id="4C4B1B8E83499133F3E59464FB9DFA61" box="[909,1031,1402,1425]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="55" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F83499133F3E59464FB9DFA61" box="[909,1031,1402,1425]" italics="true" pageId="10" pageNumber="55">B. aenulum</emphasis>
</taxonomicName>
, whereas in
<taxonomicName id="4C4B1B8E83499133F4E59464FAB6FA61" box="[1165,1324,1402,1425]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="55" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F83499133F4E59464FAB6FA61" box="[1165,1324,1402,1425]" italics="true" pageId="10" pageNumber="55">B. paraenulum</emphasis>
</taxonomicName>
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
<taxonomicName id="4C4B1B8E83499133F1CE94A1FDDDFA26" box="[422,583,1471,1494]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="55" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F83499133F1CE94A1FDDDFA26" box="[422,583,1471,1494]" italics="true" pageId="10" pageNumber="55">B. paraenulum</emphasis>
</taxonomicName>
than
<taxonomicName id="4C4B1B8E83499133F2ED94A1FC9BFA26" box="[645,769,1471,1494]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="55" phylum="Arthropoda" rank="species" species="aenulum">
<emphasis id="B93FBC1F83499133F2ED94A1FC9BFA26" box="[645,769,1471,1494]" italics="true" pageId="10" pageNumber="55">B. aenulum</emphasis>
</taxonomicName>
. C:
<taxonomicName id="4C4B1B8E83499133F35894A1FC4AFA26" box="[816,976,1471,1494]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="55" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F83499133F35894A1FC4AFA26" box="[816,976,1471,1494]" italics="true" pageId="10" pageNumber="55">B. paraenulum</emphasis>
</taxonomicName>
holotype, Virginia: Big Otter River. D:
<taxonomicName id="4C4B1B8E83499133F5ED94A1FE82FA09" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="55" phylum="Arthropoda" rank="species" species="paraenulum">
<emphasis id="B93FBC1F83499133F5ED94A1FE82FA09" italics="true" pageId="10" pageNumber="55">B. paraenulum</emphasis>
</taxonomicName>
, DRM voucher V100126, Mississippi: Perry Co., DeSoto N.F., Black Creek at Fairley Bridge Landing. All photographs to same scale; scale bar is 0.5 mm.
</paragraph>
</caption>
<paragraph id="8BF4600D83499133F0AD9747FCB7F918" blockId="10.[151,1437,1625,2000]" pageId="10" pageNumber="55">
<emphasis id="B93FBC1F83499133F0AD9747FD92F983" bold="true" box="[197,520,1625,1651]" pageId="10" pageNumber="55">Additional characteristics</emphasis>
. ABL
<quantity id="4CB3CDE883499133F2319744FC9AF984" box="[601,768,1626,1652]" metricMagnitude="-3" metricUnit="m" metricValue="3.8" metricValueMax="4.1" metricValueMin="3.5" pageId="10" pageNumber="55" unit="mm" value="3.8" valueMax="4.1" valueMin="3.5">3.5 to 4.1 mm</quantity>
, with most specimens less than 4.0 mm in length. Dorsal surface of body with metallic reflection, in most specimens golden or coppery. Appendages rufo-testaceous, with terminal two articles of maxillary palps, antennomeres 511, tarsi, and ends of tibia infuscated. Chromosome number is 2n = 22+XY (
<bibRefCitation id="EFDA1DFC83499133F23397D0FCBAF918" author="Maddison" box="[603,800,1742,1768]" pageId="10" pageNumber="60" refString="Maddison, D. R. (1985) Chromosomal diversity and evolution in the ground beetle genus Bembidion and related taxa (Coleoptera: Carabidae: Trechitae). Genetica, 66, 93 - 114." type="journal article" year="1985">Maddison, 1985</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C351338683499133F0AD97EAFDF6F873" pageId="10" pageNumber="55" type="description">
<paragraph id="8BF4600D83499133F0AD97EAFDF5F8FF" blockId="10.[151,1437,1625,2000]" box="[197,623,1780,1807]" pageId="10" pageNumber="55">
<emphasis id="B93FBC1F83499133F0AD97EAFE4BF8FE" bold="true" box="[197,465,1780,1806]" pageId="10" pageNumber="55">Geographic variation</emphasis>
. None noted.
</paragraph>
<paragraph id="8BF4600D83499133F0AD9605FDF6F873" blockId="10.[151,1437,1625,2000]" pageId="10" pageNumber="55">
<emphasis id="B93FBC1F83499133F0AD9605FEBFF8C5" bold="true" box="[197,293,1819,1845]" pageId="10" pageNumber="55">Habitat</emphasis>
. We have found this species at eight localities in Iowa and Nebraska on pure, often coarse sand of rivers (
<figureCitation id="13707C8883499133F087965CFEDBF8AC" box="[239,321,1858,1884]" 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="10" pageNumber="55">Figs. 1</figureCitation>
A), in most localities in the company of
<taxonomicName id="4C4B1B8E83499133F355965CFBA5F8AB" box="[829,1087,1858,1883]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="55" phylum="Arthropoda" rank="species" species="confusum">
<emphasis id="B93FBC1F83499133F355965CFBA5F8AB" box="[829,1087,1858,1883]" italics="true" pageId="10" pageNumber="55">Bembidion confusum</emphasis>
</taxonomicName>
, and less frequently with
<taxonomicName id="4C4B1B8E83499133F5EA965CFE02F873" authority="Hayward" authorityName="Hayward" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="55" phylum="Arthropoda" rank="species" species="robusticolle">
<emphasis id="B93FBC1F83499133F5EA965CFEB8F872" italics="true" pageId="10" pageNumber="55">B. robusticolle</emphasis>
Hayward
</taxonomicName>
,
<taxonomicName id="4C4B1B8E83499133F1CB9677FDA9F872" box="[419,563,1897,1922]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="55" phylum="Arthropoda" rank="species" species="chalceum">
<emphasis id="B93FBC1F83499133F1CB9677FDA9F872" box="[419,563,1897,1922]" italics="true" pageId="10" pageNumber="55">B. chalceum</emphasis>
</taxonomicName>
, etc.
</paragraph>
</subSubSection>
<subSubSection id="C351338683499132F0AD9691FD29FE30" lastPageId="11" lastPageNumber="56" pageId="10" pageNumber="55" type="materials_examined">
<paragraph id="8BF4600D83499132F0AD9691FD29FE30" blockId="10.[151,1437,1625,2000]" lastBlockId="11.[151,1437,151,448]" lastPageId="11" lastPageNumber="56" pageId="10" pageNumber="55">
<emphasis id="B93FBC1F83499133F0AD9691FE2EF859" bold="true" box="[197,436,1935,1961]" pageId="10" pageNumber="55">Material examined</emphasis>
(
<figureCitation id="13707C8883499133F1AD9691FD95F859" box="[453,527,1935,1961]" 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="10" pageNumber="55">Fig. 9</figureCitation>
, stars). In addition to the
<typeStatus id="54F0DEAF83499133F32A9691FC37F859" box="[834,941,1935,1961]" pageId="10" pageNumber="55" type="holotype">holotype</typeStatus>
,
<specimenCount id="9D4DAB8483499133F3D19691FBF3F859" box="[953,1129,1935,1961]" pageId="10" pageNumber="55" type="generic">148 specimens</specimenCount>
were examined, from the following localities (number of specimens seen indicated in parentheses after the locality):
<emphasis id="B93FBC1F83499132F4BC96A8FF26FF41" bold="true" lastPageId="11" lastPageNumber="56" pageId="10" pageNumber="55">
<collectionCode id="ED5AF8C883499133F4BC96A8FAB1F820" box="[1236,1323,1974,2000]" pageId="10" pageNumber="55">IOWA</collectionCode>
: Clayton Co
</emphasis>
.,
<collectingCountry id="F35C209D83489132F0BC9186FEB0FF42" box="[212,298,152,178]" name="Turkey" pageId="11" pageNumber="56">Turkey</collectingCountry>
River at Osterdock,
<quantity id="4CB3CDE883489132F2439186FDE8FF42" box="[555,626,152,178]" metricMagnitude="2" metricUnit="m" metricValue="1.95" pageId="11" pageNumber="56" unit="m" value="195.0">195m</quantity>
,
<geoCoordinate id="EE7F06CA83489132F2E89186FC9EFF42" box="[640,772,152,178]" direction="north" orientation="latitude" pageId="11" pageNumber="56" precision="5" value="42.7323">42.7323°N</geoCoordinate>
<geoCoordinate id="EE7F06CA83489132F3659186FC02FF42" box="[781,920,152,178]" direction="west" orientation="longitude" pageId="11" pageNumber="56" precision="5" value="-91.1602">91.1602°W</geoCoordinate>
(1);
<emphasis id="B93FBC1F83489132F3BD9189FBD1FF41" bold="true" box="[981,1099,151,177]" pageId="11" pageNumber="56">Jones Co</emphasis>
., Wapsipinicon River W of Oxford Junction,
<quantity id="4CB3CDE883489132F10C91A0FE33FF28" box="[356,425,190,216]" metricMagnitude="2" metricUnit="m" metricValue="2.2" pageId="11" pageNumber="56" unit="m" value="220.0">220m</quantity>
,
<geoCoordinate id="EE7F06CA83489132F1DD91A0FDAEFF28" box="[437,564,190,216]" direction="north" orientation="latitude" pageId="11" pageNumber="56" precision="5" value="41.9913">41.9913°N</geoCoordinate>
<geoCoordinate id="EE7F06CA83489132F25491A0FD59FF28" box="[572,707,190,216]" direction="west" orientation="longitude" pageId="11" pageNumber="56" precision="5" value="-91.0067">91.0067°W</geoCoordinate>
(29);
<emphasis id="B93FBC1F83489132F36191A0FC18FF28" bold="true" box="[777,898,190,216]" pageId="11" pageNumber="56">Cedar Co</emphasis>
., Cedar River at Rochester,
<quantity id="4CB3CDE883489132F4A391A0FA8AFF28" box="[1227,1296,190,216]" metricMagnitude="2" metricUnit="m" metricValue="1.95" pageId="11" pageNumber="56" unit="m" value="195.0">195m</quantity>
,
<geoCoordinate id="EE7F06CA83489132F57491A0FA01FF28" box="[1308,1435,190,216]" direction="north" orientation="latitude" pageId="11" pageNumber="56" precision="5" value="41.6715">41.6715°N</geoCoordinate>
<geoCoordinate id="EE7F06CA83489132F0FF91FBFEBFFF0F" box="[151,293,229,255]" direction="west" orientation="longitude" pageId="11" pageNumber="56" precision="5" value="-91.1599">91.1599°W</geoCoordinate>
(20); Cedar River at route
<date id="FFF546CD83489132F2E991FBFD20FF0F" box="[641,698,229,255]" pageId="11" pageNumber="56" value="1940-10">
X
<quantity id="4CB3CDE883489132F2F191FBFD20FF0F" box="[665,698,229,255]" metricMagnitude="4" metricUnit="m" metricValue="4.0198" pageId="11" pageNumber="56" unit="m" value="40198.0">40</quantity>
</date>
, 198m,
<geoCoordinate id="EE7F06CA83489132F37791FBFC02FF0F" box="[799,920,229,255]" direction="north" orientation="latitude" pageId="11" pageNumber="56" precision="55" value="41.735">41.735°N</geoCoordinate>
<geoCoordinate id="EE7F06CA83489132F3C991FBFBBAFF0F" box="[929,1056,229,255]" direction="west" orientation="longitude" pageId="11" pageNumber="56" precision="55" value="-91.247">91.247°W</geoCoordinate>
(7);
<emphasis id="B93FBC1F83489132F40891FBFB6AFF0F" bold="true" box="[1120,1264,229,255]" pageId="11" pageNumber="56">Hardin Co</emphasis>
., Iowa River, Steamboat Rock,
<quantity id="4CB3CDE883489132F1039012FE2BFED6" box="[363,433,268,294]" metricMagnitude="2" metricUnit="m" metricValue="2.9" pageId="11" pageNumber="56" unit="m" value="290.0">290m</quantity>
,
<geoCoordinate id="EE7F06CA83489132F1D69012FDDAFED6" box="[446,576,268,294]" direction="north" orientation="latitude" pageId="11" pageNumber="56" precision="5" value="42.4079">42.4079°N</geoCoordinate>
<geoCoordinate id="EE7F06CA83489132F2209012FD4BFED6" box="[584,721,268,294]" direction="west" orientation="longitude" pageId="11" pageNumber="56" precision="5" value="-93.0722">93.0722°W</geoCoordinate>
(6);
<emphasis id="B93FBC1F83489132F3649015FC3AFED5" bold="true" box="[780,928,267,293]" pageId="11" pageNumber="56">Webster Co</emphasis>
., Des Moines River near Stratford,
<quantity id="4CB3CDE883489132F5399012FA02FED6" box="[1361,1432,268,294]" metricMagnitude="2" metricUnit="m" metricValue="2.75" pageId="11" pageNumber="56" unit="m" value="275.0">275m</quantity>
,
<geoCoordinate id="EE7F06CA83489132F0FF902CFE82FEBC" box="[151,280,306,332]" direction="north" orientation="latitude" pageId="11" pageNumber="56" precision="5" value="42.3101">42.3101°N</geoCoordinate>
<geoCoordinate id="EE7F06CA83489132F177902CFE3CFEBC" box="[287,422,306,332]" direction="west" orientation="longitude" pageId="11" pageNumber="56" precision="5" value="-93.9371">93.9371°W</geoCoordinate>
(37);
<emphasis id="B93FBC1F83489132F198902CFDC2FEBC" bold="true" box="[496,600,306,332]" pageId="11" pageNumber="56">Linn Co</emphasis>
., 3.0 mi N Viola, 4291Cd,
<quantity id="4CB3CDE883489132F3F4902CFC45FEBC" box="[924,991,306,332]" metricMagnitude="2" metricUnit="m" metricValue="2.6" pageId="11" pageNumber="56" unit="m" value="260.0">260m</quantity>
(1).
<emphasis id="B93FBC1F83489132F471902CFADAFEBC" bold="true" box="[1049,1344,306,332]" pageId="11" pageNumber="56">NEBRASKA: Keith Co</emphasis>
., South Platte River at Brule,
<geoCoordinate id="EE7F06CA83489132F1CE9047FDB6FE83" box="[422,556,345,371]" direction="north" orientation="latitude" pageId="11" pageNumber="56" precision="5" value="41.0858">41.0858°N</geoCoordinate>
<geoCoordinate id="EE7F06CA83489132F25D9047FD4AFE83" box="[565,720,345,371]" direction="west" orientation="longitude" pageId="11" pageNumber="56" precision="5" value="-101.887">101.8870°W</geoCoordinate>
(43);
<emphasis id="B93FBC1F83489132F3759046FC12FE82" bold="true" box="[797,904,344,370]" pageId="11" pageNumber="56">Cass Co</emphasis>
., Louisville, Platte River,
<quantity id="4CB3CDE883489132F4B89047FA82FE83" box="[1232,1304,345,371]" metricMagnitude="2" metricUnit="m" metricValue="3.1" pageId="11" pageNumber="56" unit="m" value="310.0">310m</quantity>
,
<geoCoordinate id="EE7F06CA83489132F54E9047FA07FE83" box="[1318,1437,345,371]" direction="north" orientation="latitude" pageId="11" pageNumber="56" precision="55" value="41.003">41.003°N</geoCoordinate>
<geoCoordinate id="EE7F06CA83489132F0FF909EFE8FFE6A" box="[151,277,384,410]" direction="west" orientation="longitude" pageId="11" pageNumber="56" precision="55" value="-96.182">96.182°W</geoCoordinate>
(2);
<emphasis id="B93FBC1F83489132F13D9061FE6BFE69" bold="true" box="[341,497,383,409]" pageId="11" pageNumber="56">Merrick Co</emphasis>
., Central City, Hord Lake (1);
<emphasis id="B93FBC1F83489132F3139061FC6DFE69" bold="true" box="[891,1015,383,409]" pageId="11" pageNumber="56">Platte Co</emphasis>
., Columbus (banks Loup R.) (1).
<emphasis id="B93FBC1F83489132F0FF90B8FE19FE30" bold="true" box="[151,387,422,448]" pageId="11" pageNumber="56">
<collectionCode id="ED5AF8C883489132F0FF90B8FF64FE30" box="[151,254,422,448]" pageId="11" pageNumber="56">SOUTH</collectionCode>
DAKOTA
</emphasis>
:
<emphasis id="B93FBC1F83489132F1FB90B8FD9DFE30" bold="true" box="[403,519,422,448]" pageId="11" pageNumber="56">Spink Co</emphasis>
., Redfield (1).
</paragraph>
</subSubSection>
</treatment>
</document>