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<document id="1C3FB9D02413B38DB7E1B2B86BB5291F" ID-CLB-Dataset="299407" ID-DOI="10.5281/zenodo.11067242" ID-GBIF-Dataset="36a87cae-beb8-49c4-a561-a9e9a73afdf1" ID-ISSN="0068-547X" ID-Zenodo-Dep="11067242" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="karina" IM.materialsCitations_approvedBy="karina" IM.metadata_approvedBy="felipe" IM.tables_approvedBy="karina" IM.taxonomicNames_approvedBy="karina" IM.treatments_approvedBy="karina" checkinTime="1714052119798" checkinUser="felipe" docAuthor="Maddison, David R." docDate="2023" docId="03C82573441CA56C889D198E0ED0FE42" docLanguage="en" docName="ProcCalifAcadSci.67.19.433-450.pdf" docOrigin="Proceedings of the California Academy of Sciences 67 (19)" docStyle="DocumentStyle:E411DA009AF5E7F9A7E023431E2B7D5B.2:ProcCalifAcadSci.2015-.journal_article" docStyleId="E411DA009AF5E7F9A7E023431E2B7D5B" docStyleName="ProcCalifAcadSci.2015-.journal_article" docStyleVersion="2" docTitle="(Geocosmius) Maddison 2023, new subgenus" docType="treatment" docVersion="5" lastPageNumber="443" masterDocId="FFF15D0B4414A566892F1D770F4AFFB4" masterDocTitle="A New Subgenus of Bembidion Latreille from México and Guatemala, with Descriptions of Two New Species (Coleoptera: Carabidae)" masterLastPageNumber="450" masterPageNumber="433" pageNumber="441" updateTime="1719951180206" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:title id="81B8F5851B81910375523D1A91B70C23">A New Subgenus of Bembidion Latreille from México and Guatemala, with Descriptions of Two New Species (Coleoptera: Carabidae)</mods:title>
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<mods:namePart id="CB79E426A988B7693036F59128083D9C">Maddison, David R.</mods:namePart>
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<mods:title id="4301C3C5FA9391B4F5FD3571CF081DF7">Proceedings of the California Academy of Sciences</mods:title>
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<mods:date id="F26A17AE0753B9C9E5FB9694729FFC00">2023</mods:date>
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<treatment id="03C82573441CA56C889D198E0ED0FE42" ID-DOI="http://doi.org/10.5281/zenodo.12627925" ID-GBIF-Taxon="234411861" ID-Zenodo-Dep="12627925" LSID="urn:lsid:plazi:treatment:03C82573441CA56C889D198E0ED0FE42" httpUri="http://treatment.plazi.org/id/03C82573441CA56C889D198E0ED0FE42" lastPageId="10" lastPageNumber="443" pageId="8" pageNumber="441">
<subSubSection id="C37BC7EE441CA56E889D198E0C17FAA4" box="[434,861,1273,1297]" pageId="8" pageNumber="441" type="nomenclature">
<paragraph id="8BDE9465441CA56E889D198E0C17FAA4" blockId="8.[434,861,1273,1297]" box="[434,861,1273,1297]" pageId="8" pageNumber="441">
<heading id="D0962309441CA56E889D198E0C17FAA4" box="[434,861,1273,1297]" centered="true" fontSize="10" level="2" pageId="8" pageNumber="441" reason="4">
<emphasis id="B9154877441CA56E889D198E0C17FAA4" bold="true" box="[434,861,1273,1297]" pageId="8" pageNumber="441">
<taxonomicName id="4C61EFE6441CA56E889D198E0DF9FAA4" authority="Maddison, 2023" authorityName="Maddison" authorityYear="2023" box="[434,691,1273,1297]" kingdom="Animalia" pageId="8" pageNumber="443" rank="subGenus" status="subgen. nov." subGenus="Geocosmius">Geocosmius Maddison</taxonomicName>
,
<taxonomicNameLabel id="A226F50C441CA56E8B90198E0C17FAA4" box="[703,861,1273,1296]" pageId="8" pageNumber="441" rank="subGenus">new subgenus</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C37BC7EE441CA56E888618530C2DFA8D" box="[425,871,1315,1337]" pageId="8" pageNumber="441" type="type_taxon">
<paragraph id="8BDE9465441CA56E888618530C2DFA8D" blockId="8.[425,871,1315,1337]" box="[425,871,1315,1337]" pageId="8" pageNumber="441">
<typeStatus id="54DA2AC7441CA56E888618530E93FA8D" box="[425,473,1316,1337]" pageId="8" pageNumber="441">Type</typeStatus>
species
<taxonomicName id="4C61EFE6441CA56E8B0318540C2DFA8D" authority="Erwin, 1982" authorityName="Erwin" authorityYear="1982" box="[556,871,1315,1337]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="441" phylum="Arthropoda" rank="species" species="nahuala">
<emphasis id="B9154877441CA56E8B0318540DA3FA8D" box="[556,745,1315,1337]" italics="true" pageId="8" pageNumber="441">Bembidion nahuala</emphasis>
<bibRefCitation id="EFF0E994441CA56E8BC018540C2DFA8D" author="ERWIN, T. L." box="[751,871,1315,1337]" pageId="8" pageNumber="441" pagination="455 - 496" refId="ref7573" refString="ERWIN, T. L. 1982. Small terrestrial ground-beetles of Central America (Carabidae: Bembidiina and Anillina). Proceedings of the California Academy of Sciences, ser. 4, 42: 455 - 496." type="journal article" year="1982">Erwin, 1982</bibRefCitation>
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="C37BC7EE441CA56E898418220F8AF999" pageId="8" pageNumber="441" type="etymology">
<paragraph id="8BDE9465441CA56E898418220F8AF999" blockId="8.[123,1174,1364,1708]" pageId="8" pageNumber="441">
<emphasis id="B9154877441CA56E898418220EC7FAD8" bold="true" box="[171,397,1365,1388]" pageId="8" pageNumber="441">Derivation of name.</emphasis>
The first portion of the name, “Geo”, consists of the first three letters of George E. Ball’s given name, and is included to honor George’s long-term commitment to discovering and documenting the biodiversity of
<collectingCountry id="F376D4F5441CA56E8B6118E30DE8FA18" box="[590,674,1428,1452]" name="Mexico" pageId="8" pageNumber="441">México</collectingCountry>
, and his role in collecting most known specimens of this subgenus. In addition, “Geo” is a Greek prefix referring to the Earth, and evokes the interesting relative geographic distributions of
<taxonomicName id="4C61EFE6441CA56E8B5618A20DB1FA58" authorityName="Maddison" authorityYear="2023" box="[633,763,1493,1516]" pageId="8" pageNumber="443" rank="subGenus" subGenus="Geocosmius">
<emphasis id="B9154877441CA56E8B5618A20DB1FA58" box="[633,763,1493,1516]" italics="true" pageId="8" pageNumber="441">Geocosmius</emphasis>
</taxonomicName>
and its sister group,
<taxonomicName id="4C61EFE6441CA56E8ACC18A20B16FA58" authorityName="Casey" authorityYear="1918" box="[995,1116,1493,1516]" class="Insecta" family="Carabidae" genus="Liocosmius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="441" phylum="Arthropoda" rank="genus">
<emphasis id="B9154877441CA56E8ACC18A20B16FA58" box="[995,1116,1493,1516]" italics="true" pageId="8" pageNumber="441">Liocosmius</emphasis>
</taxonomicName>
. The last portion of the name, “cosmius”, was chosen to provide a linguistic connection to its sister group.
</paragraph>
</subSubSection>
<subSubSection id="C37BC7EE441CA56F89841B420C2DFE87" lastPageId="9" lastPageNumber="442" pageId="8" pageNumber="441" type="description">
<paragraph id="8BDE9465441CA56F89841B420C2DFE87" blockId="8.[123,1174,1364,1708]" lastBlockId="9.[123,1174,219,1725]" lastPageId="9" lastPageNumber="442" pageId="8" pageNumber="441">
<emphasis id="B9154877441CA56E89841B420E46F9F8" bold="true" box="[171,268,1589,1612]" pageId="8" pageNumber="441">Habitus.</emphasis>
Beetles of the subgenus
<taxonomicName id="4C61EFE6441CA56E8B0F1B420DE8F9F8" authorityName="Maddison" authorityYear="2023" box="[544,674,1589,1612]" pageId="8" pageNumber="441" rank="subGenus" subGenus="Geocosmius">
<emphasis id="B9154877441CA56E8B0F1B420DE8F9F8" box="[544,674,1589,1612]" italics="true" pageId="8" pageNumber="441">Geocosmius</emphasis>
</taxonomicName>
have the general appearance of typical members of non-riparian cloud forest
<taxonomicName id="4C61EFE6441CA56E88CD1B230D1CF9D8" authorityName="Latreille" authorityYear="1802" box="[482,598,1620,1644]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="441" phylum="Arthropoda" rank="genus">
<emphasis id="B9154877441CA56E88CD1B230D1CF9D8" box="[482,598,1620,1644]" italics="true" pageId="8" pageNumber="441">Bembidion</emphasis>
</taxonomicName>
; that is, they are small, dark, unspotted, and convex; most specimens lack hind wings and have rounded shoulders. A similar form has evolved multiple times within
<taxonomicName id="4C61EFE6441CA56E88261BE30E37F918" authorityName="Latreille" authorityYear="1802" box="[265,381,1684,1708]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="441" phylum="Arthropoda" rank="genus">
<emphasis id="B9154877441CA56E88261BE30E37F918" box="[265,381,1684,1708]" italics="true" pageId="8" pageNumber="441">Bembidion</emphasis>
</taxonomicName>
in similar habitats, as in the subgenus
<taxonomicName id="4C61EFE6441CA56E8A0C1BE30B10F918" authority="Moret &amp; Toledano" authorityName="Moret &amp; Toledano" authorityYear="2002" box="[803,1114,1684,1708]" class="Insecta" family="Carabidae" genus="Ecuadion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="441" phylum="Arthropoda" rank="genus">
<emphasis id="B9154877441CA56E8A0C1BE30CC3F918" box="[803,905,1684,1708]" italics="true" pageId="8" pageNumber="441">Ecuadion</emphasis>
Moret &amp; Toledano
</taxonomicName>
from Central and South America (
<bibRefCitation id="EFF0E994441DA56F88881DAC0D8EFF47" author="MORET, P. &amp; L. TOLEDANO" box="[423,708,219,243]" pageId="9" pageNumber="442" refId="ref8424" refString="MORET, P., AND L. TOLEDANO. 2002. Ecuadion, nouveau sous-genre de Bembidion Latreille, 1802 du paramo equatorien (Coleoptera, Carabidae, Bembidiini). Bollettino del Museo Civico di Storia Naturale di Venezia" type="book" year="2002">Moret and Toledano, 2002</bibRefCitation>
),
<taxonomicName id="4C61EFE6441DA56F8BF91DAC0CD2FF47" authority="Sharp" authorityName="Sharp" authorityYear="1903" box="[726,920,219,243]" class="Insecta" family="Carabidae" genus="Nesocidium" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="genus">
<emphasis id="B9154877441DA56F8BF91DAC0C1EFF47" box="[726,852,219,243]" italics="true" pageId="9" pageNumber="442">Nesocidium</emphasis>
Sharp
</taxonomicName>
from
<collectingRegion id="49A55A87441DA56F8AF71DAC0B6FFF47" box="[984,1061,219,243]" country="United States of America" name="Hawaii" pageId="9" pageNumber="442">Hawaii</collectingRegion>
(
<bibRefCitation id="EFF0E994441DA56F8D1D1DAC0FFEFEA7" author="LIEBHERR, J. K." pageId="9" pageNumber="442" pagination="31 - 78" refId="ref7834" refString="LIEBHERR, J. K. 2008. Taxonomic revision of Hawaiian Bembidion Latreille (Coleoptera: Carabidae: Bembidiini) with a discussion of their reductive and derivative evolutionary specializations. Annals of Carnegie Museum 77: 31 - 78." type="journal article" year="2008">Liebherr, 2008</bibRefCitation>
), subgenus
<taxonomicName id="4C61EFE6441DA56F881F1D8B0D45FEA7" authority="Alluaud" authorityName="Alluaud" authorityYear="1917" box="[304,527,251,275]" class="Insecta" family="Carabidae" genus="Hypsipezum" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="genus">
<emphasis id="B9154877441DA56F881F1D8B0EF8FEA7" box="[304,434,252,275]" italics="true" pageId="9" pageNumber="442">Hypsipezum</emphasis>
Alluaud
</taxonomicName>
from Africa, and some members of the
<taxonomicName id="4C61EFE6441DA56F8A961D8C0B79FEA7" box="[953,1075,251,275]" class="Insecta" family="Carabidae" genus="Ocydromus" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="genus">
<emphasis id="B9154877441DA56F8A961D8C0B79FEA7" box="[953,1075,251,275]" italics="true" pageId="9" pageNumber="442">Ocydromus</emphasis>
</taxonomicName>
complex in Central America (e.g.,
<taxonomicName id="4C61EFE6441DA56F88A31C6B0D03FE87" authority="Erwin" authorityName="Erwin" authorityYear="1982" box="[396,585,283,307]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="chiriqui">
<emphasis id="B9154877441DA56F88A31C6B0EB4FE87" box="[396,510,283,307]" italics="true" pageId="9" pageNumber="442">B. chiriqui</emphasis>
Erwin
</taxonomicName>
) (
<bibRefCitation id="EFF0E994441DA56F8B711C6C0C16FE87" author="MADDISON, D. R. &amp; J. S. SPROUL &amp; H. MENDEL" box="[606,860,283,307]" pageId="9" pageNumber="442" pagination="1155 - 1175" refId="ref8330" refString="MADDISON, D. R., J. S. SPROUL, AND H. MENDEL. 2019 b. Origin and adaptive radiation of the exceptional and threatened bembidiine beetle fauna of St Helena (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 189: 1155 - 1175." type="journal article" year="2019">
<taxonomicName id="4C61EFE6441DA56F8B711C6C0D80FE87" authority="Maddison, 2023" authorityName="Maddison" authorityYear="2023" box="[606,714,283,307]" pageId="9" pageNumber="441" rank="subGenus" status="subgen. nov." subGenus="Maddison">Maddison</taxonomicName>
et al., 2019b
</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C37BC7EE441DA56F89841C4B0E09F9AF" pageId="9" pageNumber="442" type="diagnosis">
<paragraph id="8BDE9465441DA56F89841C4B0CBEFE60" blockId="9.[123,1174,219,1725]" pageId="9" pageNumber="442">
<emphasis id="B9154877441DA56F89841C4B0E87FEE7" bold="true" box="[171,461,316,339]" pageId="9" pageNumber="442">Morphological Diagnosis.</emphasis>
Small to medium (
<quantity id="4C993980441DA56F8BB81C4C0C51FEE7" box="[663,795,315,339]" metricMagnitude="-3" metricUnit="m" metricValue="3.05" metricValueMax="3.5" metricValueMin="2.6" pageId="9" pageNumber="442" unit="mm" value="3.05" valueMax="3.5" valueMin="2.6">2.6–3.5 mm</quantity>
), shiny
<emphasis id="B9154877441DA56F8A421C4C0CA2FEE7" box="[877,1000,315,339]" italics="true" pageId="9" pageNumber="442">
<taxonomicName id="4C61EFE6441DA56F8A421C4C0CA9FEE7" authorityName="Latreille" authorityYear="1802" box="[877,995,315,339]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
,
</emphasis>
without colored markings on the elytra. Frontal furrows single, broad, shallow, well-marked (
<taxonomicName id="4C61EFE6441DA56F8A921C2B0BCCFEC0" authorityName="Erwin" authorityYear="1982" box="[957,1158,348,372]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="franiae">
<emphasis id="B9154877441DA56F8A921C2B0BCCFEC0" box="[957,1158,348,372]" italics="true" pageId="9" pageNumber="442">Bembidion franiae</emphasis>
</taxonomicName>
), or broad, shallow, and disrupted by surface rugosity (other species), not convergent or extended onto clypeus. Eyes of normal size for
<taxonomicName id="4C61EFE6441DA56F8B341CEB0DC5FE00" authorityName="Latreille" authorityYear="1802" box="[539,655,412,436]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="genus">
<emphasis id="B9154877441DA56F8B341CEB0DC5FE00" box="[539,655,412,436]" italics="true" pageId="9" pageNumber="442">Bembidion</emphasis>
</taxonomicName>
(
<taxonomicName id="4C61EFE6441DA56F8B8F1CEA0C46FE00" authorityName="Erwin" authorityYear="1982" box="[672,780,412,436]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="franiae">
<emphasis id="B9154877441DA56F8B8F1CEA0C46FE00" box="[672,780,412,436]" italics="true" pageId="9" pageNumber="442">B. franiae</emphasis>
</taxonomicName>
) or slightly reduced (other species). Mentum with anterior lateral region complete, triangular; mentum tooth triangular.
</paragraph>
<paragraph id="8BDE9465441DA56F89841CAB0EA1FDC1" blockId="9.[123,1174,219,1725]" pageId="9" pageNumber="442">
Posterior angle of pronotum with a posterolateral carina, although it is interrupted posteriorly in
<taxonomicName id="4C61EFE6441DA56F89B51C890E42FDA1" authorityName="Erwin" authorityYear="1982" box="[154,264,509,533]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="franiae">
<emphasis id="B9154877441DA56F89B51C890E42FDA1" box="[154,264,509,533]" italics="true" pageId="9" pageNumber="442">B. franiae</emphasis>
</taxonomicName>
and some specimens of other species. Posterior margin of pronotum not strongly sinuate laterally; basal transverse furrow weak, disrupted by shallow punctures, and in many specimens interrupted by the medial furrow extending posteriorly to the hind margin of the pronotum; hind angles right or slightly acute.
</paragraph>
<paragraph id="8BDE9465441DA56F89841F0A0D00FC23" blockId="9.[123,1174,219,1725]" pageId="9" pageNumber="442">
Elytron with lateral bead ending at humerus, not prolonged onto base, although in
<taxonomicName id="4C61EFE6441DA56F8D061F080BDFFD21" authorityName="Erwin" authorityYear="1982" box="[1065,1173,637,662]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="franiae">
<emphasis id="B9154877441DA56F8D061F080BDFFD21" box="[1065,1173,637,662]" italics="true" pageId="9" pageNumber="442">B. franiae</emphasis>
</taxonomicName>
a short carina extends from the end of the bead toward the center of the elytral disc at an angle, giving the impression of an angulate shoulder bead. Striae vary in depth, from anteriorly deep in
<taxonomicName id="4C61EFE6441DA56F89541FA80FACFD42" authorityName="Erwin" authorityYear="1982" box="[123,230,734,758]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="franiae">
<emphasis id="B9154877441DA56F89541FA80FACFD42" box="[123,230,734,758]" italics="true" pageId="9" pageNumber="442">B. franiae</emphasis>
</taxonomicName>
to much shallower in most other species, with lateral striae increasingly evanescent; all striae absent or nearly so in the posterior quarter of the elytra, except for the first stria; two discal setae in third stria. Elytra slightly iridescent in
<taxonomicName id="4C61EFE6441DA56F8B461E570D99FC83" authorityName="Erwin" authorityYear="1982" box="[617,723,799,823]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="franiae">
<emphasis id="B9154877441DA56F8B461E570D99FC83" box="[617,723,799,823]" italics="true" pageId="9" pageNumber="442">B. franiae</emphasis>
</taxonomicName>
, because of the transverse microsculpture, and slightly dull in
<taxonomicName id="4C61EFE6441DA56F88471E370D5DFCE3" authorityName="Maddison" authorityYear="2023" box="[360,535,831,855]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="elescarabajo">
<emphasis id="B9154877441DA56F88471E370D5DFCE3" box="[360,535,831,855]" italics="true" pageId="9" pageNumber="442">B. elescarabajo</emphasis>
</taxonomicName>
, because of the nearly isodiametric sculpticells; elytral microsculpture absent from specimens of the
<taxonomicName id="4C61EFE6441DA56F8B4D1E280DF3FCC3" authorityName="Erwin" authorityYear="1982" box="[610,697,863,887]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="nahuala">
<emphasis id="B9154877441DA56F8B4D1E280DF3FCC3" box="[610,697,863,887]" italics="true" pageId="9" pageNumber="442">nahuala</emphasis>
</taxonomicName>
group. Mesoventral process without subapical setae. Metaventral process unmargined.
</paragraph>
<paragraph id="8BDE9465441DA56F89841EE80D9BFC4C" blockId="9.[123,1174,219,1725]" pageId="9" pageNumber="442">Apex of last visible abdominal sternite with two setae in males, four setae in females. Apices of each paramere normally with three setae, but some individuals have two or four setae on the left paramere, and one has four setae on the right paramere.</paragraph>
<paragraph id="8BDE9465441DA56F898419770D57FB4D" blockId="9.[123,1174,219,1725]" pageId="9" pageNumber="442">
Within the geographic range of
<taxonomicName id="4C61EFE6441DA56F8B3B19760DDCFBAC" authorityName="Maddison" authorityYear="2023" box="[532,662,1025,1048]" pageId="9" pageNumber="443" rank="subGenus" subGenus="Geocosmius">
<emphasis id="B9154877441DA56F8B3B19760DDCFBAC" box="[532,662,1025,1048]" italics="true" pageId="9" pageNumber="442">Geocosmius</emphasis>
</taxonomicName>
live several other species of
<taxonomicName id="4C61EFE6441DA56F8AC819770B11FBAC" authorityName="Latreille" authorityYear="1802" box="[999,1115,1024,1048]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="genus">
<emphasis id="B9154877441DA56F8AC819770B11FBAC" box="[999,1115,1024,1048]" italics="true" pageId="9" pageNumber="442">Bembidion</emphasis>
</taxonomicName>
with small, convex adults without spots:
<taxonomicName id="4C61EFE6441DA56F8B2E19560DF7FB8C" authority="Erwin" authorityName="Erwin" authorityYear="1982" box="[513,701,1056,1080]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="purulha">
<emphasis id="B9154877441DA56F8B2E19560D3EFB8C" box="[513,628,1056,1080]" italics="true" pageId="9" pageNumber="442">B. purulha</emphasis>
Erwin
</taxonomicName>
(a member of the
<taxonomicName id="4C61EFE6441DA56F8AA619570B49FB8C" box="[905,1027,1056,1080]" class="Insecta" family="Carabidae" genus="Ocydromus" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="genus">
<emphasis id="B9154877441DA56F8AA619570B49FB8C" box="[905,1027,1056,1080]" italics="true" pageId="9" pageNumber="442">Ocydromus</emphasis>
</taxonomicName>
complex), as well as
<taxonomicName id="4C61EFE6441DA56F89E219350D54FBEC" authority="Bates" authorityName="Bates" authorityYear="1884" box="[205,542,1088,1113]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="cyclodes" subGenus="Cyclolopha">
<emphasis id="B9154877441DA56F89E219350FAFFBED" box="[205,229,1090,1113]" italics="true" pageId="9" pageNumber="442">B.</emphasis>
(
<emphasis id="B9154877441DA56F89DB19370E25FBEC" box="[244,367,1088,1112]" italics="true" pageId="9" pageNumber="442">Cyclolopha</emphasis>
)
<emphasis id="B9154877441DA56F88AF19370E96FBEC" box="[384,476,1088,1112]" italics="true" pageId="9" pageNumber="442">cyclodes</emphasis>
Bates
</taxonomicName>
and
<taxonomicName id="4C61EFE6441DA56F8B7C19350C8AFBEC" authority="Bates." authorityName="Bates." box="[595,960,1088,1113]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="championi" subGenus="Cyclolopha">
<emphasis id="B9154877441DA56F8B7C19350D20FBED" box="[595,618,1090,1113]" italics="true" pageId="9" pageNumber="442">B.</emphasis>
(
<emphasis id="B9154877441DA56F8B5519370DBFFBEC" box="[634,757,1088,1112]" italics="true" pageId="9" pageNumber="442">Cyclolopha</emphasis>
)
<emphasis id="B9154877441DA56F8A2A19370C3CFBEC" box="[773,886,1088,1112]" italics="true" pageId="9" pageNumber="442">championi</emphasis>
Bates.
</taxonomicName>
<taxonomicName id="4C61EFE6441DA56F8AE919370BDFFBEC" authorityName="Erwin" authorityYear="1982" box="[966,1173,1088,1112]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="purulha">
<emphasis id="B9154877441DA56F8AE919370BDFFBEC" box="[966,1173,1088,1112]" italics="true" pageId="9" pageNumber="442">Bembidion purulha</emphasis>
</taxonomicName>
can be distinguished by the elytral striae, which abruptly transition at about the halfway point from large, distinct punctures anteriorly to fully effaced striae posteriorly, as well as a fully margined metaventral process. The two
<taxonomicName id="4C61EFE6441DA56F889619D60DC2FB0D" box="[441,648,1185,1209]" pageId="9" pageNumber="442">
<emphasis id="B9154877441DA56F889619D60D7EFB0D" box="[441,564,1185,1209]" italics="true" pageId="9" pageNumber="442">Cyclolopha</emphasis>
species
</taxonomicName>
can be distinguished by the much broader pronotum with very rounded hind angles, with the posterior lateral seta notably anterior to the posterior margin of the pronotum at the midline.
</paragraph>
<paragraph id="8BDE9465441DA56F898418750D14FA2F" blockId="9.[123,1174,219,1725]" pageId="9" pageNumber="442">
All known specimens of
<taxonomicName id="4C61EFE6441DA56F889618740D6EFAAE" authorityName="Erwin" authorityYear="1982" box="[441,548,1282,1306]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="franiae">
<emphasis id="B9154877441DA56F889618740D6EFAAE" box="[441,548,1282,1306]" italics="true" pageId="9" pageNumber="442">B. franiae</emphasis>
</taxonomicName>
and
<taxonomicName id="4C61EFE6441DA56F8B7518740C49FAAE" authorityName="Maddison" authorityYear="2023" box="[602,771,1282,1306]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="elescarabajo">
<emphasis id="B9154877441DA56F8B7518740C49FAAE" box="[602,771,1282,1306]" italics="true" pageId="9" pageNumber="442">B. elescarabajo</emphasis>
</taxonomicName>
lack hind wings.
<taxonomicName id="4C61EFE6441DA56F8AEC18750BDFFAAE" authorityName="Erwin" authorityYear="1982" box="[963,1173,1282,1306]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="nahuala">
<emphasis id="B9154877441DA56F8AEC18750BDFFAAE" box="[963,1173,1282,1306]" italics="true" pageId="9" pageNumber="442">Bembidion nahuala</emphasis>
</taxonomicName>
and
<emphasis id="B9154877441DA56F898518540F88FA8E" box="[170,194,1315,1338]" italics="true" pageId="9" pageNumber="442">B.</emphasis>
“
<collectingRegion id="49A55A87441DA56F89FA18550E65FA8E" box="[213,303,1314,1338]" country="Mexico" name="Chiapas" pageId="9" pageNumber="442">Chiapas</collectingRegion>
” are wing dimorphic, with most specimens lacking hind wings, and with more rounded elytra and notably rounded shoulders (
<figureCitation id="135A88E0441DA56F8B5618350D85FAEE" box="[633,719,1346,1370]" captionStart="FIGURE" captionStartId="1.[155,167,1563,1582]" captionTargetBox="[156,1171,245,1547]" captionTargetId="figure-14@1.[120,1176,216,1551]" captionTargetPageId="1" captionText="FIGURE 1. Habitus of Geocosmius species. (A) Bembidion “Chiapas”, brachypterous form, voucher V101274. (B) B. “Chiapas”, fully winged form, voucher V101273. (C) B. elescarabajo, paratype, voucher V101491. (D) B. franiae, specimen UASM371160. Note: B. nahuala (not shown) is extremely similar externally to B. “Chiapas”. Scale bar 1 mm." figureDoi="http://doi.org/10.5281/zenodo.11067244" httpUri="https://zenodo.org/record/11067244/files/figure.png" pageId="9" pageNumber="442">Fig. 1A</figureCitation>
). A minority of specimens of both species are fully winged, with more parallel-sided elytra and less rounded shoulders, giving these specimens a rather different appearance (
<figureCitation id="135A88E0441DA56F88D218F40D1AFA2F" box="[509,592,1411,1435]" captionStart="FIGURE" captionStartId="1.[155,167,1563,1582]" captionTargetBox="[156,1171,245,1547]" captionTargetId="figure-14@1.[120,1176,216,1551]" captionTargetPageId="1" captionText="FIGURE 1. Habitus of Geocosmius species. (A) Bembidion “Chiapas”, brachypterous form, voucher V101274. (B) B. “Chiapas”, fully winged form, voucher V101273. (C) B. elescarabajo, paratype, voucher V101491. (D) B. franiae, specimen UASM371160. Note: B. nahuala (not shown) is extremely similar externally to B. “Chiapas”. Scale bar 1 mm." figureDoi="http://doi.org/10.5281/zenodo.11067244" httpUri="https://zenodo.org/record/11067244/files/figure.png" pageId="9" pageNumber="442">Fig. 1B</figureCitation>
).
</paragraph>
<paragraph id="8BDE9465441DA56F898418D30E09F9AF" blockId="9.[123,1174,219,1725]" pageId="9" pageNumber="442">
<emphasis id="B9154877441DA56F898418D30E43FA0F" bold="true" box="[171,265,1444,1467]" pageId="9" pageNumber="442">Habitat.</emphasis>
<collectorName id="2694F1B3441DA56F883A18D40E1CFA0F" box="[277,342,1443,1467]" pageId="9" pageNumber="442">Based</collectorName>
upon the available label data, species in this group live in cloud forests (
<collectingCountry id="F376D4F5441DA56F89AD18B40FB0FA6F" box="[130,250,1475,1499]" name="Guatemala" pageId="9" pageNumber="442">Guatemala</collectingCountry>
, Chiapas, Oaxaca,
<collectingMunicipality id="6BBA0E1F441DA56F88E818B40D60FA6F" box="[455,554,1475,1499]" pageId="9" pageNumber="442">Guerrero</collectingMunicipality>
, and
<location id="8EBEC2BE441DA56F8B4C18B40DFAFA6F" LSID="urn:lsid:plazi:treatment:03C82573441CA56C889D198E0ED0FE42:8EBEC2BE441DA56F8B4C18B40DFAFA6F" box="[611,688,1475,1499]" country="Guatemala" municipality="Guerrero" name="Puebla" pageId="9" pageNumber="442" stateProvince="Oaxaca">Puebla</location>
), oak-pine forests (
<collectingCountry id="F376D4F5441DA56F8A5218B40CBFFA6F" box="[893,1013,1475,1499]" name="Guatemala" pageId="9" pageNumber="442">Guatemala</collectingCountry>
,
<collectingCountry id="F376D4F5441DA56F8D2E18B40B1FFA6F" box="[1025,1109,1475,1499]" name="Mexico" pageId="9" pageNumber="442">México</collectingCountry>
,
<collectingRegion id="49A55A87441DA56F8D4F18B40FE8FA4F" country="Mexico" name="Oaxaca" pageId="9" pageNumber="442">Oaxaca</collectingRegion>
), pine forests (
<collectingRegion id="49A55A87441DA56F881118940EE9FA4F" box="[318,419,1507,1531]" country="Mexico" name="Veracruz de Ignacio de la Llave" pageId="9" pageNumber="442">Veracruz</collectingRegion>
), pine-alder forests (
<collectingRegion id="49A55A87441DA56F8B5218940DA8FA4F" box="[637,738,1507,1531]" country="Mexico" name="Veracruz de Ignacio de la Llave" pageId="9" pageNumber="442">Veracruz</collectingRegion>
), wet oak forests (
<collectingRegion id="49A55A87441DA56F8A8C18940CBBFA4F" box="[931,1009,1507,1531]" country="Mexico" name="Puebla" pageId="9" pageNumber="442">Puebla</collectingRegion>
), and oak-alder forests (
<collectingRegion id="49A55A87441DA56F89FF1B740E7DF9AF" box="[208,311,1539,1563]" country="Mexico" name="Guerrero" pageId="9" pageNumber="442">Guerrero</collectingRegion>
).
</paragraph>
</subSubSection>
<subSubSection id="C37BC7EE441DA56F89841B530F98F9E8" pageId="9" pageNumber="442" type="distribution">
<paragraph id="8BDE9465441DA56F89841B530F98F9E8" blockId="9.[123,1174,219,1725]" pageId="9" pageNumber="442">
<emphasis id="B9154877441DA56F89841B530E83F98F" bold="true" box="[171,457,1572,1595]" pageId="9" pageNumber="442">Geographic distribution.</emphasis>
This subgenus is known from southern
<collectingCountry id="F376D4F5441DA56F8ABC1B530CAEF988" box="[915,996,1572,1596]" name="Mexico" pageId="9" pageNumber="442">México</collectingCountry>
and
<collectingCountry id="F376D4F5441DA56F8D0D1B530BDCF988" box="[1058,1174,1572,1596]" name="Guatemala" pageId="9" pageNumber="442">Guatemala</collectingCountry>
(
<figureCitation id="135A88E0441DA56F89AC1B330F8FF9E8" box="[131,197,1604,1628]" captionStart="FIGURE" captionStartId="2.[155,167,821,840]" captionTargetBox="[140,1175,216,799]" captionTargetId="figure-313@2.[120,1176,216,811]" captionTargetPageId="2" captionText="FIGURE 2. Geographic distribution of Geocosmius in México and Guatemala." figureDoi="http://doi.org/10.5281/zenodo.11067246" httpUri="https://zenodo.org/record/11067246/files/figure.png" pageId="9" pageNumber="442">Fig. 2</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C37BC7EE441DA56C89841B120ED0FE42" lastPageId="10" lastPageNumber="443" pageId="9" pageNumber="442" type="description">
<paragraph id="8BDE9465441DA56C89841B120D5EFEA7" blockId="9.[123,1174,219,1725]" lastBlockId="10.[123,1174,219,502]" lastPageId="10" lastPageNumber="443" pageId="9" pageNumber="442">
<emphasis id="B9154877441DA56F89841B120E09F9C8" bold="true" box="[171,323,1637,1660]" pageId="9" pageNumber="442">Composition.</emphasis>
The known species of subgenus
<taxonomicName id="4C61EFE6441DA56F8B911B120C0AF9C8" authorityName="Maddison" authorityYear="2023" box="[702,832,1637,1660]" pageId="9" pageNumber="442" rank="subGenus" subGenus="Geocosmius">
<emphasis id="B9154877441DA56F8B911B120C0AF9C8" box="[702,832,1637,1660]" italics="true" pageId="9" pageNumber="442">Geocosmius</emphasis>
</taxonomicName>
are
<taxonomicName id="4C61EFE6441DA56F8A581B130BDBF9C8" authority="Erwin" authorityName="Erwin" authorityYear="1982" box="[887,1169,1636,1660]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="franiae">
<emphasis id="B9154877441DA56F8A581B130B08F9C8" box="[887,1090,1636,1660]" italics="true" pageId="9" pageNumber="442">Bembidion franiae</emphasis>
Erwin
</taxonomicName>
,
<taxonomicName id="4C61EFE6441DA56F89541BF30E34F928" authorityName="Maddison" authorityYear="2023" box="[123,382,1668,1692]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="elescarabajo">
<emphasis id="B9154877441DA56F89541BF30E34F928" box="[123,382,1668,1692]" italics="true" pageId="9" pageNumber="442">Bembidion elescarabajo</emphasis>
</taxonomicName>
Maddison,
<taxonomicNameLabel id="A226F50C441DA56F88D41BF20D1AF928" box="[507,592,1669,1692]" pageId="9" pageNumber="442" rank="species">sp. nov.</taxonomicNameLabel>
, and then a group of similar forms (the
<taxonomicName id="4C61EFE6441DA56F8ADE1BF30B02F928" authorityName="Erwin" authorityYear="1982" box="[1009,1096,1668,1692]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="nahuala">
<emphasis id="B9154877441DA56F8ADE1BF30B02F928" box="[1009,1096,1668,1692]" italics="true" pageId="9" pageNumber="442">nahuala</emphasis>
</taxonomicName>
group) including
<taxonomicName id="4C61EFE6441DA56F89C81BD30D49F908" authority="Erwin" authorityName="Erwin" authorityYear="1982" box="[231,515,1700,1724]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="nahuala">
<emphasis id="B9154877441DA56F89C81BD30EF0F908" box="[231,442,1700,1724]" italics="true" pageId="9" pageNumber="442">Bembidion nahuala</emphasis>
Erwin
</taxonomicName>
and
<emphasis id="B9154877441DA56F8B151BD10D1BF909" box="[570,593,1702,1725]" italics="true" pageId="9" pageNumber="442">B.</emphasis>
“
<collectingRegion id="49A55A87441DA56F8B4C1BD30D8AF908" box="[611,704,1700,1724]" country="Mexico" name="Chiapas" pageId="9" pageNumber="442">Chiapas</collectingRegion>
”. I have examined
<specimenCount id="9D675FEC441DA56F8AA01BD30B5EF908" box="[911,1044,1700,1724]" count="5" pageId="9" pageNumber="442" type="generic">5 specimens</specimenCount>
of
<taxonomicName id="4C61EFE6441DA56C8D151BD10FDEFF47" authorityName="Erwin" authorityYear="1982" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" lastPageId="10" lastPageNumber="443" order="Coleoptera" pageId="9" pageNumber="442" phylum="Arthropoda" rank="species" species="franiae">
<emphasis id="B9154877441DA56C8D151BD10FDEFF47" italics="true" lastPageId="10" lastPageNumber="443" pageId="9" pageNumber="442">B. franiae</emphasis>
</taxonomicName>
, 13 of
<taxonomicName id="4C61EFE6441EA56C89C11DAB0ED0FF47" authorityName="Maddison" authorityYear="2023" box="[238,410,219,243]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="443" phylum="Arthropoda" rank="species" species="elescarabajo">
<emphasis id="B9154877441EA56C89C11DAB0ED0FF47" box="[238,410,219,243]" italics="true" pageId="10" pageNumber="443">B. elescarabajo</emphasis>
</taxonomicName>
, and
<specimenCount id="9D675FEC441EA56C88F11DAC0D35FF47" box="[478,639,219,243]" count="311" pageId="10" pageNumber="443" type="generic">311 specimens</specimenCount>
of the
<taxonomicName id="4C61EFE6441EA56C8BF71DAC0C65FF47" authorityName="Erwin" authorityYear="1982" box="[728,815,219,243]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="443" phylum="Arthropoda" rank="species" species="nahuala">
<emphasis id="B9154877441EA56C8BF71DAC0C65FF47" box="[728,815,219,243]" italics="true" pageId="10" pageNumber="443">nahuala</emphasis>
</taxonomicName>
group (
<collectionCode id="ED700CA0441EA56C8AA31DAC0C97FF47" box="[908,989,219,243]" country="Canada" lsid="urn:lsid:biocol.org:col:34404" name="University of Alberta, E.H. Strickland Entomological Museum" pageId="10" pageNumber="443">UASM</collectionCode>
,
<collectionCode id="ED700CA0441EA56C8AC21DAC0B65FF47" box="[1005,1071,219,243]" country="USA" name="Cornell University Insect Collection" pageId="10" pageNumber="443" type="University or college">CUIC</collectionCode>
,
<collectionCode id="ED700CA0441EA56C8D101DAC0BD8FF47" box="[1087,1170,219,243]" country="USA" name="The Cleveland Museum of Natural History" pageId="10" pageNumber="443" type="Museum">CMNH</collectionCode>
,
<collectionCode id="ED700CA0441EA56C89541D8B0F85FEA7" box="[123,207,252,275]" country="USA" httpUri="http://grbio.org/cool/142r-0w94" name="Smithsonian Institution, National Museum of Natural History" pageId="10" pageNumber="443" type="Museum">NMNH</collectionCode>
,
<collectionCode id="ED700CA0441EA56C89F51D8C0E64FEA7" box="[218,302,251,275]" country="Mexico" name="Universidad Nacional Autonoma de Mexico" pageId="10" pageNumber="443">UNAM</collectionCode>
,
<collectionCode id="ED700CA0441EA56C88151D8C0ECEFEA7" box="[314,388,251,275]" pageId="10" pageNumber="443">CTVR</collectionCode>
, and
<collectionCode id="ED700CA0441EA56C88921D8C0D43FEA7" box="[445,521,251,275]" country="USA" name="Oregon State Arthropod Collection" pageId="10" pageNumber="443" type="University or college">OSAC</collectionCode>
).
</paragraph>
<paragraph id="8BDE9465441EA56C89841C6C0ED0FE42" blockId="10.[123,1174,219,502]" pageId="10" pageNumber="443">
Members of the
<taxonomicName id="4C61EFE6441EA56C88711C6C0EFFFE87" authorityName="Erwin" authorityYear="1982" box="[350,437,283,307]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="443" phylum="Arthropoda" rank="species" species="nahuala">
<emphasis id="B9154877441EA56C88711C6C0EFFFE87" box="[350,437,283,307]" italics="true" pageId="10" pageNumber="443">nahuala</emphasis>
</taxonomicName>
group are very similar to one another. With the exception of a form with deeper elytral stria from Volcán de
<collectingRegion id="49A55A87441EA56C8B0C1C4B0D38FEE0" box="[547,626,316,340]" country="Mexico" name="Colima" pageId="10" pageNumber="443">Colima</collectingRegion>
in
<collectingRegion id="49A55A87441EA56C8BBC1C4B0D94FEE0" box="[659,734,316,340]" country="Mexico" name="Jalisco" pageId="10" pageNumber="443">Jalisco</collectingRegion>
, and the differences associated with wing dimorphism and gender, this widespread complex shows no obvious patterns of external morphological variation.
<taxonomicName id="4C61EFE6441EA56C88181C0A0D1BFE21" authority="Erwin" authorityName="Erwin" authorityYear="1982" box="[311,593,381,405]" class="Insecta" family="Carabidae" genus="Bembidion" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="443" phylum="Arthropoda" rank="species" species="nahuala">
<emphasis id="B9154877441EA56C88181C0A0D42FE21" box="[311,520,381,405]" italics="true" pageId="10" pageNumber="443">Bembidion nahuala</emphasis>
Erwin
</taxonomicName>
and
<emphasis id="B9154877441EA56C8BA91C090DD7FE21" box="[646,669,382,405]" italics="true" pageId="10" pageNumber="443">B.</emphasis>
“
<collectingRegion id="49A55A87441EA56C8B821C0A0C4DFE21" box="[685,775,381,405]" country="Mexico" name="Chiapas" pageId="10" pageNumber="443">Chiapas</collectingRegion>
” appear to be different species based upon DNA sequences (
<figureCitation id="135A88E0441EA56C885D1CEA0EF8FE01" box="[370,434,413,437]" captionStart="FIGURE" captionStartId="8.[155,167,938,957]" captionTargetBox="[151,1176,216,924]" captionTargetId="figure-172@8.[120,1176,216,924]" captionTargetPageId="8" captionText="FIGURE 4. Phylogenetic results, focusing on subgenera Geocosmius, Liocosmius, and Trechonepha. (A) A portion of the maximum likelihood tree inferred from the concatenated, eight-gene matrix. Numbers of the branches are maximum likelihood bootstrap percentages; bootstrap values within most of Liocosmius not shown. Other bembidiines omitted from the image. Scale bar 0.1 units, as calculated by IQ-TREE. (B) Support for various clades, measured with bootstrap support percentages from maximum likelihood phylogenetic analyses. One or two values are given in each cell. If the bootstrap support percentage is 90 or more, only that value is listed. If bootstrap support is less than 90, two values are listed: the bootstrap support for the clade, followed by a negative value which is the bootstrap support against the clade, as measured by the bootstrap value for the contradictory clade with the highest support value. The “8g” column shows the support values for the concatenated, eight-gene matrix. “-” indicates only one species is included, and thus support for that clade cannot be measured." figureDoi="http://doi.org/10.5281/zenodo.11067250" httpUri="https://zenodo.org/record/11067250/files/figure.png" pageId="10" pageNumber="443">Fig. 4</figureCitation>
) and male genitalia (
<figureCitation id="135A88E0441EA56C8BBA1CEA0D9CFE01" box="[661,726,413,437]" captionStart="FIGURE" captionStartId="11.[155,167,1212,1231]" captionTargetBox="[120,1164,236,1178]" captionTargetId="figure-170@11.[120,1176,216,1200]" captionTargetPageId="11" captionText="FIGURE 5. Aedeagi of Geocosmius. (A) Bembidion “Chiapas”, voucher DNA5317. (B) B. “Chiapas”, voucher DNA4882.Arrow indicates the prominent ridge characteristic of this species. (C) B. elescarabajo, voucher V101490.Arrow indicates the less curved sclerite margin characteristic of this species. (D) B. elescarabajo, voucher V101489. (E) B. nahuala voucher DNA5033. (F) B. nahuala voucher DNA5034. (G) B. nahuala, holotype. (H) B. franiae, voucher DNA4623. Scale bar 100 µm." figureDoi="http://doi.org/10.5281/zenodo.11067252" httpUri="https://zenodo.org/record/11067252/files/figure.png" pageId="10" pageNumber="443">Fig. 5</figureCitation>
). The two forms differ by 4 bases in 28S, and by 27 bases or 4.1% in COI; these nucleotide differences in COI imply two amino acid differences between the species.
</paragraph>
</subSubSection>
</treatment>
</document>
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