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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.416.7706" ID-GBIF-Dataset="47852319-33aa-4966-89af-68e6352e05ae" ID-PMC="PMC4109512" ID-Pensoft-Pub="1313-2970-416-113" ID-PubMed="25061349" ID-ZBK="CE0561FB5EE4498BA2C2EDF9B14F241D" ModsDocAuthor="" ModsDocDate="2014" ModsDocID="1313-2970-416-113" ModsDocOrigin="ZooKeys 416" ModsDocTitle="An unexpected clade of South American ground beetles (Coleoptera, Carabidae, Bembidion)" checkinTime="1451245749490" checkinUser="pensoft" docAuthor="Maddison, David R." docDate="2014" docId="BC5FFC0B030ABA4EEC5B96A9670AA197" docLanguage="en" docName="ZooKeys 416: 113-155" docOrigin="ZooKeys 416" docSource="http://dx.doi.org/10.3897/zookeys.416.7706" docTitle="Notholopha (Ecuadion) Moret &amp; Toledano 2002" docType="treatment" docVersion="3" lastPageNumber="131" masterDocId="7643F6149609155E6212FF95A85FFFBF" masterDocTitle="An unexpected clade of South American ground beetles (Coleoptera, Carabidae, Bembidion)" masterLastPageNumber="155" masterPageNumber="113" pageNumber="131" updateTime="1668158654224" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>An unexpected clade of South American ground beetles (Coleoptera, Carabidae, Bembidion)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Maddison, David R.</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2014</mods:date>
<mods:detail type="volume">
<mods:number>416</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>113</mods:start>
<mods:end>155</mods:end>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.416.7706</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.416.7706</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-416-113</mods:identifier>
<mods:identifier type="ZBK">CE0561FB5EE4498BA2C2EDF9B14F241D</mods:identifier>
<mods:identifier type="ZooBank">CE0561FB5EE4498BA2C2EDF9B14F241D</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="152053566" LSID="urn:lsid:plazi:treatment:BC5FFC0B030ABA4EEC5B96A9670AA197" httpUri="http://treatment.plazi.org/id/BC5FFC0B030ABA4EEC5B96A9670AA197" lastPageNumber="131" pageId="18" pageNumber="131">
<subSubSection pageId="18" pageNumber="131" type="multiple">
<paragraph pageId="18" pageNumber="131">Taxon classification Animalia Coleoptera Carabidae</paragraph>
</subSubSection>
<subSubSection pageId="18" pageNumber="131" type="nomenclature">
<paragraph pageId="18" pageNumber="131">
Subgenus
<taxonomicName authority="Moret &amp; Toledano, 2002" authorityName="Moret &amp; Toledano" authorityYear="2002" class="Insecta" family="Carabidae" genus="Notholopha" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Notholopha (Ecuadion)" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="subGenus" subGenus="Ecuadion">Ecuadion Moret &amp; Toledano, 2002</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection pageId="18" pageNumber="131" type="reference_group">
<paragraph pageId="18" pageNumber="131">
<taxonomicName class="Insecta" family="Carabidae" genus="Ecuadion" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Ecuadion" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="genus">Ecuadion</taxonomicName>
Moret &amp; Toledano, 2002; type species
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion fulvocinctum" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="fulvocinctum">Bembidion fulvocinctum</taxonomicName>
Bates, by original designation.
</paragraph>
</subSubSection>
<subSubSection pageId="18" pageNumber="131" type="remarks">
<paragraph pageId="18" pageNumber="131">Remarks.</paragraph>
<paragraph pageId="18" pageNumber="131">
<taxonomicName class="Insecta" family="Carabidae" genus="Ecuadion" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Ecuadion" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="genus">Ecuadion</taxonomicName>
is the largest subgenus in the
<taxonomicName class="Insecta" family="Carabidae" genus="Antiperyphanes" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Antiperyphanes" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="genus">Antiperyphanes</taxonomicName>
Complex, with over 50 described species (
<bibRefCitation author="Maddison, DR" journalOrPublisher="A new species of Bembidion (Ecuadion) from Ecuador (Coleoptera, Carabidae, Bembidiini), with a key to members of the georgeballi species group." pageId="27" pageNumber="140" pagination="51 - 60" url="10.3897/zookeys.249.4149" volume="249" year="2012">Maddison and Toledano 2012</bibRefCitation>
; Moret and
<bibRefCitation author="Toledano, L" journalOrPublisher="Koleopterologische Rundschau" pageId="27" pageNumber="140" pagination="1 - 14" title="Nomenclatorial revision of the supraspecific taxa of Bembidiini s. str. of South America described by Jeannel (1962) and related taxa with some descriptions of the fauna of South America (Coleoptera: Carabidae)." volume="72" year="2002">Toledano 2002</bibRefCitation>
;
<bibRefCitation author="Toledano, L" journalOrPublisher="Memoirs on Biodiversity" pageId="27" pageNumber="140" pagination="81 - 130" title="Systematic notes on the Bembidiina of the northern Andes with particular reference to the fauna of Ecuador (Coleoptera, Carabidae)." volume="1" year="2008">Toledano 2008</bibRefCitation>
;
<bibRefCitation author="Vigna Taglianti, A" journalOrPublisher="Memoirs on Biodiversity" pageId="27" pageNumber="140" pagination="77 - 80" title="Bembidion (Ecuadion) agonoides sp. n. from Ecuador (Coleoptera, Carabidae, Bembidiina)." volume="1" year="2008">Vigna Taglianti and Toledano 2008</bibRefCitation>
) and likely many undescribed. It occurs from Costa Rica south to the mountains near Mendoza, Argentina. There are no known exoskeletal synapomorphies of the group, but it is well-supported by the molecular data, with bootstrap support in six of the seven genes examined (Table 3).
</paragraph>
<paragraph pageId="18" pageNumber="131">
<taxonomicName class="Insecta" family="Carabidae" genus="Ecuadion" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Ecuadion" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="genus">Ecuadion</taxonomicName>
falls into two distinct clades among the sampled species: (1) the
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion chimborazonum" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="chimborazonum">Bembidion chimborazonum</taxonomicName>
group (clade m in Fig. 8), consisting mostly of larger, long-legged species (Figs 3A, B); (2) remaining
<taxonomicName class="Insecta" family="Carabidae" genus="Ecuadion" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Ecuadion" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="genus">Ecuadion</taxonomicName>
(clade n in Fig. 8), consisting of mostly smaller species with shorter appendages (Figs 3
<normalizedToken originalValue="CF">C-F</normalizedToken>
). The
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion chimborazonum" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="chimborazonum">Bembidion chimborazonum</taxonomicName>
group is supported by all genes examined (Table 3); support for the complementary clade is not quite as strong, with the clearest evidence coming from ribosomal genes (Table 3).
</paragraph>
<paragraph pageId="18" pageNumber="131">
Unlike most
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
, this subgenus has radiated in habitats away from water. Some occur in leaf litter in cloud forests (e.g.,
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion andersoni" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="andersoni">Bembidion andersoni</taxonomicName>
,
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion georgeballi" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="georgeballi">Bembidion georgeballi</taxonomicName>
,
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion onorei" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="onorei">Bembidion onorei</taxonomicName>
,
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
sp.
<normalizedToken originalValue="“Papallacta”">&quot;Papallacta&quot;</normalizedToken>
; Fig. 4D), in habitats that would typically be occupied by the genus
<taxonomicName class="Insecta" family="Carabidae" genus="Trechus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Trechus" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="genus">Trechus</taxonomicName>
Clairville in North America. A number of species are found in open, high-elevation grasslands (e.g.,
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion chimborazonum" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="chimborazonum">Bembidion chimborazonum</taxonomicName>
,
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion guamani" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="guamani">Bembidion guamani</taxonomicName>
,
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion humboldti" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="humboldti">Bembidion humboldti</taxonomicName>
; Fig. 4C). Some species occur on the upper banks of creek shores (e.g.,
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion sanctaemarthae" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="sanctaemarthae">Bembidion sanctaemarthae</taxonomicName>
,
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion ricei" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="ricei">Bembidion ricei</taxonomicName>
); others are found on clay or silt cliffs (e.g.,
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion agonoides" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="agonoides">Bembidion agonoides</taxonomicName>
,
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion walterrossii" order="Coleoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="walterrossii">Bembidion walterrossii</taxonomicName>
).
</paragraph>
</subSubSection>
</treatment>
</document>