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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.662.12124" ID-GBIF-Dataset="85e41c76-28ec-4f0d-86cc-6a6ffd5f3784" ID-PMC="PMC5539362" ID-Pensoft-Pub="1313-2970-662-101" ID-PubMed="28769612" ID-ZBK="0250ADB4740A4DB283FA92E3BA0363D2" ModsDocAuthor="" ModsDocDate="2017" ModsDocID="1313-2970-662-101" ModsDocOrigin="ZooKeys 662" ModsDocTitle="The ground beetle genus Bembidion Latreille in Baltic amber: Review of preserved specimens and first 3D reconstruction of endophallic structures using X-ray microscopy (Coleoptera, Carabidae, Bembidiini)" checkinTime="1490252562655" checkinUser="pensoft" docAuthor="Schmidt, Joachim &amp; Michalik, Peter" docDate="2017" docId="EE50980CEEC849CA597155CA1DAF485D" docLanguage="en" docName="ZooKeys 662: 101-126" docOrigin="ZooKeys 662" docSource="http://dx.doi.org/10.3897/zookeys.662.12124" docTitle="Bembidion christelae Ortuno &amp; Arillo" docType="treatment" docVersion="4" lastPageNumber="103" masterDocId="4035B67CBA16C000FFD2B459FFD5FF83" masterDocTitle="The ground beetle genus Bembidion Latreille in Baltic amber: Review of preserved specimens and first 3 D reconstruction of endophallic structures using X-ray microscopy (Coleoptera, Carabidae, Bembidiini)" masterLastPageNumber="126" masterPageNumber="101" pageNumber="103" updateTime="1668164303151" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>The ground beetle genus Bembidion Latreille in Baltic amber: Review of preserved specimens and first 3 D reconstruction of endophallic structures using X-ray microscopy (Coleoptera, Carabidae, Bembidiini)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Schmidt, Joachim</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Michalik, Peter</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:title>ZooKeys</mods:title>
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<mods:date>2017</mods:date>
<mods:detail type="volume">
<mods:number>662</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>101</mods:start>
<mods:end>126</mods:end>
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<mods:url>http://dx.doi.org/10.3897/zookeys.662.12124</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.662.12124</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-662-101</mods:identifier>
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<mods:identifier type="ZooBank">0250ADB4740A4DB283FA92E3BA0363D2</mods:identifier>
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<treatment ID-GBIF-Taxon="127879721" LSID="urn:lsid:plazi:treatment:EE50980CEEC849CA597155CA1DAF485D" httpUri="http://treatment.plazi.org/id/EE50980CEEC849CA597155CA1DAF485D" lastPageId="3" lastPageNumber="103" pageId="2" pageNumber="103">
<subSubSection pageId="2" pageNumber="103" type="nomenclature">
<paragraph pageId="2" pageNumber="103">
<taxonomicName authority="Ortuno &amp; Arillo" class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion christelae" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="species" species="christelae">
<pageBreakToken pageId="2" pageNumber="103" start="start">Bembidion</pageBreakToken>
christelae
<normalizedToken originalValue="Ortuño">Ortuno</normalizedToken>
&amp; Arillo
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="103" type="reference_group">
<paragraph pageId="2" pageNumber="103">
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion (Archaeophilochthus) christelae" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="species" species="christelae" subGenus="Archaeophilochthus">Bembidion (Archaeophilochthus) christelae</taxonomicName>
<normalizedToken originalValue="Ortuño">Ortuno</normalizedToken>
&amp; Arillo, 2010: 190.
</paragraph>
</subSubSection>
<subSubSection lastPageId="3" lastPageNumber="104" pageId="2" pageNumber="103" type="remarks">
<paragraph pageId="2" pageNumber="103">Remarks on taxonomic position.</paragraph>
<paragraph pageId="2" pageNumber="103">
A monotypic subgenus
<taxonomicName class="Insecta" family="Carabidae" genus="Tachyina" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Tachyina (Archaeophilochthus)" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="subGenus" subGenus="Archaeophilochthus">Archaeophilochthus</taxonomicName>
<normalizedToken originalValue="Ortuño">Ortuno</normalizedToken>
&amp; Arillo, 2010 was established for the fossil
<taxonomicName lsidName="B. christelae" pageId="2" pageNumber="103" rank="species" species="christelae">B. christelae</taxonomicName>
. In the subgeneric diagnosis, the authors mentioned the following six features to be important for assigning the taxon: (i) elytral discal setae each are inserted in the third interval separated from the third stria; (ii) the groove of the rounded humeral margin ends &quot;close to the 4th or 5th striae&quot; (
<normalizedToken originalValue="Ortuño">Ortuno</normalizedToken>
and Arillo, 2010: 191, subgeneric diagnosis) and &quot;at the base of 6th interstriae&quot; (p. 190, species diagnosis), respectively; (iii) humeral setae of the elytral umbilicate series grouped and +/- equidistant; (iv) pronotal basolateral fovea poorly delimited; (v) pronotal laterobasal angles with carina well developed; vi) pronotal laterobasal angles formed oblique.
</paragraph>
<paragraph pageId="2" pageNumber="103">
Because the first five of these features are similarly developed in
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Philochthus" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Philochthus</taxonomicName>
Stephens, 1828,
<bibRefCitation author="Ortuno, VM" journalOrPublisher="Annales de la Societe Entomologique de France (Nouvelle serie)" pageId="9" pageNumber="110" pagination="189 - 192" title="Fossil carabids from Baltic amber - II - A new subgenus of Bembidion Latreille 1802 (Coleoptera: Carabidae: Bembidiini)." url="https://doi.org/10.1080/00379271.2010.10697656" volume="46" year="2010">
<normalizedToken originalValue="Ortuño">Ortuno</normalizedToken>
and Arillo (2010)
</bibRefCitation>
assume a close relationship of the fossil with this
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
subgenus. Character state (vi) was considered plesiomorphic by the authors, and because the apomorphic state 'pedunculated pronotal basal
<normalizedToken originalValue="margin">margin'</normalizedToken>
is developed in
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Philochthus" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Philochthus</taxonomicName>
the authors proposed a separate subgenus for the
<taxonomicName lsidName="B. christelae" pageId="2" pageNumber="103" rank="species" species="christelae">B. christelae</taxonomicName>
.
</paragraph>
<paragraph lastPageId="3" lastPageNumber="104" pageId="2" pageNumber="103">
Two extant
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
subgenera have been identified to be closely related to
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Philochthus" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Philochthus</taxonomicName>
(
<bibRefCitation author="Maddison, DR" journalOrPublisher="Molecular Phylogenetics and Evolution," pageId="9" pageNumber="110" pagination="533 - 576" title="Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidiini: Bembidiina)." url="http://dx.doi.org/10.1016/j.ympev.2012.01.015" volume="63" year="2012">Maddison 2012</bibRefCitation>
) and which should be therefore considered when discussing the probable relationships of
<taxonomicName lsidName="B. christelae" pageId="2" pageNumber="103" rank="species" species="christelae">B. christelae</taxonomicName>
:
<taxonomicName class="Insecta" family="Carabidae" genus="Lindrochthus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Lindrochthus" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Lindrochthus</taxonomicName>
Maddison, 2012, and
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthemphanes" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Philochthemphanes" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Philochthemphanes</taxonomicName>
Netolitzky, 1943. The former is monotypic and only includes
<taxonomicName lsidName="B. wickhami" pageId="2" pageNumber="103" rank="species" species="wickhami">B. wickhami</taxonomicName>
Hayward, 1897, a species endemic to north-western North America.
<taxonomicName class="Insecta" family="Carabidae" genus="Lindrochthus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Lindrochthus" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Lindrochthus</taxonomicName>
is distinguished from
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Philochthus" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Philochthus</taxonomicName>
&quot;by the less abruptly sinuate hind margin of the prothorax, and the reduced number of elytral striae&quot; (
<bibRefCitation author="Maddison, DR" journalOrPublisher="Molecular Phylogenetics and Evolution," pageId="9" pageNumber="110" pagination="533 - 576" title="Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidiini: Bembidiina)." url="http://dx.doi.org/10.1016/j.ympev.2012.01.015" volume="63" year="2012">Maddison 2012</bibRefCitation>
: 570). The pronotal margin is, however, more markedly sinuate in
<taxonomicName lsidName="B. wickhami" pageId="2" pageNumber="103" rank="species" species="wickhami">B. wickhami</taxonomicName>
than in the fossil
<taxonomicName lsidName="B. christelae" pageId="2" pageNumber="103" rank="species" species="christelae">B. christelae</taxonomicName>
, and the latter possesses eight completely developed elytral striae (
<bibRefCitation author="Ortuno, VM" journalOrPublisher="Annales de la Societe Entomologique de France (Nouvelle serie)" pageId="9" pageNumber="110" pagination="189 - 192" title="Fossil carabids from Baltic amber - II - A new subgenus of Bembidion Latreille 1802 (Coleoptera: Carabidae: Bembidiini)." url="https://doi.org/10.1080/00379271.2010.10697656" volume="46" year="2010">
<normalizedToken originalValue="Ortuño">Ortuno</normalizedToken>
and Arillo 2010
</bibRefCitation>
), which represents the plesiomorphic state.
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthemphanes" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Philochthemphanes" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Philochthemphanes</taxonomicName>
Netolitzky, 1943 (type species:
<taxonomicName lsidName="B. exquisitum" pageId="2" pageNumber="103" rank="species" species="exquisitum">B. exquisitum</taxonomicName>
Andrewes, 1923 from the Central Himalaya) is a
<taxonomicName class="Insecta" family="Carabidae" genus="Bembidion" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bembidion" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Bembidion</taxonomicName>
subgenus containing little more than ten species distributed in East and Southeast Asia (
<bibRefCitation author="Lorenz, W" journalOrPublisher="Published by the author, Tutzing" pageId="9" pageNumber="110" title="Systematic list of extant ground beetles of the world. Second edition." year="2005">Lorenz 2005</bibRefCitation>
,
<bibRefCitation author="Toledano, L" journalOrPublisher="Memorie del Museo Civico di Storia Naturale di Verona (II serie), Sezione Scienze della vita" pageId="9" pageNumber="110" pagination="95 - 100" title="Checklist of the Bembidiina of China (ColeopteraCarabidae)." volume="18" year="2008">Toledano 2008</bibRefCitation>
, database of J.S.) and is considered to be the sister group of
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Philochthus" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Philochthus</taxonomicName>
based on molecular data (
<bibRefCitation author="Maddison, DR" journalOrPublisher="Molecular Phylogenetics and Evolution," pageId="9" pageNumber="110" pagination="533 - 576" title="Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidiini: Bembidiina)." url="http://dx.doi.org/10.1016/j.ympev.2012.01.015" volume="63" year="2012">Maddison 2012</bibRefCitation>
). Species of
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthemphanes" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Philochthemphanes" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Philochthemphanes</taxonomicName>
possess a moderately sinuate pronotal basal margin as developed in
<taxonomicName lsidName="B. christelae" pageId="2" pageNumber="103" rank="species" species="christelae">B. christelae</taxonomicName>
. Most noteworthy,
<taxonomicName lsidName="B. christelae" pageId="2" pageNumber="103" rank="species" species="christelae">B. christelae</taxonomicName>
shows all diagnostic features of the exoskeleton compared to
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthemphanes" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Philochthemphanes" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Philochthemphanes</taxonomicName>
. It is thus very likely that the fossil taxon
<taxonomicName class="Insecta" family="Carabidae" genus="Archaeophilochthus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Archaeophilochthus" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Archaeophilochthus</taxonomicName>
is a junior synonym of
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthemphanes" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Philochthemphanes" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Philochthemphanes</taxonomicName>
. This assumption is further supported biogeographically as the flora and fauna of the Baltic amber forests is very similar to the modern biota of Indo-Malaya where
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthemphanes" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Philochthemphanes" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Philochthemphanes</taxonomicName>
is distributed nowadays (
<bibRefCitation author="Wolfe, JA" journalOrPublisher="Annals of the Missouri Botanical Garden" pageId="9" pageNumber="110" pagination="264 - 279" title="Some aspects of the plant geography of the Northern Hemisphere during the Late Cretaceous and Tertiary." url="https://doi.org/10.2307/2395198" volume="62" year="1975">Wolfe 1975</bibRefCitation>
,
<bibRefCitation author="Schmidt, J" journalOrPublisher="Zootaxa," pageId="9" pageNumber="110" pagination="573 - 581" title="On the Eocene age of Limodromus Motschulsky, 1850, with description of L. hoffeinsorum sp. n. from Baltic Amber (Coleoptera, Carabidae, Platynini)." url="https://doi.org/10.11646/zootaxa.3974.4.8" volume="3974" year="2015">Schmidt 2015</bibRefCitation>
). Here, extant species of
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthemphanes" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Philochthemphanes" order="Coleoptera" pageId="2" pageNumber="103" phylum="Arthropoda" rank="genus">Philochthemphanes</taxonomicName>
inhabit cloud forests and
<normalizedToken originalValue="forests">forest's</normalizedToken>
edges and the adults can be found on the ground as well as on the mossy branches of shrubs and trees, even though these species lack the common features present among most of the arboreal ground beetles.
<pageBreakToken pageId="3" pageNumber="104" start="start">Therefore</pageBreakToken>
and in contrast to the suggestion of
<bibRefCitation author="Ortuno, VM" journalOrPublisher="Annales de la Societe Entomologique de France (Nouvelle serie)" pageId="9" pageNumber="110" pagination="189 - 192" title="Fossil carabids from Baltic amber - II - A new subgenus of Bembidion Latreille 1802 (Coleoptera: Carabidae: Bembidiini)." url="https://doi.org/10.1080/00379271.2010.10697656" volume="46" year="2010">
<normalizedToken originalValue="Ortuño">Ortuno</normalizedToken>
and Arillo (2010)
</bibRefCitation>
,
<taxonomicName lsidName="B. christelae" pageId="3" pageNumber="104" rank="species" species="christelae">B. christelae</taxonomicName>
maybe had a semi-arboreal life style very similar to extant
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthemphanes" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Philochthemphanes" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Philochthemphanes</taxonomicName>
species. On the other hand,
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthemphanes" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Philochthemphanes" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Philochthemphanes</taxonomicName>
is not defined by clear synapomorphies. Thus, a careful revision and character analysis of
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthemphanes" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Philochthemphanes" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Philochthemphanes</taxonomicName>
is needed before a well-funded taxonomic conclusions regarding the synonymization of
<taxonomicName class="Insecta" family="Carabidae" genus="Archaeophilochthus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Archaeophilochthus" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Archaeophilochthus</taxonomicName>
and
<taxonomicName class="Insecta" family="Carabidae" genus="Philochthemphanes" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Philochthemphanes" order="Coleoptera" pageId="3" pageNumber="104" phylum="Arthropoda" rank="genus">Philochthemphanes</taxonomicName>
can be drawn.
</paragraph>
</subSubSection>
</treatment>
</document>