459 lines
72 KiB
XML
459 lines
72 KiB
XML
<document ID-DOI="http://doi.org/10.5281/zenodo.5943535" ID-GBIF-Dataset="775fe351-8dad-4490-9d4d-34701f62b5c1" ID-GBIF-Taxon="151742855" ID-Zenodo-Dep="5943535" checkinTime="1548713912964" checkinUser="plazi" docAuthor="Evgeny Viktorovich Yana, Rolf Georg Beutel, John Francis Lawrence, Margarita Igorevna Yavorskaya, Thomas Hörnschemeyer, Hans Pohl, Dmitry Vladimirovich Vassilenko, Alexey Semenovich Bashkuev & Alexander Georgievich Ponomarenko" docDate="2019" docId="038E878E22492211F3F2FD35FE90F835" docLanguage="en" docName="historicalBiology.08912963.2018.1561672.pdf" docOrigin="Historical Biology 1561672" docStyle="DocumentStyle{}" docTitle="Archaeomalthus synoriakos Yan et al. 2019, gen. et sp. nov." docType="treatment" docVersion="4" lastPageId="6" lastPageNumber="7" masterDocId="FFB7FFF6224D2217F362FFCBFFB0FFD3" masterDocTitle="Archaeomalthus - (Coleoptera, Archostemata) a ‘ ghost adult ’ of Micromalthidae from Upper Permian deposits of Siberia?" masterLastPageNumber="9" masterPageNumber="1" originalUpdateDomain="Boston" originalUpdateTime="1547027548802" originalUpdateUser="plazi" pageId="4" pageNumber="5" updateTime="1643761532324" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>Archaeomalthus - (Coleoptera, Archostemata) a ‘ ghost adult ’ of Micromalthidae from Upper Permian deposits of Siberia?</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Evgeny Viktorovich Yana</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Rolf Georg Beutel</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>John Francis Lawrence</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Margarita Igorevna Yavorskaya</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Thomas Hörnschemeyer</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Hans Pohl</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Dmitry Vladimirovich Vassilenko</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Alexey Semenovich Bashkuev</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Alexander Georgievich Ponomarenko</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>Historical Biology</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2019</mods:date>
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<mods:detail type="pubDate">
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<mods:number>2019-01-07</mods:number>
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</mods:detail>
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<mods:detail type="volume">
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<mods:number>1561672</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>1</mods:start>
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<mods:end>9</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">10.1080/08912963.2018.1561672</mods:identifier>
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<mods:identifier type="GBIF-Dataset">775fe351-8dad-4490-9d4d-34701f62b5c1</mods:identifier>
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<mods:identifier type="Zenodo-Dep">2536029</mods:identifier>
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</mods:mods>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.5943535" ID-GBIF-Taxon="151742855" ID-Zenodo-Dep="5943535" LSID="urn:lsid:plazi:treatment:038E878E22492211F3F2FD35FE90F835" httpUri="http://treatment.plazi.org/id/038E878E22492211F3F2FD35FE90F835" lastPageId="6" lastPageNumber="7" pageId="4" pageNumber="5">
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<heading centered="true" fontSize="10" level="4" pageId="4" pageNumber="5" reason="4">
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||
<subSubSection box="[144,781,766,790]" pageId="4" pageNumber="5" type="nomenclature">
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||
<paragraph blockId="4.[144,781,766,822]" box="[144,781,766,790]" pageId="4" pageNumber="5">
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<emphasis bold="true" box="[144,483,766,790]" pageId="4" pageNumber="5">
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†
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<emphasis box="[158,775,766,790]" italics="true" pageId="4" pageNumber="5">
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<taxonomicName authority="Yan et al." authorityName="Yan et al." authorityYear="2019" box="[158,599,766,790]" class="Insecta" family="Micromalthidae" genus="Archaeomalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="synoriakos" status="gen. et sp. nov.">Archaeomalthus synoriakos Yan et al.</taxonomicName>
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,
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<taxonomicNameLabel box="[612,781,766,790]" pageId="4" pageNumber="5" rank="species">gen. et sp. nov.</taxonomicNameLabel>
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</emphasis>
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</emphasis>
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</paragraph>
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</subSubSection>
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<subSubSection box="[381,544,798,822]" pageId="4" pageNumber="5" type="description">
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<paragraph blockId="4.[144,781,766,822]" box="[381,544,798,822]" pageId="4" pageNumber="5">
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(
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<figureCitation box="[389,462,798,822]" captionStart="Figure 1" captionStartId="2.[128,183,987,1006]" captionTargetBox="[302,1325,140,960]" captionTargetId="figure@2.[301,1325,139,962]" captionTargetPageId="2" captionText="Figure 1. Archaeomalthus synoriacos sp. nov., Habitus. (a) Scanning electron micrograph of the holotype 5381/32. (b) Interpretative drawing." httpUri="https://zenodo.org/record/2536031/files/figure.png" pageId="4" pageNumber="5">Figs. 1</figureCitation>
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and
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<figureCitation box="[522,536,798,822]" captionStart="Figure 2" captionStartId="3.[128,183,1045,1064]" captionTargetBox="[253,1373,156,1020]" captionTargetId="figure@3.[253,1373,139,1020]" captionTargetPageId="3" captionText="Figure 2. Details of the organization in extinct and recent Micromalthidae. (a) Archaeomalthus synoriacos sp. nov. head, scanning electron micrograph. (b) A. synoriacos sp. nov. head interpretative drawing. (c) Micromalthus debilis, head dorsal aspect, scanning electron micrograph. (d) M. debilis interpretative drawing of head dorsal aspect. (e) M. debilis, head ventral aspect, scanning electron micrograph. (f) M. debilis interpretative drawing of head ventral aspect. (g) A. synoriacos sp. nov. pronotum, scanning electron micrograph. (h) A. synoriacos sp. nov. pronotum interpretative drawing. (i) A. synoriacos sp. nov. metathorax, scanning electron micrograph. (j) A. synoriacos sp. nov. metathorax interpretative drawing. (k) M. debilis, thoracic dorsal aspect, scanning electron micrograph. (l) M. debilis interpretative drawing of thoracic dorsal aspect. (m) M. debilis prothoracic lateral aspect, scanning electron micrograph. (n) M. debilis photograph of thoracic ventrall aspect. (m) M. debilis interpretative drawing of thoracic ventral aspect." httpUri="https://zenodo.org/record/2536033/files/figure.png" pageId="4" pageNumber="5">2</figureCitation>
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)
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</paragraph>
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</subSubSection>
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</heading>
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<subSubSection box="[128,317,877,935]" pageId="4" pageNumber="5" type="diagnosis">
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<paragraph blockId="4.[128,317,877,935]" box="[128,238,877,902]" pageId="4" pageNumber="5">
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<heading bold="true" box="[128,238,877,902]" fontSize="10" level="3" pageId="4" pageNumber="5" reason="6">
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<emphasis bold="true" box="[128,238,877,902]" italics="true" pageId="4" pageNumber="5">Diagnosis</emphasis>
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</heading>
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</paragraph>
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<paragraph blockId="4.[128,317,877,935]" box="[128,317,910,935]" pageId="4" pageNumber="5">As for the genus.</paragraph>
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</subSubSection>
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<subSubSection pageId="4" pageNumber="5" type="etymology">
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<paragraph blockId="4.[128,797,973,1094]" box="[128,351,973,998]" pageId="4" pageNumber="5">
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<heading bold="true" box="[128,351,973,998]" fontSize="10" level="3" pageId="4" pageNumber="5" reason="6">
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<emphasis bold="true" box="[128,351,973,998]" italics="true" pageId="4" pageNumber="5">Derivation of name</emphasis>
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</heading>
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</paragraph>
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<paragraph blockId="4.[128,797,973,1094]" pageId="4" pageNumber="5">The Greek-specific name ‘synoriakos’ – ‘borderline’, refers to the proximity of Babiy Kamen`locality to Permian-Triassic boundary.</paragraph>
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</subSubSection>
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<subSubSection pageId="4" pageNumber="5" type="materials_examined">
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<paragraph blockId="4.[128,797,1133,1286]" box="[128,222,1133,1158]" pageId="4" pageNumber="5">
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<heading bold="true" box="[128,222,1133,1158]" fontSize="10" level="3" pageId="4" pageNumber="5" reason="6">
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<emphasis bold="true" box="[128,222,1133,1158]" italics="true" pageId="4" pageNumber="5">Material</emphasis>
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</heading>
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</paragraph>
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<materialsCitation ID-GBIF-Occurrence="1981441626" collectionCode="Paleontological Institute, Russian Academy of Sciences" collectorName="Prof. Dr A. P. Rasnitsyn" location="Kamen`locality, Tom`river, Kuznetsk Basin, S. Siberian, in the upper part of the Maltsevo Formation" pageId="4" pageNumber="5" specimenCode="PIN 4887/102" specimenCount="1" typeStatus="holotype">
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<paragraph blockId="4.[128,797,1133,1286]" pageId="4" pageNumber="5">
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<typeStatus box="[128,231,1166,1190]" pageId="4" pageNumber="5">Holotype</typeStatus>
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:
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<specimenCode box="[246,403,1166,1190]" pageId="4" pageNumber="5">PIN 4887/102</specimenCode>
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, single
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<specimenCount box="[494,597,1166,1190]" pageId="4" pageNumber="5" type="generic">specimen</specimenCount>
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, counter print of the complete body impression with antennae and all legs missing. Repository:
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<collectionCode pageId="4" pageNumber="5">Paleontological Institute, Russian Academy of Sciences</collectionCode>
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(curator
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<collectorName box="[463,726,1262,1286]" pageId="4" pageNumber="5">Prof. Dr A.P. Rasnitsyn</collectorName>
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).
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</paragraph>
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<paragraph blockId="4.[128,797,1325,1414]" box="[128,254,1325,1350]" pageId="4" pageNumber="5">
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<heading bold="true" box="[128,254,1325,1350]" fontSize="10" level="3" pageId="4" pageNumber="5" reason="6">
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<emphasis bold="true" box="[128,254,1325,1350]" italics="true" pageId="4" pageNumber="5">Occurrence</emphasis>
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</heading>
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</paragraph>
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<paragraph blockId="4.[128,797,1325,1414]" pageId="4" pageNumber="5">
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Babiy
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<location LSID="urn:lsid:plazi:treatment:038E878E22492211F3F2FD35FE90F835:8EF8604322492213F3B4FA85FD41FA55" name="Kamen`locality, Tom`river, Kuznetsk Basin, S. Siberian, in the upper part of the Maltsevo Formation" pageId="4" pageNumber="5">Kamen`locality, Tom`river, Kuznetsk Basin, S. Siberian, in the upper part of the Maltsevo Formation</location>
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.
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</paragraph>
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</materialsCitation>
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</subSubSection>
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<subSubSection pageId="4" pageNumber="5" type="description">
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<paragraph blockId="4.[128,797,1453,2022]" box="[128,256,1453,1478]" pageId="4" pageNumber="5">
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<heading bold="true" box="[128,256,1453,1478]" fontSize="10" level="3" pageId="4" pageNumber="5" reason="6">
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<emphasis bold="true" box="[128,256,1453,1478]" italics="true" pageId="4" pageNumber="5">Description</emphasis>
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</heading>
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</paragraph>
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<paragraph blockId="4.[128,797,1453,2022]" pageId="4" pageNumber="5">Head slightly, ca. 0.23 mm, long and about as wide at the ocular region. Compound eyes round, located at anterior third of head, half as long as temporal region. Antennal insertion areas round, with outer margin bordered by fine carina. Antenna with small scapus, larger pedicellus with slightly rounded edges, and stout flagellomeres, apparently moniliform. The arrangement of apical mandibular teeth not recognizable. Prementum rather large, at least as wide as eye.</paragraph>
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<paragraph blockId="4.[128,797,1453,2022]" pageId="4" pageNumber="5">Anterior angles of pronotum blunt, not protruding forward; pronotal disc distinctly convex; anterior pronotal margin with median subtriangular depression; posterior angles slightly protruding laterally. Propleuron narrow, triangular, obliterating on anterior third of prothorax.</paragraph>
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<paragraph blockId="4.[128,797,1453,2022]" pageId="4" pageNumber="5">Elytra with fine lateral carina; epipleuron widest near humeral bulge, moderately wide on basal half, strongly narrowed posteriorly.</paragraph>
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<paragraph blockId="4.[829,1498,142,678]" pageId="4" pageNumber="5">Mesepimeron oblique, with slightly sinuated anterior and posterior margins. Mesocoxal pits closed laterally by mesepimeron and metanepisternum. Metanotum with distinct alacristae. Metanepisternum parallelogram-shaped. Metaventrite about as long as width between anterolateral edges, posterior edge about 1.5 times as wide.</paragraph>
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<paragraph blockId="4.[829,1498,142,678]" pageId="4" pageNumber="5">Procoxae transverse, mesocoxae almost globular, with inconspicuous lateral extension. Metacoxae short, with mesal part posteriorly rounded and extending through approximately two-thirds of first abdominal ventrite, only slightly narrowing laterally.</paragraph>
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<paragraph blockId="4.[829,1498,142,678]" pageId="4" pageNumber="5">Sternites II-IV distinctly longer than V-VII, gradually narrowing towards the abdominal apex. Short sternite VII with round median structure with unclear homology (apparently glandular opening). Terminal sternite VIII as long as VI and VII combined, with rounded hind margin. At least posterior abdominal sternites with narrow laterosternites..</paragraph>
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</subSubSection>
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<subSubSection lastPageId="6" lastPageNumber="7" pageId="4" pageNumber="5" type="discussion">
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<paragraph blockId="4.[829,962,749,774]" box="[829,962,749,774]" pageId="4" pageNumber="5">
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<heading bold="true" box="[829,962,749,774]" fontSize="11" level="1" pageId="4" pageNumber="5" reason="0">
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<emphasis bold="true" box="[829,962,749,774]" pageId="4" pageNumber="5">Discussion</emphasis>
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</heading>
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</paragraph>
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<paragraph blockId="4.[829,1291,798,822]" box="[829,1291,798,822]" pageId="4" pageNumber="5">
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<emphasis bold="true" box="[829,1291,798,822]" italics="true" pageId="4" pageNumber="5">
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Taxonomic position of
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<taxonomicName authority="Evgeny Viktorovich Yana & Rolf Georg Beutel & John Francis Lawrence & Margarita Igorevna Yavorskaya & Thomas Hörnschemeyer & Hans Pohl & Dmitry Vladimirovich Vassilenko & Alexey Semenovich Bashkuev & Alexander Georgievich Ponomarenko, 2019" authorityName="Evgeny Viktorovich Yana & Rolf Georg Beutel & John Francis Lawrence & Margarita Igorevna Yavorskaya & Thomas Hörnschemeyer & Hans Pohl & Dmitry Vladimirovich Vassilenko & Alexey Semenovich Bashkuev & Alexander Georgievich Ponomarenko" authorityYear="2019" box="[1101,1291,798,822]" class="Insecta" family="Micromalthidae" genus="Archaeomalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="3" phylum="Arthropoda" rank="genus" status="gen. nov.">Archaeomalthus</taxonomicName>
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</emphasis>
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</paragraph>
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<paragraph blockId="4.[829,1498,846,2022]" pageId="4" pageNumber="5">
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In contrast to specimens of
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<taxonomicName authorityName="LeConte" authorityYear="1878" box="[1164,1313,846,870]" class="Insecta" family="Micromalthidae" genus="Micromalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
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<emphasis box="[1164,1313,846,870]" italics="true" pageId="4" pageNumber="5">Micromalthus</emphasis>
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</taxonomicName>
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described from Miocene amber,which were not very different from the recent
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<taxonomicName authority="Hornschemeyer et al. 2010" authorityName="Hornschemeyer et al." authorityYear="2010" box="[915,1379,910,934]" class="Insecta" family="Micromalthidae" genus="Micromalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="debilis">
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<emphasis box="[915,1032,910,934]" italics="true" pageId="4" pageNumber="5">M. debilis</emphasis>
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<bibRefCitation author="Hornschemeyer T & Wedman S & Poinar G." box="[1052,1379,910,934]" journalOrPublisher="Zool JLinn Soc." pageId="4" pageNumber="5" pagination="300 - 311" part="158" refId="ref6273" refString="Hornschemeyer T, Wedman S, Poinar G. 2010. How long can insect species exist? Evidence from extant and fossil Micromalthus beetles (Insecta: Coleoptera). Zool JLinn Soc. 158: 300 - 311." title="How long can insect species exist? Evidence from extant and fossil Micromalthus beetles (Insecta: Coleoptera)" type="journal article" year="2010">Hörnschemeyer et al. 2010</bibRefCitation>
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</taxonomicName>
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, Permian
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<taxonomicName authority="Evgeny Viktorovich Yana & Rolf Georg Beutel & John Francis Lawrence & Margarita Igorevna Yavorskaya & Thomas Hörnschemeyer & Hans Pohl & Dmitry Vladimirovich Vassilenko & Alexey Semenovich Bashkuev & Alexander Georgievich Ponomarenko, 2019" authorityName="Evgeny Viktorovich Yana & Rolf Georg Beutel & John Francis Lawrence & Margarita Igorevna Yavorskaya & Thomas Hörnschemeyer & Hans Pohl & Dmitry Vladimirovich Vassilenko & Alexey Semenovich Bashkuev & Alexander Georgievich Ponomarenko" authorityYear="2019" box="[829,1002,942,966]" class="Insecta" family="Micromalthidae" genus="Archaeomalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="3" phylum="Arthropoda" rank="genus" status="gen. nov.">
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<emphasis box="[829,1002,942,966]" italics="true" pageId="4" pageNumber="5">Archaeomalthus</emphasis>
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</taxonomicName>
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shows distinct differences compared to the Recent species and Cenozoic fossils of the family. Nevertheless, the affinities of extant
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<taxonomicName authorityName=", Barber" authorityYear="1913" box="[1222,1395,1006,1030]" class="Insecta" family="Micromalthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="family">Micromalthidae</taxonomicName>
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are suffi- cient for justified placement in this family. This includes a number of diagnostic features: a small and slender body, the large relative size of the head capsule, and well-developed, round compound eyes. Clear apomorphies shared with fossil and extant
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<taxonomicName authorityName="LeConte" authorityYear="1878" box="[963,1112,1166,1190]" class="Insecta" family="Micromalthidae" genus="Micromalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
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<emphasis box="[963,1112,1166,1190]" italics="true" pageId="4" pageNumber="5">Micromalthus</emphasis>
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</taxonomicName>
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are shortened, quadrangular and posteriorly truncated elytra, leaving abdominal tergites V-VIII exposed, and the increased number of abdominal sternites, in contrast to five in ancestral Permian species (Stem group
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<taxonomicName authorityName=", Linnaeus" authorityYear="1758" box="[986,1106,1294,1318]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="order">Coleoptera</taxonomicName>
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) and the vast majority of extant beetles (
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<bibRefCitation author="Ponomarenko AG" box="[923,1144,1326,1350]" journalOrPublisher="Tr Palaeontol Inst Akad Ski SSSR" pageId="4" pageNumber="5" pagination="1 - 233" refId="ref7091" refString="Ponomarenko AG. 1969. Historical development of archostematan beetles. Tr Palaeontol Inst Akad Ski SSSR. 1 - 233." title="Historical development of archostematan beetles" type="book chapter" year="1969">Ponomarenko 1969</bibRefCitation>
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;
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<bibRefCitation author="Beutel RG & Ge SQ & Hornschemeyer T." box="[1166,1373,1326,1350]" journalOrPublisher="Cladistics." pageId="4" pageNumber="5" pagination="270 - 298" part="24" refId="ref5728" refString="Beutel RG, Ge SQ, Hornschemeyer T. 2008. On the head morphology of Tetraphalerus, the phylogeny of Archostemata and the basal branching events in Coleoptera. Cladistics. 24: 270 - 298." title="On the head morphology of Tetraphalerus, the phylogeny of Archostemata and the basal branching events in Coleoptera" type="journal article" year="2008">Beutel et al. 2008</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Lawrence JF & Slipinski A & Seago AE & Thayer MK & Newton AF & Marvaldi AE" journalOrPublisher="Ann Zool." pageId="4" pageNumber="5" pagination="1 - 217" part="61" refId="ref6630" refString="Lawrence JF, Slipinski A, Seago AE, Thayer MK, Newton AF, Marvaldi AE. 2011. Phylogeny of the Coleoptera based on morphological characters of adults and larvae. Ann Zool. 61: 1 - 217." title="Phylogeny of the Coleoptera based on morphological characters of adults and larvae" type="journal article" year="2011">Lawrence et al. 2011</bibRefCitation>
|
||
). Additional potential apomorphies are the fusion of the labrum and clypeus, the long subparallel temporal region, the stout antennal flagellomeres, the semioval scutellar shield (
|
||
<figureCitation box="[910,998,1454,1478]" captionStart="Figure 1" captionStartId="2.[128,183,987,1006]" captionTargetBox="[302,1325,140,960]" captionTargetId="figure@2.[301,1325,139,962]" captionTargetPageId="2" captionText="Figure 1. Archaeomalthus synoriacos sp. nov., Habitus. (a) Scanning electron micrograph of the holotype 5381/32. (b) Interpretative drawing." httpUri="https://zenodo.org/record/2536031/files/figure.png" pageId="4" pageNumber="5">Figure 1</figureCitation>
|
||
(b)), and probably, weak sclerotization of the body. Considering the placement in
|
||
<taxonomicName authorityName="Kolbe" authorityYear="1908" box="[1249,1399,1486,1510]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="subOrder" subOrder="Archostemata">Archostemata</taxonomicName>
|
||
and the ground plan of this suborder (and
|
||
<taxonomicName authorityName=", Linnaeus" authorityYear="1758" box="[1246,1364,1518,1542]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="order">Coleoptera</taxonomicName>
|
||
s.l. (
|
||
<bibRefCitation author="Beutel RG" journalOrPublisher="Verh Naturwiss Ver Hamburg (NF)." pageId="4" pageNumber="5" pagination="1 - 64" part="31" refId="ref5695" refString="Beutel RG. 1997. Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga. Verh Naturwiss Ver Hamburg (NF). 31: 1 - 64." title="Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga" type="journal article" year="1997">Beutel 1997</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Beutel RG & Ge SQ & Hornschemeyer T." box="[899,1099,1550,1574]" journalOrPublisher="Cladistics." pageId="4" pageNumber="5" pagination="270 - 298" part="24" refId="ref5728" refString="Beutel RG, Ge SQ, Hornschemeyer T. 2008. On the head morphology of Tetraphalerus, the phylogeny of Archostemata and the basal branching events in Coleoptera. Cladistics. 24: 270 - 298." title="On the head morphology of Tetraphalerus, the phylogeny of Archostemata and the basal branching events in Coleoptera" type="journal article" year="2008">Beutel et al. 2008</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Friedrich F & Farrell BD & Beutel RG" box="[1118,1351,1550,1574]" journalOrPublisher="Cladistics." pageId="4" pageNumber="5" pagination="1 - 37" part="25" refId="ref6026" refString="Friedrich F, Farrell BD, Beutel RG. 2009. The thoracic morphology of Archostemata and the relationships of the extant suborders of Coleoptera (Hexapoda). Cladistics. 25: 1 - 37." title="The thoracic morphology of Archostemata and the relationships of the extant suborders of Coleoptera (Hexapoda)" type="journal article" year="2009">Friedrich et al. 2009</bibRefCitation>
|
||
)), additional apomorphies shared with
|
||
<taxonomicName authorityName="LeConte" authorityYear="1878" box="[1126,1275,1582,1606]" class="Insecta" family="Micromalthidae" genus="Micromalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[1126,1275,1582,1606]" italics="true" pageId="4" pageNumber="5">Micromalthus</emphasis>
|
||
</taxonomicName>
|
||
are the absence of cuticular tubercles and scales, the complete lack of dorsal cephalic protuberances, the absence of a constricted neck region, evenly sclerotized elytra lacking window punctures, the absence of the transverse ridge on the mesoventrite, the concealed metatrochantins (
|
||
<bibRefCitation author="Beutel RG & Ge SQ & Hornschemeyer T." box="[1201,1491,1742,1766]" journalOrPublisher="Cladistics." pageId="4" pageNumber="5" pagination="270 - 298" part="24" refId="ref5728" refString="Beutel RG, Ge SQ, Hornschemeyer T. 2008. On the head morphology of Tetraphalerus, the phylogeny of Archostemata and the basal branching events in Coleoptera. Cladistics. 24: 270 - 298." title="On the head morphology of Tetraphalerus, the phylogeny of Archostemata and the basal branching events in Coleoptera" type="journal article" year="2008">Beutel et al. 2008</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Hornschemeyer T." box="[829,1060,1774,1798]" journalOrPublisher="Syst Entomol." pageId="4" pageNumber="5" pagination="533 - 558" part="32" refId="ref6169" refString="Hornschemeyer T. 2009. The species-level phylogeny of archostematan beetles - where do Micromalthus debilis and Crowsoniella relicta belong? Syst Entomol. 32: 533 - 558." title="The species-level phylogeny of archostematan beetles - where do Micromalthus debilis and Crowsoniella relicta belong?" type="journal article" year="2009">Hörnschemeyer 2009</bibRefCitation>
|
||
) (
|
||
<figureCitation box="[1086,1186,1774,1798]" captionStart="Figure 1" captionStartId="2.[128,183,987,1006]" captionTargetBox="[302,1325,140,960]" captionTargetId="figure@2.[301,1325,139,962]" captionTargetPageId="2" captionText="Figure 1. Archaeomalthus synoriacos sp. nov., Habitus. (a) Scanning electron micrograph of the holotype 5381/32. (b) Interpretative drawing." httpUri="https://zenodo.org/record/2536031/files/figure.png" pageId="4" pageNumber="5">Figures 1</figureCitation>
|
||
and
|
||
<figureCitation box="[1243,1256,1774,1798]" captionStart="Figure 2" captionStartId="3.[128,183,1045,1064]" captionTargetBox="[253,1373,156,1020]" captionTargetId="figure@3.[253,1373,139,1020]" captionTargetPageId="3" captionText="Figure 2. Details of the organization in extinct and recent Micromalthidae. (a) Archaeomalthus synoriacos sp. nov. head, scanning electron micrograph. (b) A. synoriacos sp. nov. head interpretative drawing. (c) Micromalthus debilis, head dorsal aspect, scanning electron micrograph. (d) M. debilis interpretative drawing of head dorsal aspect. (e) M. debilis, head ventral aspect, scanning electron micrograph. (f) M. debilis interpretative drawing of head ventral aspect. (g) A. synoriacos sp. nov. pronotum, scanning electron micrograph. (h) A. synoriacos sp. nov. pronotum interpretative drawing. (i) A. synoriacos sp. nov. metathorax, scanning electron micrograph. (j) A. synoriacos sp. nov. metathorax interpretative drawing. (k) M. debilis, thoracic dorsal aspect, scanning electron micrograph. (l) M. debilis interpretative drawing of thoracic dorsal aspect. (m) M. debilis prothoracic lateral aspect, scanning electron micrograph. (n) M. debilis photograph of thoracic ventrall aspect. (m) M. debilis interpretative drawing of thoracic ventral aspect." httpUri="https://zenodo.org/record/2536033/files/figure.png" pageId="4" pageNumber="5">2</figureCitation>
|
||
(m,n)) and abdominal glands (
|
||
<figureCitation box="[916,1014,1806,1830]" captionStart="Figure 1" captionStartId="2.[128,183,987,1006]" captionTargetBox="[302,1325,140,960]" captionTargetId="figure@2.[301,1325,139,962]" captionTargetPageId="2" captionText="Figure 1. Archaeomalthus synoriacos sp. nov., Habitus. (a) Scanning electron micrograph of the holotype 5381/32. (b) Interpretative drawing." httpUri="https://zenodo.org/record/2536031/files/figure.png" pageId="4" pageNumber="5">Figures 1</figureCitation>
|
||
(b) and 3)
|
||
</paragraph>
|
||
<paragraph blockId="4.[829,1498,846,2022]" lastBlockId="5.[128,797,782,2022]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
|
||
Shortened antennae, wings and legs of a Miocene micromalthid (lately
|
||
<emphasis box="[1025,1268,1870,1894]" italics="true" pageId="4" pageNumber="5">
|
||
<taxonomicName authorityName="LeConte" authorityYear="1878" box="[1025,1174,1870,1894]" class="Insecta" family="Micromalthidae" genus="Micromalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">Micromalthus</taxonomicName>
|
||
anansi
|
||
</emphasis>
|
||
was synonymized with
|
||
<taxonomicName box="[888,996,1902,1926]" class="Insecta" family="Micromalthidae" genus="Micromalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="debilis">
|
||
<emphasis box="[888,996,1902,1926]" italics="true" pageId="4" pageNumber="5">M. debilis</emphasis>
|
||
</taxonomicName>
|
||
by (
|
||
<bibRefCitation author="Hornschemeyer T & Wedman S & Poinar G." box="[1052,1356,1902,1926]" journalOrPublisher="Zool JLinn Soc." pageId="4" pageNumber="5" pagination="300 - 311" part="158" refId="ref6273" refString="Hornschemeyer T, Wedman S, Poinar G. 2010. How long can insect species exist? Evidence from extant and fossil Micromalthus beetles (Insecta: Coleoptera). Zool JLinn Soc. 158: 300 - 311." title="How long can insect species exist? Evidence from extant and fossil Micromalthus beetles (Insecta: Coleoptera)" type="journal article" year="2010">Hörnschemeyer et al. 2010</bibRefCitation>
|
||
)) were considered as a step of progressing morphological simplification of adults (
|
||
<bibRefCitation author="Perkovsky EE" box="[948,1126,1966,1990]" journalOrPublisher="Paleontol J." pageId="4" pageNumber="5" pagination="626 - 628" part="41" refId="ref6846" refString="Perkovsky EE. 2007. On the find of adult Micromalthus (Coleoptera, Micromalthidae) in Dominican amber. Paleontol J. 41: 626 - 628." title="On the find of adult Micromalthus (Coleoptera, Micromalthidae) in Dominican amber" type="journal article" year="2007">Perkovsky 2007</bibRefCitation>
|
||
). However, this term is problematic, as these features have obviously nothing to do with regressive character conditions occurring in beetles (e.g.
|
||
<bibRefCitation author="Rempel JG & Church NS" box="[128,453,813,838]" journalOrPublisher="Can J Zool." pageId="5" pageNumber="6" pagination="915 - 925" part="43" refId="ref7312" refString="Rempel JG, Church NS. 1965. The embryology of Lytta viridiana Le Conte (Coleoptera: Meloidae): I. Maturation, fertilization and cleavage. Can J Zool. 43: 915 - 925." title="The embryology of Lytta viridiana Le Conte (Coleoptera: Meloidae): I. Maturation, fertilization and cleavage" type="journal article" year="1965">Rempel and Church 1965</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Rempel JG & Church NS" box="[482,533,814,838]" journalOrPublisher="Can J Zool." pageId="5" pageNumber="6" pagination="1157 - 1171" part="47" refId="ref7351" refString="Rempel JG, Church NS. 1969. The embryology of Lytta viridana Le Conte (Coleoptera: Meloidae). V. The blastoderm, germ layers, and body segments. Can J Zool. 47: 1157 - 1171." title="The embryology of Lytta viridana Le Conte (Coleoptera: Meloidae). V. The blastoderm, germ layers, and body segments" type="journal article" year="1969">1969</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Rempel JG & Church NS" box="[563,614,814,838]" journalOrPublisher="Can J Zool." pageId="5" pageNumber="6" pagination="1571 - 1581" part="49" refId="ref7394" refString="Rempel JG, Church NS. 1971. The embryology of Lytta viridana Le Conte (Coleoptera: Meloidae). VII. Eighty-eight to 132 h: the appendages, the cephalic apodemes, and head segmentation. Can J Zool. 49: 1571 - 1581." title="The embryology of Lytta viridana Le Conte (Coleoptera: Meloidae). VII. Eighty-eight to 132 h: the appendages, the cephalic apodemes, and head segmentation" type="journal article" year="1971">1971</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Kuhne H." box="[644,790,814,838]" journalOrPublisher="JAppl Entomol." pageId="5" pageNumber="6" pagination="157 - 168" part="72" refId="ref6599" refString="Kuhne H. 1972. Entwicklungsablauf und-stadien von Micromalthus debilis LeConte (Col., Micromalthidae) aus einer Laboratoriums- Population. JAppl Entomol. 72: 157 - 168." title="Entwicklungsablauf und-stadien von Micromalthus debilis LeConte (Col., Micromalthidae) aus einer Laboratoriums- Population" type="journal article" year="1972">Kühne 1972</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Kobayashi Y & Niikura K & Oosawa Y & Takami Y." box="[128,361,846,870]" journalOrPublisher="J Morphol." pageId="5" pageNumber="6" pagination="1323 - 1352" part="274" refId="ref6419" refString="Kobayashi Y, Niikura K, Oosawa Y, Takami Y. 2013. Embryonic development of Carabus insulicola (Insecta, Coleoptera, Carabidae) with special reference to external morphology and tangible evidence for the subcoxal theory. J Morphol. 274: 1323 - 1352." title="Embryonic development of Carabus insulicola (Insecta, Coleoptera, Carabidae) with special reference to external morphology and tangible evidence for the subcoxal theory" type="journal article" year="2013">Kobayashi et al. 2013</bibRefCitation>
|
||
). It appears likely that small size, shortened elytra, and weakly sclerotized cuticle are related to the short lifespan and sporadic occurrence of adults, and their ‘vestigial’ status in the context of reproduction (
|
||
<bibRefCitation author="Pollock DA & Normark BB" journalOrPublisher="J Zool Syst Evol Res." pageId="5" pageNumber="6" pagination="105 - 112" part="40" refId="ref7050" refString="Pollock DA, Normark BB. 2002. The life cycle of Micromalthus debilis LeConte (Coleoptera: Archostemata: Micromalthidae): historical review and evolutionary perspective. J Zool Syst Evol Res. 40: 105 - 112." title="The life cycle of Micromalthus debilis LeConte (Coleoptera: Archostemata: Micromalthidae): historical review and evolutionary perspective" type="journal article" year="2002">Pollock and Normark 2002</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Hornschemeyer T & Wedman S & Poinar G." box="[305,601,974,998]" journalOrPublisher="Zool JLinn Soc." pageId="5" pageNumber="6" pagination="300 - 311" part="158" refId="ref6273" refString="Hornschemeyer T, Wedman S, Poinar G. 2010. How long can insect species exist? Evidence from extant and fossil Micromalthus beetles (Insecta: Coleoptera). Zool JLinn Soc. 158: 300 - 311." title="How long can insect species exist? Evidence from extant and fossil Micromalthus beetles (Insecta: Coleoptera)" type="journal article" year="2010">Hörnschemeyer et al. 2010</bibRefCitation>
|
||
). Relatively short legs are also common in other extant archostematan beetles including Permian forms (e.g.
|
||
<bibRefCitation author="Ponomarenko AG" box="[477,693,1038,1062]" journalOrPublisher="Tr Palaeontol Inst Akad Ski SSSR" pageId="5" pageNumber="6" pagination="1 - 233" refId="ref7091" refString="Ponomarenko AG. 1969. Historical development of archostematan beetles. Tr Palaeontol Inst Akad Ski SSSR. 1 - 233." title="Historical development of archostematan beetles" type="book chapter" year="1969">Ponomarenko 1969</bibRefCitation>
|
||
; see also
|
||
<bibRefCitation author="Yan EV & Lawrence JF & Beattie R & Beutel RG" box="[128,309,1070,1094]" journalOrPublisher="J Syst Palaeontol." pageId="5" pageNumber="6" pagination="1 - 9" part="16" refId="ref7810" refString="Yan EV, Lawrence JF, Beattie R, Beutel RG. 2017 a. At the dawn of the great rise: † Ponomarenkia belmonthensis (Insecta: Coleoptera), a remarkable new Late Permian beetle from the Southern Hemisphere. J Syst Palaeontol. 16: 1 - 9." title="At the dawn of the great rise: † Ponomarenkia belmonthensis (Insecta: Coleoptera), a remarkable new Late Permian beetle from the Southern Hemisphere" type="journal article" year="2017">Yan et al. 2017a</bibRefCitation>
|
||
) and moniliform antennae characterized as a plesiomorphic character by
|
||
<bibRefCitation author="Perkovsky EE" box="[455,646,1102,1126]" journalOrPublisher="Paleontol J." pageId="5" pageNumber="6" pagination="626 - 628" part="41" refId="ref6846" refString="Perkovsky EE. 2007. On the find of adult Micromalthus (Coleoptera, Micromalthidae) in Dominican amber. Paleontol J. 41: 626 - 628." title="On the find of adult Micromalthus (Coleoptera, Micromalthidae) in Dominican amber" type="journal article" year="2007">Perkovsky (2007)</bibRefCitation>
|
||
.
|
||
</paragraph>
|
||
<caption box="[128,1489,705,725]" httpUri="https://zenodo.org/record/2536035/files/figure.png" pageId="5" pageNumber="6" startId="5.[128,183,706,725]" targetBox="[412,1213,141,680]" targetPageId="5">
|
||
<paragraph blockId="5.[128,1489,705,725]" box="[128,1489,705,725]" pageId="5" pageNumber="6">
|
||
<emphasis bold="true" box="[128,206,706,725]" pageId="5" pageNumber="6">Figure 3.</emphasis>
|
||
Abdominal glands of
|
||
<taxonomicName box="[396,563,705,725]" class="Insecta" family="Micromalthidae" genus="Micromalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="species" species="debilis">
|
||
<emphasis box="[396,563,705,725]" italics="true" pageId="5" pageNumber="6">
|
||
Micromalthus
|
||
<taxonomicName box="[513,563,705,725]" pageId="5" pageNumber="6" rank="species" species="debilis">debilis</taxonomicName>
|
||
</emphasis>
|
||
</taxonomicName>
|
||
. (a) Ventrites III – V with median glandular pits. (b) Scanning electron micrograph of the glandular opening.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="5.[128,797,782,2022]" pageId="5" pageNumber="6">
|
||
One of several distinct features of
|
||
<taxonomicName authority="Evgeny Viktorovich Yana & Rolf Georg Beutel & John Francis Lawrence & Margarita Igorevna Yavorskaya & Thomas Hörnschemeyer & Hans Pohl & Dmitry Vladimirovich Vassilenko & Alexey Semenovich Bashkuev & Alexander Georgievich Ponomarenko, 2019" authorityName="Evgeny Viktorovich Yana & Rolf Georg Beutel & John Francis Lawrence & Margarita Igorevna Yavorskaya & Thomas Hörnschemeyer & Hans Pohl & Dmitry Vladimirovich Vassilenko & Alexey Semenovich Bashkuev & Alexander Georgievich Ponomarenko" authorityYear="2019" box="[525,698,1134,1158]" class="Insecta" family="Micromalthidae" genus="Archaeomalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="3" phylum="Arthropoda" rank="genus" status="gen. nov.">
|
||
<emphasis box="[525,698,1134,1158]" italics="true" pageId="5" pageNumber="6">Archaeomalthus</emphasis>
|
||
</taxonomicName>
|
||
differing from
|
||
<taxonomicName authorityName="LeConte" authorityYear="1878" box="[189,338,1166,1190]" class="Insecta" family="Micromalthidae" genus="Micromalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[189,338,1166,1190]" italics="true" pageId="5" pageNumber="6">Micromalthus</emphasis>
|
||
</taxonomicName>
|
||
is the presence of an exposed propleuron separated by distinct sutures. This is clearly a plesiomorphy corresponding to the autapomorphic fusion of all prothoracic sclerites in extant or Cenozoic species (e.g.
|
||
<bibRefCitation author="Beutel RG & Ge SQ & Hornschemeyer T." box="[603,790,1262,1286]" journalOrPublisher="Cladistics." pageId="5" pageNumber="6" pagination="270 - 298" part="24" refId="ref5728" refString="Beutel RG, Ge SQ, Hornschemeyer T. 2008. On the head morphology of Tetraphalerus, the phylogeny of Archostemata and the basal branching events in Coleoptera. Cladistics. 24: 270 - 298." title="On the head morphology of Tetraphalerus, the phylogeny of Archostemata and the basal branching events in Coleoptera" type="journal article" year="2008">Beutel et al. 2008</bibRefCitation>
|
||
; Friedrich et al. 2008). Two other conspicuous features are the presence of a very distinctly delimited groove on the mesoventrite and a corresponding anteromedian process of the metaventrite. Both features resemble conditions commonly found in Adephaga, notably the aquatic groups (excluding
|
||
<taxonomicName box="[680,789,1422,1446]" class="Insecta" family="Gyrinidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="family">Gyrinidae</taxonomicName>
|
||
) and basal grade
|
||
<taxonomicName authority="(Beutel 1992)" authorityName="Beutel" authorityYear="1992" baseAuthorityName="Beutel" baseAuthorityYear="1992" box="[361,662,1454,1478]" class="Insecta" family="Carabidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="family">
|
||
Carabidae (
|
||
<bibRefCitation author="Beutel RG" box="[504,651,1454,1478]" journalOrPublisher="Zool Syst Evol Forsch." pageId="5" pageNumber="6" pagination="53 - 74" part="30" refId="ref5668" refString="Beutel RG. 1992. Phylogenetic analysis of thoracic structures of Carabidae (Coleoptera). Zool Syst Evol Forsch. 30: 53 - 74." title="Phylogenetic analysis of thoracic structures of Carabidae (Coleoptera)" type="journal article" year="1992">Beutel 1992</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. However, a placement of
|
||
<taxonomicName authority="Evgeny Viktorovich Yana & Rolf Georg Beutel & John Francis Lawrence & Margarita Igorevna Yavorskaya & Thomas Hörnschemeyer & Hans Pohl & Dmitry Vladimirovich Vassilenko & Alexey Semenovich Bashkuev & Alexander Georgievich Ponomarenko, 2019" authorityName="Evgeny Viktorovich Yana & Rolf Georg Beutel & John Francis Lawrence & Margarita Igorevna Yavorskaya & Thomas Hörnschemeyer & Hans Pohl & Dmitry Vladimirovich Vassilenko & Alexey Semenovich Bashkuev & Alexander Georgievich Ponomarenko" authorityYear="2019" box="[312,485,1486,1510]" class="Insecta" family="Micromalthidae" genus="Archaeomalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="3" phylum="Arthropoda" rank="genus" status="gen. nov.">
|
||
<emphasis box="[312,485,1486,1510]" italics="true" pageId="5" pageNumber="6">Archaeomalthus</emphasis>
|
||
</taxonomicName>
|
||
in these suborders can be clearly excluded based on the short metacoxae without enlarged coxal plates, the lack of a prosternal process, and the increased number of abdominal sternites (e.g.
|
||
<bibRefCitation author="Beutel RG" box="[660,790,1582,1606]" journalOrPublisher="Zool Syst Evol Forsch." pageId="5" pageNumber="6" pagination="53 - 74" part="30" refId="ref5668" refString="Beutel RG. 1992. Phylogenetic analysis of thoracic structures of Carabidae (Coleoptera). Zool Syst Evol Forsch. 30: 53 - 74." title="Phylogenetic analysis of thoracic structures of Carabidae (Coleoptera)" type="journal article" year="1992">Beutel 1992</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Beutel RG & Haas F." box="[128,363,1614,1638]" journalOrPublisher="Cladistics." pageId="5" pageNumber="6" pagination="103 - 141" part="16" refId="ref5766" refString="Beutel RG, Haas F. 2000. Phylogenetic relationships of the suborders of Coleoptera (Insecta). Cladistics. 16: 103 - 141." title="Phylogenetic relationships of the suborders of Coleoptera (Insecta)" type="journal article" year="2000">Beutel and Haas 2000</bibRefCitation>
|
||
). The strongly developed process of the metaventrite, which widely separates the mesocoxae, is uncommon in recent
|
||
<taxonomicName authorityName="Kolbe" authorityYear="1908" box="[366,518,1678,1702]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="subOrder" subOrder="Archostemata">Archostemata</taxonomicName>
|
||
, even though it occurs in some extinct groups of this suborder, such as
|
||
<taxonomicName authorityName="Ponomarenko" authorityYear="1968" box="[631,797,1710,1734]" class="Insecta" family="Schizophoridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="family">Schizophoridae</taxonomicName>
|
||
(e.g
|
||
<taxonomicName authority="Martynov, 1926" authorityName="Martynov" authorityYear="1926" box="[181,442,1742,1766]" class="Insecta" family="Schizophoridae" genus="Tersus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[181,250,1742,1765]" italics="true" pageId="5" pageNumber="6">Tersus</emphasis>
|
||
Martynov, 1926
|
||
</taxonomicName>
|
||
) or
|
||
<taxonomicName authority="(Yan et al. 2017 b)" authorityName="Yan et al." authorityYear="2017" class="Insecta" family="Ademosynidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="family">
|
||
Ademosynidae (
|
||
<bibRefCitation author="Yan EV & Beutel RG & Ponomarenko AG" journalOrPublisher="Palaeontologia Electronica" pageId="5" pageNumber="6" refId="ref7766" refString="Yan EV, Beutel RG, Ponomarenko AG. 2017 b. Ademosynidae (Insecta: Coleoptera): A new concept for a coleopteran key taxon and its phylogenetic affinities to the extant suborders. Palaeontologia Electronica. doi: 10.26879 / 739" title="Ademosynidae (Insecta: Coleoptera): A new concept for a coleopteran key taxon and its phylogenetic affinities to the extant suborders" type="book" year="2017">Yan et al. 2017b</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. It is likely that this condition has evolved several times independently, in scattered groups of
|
||
<taxonomicName authorityName="Kolbe" authorityYear="1908" box="[647,797,1806,1830]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="subOrder" subOrder="Archostemata">Archostemata</taxonomicName>
|
||
including
|
||
<taxonomicName authorityName=", Barber" authorityYear="1913" box="[239,414,1838,1862]" class="Insecta" family="Micromalthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="family">Micromalthidae</taxonomicName>
|
||
, and also in Adephaga (e.g.
|
||
<bibRefCitation author="Beutel RG" journalOrPublisher="Stutt Beitrage zur Nat kd." pageId="5" pageNumber="6" pagination="1 - 54" part="388" refId="ref5622" refString="Beutel RG. 1986. Skelet und Muskulatur des Kopfes und Thorax von Hygrobia tarda (Herbst). Ein Beitrag zur Klarung der phylogenetischen Beziehungen der Hydradephaga (Insecta: Coleoptera). Stutt Beitrage zur Nat kd. 388: 1 - 54." title="Skelet und Muskulatur des Kopfes und Thorax von Hygrobia tarda (Herbst). Ein Beitrag zur Klarung der phylogenetischen Beziehungen der Hydradephaga (Insecta: Coleoptera)" type="journal article" year="1986">Beutel 1986</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Beutel RG" box="[210,261,1870,1894]" journalOrPublisher="Zool Syst Evol Forsch." pageId="5" pageNumber="6" pagination="53 - 74" part="30" refId="ref5668" refString="Beutel RG. 1992. Phylogenetic analysis of thoracic structures of Carabidae (Coleoptera). Zool Syst Evol Forsch. 30: 53 - 74." title="Phylogenetic analysis of thoracic structures of Carabidae (Coleoptera)" type="journal article" year="1992">1992</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Beutel RG & Haas F." box="[291,576,1870,1894]" journalOrPublisher="Cladistics." pageId="5" pageNumber="6" pagination="103 - 141" part="16" refId="ref5766" refString="Beutel RG, Haas F. 2000. Phylogenetic relationships of the suborders of Coleoptera (Insecta). Cladistics. 16: 103 - 141." title="Phylogenetic relationships of the suborders of Coleoptera (Insecta)" type="journal article" year="2000">Beutel and Haas 2000</bibRefCitation>
|
||
). In contrast to
|
||
<taxonomicName authority="Evgeny Viktorovich Yana & Rolf Georg Beutel & John Francis Lawrence & Margarita Igorevna Yavorskaya & Thomas Hörnschemeyer & Hans Pohl & Dmitry Vladimirovich Vassilenko & Alexey Semenovich Bashkuev & Alexander Georgievich Ponomarenko, 2019" authorityName="Evgeny Viktorovich Yana & Rolf Georg Beutel & John Francis Lawrence & Margarita Igorevna Yavorskaya & Thomas Hörnschemeyer & Hans Pohl & Dmitry Vladimirovich Vassilenko & Alexey Semenovich Bashkuev & Alexander Georgievich Ponomarenko" authorityYear="2019" box="[128,301,1902,1926]" class="Insecta" family="Micromalthidae" genus="Archaeomalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="3" phylum="Arthropoda" rank="genus" status="gen. nov.">
|
||
<emphasis box="[128,301,1902,1926]" italics="true" pageId="5" pageNumber="6">Archaeomalthus</emphasis>
|
||
</taxonomicName>
|
||
, the thoracic venter of
|
||
<taxonomicName authorityName="LeConte" authorityYear="1878" box="[557,706,1902,1926]" class="Insecta" family="Micromalthidae" genus="Micromalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[557,706,1902,1926]" italics="true" pageId="5" pageNumber="6">Micromalthus</emphasis>
|
||
</taxonomicName>
|
||
appears distinctly simplified (
|
||
<figureCitation box="[356,454,1934,1958]" captionStart="Figure 2" captionStartId="3.[128,183,1045,1064]" captionTargetBox="[253,1373,156,1020]" captionTargetId="figure@3.[253,1373,139,1020]" captionTargetPageId="3" captionText="Figure 2. Details of the organization in extinct and recent Micromalthidae. (a) Archaeomalthus synoriacos sp. nov. head, scanning electron micrograph. (b) A. synoriacos sp. nov. head interpretative drawing. (c) Micromalthus debilis, head dorsal aspect, scanning electron micrograph. (d) M. debilis interpretative drawing of head dorsal aspect. (e) M. debilis, head ventral aspect, scanning electron micrograph. (f) M. debilis interpretative drawing of head ventral aspect. (g) A. synoriacos sp. nov. pronotum, scanning electron micrograph. (h) A. synoriacos sp. nov. pronotum interpretative drawing. (i) A. synoriacos sp. nov. metathorax, scanning electron micrograph. (j) A. synoriacos sp. nov. metathorax interpretative drawing. (k) M. debilis, thoracic dorsal aspect, scanning electron micrograph. (l) M. debilis interpretative drawing of thoracic dorsal aspect. (m) M. debilis prothoracic lateral aspect, scanning electron micrograph. (n) M. debilis photograph of thoracic ventrall aspect. (m) M. debilis interpretative drawing of thoracic ventral aspect." httpUri="https://zenodo.org/record/2536033/files/figure.png" pageId="5" pageNumber="6">Figures 2</figureCitation>
|
||
(g–j,m,n)). It is conceivable that this is a reduction linked with the very limited role of the adults in the life cycle of the extant species.
|
||
</paragraph>
|
||
<paragraph blockId="5.[829,1498,781,1222]" pageId="5" pageNumber="6">
|
||
Despite the marked differences between
|
||
<taxonomicName authority="Evgeny Viktorovich Yana & Rolf Georg Beutel & John Francis Lawrence & Margarita Igorevna Yavorskaya & Thomas Hörnschemeyer & Hans Pohl & Dmitry Vladimirovich Vassilenko & Alexey Semenovich Bashkuev & Alexander Georgievich Ponomarenko, 2019" authorityName="Evgeny Viktorovich Yana & Rolf Georg Beutel & John Francis Lawrence & Margarita Igorevna Yavorskaya & Thomas Hörnschemeyer & Hans Pohl & Dmitry Vladimirovich Vassilenko & Alexey Semenovich Bashkuev & Alexander Georgievich Ponomarenko" authorityYear="2019" box="[1324,1497,782,806]" class="Insecta" family="Micromalthidae" genus="Archaeomalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="3" phylum="Arthropoda" rank="genus" status="gen. nov.">
|
||
<emphasis box="[1324,1497,782,806]" italics="true" pageId="5" pageNumber="6">Archaeomalthus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="LeConte" authorityYear="1878" box="[881,1030,814,838]" class="Insecta" family="Micromalthidae" genus="Micromalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[881,1030,814,838]" italics="true" pageId="5" pageNumber="6">Micromalthus</emphasis>
|
||
</taxonomicName>
|
||
, the former already show a distinct and apparently very stable degree of deviation from the ground plan of
|
||
<taxonomicName authorityName=", Linnaeus" authorityYear="1758" box="[1011,1129,878,902]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="order">Coleoptera</taxonomicName>
|
||
s.str. and s.l. (e.g.
|
||
<bibRefCitation author="Beutel RG" box="[1352,1482,878,902]" journalOrPublisher="Verh Naturwiss Ver Hamburg (NF)." pageId="5" pageNumber="6" pagination="1 - 64" part="31" refId="ref5695" refString="Beutel RG. 1997. Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga. Verh Naturwiss Ver Hamburg (NF). 31: 1 - 64." title="Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga" type="journal article" year="1997">Beutel 1997</bibRefCitation>
|
||
). The remarkable series of reductional features shared by the Permian fossil and the extant species include a lightly sclerotized body without any distinct surface sculpture, a head lacking dorsal protuberances, shortened and apparently thin elytra, exposed membranous areas behind the procoxae and between at least two basal visible abdominal sternites, and the reversal of the invagination of the terminal abdominal segments. It is likely that the reduced role of adults of
|
||
<taxonomicName authority="(Perotti et al. 2016)" authorityName="Perotti et al." authorityYear="2016" box="[829,1219,1166,1190]" class="Insecta" family="Micromalthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="family">
|
||
Micromalthidae (
|
||
<bibRefCitation author="Perotti MA & Young DK & Braig HR" box="[1016,1210,1166,1190]" journalOrPublisher="Sci Rep." pageId="5" pageNumber="6" pagination="27364" part="6" refId="ref6947" refString="Perotti MA, Young DK, Braig HR. 2016. The ghost sex-life of the paedogenetic beetle Micromalthus debilis. Sci Rep. 6: 27364." title="The ghost sex-life of the paedogenetic beetle Micromalthus debilis" type="journal article" year="2016">Perotti et al. 2016</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
was a syndrome of vestigial characters acquired at least 255 million years ago.
|
||
</paragraph>
|
||
<paragraph blockId="5.[829,1305,1278,1302]" box="[829,1305,1278,1302]" pageId="5" pageNumber="6">
|
||
<emphasis bold="true" box="[829,1305,1278,1302]" italics="true" pageId="5" pageNumber="6">
|
||
Evolutionary stability of
|
||
<taxonomicName box="[1121,1305,1278,1302]" class="Insecta" family="Micromalthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="family">Micromalthidae</taxonomicName>
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="5.[829,1498,1326,2022]" lastBlockId="6.[128,797,142,2022]" lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6">
|
||
There are two main competing hypotheses to explain clades that accumulate phenotypic variation at a slow rate: the first one invokes genetic and developmental constraints that restrict the production of phenotypic variation (
|
||
<bibRefCitation author="Smith JM" box="[1364,1491,1422,1446]" journalOrPublisher="Nature." pageId="5" pageNumber="6" pagination="13 - 14" part="289" refId="ref7668" refString="Smith JM. 1981. Macroevolution. Nature. 289: 13 - 14." title="Macroevolution" type="journal article" year="1981">Smith 1981</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Raff RA" box="[829,931,1454,1478]" journalOrPublisher="Chic: Univ of Chic Press" pageId="5" pageNumber="6" pagination="1 - 544" refId="ref7207" refString="Raff RA. 1996. The shape of life: genes, development, and the evolution of animal form. Chic: Univ of Chic Press; p. 1 - 544." title="The shape of life: genes, development, and the evolution of animal form" type="book chapter" year="1996">Raff 1996</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Gould SJ" box="[946,1073,1454,1478]" journalOrPublisher="Cambridge: Harv Univ Press" pageId="5" pageNumber="6" pagination="1 - 1433" refId="ref6117" refString="Gould SJ. 2002. The structure of evolutionary theory. Cambridge: Harv Univ Press. p. 1 - 1433." title="The structure of evolutionary theory" type="book chapter" year="2002">Gould 2002</bibRefCitation>
|
||
). The second explanation is that stasis is a result of stabilizing selection (
|
||
<bibRefCitation author="Charlesworth B & Lande R & Slatkin M." box="[1215,1491,1486,1510]" journalOrPublisher="Evol." pageId="5" pageNumber="6" pagination="474 - 498" part="36" refId="ref5849" refString="Charlesworth B, Lande R, Slatkin M. 1982. A Neo - Darwinian commentary on macroevolution. Evol. 36: 474 - 498." title="A Neo - Darwinian commentary on macroevolution" type="journal article" year="1982">Charlesworth et al. 1982</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Kirkpatrick M." box="[829,1011,1518,1542]" journalOrPublisher="Am Nat." pageId="5" pageNumber="6" pagination="833 - 848" part="119" refId="ref6395" refString="Kirkpatrick M. 1982. Quantum evolution and punctuated equilibria in continuous genetic characters. Am Nat. 119: 833 - 848." title="Quantum evolution and punctuated equilibria in continuous genetic characters" type="journal article" year="1982">Kirkpatrick 1982</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Estes S & Arnold SJ" box="[1025,1272,1518,1542]" journalOrPublisher="Am Nat." pageId="5" pageNumber="6" pagination="227 - 244" part="169" refId="ref5992" refString="Estes S, Arnold SJ. 2007. Resolving the paradox of stasis: models with stabilizing selection explain evolutionary divergence on all timescales. Am Nat. 169: 227 - 244." title="Resolving the paradox of stasis: models with stabilizing selection explain evolutionary divergence on all timescales" type="journal article" year="2007">Estes and Arnold 2007</bibRefCitation>
|
||
). Exceptional evolutionary stability, bradytely, or ‘arrested evolution’ (
|
||
<bibRefCitation author="Simpson GG" journalOrPublisher="New York: Columbia University Press" pageId="5" pageNumber="6" pagination="1 - 217" refId="ref7644" refString="Simpson GG. 1944. Tempo and mode in evolution. New York: Columbia University Press; p. 1 - 217." title="Tempo and mode in evolution" type="book chapter" year="1944">Simpson 1944</bibRefCitation>
|
||
) of
|
||
<taxonomicName box="[925,1100,1582,1606]" class="Insecta" family="Micromalthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="family">Micromalthidae</taxonomicName>
|
||
, resulted in very slow morphological change over time, which characterizes ‘living fossils’. On principle, this could be due to very long periods of stable environmental conditions (
|
||
<bibRefCitation author="Hornschemeyer T & Wedman S & Poinar G." box="[1135,1445,1678,1702]" journalOrPublisher="Zool JLinn Soc." pageId="5" pageNumber="6" pagination="300 - 311" part="158" refId="ref6273" refString="Hornschemeyer T, Wedman S, Poinar G. 2010. How long can insect species exist? Evidence from extant and fossil Micromalthus beetles (Insecta: Coleoptera). Zool JLinn Soc. 158: 300 - 311." title="How long can insect species exist? Evidence from extant and fossil Micromalthus beetles (Insecta: Coleoptera)" type="journal article" year="2010">Hörnschemeyer et al. 2010</bibRefCitation>
|
||
, see historical analysis of bradytely in
|
||
<bibRefCitation author="BuslovClarke D & Chatzimanolis S." journalOrPublisher="Cretac Res." pageId="5" pageNumber="6" pagination="1426 - 1434" part="30" refId="ref5793" refString="BuslovClarke D, Chatzimanolis S. 2009. Antiquity and long-term morphological stasis in a group of rove beetles (Coleoptera: staphylinidae): description of the oldest Octavius species, from Cretaceous Burmese amber and review of the " Euaesthetine subgroup " fossil record. Cretac Res. 30: 1426 - 1434." title="Antiquity and long-term morphological stasis in a group of rove beetles (Coleoptera: staphylinidae): description of the oldest Octavius species, from Cretaceous Burmese amber and review of the " Euaesthetine subgroup " fossil record" type="journal article" year="2009">Clarke and Chatzimanolis 2009</bibRefCitation>
|
||
). In the case of
|
||
<taxonomicName box="[1073,1246,1742,1766]" class="Insecta" family="Micromalthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="family">Micromalthidae</taxonomicName>
|
||
this is the subcortical space of fungus-infested wood, which may have been one factor that buffered micromalthids from strong selection for morphological change (
|
||
<bibRefCitation author="BuslovClarke D & Chatzimanolis S." journalOrPublisher="Cretac Res." pageId="5" pageNumber="6" pagination="1426 - 1434" part="30" refId="ref5793" refString="BuslovClarke D, Chatzimanolis S. 2009. Antiquity and long-term morphological stasis in a group of rove beetles (Coleoptera: staphylinidae): description of the oldest Octavius species, from Cretaceous Burmese amber and review of the " Euaesthetine subgroup " fossil record. Cretac Res. 30: 1426 - 1434." title="Antiquity and long-term morphological stasis in a group of rove beetles (Coleoptera: staphylinidae): description of the oldest Octavius species, from Cretaceous Burmese amber and review of the " Euaesthetine subgroup " fossil record" type="journal article" year="2009">BuslovClarke and Chatzimanolis 2009</bibRefCitation>
|
||
). Almost any change, but particularly a major change, of the phenotype in such a well-balanced system will be deleterious. Although minor gene substitution may be frequent, the well-buffered system of developmental canalizations shields the phenotype from major changes. There is an opportunity for speciation, but a major alteration of the morphotype is impossible as long as the epigenotype is intact (
|
||
<bibRefCitation author="Mayr E." box="[135,255,206,230]" journalOrPublisher="Camb: Belknap" pageId="6" pageNumber="7" pagination="1 - 454" refId="ref6672" refString="Mayr E. 1970. Populations, species, and evolution. Camb: Belknap; p. 1 - 454." title="Populations, species, and evolution" type="book chapter" year="1970">Mayr 1970</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="6.[128,797,142,2022]" pageId="6" pageNumber="7">
|
||
Association with prokaryotic endosymbionts or parasites can result in genome reduction and also an overcomplicated lifecycle including hypermetamorphosis and parthenogenesis. A long co-evolution with
|
||
<taxonomicName authority="Hertig, 1936" authorityName="Hertig" authorityYear="1936" box="[412,670,334,358]" class="Alphaproteobacteria" family="Rickettsiaceae" genus="Wolbachia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rickettsiales" pageId="6" pageNumber="7" phylum="Proteobacteria" rank="genus">
|
||
<emphasis box="[412,527,334,358]" italics="true" pageId="6" pageNumber="7">Wolbachia</emphasis>
|
||
Hertig, 1936
|
||
</taxonomicName>
|
||
is assumed in the case of
|
||
<taxonomicName box="[279,383,366,390]" class="Insecta" family="Micromalthidae" genus="Micromalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="debilis">
|
||
<emphasis box="[279,383,366,390]" italics="true" pageId="6" pageNumber="7">M. debilis</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Grimaldi D & Engel MS" box="[398,672,366,390]" journalOrPublisher="New York (NY): Cambridge University Press" pageId="6" pageNumber="7" pagination="1 - 755" refId="ref6140" refString="Grimaldi D, Engel MS. 2005. Evolution of the insects. New York (NY): Cambridge University Press; p. 1 - 755." title="Evolution of the insects" type="book chapter" year="2005">Grimaldi and Engel 2005</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Perkovsky EE" journalOrPublisher="Vestnik Zoologii." pageId="6" pageNumber="7" pagination="51 - 58" part="46" refId="ref6875" refString="Perkovsky EE. 2012. On the effect of latitudinal changes in summer temperatures on the Late Eocene aphid Germaraphis (Homoptera, Aphidoidea) and their symbiosis with ants. Vestnik Zoologii. 46: 51 - 58." title="On the effect of latitudinal changes in summer temperatures on the Late Eocene aphid Germaraphis (Homoptera, Aphidoidea) and their symbiosis with ants" type="journal article" year="2012">Perkovsky 2012</bibRefCitation>
|
||
). One of the results is apparently the ‘ghost sex life’, linked with a marginalization and vestigialization of the ‘ghost adults’ (
|
||
<bibRefCitation author="Perotti MA & Young DK & Braig HR" box="[284,474,462,486]" journalOrPublisher="Sci Rep." pageId="6" pageNumber="7" pagination="27364" part="6" refId="ref6947" refString="Perotti MA, Young DK, Braig HR. 2016. The ghost sex-life of the paedogenetic beetle Micromalthus debilis. Sci Rep. 6: 27364." title="The ghost sex-life of the paedogenetic beetle Micromalthus debilis" type="journal article" year="2016">Perotti et al. 2016</bibRefCitation>
|
||
). It is likely that the distinctly simplified adult morphology of
|
||
<taxonomicName authorityName="LeConte" authorityYear="1878" box="[471,620,494,518]" class="Insecta" family="Micromalthidae" genus="Micromalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[471,620,494,518]" italics="true" pageId="6" pageNumber="7">Micromalthus</emphasis>
|
||
</taxonomicName>
|
||
is closely linked with the obsolete role in the life cycle, and that this applies also to the lesser but already distinct reduction in
|
||
<taxonomicName authority="Evgeny Viktorovich Yana & Rolf Georg Beutel & John Francis Lawrence & Margarita Igorevna Yavorskaya & Thomas Hörnschemeyer & Hans Pohl & Dmitry Vladimirovich Vassilenko & Alexey Semenovich Bashkuev & Alexander Georgievich Ponomarenko, 2019" authorityName="Evgeny Viktorovich Yana & Rolf Georg Beutel & John Francis Lawrence & Margarita Igorevna Yavorskaya & Thomas Hörnschemeyer & Hans Pohl & Dmitry Vladimirovich Vassilenko & Alexey Semenovich Bashkuev & Alexander Georgievich Ponomarenko" authorityYear="2019" box="[128,301,590,614]" class="Insecta" family="Micromalthidae" genus="Archaeomalthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="3" phylum="Arthropoda" rank="genus" status="gen. nov.">
|
||
<emphasis box="[128,301,590,614]" italics="true" pageId="6" pageNumber="7">Archaeomalthus</emphasis>
|
||
</taxonomicName>
|
||
. The apparent stability of morphological simplification in the adult stage suggests a very old origin of a specialized mode of reproduction and possibly an equally old association with endosymbiotic organisms. Selective pressure on mechanical protection and a highly efficient locomotor apparatus in ‘ghost adults’ is likely minimal. Apparently, the resulting structural simplifications are irreversible once accomplished, even though the sex-life of adults can be revived as shown by
|
||
<bibRefCitation author="Perotti MA & Young DK & Braig HR" box="[381,612,846,870]" journalOrPublisher="Sci Rep." pageId="6" pageNumber="7" pagination="27364" part="6" refId="ref6947" refString="Perotti MA, Young DK, Braig HR. 2016. The ghost sex-life of the paedogenetic beetle Micromalthus debilis. Sci Rep. 6: 27364." title="The ghost sex-life of the paedogenetic beetle Micromalthus debilis" type="journal article" year="2016">Perotti et al. (2016)</bibRefCitation>
|
||
. In the case of
|
||
<taxonomicName box="[128,303,878,902]" class="Insecta" family="Micromalthidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="family">Micromalthidae</taxonomicName>
|
||
, the increasing degree of reduction from the Permian to the Miocene is likely a matter of reduced investment in adult structures, and therefore a result of ‘evolutionary economy’.
|
||
</paragraph>
|
||
<paragraph blockId="6.[128,797,142,2022]" pageId="6" pageNumber="7">
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A side effect of an association with symbionts can be a reduced ability to survive overheating (
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<bibRefCitation author="Dunbar HE & Wilson ACC & Ferguson NR & Moran NA" box="[582,790,1038,1062]" journalOrPublisher="PLoS Biol." pageId="6" pageNumber="7" pagination="96" part="5" refId="ref5957" refString="Dunbar HE, Wilson ACC, Ferguson NR, Moran NA. 2007. Aphid thermal tolerance is governed by a point mutation in bacterial symbionts. PLoS Biol. 5: e 96." title="Aphid thermal tolerance is governed by a point mutation in bacterial symbionts" type="journal article" year="2007">Dunbar et al. 2007</bibRefCitation>
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;
|
||
<bibRefCitation author="Perkovsky EE" box="[128,300,1070,1094]" journalOrPublisher="Vestnik Zoologii." pageId="6" pageNumber="7" pagination="51 - 58" part="46" refId="ref6875" refString="Perkovsky EE. 2012. On the effect of latitudinal changes in summer temperatures on the Late Eocene aphid Germaraphis (Homoptera, Aphidoidea) and their symbiosis with ants. Vestnik Zoologii. 46: 51 - 58." title="On the effect of latitudinal changes in summer temperatures on the Late Eocene aphid Germaraphis (Homoptera, Aphidoidea) and their symbiosis with ants" type="journal article" year="2012">Perkovsky 2012</bibRefCitation>
|
||
). It is likely that the immediate ancestors of extant micromalthid species were also confined to extratropical areas, while the climate of Western Europe in the Ypresian (Early Eocene) was undoubtedly macrothermal, which is confirmed by, among other things, the absence of any Holarctic elements in the amber, in particular, in the myrmecofauna of Oise (
|
||
<bibRefCitation author="Aria C & Perrichot V & Nel A." box="[192,363,1262,1286]" journalOrPublisher="Zootaxa." pageId="6" pageNumber="7" pagination="53 - 62" part="2870" refId="ref5480" refString="Aria C, Perrichot V, Nel A. 2011. Fossil ponerinae (Hymenoptera: Formicidae) in early Eocene amber of France. Zootaxa. 2870: 53 - 62." title="Fossil ponerinae (Hymenoptera: Formicidae) in early Eocene amber of France" type="journal article" year="2011">Aria et al. 2011</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Perkovsky EE" box="[378,548,1262,1286]" journalOrPublisher="Paleontol J." pageId="6" pageNumber="7" pagination="293 - 296" part="50" refId="ref6915" refString="Perkovsky EE. 2016. A new species of Micromalthidae (Coleoptera) from the Rovno Amber: 1. Adult morphology. Paleontol J. 50: 293 - 296." title="A new species of Micromalthidae (Coleoptera) from the Rovno Amber: 1. Adult morphology" type="journal article" year="2016">Perkovsky 2016</bibRefCitation>
|
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).
|
||
</paragraph>
|
||
<paragraph blockId="6.[128,797,142,2022]" pageId="6" pageNumber="7">
|
||
There are no specific paleoclimatic studies on Babiy Kamen`locality. However, some conclusions about climate during sediment accumulation could be made on the basis of paleofloristic analyses. The Kuznetsk Basin is mostly coal bearing, comprising industrial reserves of hard and brown coal. The process of coal-forming occurred on the former territory of a Carboniferous shallow marine bay, which was replaced by rather swampy lowland plains during the Permian. Alteration of coal and paleosols indicate sediment accumulation in rather humid environments, necessary for the increased accumulation of coal-forming biomass. For the upper part of the section, in which the fossil insects have been found (Dr E. Karasev PIN RAS, 2018 pers. comm. to DV), it is possible to assume alteration of humid and arid climatic conditions. Temperatures were not yet reconstructed for Babiy Kamen`locality. However, significant abundance of conifers (e.g.
|
||
<taxonomicName authority=", Elatocladus, Voltzia" authorityName="Elatocladus, Voltzia" box="[279,659,1806,1830]" class="Pinopsida" family="Pinopsida (awaiting allocation)" genus="Quadrocladus" higherTaxonomySource="GBIF" kingdom="Plantae" order="Pinopsida (awaiting allocation)" pageId="6" pageNumber="7" phylum="Tracheophyta" rank="genus">
|
||
<emphasis box="[279,659,1806,1830]" italics="true" pageId="6" pageNumber="7">Quadrocladus, Elatocladus, Voltzia</emphasis>
|
||
</taxonomicName>
|
||
) is reported from the upper part of the Maltsevo Formation (including insect-bearing layers) (
|
||
<bibRefCitation author="Betechtina OA & Mogutcheva NK & Batyaeva SK & Kushnarev MP" box="[380,622,1870,1894]" editor="Golbert AV" journalOrPublisher="Novosibirsk: Nauka" pageId="6" pageNumber="7" pagination="31 - 38" refId="ref5569" refString="Betechtina OA, Mogutcheva NK, Batyaeva SK, Kushnarev MP. 1986. Permian-Triassic boundary in the stratotype of the Maltsevo formation of Kuzbass. In: Golbert AV, editor. Biostratigraphy of the Mesozoic of Siberia and far east. Novosibirsk: Nauka; p. 31 - 38." title="Permian-Triassic boundary in the stratotype of the Maltsevo formation of Kuzbass" type="book chapter" volumeTitle="Biostratigraphy of the Mesozoic of Siberia and far east" year="1986">Betechtina et al. 1986</bibRefCitation>
|
||
). It was shown for Permian-Triassic deposits of the Severodvinian Basin that climatic cooling was also characterized by an increased proportion of conifers in Babiy Kamen`locality (
|
||
<bibRefCitation author="Krassilov V & Karasev E." journalOrPublisher="Paleogeogr Paleoclimatol Palaeoecol." pageId="6" pageNumber="7" pagination="326 - 336" part="284" refId="ref6569" refString="Krassilov V, Karasev E. 2009. Paleofloristic evidence of climate change near and beyond the Permian-Triassic boundary. Paleogeogr Paleoclimatol Palaeoecol. 284: 326 - 336." title="Paleofloristic evidence of climate change near and beyond the Permian-Triassic boundary" type="journal article" year="2009">Krassilov and Karasev 2009</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |