813 lines
115 KiB
XML
813 lines
115 KiB
XML
<document ID-DOI="http://doi.org/10.5281/zenodo.4328174" ID-GBIF-Dataset="faefcf9b-3673-4f50-b6a8-8a5d0cbd01de" ID-GBIF-Taxon="170718954" ID-Zenodo-Dep="4328174" approvalRequired="16" approvalRequired_for_document="1" approvalRequired_for_textStreams="15" checkinTime="1607453490458" checkinUser="plazi" docAuthor="Spangler, Paul J. & Steiner, Warren E." docDate="2005" docId="03C5879CFFC6FF8DFC96FF3BBD31F8FC" docLanguage="en" docName="systematicEntomology.30.3.339-357.pdf" docOrigin="Systematic Entomology 30 (3)" docSource="http://dx.doi.org/10.1111/j.1365-3113.2005.00288.x" docTitle="Meruidae Spangler & Steiner 2005, fam. nov." docType="treatment" docVersion="5" lastPageId="16" lastPageNumber="355" masterDocId="FFFCFFE4FFCAFF9DFFA3FFA0B963FFCF" masterDocTitle="A new aquatic beetle family, Meruidae, from Venezuela (Coleoptera: Adephaga)" masterLastPageNumber="357" masterPageNumber="339" pageId="12" pageNumber="351" updateTime="1643586909968" 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>A new aquatic beetle family, Meruidae, from Venezuela (Coleoptera: Adephaga)</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:namePart>Spangler, Paul J.</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>Steiner, Warren E.</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:titleInfo>
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<mods:title>Systematic Entomology</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2005</mods:date>
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<mods:detail type="pubDate">
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<mods:number>2005-03-07</mods:number>
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</mods:detail>
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<mods:detail type="volume">
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<mods:number>30</mods:number>
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</mods:detail>
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<mods:detail type="issue">
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<mods:number>3</mods:number>
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<mods:extent unit="page">
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<mods:start>339</mods:start>
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<mods:end>357</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:location>
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<mods:url>http://dx.doi.org/10.1111/j.1365-3113.2005.00288.x</mods:url>
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</mods:location>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">10.1111/j.1365-3113.2005.00288.x</mods:identifier>
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<mods:identifier type="GBIF-Dataset">faefcf9b-3673-4f50-b6a8-8a5d0cbd01de</mods:identifier>
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<mods:identifier type="Zenodo-Dep">4311852</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.4328174" ID-GBIF-Taxon="170718954" ID-Zenodo-Dep="4328174" LSID="urn:lsid:plazi:treatment:03C5879CFFC6FF8DFC96FF3BBD31F8FC" httpUri="http://treatment.plazi.org/id/03C5879CFFC6FF8DFC96FF3BBD31F8FC" lastPageId="16" lastPageNumber="355" pageId="12" pageNumber="351">
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<subSubSection box="[821,1040,155,177]" pageId="12" pageNumber="351" type="nomenclature">
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<paragraph blockId="12.[821,1040,155,177]" box="[821,1040,155,177]" pageId="12" pageNumber="351">
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<heading bold="true" box="[821,1040,155,177]" fontSize="9" level="1" pageId="12" pageNumber="351" reason="2">
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<emphasis bold="true" box="[821,1040,155,177]" pageId="12" pageNumber="351">
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<taxonomicName authority="Spangler & Steiner, 2005" authorityName="Spangler & Steiner" authorityYear="2005" box="[821,926,155,177]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="family" status="fam. nov.">Meruidae</taxonomicName>
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<taxonomicNameLabel box="[934,1040,155,177]" pageId="12" pageNumber="351" rank="family">fam. nov.</taxonomicNameLabel>
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</emphasis>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection box="[821,1038,214,235]" pageId="12" pageNumber="351" type="description">
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<paragraph blockId="12.[821,1038,214,235]" box="[821,1038,214,235]" pageId="12" pageNumber="351">
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<emphasis box="[821,1038,214,235]" italics="true" pageId="12" pageNumber="351">Systematic placement</emphasis>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="12" pageNumber="351" type="discussion">
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<paragraph blockId="12.[821,1427,273,850]" pageId="12" pageNumber="351">
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The combination of skeletal characters, habits and minute size in
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<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[895,1021,302,323]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="species" species="phyllisae">
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<emphasis box="[895,1021,302,323]" italics="true" pageId="12" pageNumber="351">M. phyllisae</emphasis>
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</taxonomicName>
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has offered a perplexing challenge for its classification. With the fusion of the hind coxae and metaventrite, the division of the first (of six) visible abdominal sternite (ventrite II divided by metacoxae) and fusion of visible sternites 1–3 (ventrites II–IV), the large propleuron and visible notopleural and pleurosternal sutures, the broad, lobed mentum, and glabrous palpiform two-segmented galea, suggest placement of
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<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[966,1021,536,557]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="genus">
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<emphasis box="[966,1021,536,557]" italics="true" pageId="12" pageNumber="351">Meru</emphasis>
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</taxonomicName>
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in the suborder
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<taxonomicName box="[1205,1312,536,558]" pageId="12" pageNumber="351" rank="subOrder" subOrder="Adephaga">Adephaga</taxonomicName>
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, according to the adult synapomorphies summarized by recent phylogenetic studies (
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<bibRefCitation author="Beutel, R. G." box="[1048,1181,594,616]" journalOrPublisher="Muzeum i Instytut Zoologii PAN, Warszawa" pageId="12" pageNumber="351" refId="ref9728" refString="Beutel, R. G. (1995) The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski), pp. 503 - 539. Muzeum i Instytut Zoologii PAN, Warszawa." title="The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski)" type="book" year="1995">Beutel, 1995</bibRefCitation>
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,
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<bibRefCitation author="Beutel, R." box="[1197,1247,594,616]" journalOrPublisher="Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF)" pageId="12" pageNumber="351" pagination="1 - 164" part="31" refId="ref9797" refString="Beutel, R. (1997) Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga. Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF), 31, 1 - 164." title="Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga" type="journal article" year="1997">1997</bibRefCitation>
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;
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<bibRefCitation author="Beutel, R. G. & Haas, A." journalOrPublisher="Cladistics" pageId="12" pageNumber="351" pagination="103 - 141" part="16" refId="ref9908" refString="Beutel, R. G. & Haas, A. (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 & Haas, 2000</bibRefCitation>
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). Glabrous antennomeres place it among the Hydradephaga, an aquatic adephagan beetle assemblage for which monophyly is still open to question (
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<bibRefCitation author="Beutel, R." journalOrPublisher="Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF)" pageId="12" pageNumber="351" pagination="1 - 164" part="31" refId="ref9797" refString="Beutel, R. (1997) Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga. Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF), 31, 1 - 164." title="Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga" type="journal article" year="1997">Beutel, 1997</bibRefCitation>
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;
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<bibRefCitation author="Ribera, I. & Hogan, J. E. & Vogler, A. P." box="[893,1134,711,733]" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="12" pageNumber="351" pagination="43 - 62" part="23" refId="ref10474" refString="Ribera, I., Hogan, J. E. & Vogler, A. P. (2002 a) Phylogeny of hydradephagan water beetles inferred from 18 S rRNA sequences. Molecular Phylogenetics and Evolution, 23, 43 - 62." title="Phylogeny of hydradephagan water beetles inferred from 18 S rRNA sequences" type="journal article" year="2002">
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Ribera
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<emphasis box="[981,1036,711,732]" italics="true" pageId="12" pageNumber="351">et al</emphasis>
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., 2002a
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</bibRefCitation>
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,b). For reasons discussed below, however, it has not been possible to place the genus in any known family taxon. We propose that it be placed in a new family,
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<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[1125,1226,799,821]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
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, the ‘comb-clawed cascade beetles’, based on
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<taxonomicName authority="Spangler & Steiner, 2005" authorityName="Spangler & Steiner" authorityYear="2005" box="[1107,1234,828,849]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="species" species="phyllisae" status="gen. et sp.n.">
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<emphasis box="[1107,1234,828,849]" italics="true" pageId="12" pageNumber="351">M. phyllisae</emphasis>
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</taxonomicName>
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<taxonomicNameLabel box="[1245,1371,828,850]" pageId="12" pageNumber="351" rank="species">
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<emphasis bold="true" box="[1245,1285,828,850]" pageId="12" pageNumber="351">gen.</emphasis>
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et
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<emphasis bold="true" box="[1325,1371,828,850]" pageId="12" pageNumber="351">sp.n.</emphasis>
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</taxonomicNameLabel>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="12" pageNumber="351" type="diagnosis">
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<paragraph blockId="12.[821,919,940,961]" box="[821,919,940,961]" pageId="12" pageNumber="351">
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<emphasis box="[821,919,940,961]" italics="true" pageId="12" pageNumber="351">Diagnosis</emphasis>
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</paragraph>
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<paragraph blockId="12.[821,1426,998,1049]" pageId="12" pageNumber="351">
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<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[845,944,998,1020]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
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can readily be identified by the following combination of characters:
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</paragraph>
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<paragraph blockId="12.[821,1426,1072,1387]" box="[821,1166,1072,1094]" pageId="12" pageNumber="351">1. Very small adephagan species.</paragraph>
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<paragraph blockId="12.[821,1426,1072,1387]" box="[821,1243,1102,1124]" pageId="12" pageNumber="351">2. Habitus resembling a minute haliplid.</paragraph>
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<paragraph blockId="12.[821,1426,1072,1387]" pageId="12" pageNumber="351">3. Rough body sculpture with flat, wrinkled setae (possibly respiratory structures).</paragraph>
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<paragraph blockId="12.[821,1426,1072,1387]" box="[821,1080,1189,1211]" pageId="12" pageNumber="351">4. Pectinate tarsal claws.</paragraph>
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<paragraph blockId="12.[821,1426,1072,1387]" box="[821,1374,1218,1240]" pageId="12" pageNumber="351">5. Trident tibial spurs (inner metatibial spur serrate).</paragraph>
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<paragraph blockId="12.[821,1426,1072,1387]" box="[821,1373,1248,1270]" pageId="12" pageNumber="351">6. Pair of overlapping spatulate setae on the labrum.</paragraph>
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<paragraph blockId="12.[821,1426,1072,1387]" box="[821,1229,1277,1299]" pageId="12" pageNumber="351">7. Lack of swimming hairs on the legs.</paragraph>
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<paragraph blockId="12.[821,1426,1072,1387]" box="[821,1091,1306,1328]" pageId="12" pageNumber="351">8. Small, rounded scapus.</paragraph>
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<paragraph blockId="12.[821,1426,1072,1387]" pageId="12" pageNumber="351">9. Alternating large and small segments of the antennal flagellum.</paragraph>
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</subSubSection>
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<subSubSection pageId="12" pageNumber="351" type="key">
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<paragraph blockId="12.[821,1427,1410,1578]" pageId="12" pageNumber="351">
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In the most recent key to adephagan families (
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<bibRefCitation author="Balke, M. & Ribera, I. & Beutel, R. G." journalOrPublisher="Zoologisch-Botanische Gesellschaft in Osterreich and Wiener Coleopterologenverein, Vienna" pageId="12" pageNumber="351" refId="ref9521" refString="Balke, M., Ribera, I. & Beutel, R. G. (2003) Aspidytidae: on the discovery of a new beetle family: detailed morphological analysis, description of a second species, and a key to fossil and extant adephagan families. Water Beetles of China, III (ed. by M. A. Jach and L. Ji), pp. 53 - 66. Zoologisch-Botanische Gesellschaft in Osterreich and Wiener Coleopterologenverein, Vienna." title="Aspidytidae: on the discovery of a new beetle family: detailed morphological analysis, description of a second species, and a key to fossil and extant adephagan families. Water Beetles of China, III (ed. by M. A. Jach and L. Ji)" type="book" year="2003">
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Balke
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<emphasis box="[1369,1413,1410,1431]" italics="true" pageId="12" pageNumber="351">et al</emphasis>
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., 2003
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</bibRefCitation>
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),
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<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[892,1018,1439,1460]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="species" species="phyllisae">
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<emphasis box="[892,1018,1439,1460]" italics="true" pageId="12" pageNumber="351">M. phyllisae</emphasis>
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</taxonomicName>
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runs to couplet 11,
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<taxonomicName box="[1234,1345,1439,1461]" class="Insecta" family="Dytiscidae" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="family">Dytiscidae</taxonomicName>
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, having obsolete metacoxal plates, but in
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<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[1167,1222,1468,1489]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="genus">
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<emphasis box="[1167,1222,1468,1489]" italics="true" pageId="12" pageNumber="351">Meru</emphasis>
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</taxonomicName>
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, the anterior extension of the metacoxae is obscured by complete fusion. Insertion of a couplet ‘11a’ at this point would separate
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<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[821,923,1556,1578]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
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:
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</paragraph>
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<paragraph blockId="12.[821,1427,1601,1681]" pageId="12" pageNumber="351">
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11a. Beetles less than
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long. Tarsal claws pectinate ................................................................
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<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[1328,1427,1630,1652]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
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– Beetles larger than
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<quantity box="[1049,1104,1659,1681]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="12" pageNumber="351" unit="mm" value="1.0">1 mm</quantity>
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long. Tarsal claws simple......11
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="16" lastPageNumber="355" pageId="12" pageNumber="351" type="discussion">
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<paragraph blockId="12.[821,1427,1705,1843]" pageId="12" pageNumber="351">
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Being less than
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<quantity box="[1016,1072,1705,1727]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="12" pageNumber="351" unit="mm" value="1.0">1 mm</quantity>
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in length,
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<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[1191,1320,1705,1726]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="species" species="phyllisae">
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<emphasis box="[1191,1320,1705,1726]" italics="true" pageId="12" pageNumber="351">M. phyllisae</emphasis>
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</taxonomicName>
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is among the smallest known members of
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<taxonomicName box="[1234,1344,1734,1756]" pageId="12" pageNumber="357" rank="subOrder" subOrder="Adephaga">Adephaga</taxonomicName>
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. Some
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<taxonomicName authorityName="Sharp" authorityYear="1882" box="[821,939,1763,1784]" class="Insecta" family="Noteridae" genus="Notomicrus" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="genus">
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||
<emphasis box="[821,939,1763,1784]" italics="true" pageId="12" pageNumber="351">Notomicrus</emphasis>
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</taxonomicName>
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(
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<taxonomicName baseAuthorityName="Beutel" baseAuthorityYear="1997" box="[960,1067,1763,1785]" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="family">Noteridae</taxonomicName>
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) species are barely longer than
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<quantity box="[821,880,1792,1814]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="12" pageNumber="351" unit="mm" value="1.0">1 mm</quantity>
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||
, as are the smallest
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<taxonomicName authorityName="Latreille" authorityYear="1802" box="[1120,1230,1792,1814]" class="Insecta" family="Carabidae" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="family">Carabidae</taxonomicName>
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, e.g. members of
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<taxonomicName authority="Jeannel." authorityName="Jeannel." box="[821,1048,1821,1843]" class="Insecta" family="Carabidae" genus="Geocharidius" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="genus">
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<emphasis box="[821,953,1822,1843]" italics="true" pageId="12" pageNumber="351">Geocharidius</emphasis>
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Jeannel.
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</taxonomicName>
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</paragraph>
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<caption ID-DOI="http://doi.org/10.5281/zenodo.4311886" ID-Zenodo-Dep="4311886" box="[151,610,1816,1836]" httpUri="https://zenodo.org/record/4311886/files/figure.png" pageId="12" pageNumber="351" startId="12.[151,185,1816,1836]" targetBox="[151,756,1070,1785]" targetPageId="12">
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||
<paragraph blockId="12.[151,610,1816,1836]" box="[151,610,1816,1836]" pageId="12" pageNumber="351">
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||
<emphasis bold="true" box="[151,222,1816,1836]" pageId="12" pageNumber="351">Fig. 16.</emphasis>
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<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[233,366,1817,1836]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="351" phylum="Arthropoda" rank="species" species="phyllisae">
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<emphasis box="[233,366,1817,1836]" italics="true" pageId="12" pageNumber="351">Meru phyllisae</emphasis>
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</taxonomicName>
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, proventriculus, end view.
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</paragraph>
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</caption>
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<paragraph blockId="13.[161,659,156,177]" box="[161,659,156,177]" pageId="13" pageNumber="352">
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<emphasis box="[161,659,156,177]" italics="true" pageId="13" pageNumber="352">
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Characters of Hydradephaga coded for
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[563,659,156,177]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="352" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="13.[161,767,214,762]" pageId="13" pageNumber="352">
|
||
In their placement of the newly proposed family
|
||
<taxonomicName authority=", Ribera et al. (2002 b)" authorityName="Ribera" authorityYear="2002" class="Insecta" family="Aspidytidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="352" phylum="Arthropoda" rank="family">
|
||
Aspidytidae,
|
||
<bibRefCitation author="Ribera, I. & Beutel, R. G. & Balke, M. & Vogler, A. P." box="[224,435,243,265]" journalOrPublisher="Proceedings of the Royal Society of London (B" pageId="13" pageNumber="352" pagination="2351 - 2356" part="269" refId="ref10419" refString="Ribera, I., Beutel, R. G., Balke, M. & Vogler, A. P. (2002 b) Discovery of Aspidytidae, a new family of aquatic Coleoptera. Proceedings of the Royal Society of London (B), 269, 2351 - 2356." title="Discovery of Aspidytidae, a new family of aquatic Coleoptera" type="journal article" year="2002">
|
||
Ribera
|
||
<emphasis box="[301,344,244,265]" italics="true" pageId="13" pageNumber="352">et al</emphasis>
|
||
. (2002b)
|
||
</bibRefCitation>
|
||
</taxonomicName>
|
||
listed forty adult morphological characters used in analyses to hypothesize relationships among the family taxa of Hydradephaga. Although a full cladistic analysis to determine the sister group of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[668,767,331,353]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="352" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
|
||
is beyond the scope of this paper (but will be the subject of ongoing and future studies), coding these characters at this point will assist in supporting the exclusion of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[653,708,419,440]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="352" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[653,708,419,440]" italics="true" pageId="13" pageNumber="352">Meru</emphasis>
|
||
</taxonomicName>
|
||
from other adephagan families, and point out some character systems that need additional scrutiny among many taxa. We were able to score most of these characters for
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[668,767,506,528]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="352" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
|
||
using character states defined by
|
||
<bibRefCitation author="Ribera, I. & Beutel, R. G. & Balke, M. & Vogler, A. P." box="[513,733,536,558]" journalOrPublisher="Proceedings of the Royal Society of London (B" pageId="13" pageNumber="352" pagination="2351 - 2356" part="269" refId="ref10419" refString="Ribera, I., Beutel, R. G., Balke, M. & Vogler, A. P. (2002 b) Discovery of Aspidytidae, a new family of aquatic Coleoptera. Proceedings of the Royal Society of London (B), 269, 2351 - 2356." title="Discovery of Aspidytidae, a new family of aquatic Coleoptera" type="journal article" year="2002">
|
||
Ribera
|
||
<emphasis box="[591,637,536,557]" italics="true" pageId="13" pageNumber="352">et al</emphasis>
|
||
. (2002b)
|
||
</bibRefCitation>
|
||
, as follows, with (0) being plesiomorphic:
|
||
</paragraph>
|
||
<paragraph blockId="13.[161,767,214,762]" lastBlockId="13.[830,1437,156,762]" pageId="13" pageNumber="352">Character 1, body shape: with distinct pronoto-elytral angle (0); 2, head shape: not shortened and laterally rounded, eyes protruding (0); 3, compound eyes: undivided (0); 4, shape of scapus: strongly shortened but without enlarged globular part (3); 5, pedicellus: not enlarged, not strongly shortened or enclosed by scapus (0); 6, flagellomeres of males: antennomeres 5 and more than one of the following segments distinctly broadened (2); 7, galea: two-segmented (0); 8, elongate sensorial field of distal labial palpomere: absent (0); 9, sensorial field on a protuberance of the dorsal side of the distal labial palpomere: absent (0); 10, shape of the prosternal process: strongly broadened and apically truncate (3); 11, ventral procoxal joint: with distinct coxal condyle (0); 12, profemoral cleaning device: absent (0); 13, protibial burrowing spur: absent (0); 14, row of flattened thorns on apical part of protibia: absent (0); 15, outer edge of protibia: not rounded (0); 16, tibial groove or concavity for reception of protibial burrowing spur: absent (0); 17, curved spurs on ventral side of protarsomeres 1–3: absent (0); 18, prothoracic defence gland: (0, probably; needs further study); 19, mesoventrite: short, with hexagonal groove and anterolateral grooves for reception of procoxae (0); 20, mesocoxae: globular (1); 21, mesocoxal cavity: laterally bordered by mesepimeron and metathoracic anepisternum (1, but difficult to interpret with certainty due to extensive fusion); 22, proximal pro- and mesotarsomeres: not elongated and broadened (0); 23, middle and hind legs: elongate (0); 24,</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.4311888" ID-Zenodo-Dep="4311888" httpUri="https://zenodo.org/record/4311888/files/figure.png" pageId="13" pageNumber="352" startId="13.[161,195,1710,1730]" targetBox="[161,1436,817,1681]" targetPageId="13">
|
||
<paragraph blockId="13.[161,1437,1710,1836]" pageId="13" pageNumber="352">
|
||
<emphasis bold="true" box="[161,232,1710,1730]" pageId="13" pageNumber="352">Fig. 17.</emphasis>
|
||
Biotope at El Tobogán, the type locality of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[653,787,1711,1730]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="352" phylum="Arthropoda" rank="species" species="phyllisae">
|
||
<emphasis box="[653,787,1711,1730]" italics="true" pageId="13" pageNumber="352">Meru phyllisae</emphasis>
|
||
</taxonomicName>
|
||
. A, The upper cascade looking upstream from the dam; B, the upper cascade looking downstream from the upper edge. C, The edge of the upper cascade showing the wet rock face and stream edge; D, the stream edge at the upper cascade where the root mat habitat was sampled; E, Biotope at El Tobogán, the type locality of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[1186,1317,1764,1783]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="352" phylum="Arthropoda" rank="species" species="phyllisae">
|
||
<emphasis box="[1186,1317,1764,1783]" italics="true" pageId="13" pageNumber="352">Meru phyllisae</emphasis>
|
||
</taxonomicName>
|
||
. A, The dam and pool below the upper cascade looking upstream; F, the pool and forested stream below the cascade looking downstream; G, Section of forested stream below the cascade, looking downstream; H, the open rock surface and edge of a rapid section of the cascade.
|
||
</paragraph>
|
||
</caption>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.4311890" ID-Zenodo-Dep="4311890" box="[151,329,1070,1090]" httpUri="https://zenodo.org/record/4311890/files/figure.png" pageId="14" pageNumber="353" startId="14.[151,185,1070,1090]" targetBox="[151,1426,608,1041]" targetPageId="14">
|
||
<paragraph blockId="14.[151,329,1070,1090]" box="[151,329,1070,1090]" pageId="14" pageNumber="353">
|
||
<emphasis bold="true" box="[151,222,1070,1090]" pageId="14" pageNumber="353">
|
||
<figureCitation box="[151,219,1070,1090]" captionStart="Fig" captionStartId="13.[161,195,1710,1730]" captionTargetBox="[161,1436,817,1681]" captionTargetPageId="13" captionText="Fig. 17. Biotope at El Tobogán, the type locality of Meru phyllisae. A, The upper cascade looking upstream from the dam; B, the upper cascade looking downstream from the upper edge. C, The edge of the upper cascade showing the wet rock face and stream edge; D, the stream edge at the upper cascade where the root mat habitat was sampled; E, Biotope at El Tobogán, the type locality of Meru phyllisae. A, The dam and pool below the upper cascade looking upstream; F, the pool and forested stream below the cascade looking downstream; G, Section of forested stream below the cascade, looking downstream; H, the open rock surface and edge of a rapid section of the cascade." figureDoi="http://doi.org/10.5281/zenodo.4311888" httpUri="https://zenodo.org/record/4311888/files/figure.png" pageId="14" pageNumber="353">Fig. 17</figureCitation>
|
||
,
|
||
</emphasis>
|
||
<emphasis box="[233,323,1071,1090]" italics="true" pageId="14" pageNumber="353">Continued</emphasis>
|
||
.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="14.[151,758,1176,1607]" pageId="14" pageNumber="353">swimming hairs on meso- and metathoracic legs: absent (0); 25, noterid platform of metaventrite: absent (0); 26, transverse ridge of metaventrite: absent (2); 27, contact between pro- and metasternal process: present (1); 28, metafurca: originates from intercoxal septum (1); 29, size of metafurca: narrow, with reduced lateral arms (1); 30, mesal walls of metacoxae: with extensive contact area and intercoxal septum (3); 31, anterior margin of metacoxa: undetermined due to extensive fusion; 32, metacoxal plates: largely reduced (4); 33, lateral margin of metacoxal plates: indistinct anteriorly (4); 34–35, muscle characters (not studied); 36, abdominal segments III and IV: completely fused (1); 37, bulges on anterior abdominal sternites: absent (0); 38, gonocoxosterna VIII: not exposed (0); 39–40, ramen and laterotergite: (unsclerotized; not observed).</paragraph>
|
||
<paragraph blockId="14.[151,608,1646,1667]" box="[151,608,1646,1667]" pageId="14" pageNumber="353">
|
||
<emphasis box="[151,608,1646,1667]" italics="true" pageId="14" pageNumber="353">Exclusion from other hydradephagan families</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="14.[151,757,1705,1844]" lastBlockId="14.[821,1427,1176,1841]" pageId="14" pageNumber="353">
|
||
Although
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[282,337,1705,1726]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="353" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[282,337,1705,1726]" italics="true" pageId="14" pageNumber="353">Meru</emphasis>
|
||
</taxonomicName>
|
||
keys to
|
||
<taxonomicName box="[430,541,1705,1727]" class="Insecta" family="Dytiscidae" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="353" phylum="Arthropoda" rank="family">Dytiscidae</taxonomicName>
|
||
, it must be excluded from that family because of the undifferentiated legs, lack of swimming fringes, and alternating leg movement during swimming (
|
||
<bibRefCitation author="Ribera, I. & Beutel, R. G. & Balke, M. & Vogler, A. P." box="[269,478,1792,1814]" journalOrPublisher="Proceedings of the Royal Society of London (B" pageId="14" pageNumber="353" pagination="2351 - 2356" part="269" refId="ref10419" refString="Ribera, I., Beutel, R. G., Balke, M. & Vogler, A. P. (2002 b) Discovery of Aspidytidae, a new family of aquatic Coleoptera. Proceedings of the Royal Society of London (B), 269, 2351 - 2356." title="Discovery of Aspidytidae, a new family of aquatic Coleoptera" type="journal article" year="2002">
|
||
Ribera
|
||
<emphasis box="[347,391,1793,1814]" italics="true" pageId="14" pageNumber="353">et al</emphasis>
|
||
., 2002b
|
||
</bibRefCitation>
|
||
). The oarlike hind legs and often highly streamlined body of ‘diving beetles’ separate
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[821,920,1176,1198]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="353" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
|
||
and other aquatic taxa that are less equipped for rapid and evasive swimming.
|
||
<bibRefCitation author="Roughley, R. E. & Larson, J. D." box="[1140,1426,1205,1227]" journalOrPublisher="American Beetles, Vol. 1 (ed. by R. H. Arnett, Jr and M. C. Thomas), pp. 156 - 186. CRC Press, Boca Raton, Florida" pageId="14" pageNumber="353" refId="ref10712" refString="Roughley, R. E. & Larson, J. D. (2001) Dytiscidae Leach, 1815. American Beetles, Vol. 1 (ed. by R. H. Arnett, Jr and M. C. Thomas), pp. 156 - 186. CRC Press, Boca Raton, Florida." title="Dytiscidae Leach" type="book" year="2001">Roughley & Larson (2001)</bibRefCitation>
|
||
reviewed the characters, classification and literature on the
|
||
<taxonomicName box="[821,932,1264,1286]" class="Insecta" family="Dytiscidae" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="353" phylum="Arthropoda" rank="family">Dytiscidae</taxonomicName>
|
||
;
|
||
<bibRefCitation author="Miller, K. B." box="[942,1076,1264,1286]" journalOrPublisher="Insect Systematics and Evolution" pageId="14" pageNumber="353" pagination="45 - 92" part="32" refId="ref10321" refString="Miller, K. B. (2001) On the phylogeny of the Dytiscidae (Insecta: Coleoptera) with emphasis on the morphology of the female reproductive system. Insect Systematics and Evolution, 32, 45 - 92." title="On the phylogeny of the Dytiscidae (Insecta: Coleoptera) with emphasis on the morphology of the female reproductive system" type="journal article" year="2001">Miller (2001)</bibRefCitation>
|
||
provided the most recent analysis of morphology and phylogeny.
|
||
</paragraph>
|
||
<paragraph blockId="14.[821,1427,1176,1841]" pageId="14" pageNumber="353">
|
||
Apomorphic features of
|
||
<taxonomicName box="[1098,1201,1322,1344]" class="Insecta" family="Gyrinidae" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="353" phylum="Arthropoda" rank="family">Gyrinidae</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Steiner, W. E. Jr & Anderson, J. J." journalOrPublisher="Pan-Pacific Entomologist" pageId="14" pageNumber="353" pagination="124 - 132" part="57" refId="ref11013" refString="Steiner, W. E. Jr & Anderson, J. J. (1981) Notes on the natural history of Spanglerogyrus albiventris Folkerts, with a new distribution record (Coleoptera: Gyrinidae). Pan-Pacific Entomologist, 57, 124 - 132." title="Notes on the natural history of Spanglerogyrus albiventris Folkerts, with a new distribution record (Coleoptera: Gyrinidae)" type="journal article" year="1981">Steiner & Anderson, 1981</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Beutel, R. G. & Roughley, R. E." box="[878,1126,1351,1373]" journalOrPublisher="Zeitschrift fur Zoologische Systematik und Evolutionsforschung" pageId="14" pageNumber="353" pagination="380 - 400" part="26" refId="ref9983" refString="Beutel, R. G. & Roughley, R. E. (1988) On the systematic position of the family Gyrinidae (Coleoptera: Adephaga). Zeitschrift fur Zoologische Systematik und Evolutionsforschung, 26, 380 - 400." title="On the systematic position of the family Gyrinidae (Coleoptera: Adephaga)" type="journal article" year="1988">Beutel & Roughley, 1988</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Beutel, R. G." box="[1135,1258,1351,1373]" journalOrPublisher="Quaestiones Entomologicae" pageId="14" pageNumber="353" pagination="163 - 191" part="26" refId="ref9694" refString="Beutel, R. G. (1990) Phylogenetic analysis of the family Gyrinidae (Coleoptera) based on meso- and metathoracic characters. Quaestiones Entomologicae, 26, 163 - 191." title="Phylogenetic analysis of the family Gyrinidae (Coleoptera) based on meso- and metathoracic characters" type="journal article" year="1990">Beutel, 1990</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Beutel, R. G." box="[1266,1315,1351,1373]" journalOrPublisher="Muzeum i Instytut Zoologii PAN, Warszawa" pageId="14" pageNumber="353" refId="ref9728" refString="Beutel, R. G. (1995) The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski), pp. 503 - 539. Muzeum i Instytut Zoologii PAN, Warszawa." title="The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski)" type="book" year="1995">1995</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Roughley, R. E." journalOrPublisher="American Beetles, Vol. 1 (ed. by R. H. Arnett, Jr and M. C. Thomas), pp. 133 - 137. CRC Press, Boca Raton, Florida" pageId="14" pageNumber="353" refId="ref10546" refString="Roughley, R. E. (2001 a) Gyrinidae Latreille, 1810. American Beetles, Vol. 1 (ed. by R. H. Arnett, Jr and M. C. Thomas), pp. 133 - 137. CRC Press, Boca Raton, Florida." title="Gyrinidae Latreille" type="book" year="2001">Roughley, 2001a</bibRefCitation>
|
||
) include divided eyes (for surface swimming), highly modified legs for swimming, stout and highly specialized antennae, and a single protibial spur.
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[1371,1426,1439,1460]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="353" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[1371,1426,1439,1460]" italics="true" pageId="14" pageNumber="353">Meru</emphasis>
|
||
</taxonomicName>
|
||
shares none of these features but has synapomorphies with the remaining families of
|
||
<taxonomicName box="[1160,1267,1497,1519]" pageId="14" pageNumber="357" rank="subOrder" subOrder="Adephaga">Adephaga</taxonomicName>
|
||
, e.g. the short mesosternite, form of the male genitalia (curved asymmetric base, torsion) and other characters that would exclude it from
|
||
<taxonomicName box="[1067,1170,1585,1607]" class="Insecta" family="Gyrinidae" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="353" phylum="Arthropoda" rank="family">Gyrinidae</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Beutel, R. G." box="[1187,1320,1585,1607]" journalOrPublisher="Muzeum i Instytut Zoologii PAN, Warszawa" pageId="14" pageNumber="353" refId="ref9728" refString="Beutel, R. G. (1995) The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski), pp. 503 - 539. Muzeum i Instytut Zoologii PAN, Warszawa." title="The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski)" type="book" year="1995">Beutel, 1995</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Beutel, R." box="[1335,1387,1585,1607]" journalOrPublisher="Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF)" pageId="14" pageNumber="353" pagination="1 - 164" part="31" refId="ref9797" refString="Beutel, R. (1997) Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga. Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF), 31, 1 - 164." title="Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga" type="journal article" year="1997">1997</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="14.[821,1427,1176,1841]" lastBlockId="15.[161,767,156,1464]" lastPageId="15" lastPageNumber="354" pageId="14" pageNumber="353">
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[845,993,1615,1636]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="353" phylum="Arthropoda" rank="species" species="phyllisae">
|
||
<emphasis box="[845,993,1615,1636]" italics="true" pageId="14" pageNumber="353">Meru phyllisae</emphasis>
|
||
</taxonomicName>
|
||
is the ‘undescribed adephagan from South America’ mentioned by
|
||
<bibRefCitation author="Lawrence, J. F. & Newton, A. F. Jr" box="[1088,1392,1643,1665]" journalOrPublisher="Muzeum i Instytut Zoologii PAN, Warszawa" pageId="14" pageNumber="353" refId="ref10235" refString="Lawrence, J. F. & Newton, A. F. Jr (1995) Families and subfamilies of Coleoptera (with selected genera, notes, references and data on family-group names). Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski), pp. 779 - 1006. Muzeum i Instytut Zoologii PAN, Warszawa." title="Families and subfamilies of Coleoptera (with selected genera, notes, references and data on family-group names). Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski)" type="book" year="1995">Lawrence & Newton (1995)</bibRefCitation>
|
||
as having haliplidlike features. The general body form is reminiscent of
|
||
<taxonomicName box="[978,1087,1702,1724]" class="Insecta" family="Haliplidae" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="353" phylum="Arthropoda" rank="family">Haliplidae</taxonomicName>
|
||
, but the very small size, surface sculpture and setae of the body, the pectinate claws, the lack of expanded coxal plates, the legs lacking swimming hairs, the differing shape of the hind femur (
|
||
<bibRefCitation author="Kavanaugh, D. H." box="[1234,1420,1790,1812]" journalOrPublisher="Proceedings of the California Academy of Sciences" pageId="14" pageNumber="353" pagination="67 - 109" part="44" refId="ref10137" refString="Kavanaugh, D. H. (1986) A systematic review of amphizoid beetles (Amphizoidae: Coleoptera) and their phylogenetic relationships to other Adephaga. Proceedings of the California Academy of Sciences, 44, 67 - 109." title="A systematic review of amphizoid beetles (Amphizoidae: Coleoptera) and their phylogenetic relationships to other Adephaga" type="journal article" year="1986">Kavanaugh, 1986</bibRefCitation>
|
||
) and trochanter, and the lack of a specialized ovipositor, readily separate
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[333,432,156,178]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
|
||
from
|
||
<taxonomicName box="[500,610,156,178]" class="Insecta" family="Haliplidae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Haliplidae</taxonomicName>
|
||
. Most of these characters are autapomorphies for one or the other taxon; the greatly expanded hind coxal plates are unique to
|
||
<taxonomicName class="Insecta" family="Haliplidae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Haliplidae</taxonomicName>
|
||
, not known from any other extant group of beetles (
|
||
<bibRefCitation author="Beutel, R. G. & Ruhnau, S." box="[168,414,273,295]" journalOrPublisher="Aquatic Insects" pageId="15" pageNumber="354" pagination="1 - 17" part="12" refId="ref10026" refString="Beutel, R. G. & Ruhnau, S. (1990) Phylogenetic analysis of the genera of Haliplidae (Coleoptera) based on characters of adults. Aquatic Insects, 12, 1 - 17." title="Phylogenetic analysis of the genera of Haliplidae (Coleoptera) based on characters of adults" type="journal article" year="1990">Beutel & Ruhnau, 1990</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Beutel, R. G." box="[425,553,273,295]" journalOrPublisher="Muzeum i Instytut Zoologii PAN, Warszawa" pageId="15" pageNumber="354" refId="ref9728" refString="Beutel, R. G. (1995) The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski), pp. 503 - 539. Muzeum i Instytut Zoologii PAN, Warszawa." title="The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski)" type="book" year="1995">Beutel, 1995</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Roughley, R. E." box="[564,742,273,295]" journalOrPublisher="American Beetles, Vol. 1 (ed. by R. H. Arnett, Jr and M. C. Thomas), pp. 138 - 143. CRC Press, Boca Raton, Florida" pageId="15" pageNumber="354" refId="ref10600" refString="Roughley, R. E. (2001 b) Haliplidae Aube', 1836. American Beetles, Vol. 1 (ed. by R. H. Arnett, Jr and M. C. Thomas), pp. 138 - 143. CRC Press, Boca Raton, Florida." title="Haliplidae Aube'" type="book" year="2001">Roughley, 2001b</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="15.[161,767,156,1464]" pageId="15" pageNumber="354">
|
||
In
|
||
<taxonomicName baseAuthorityName="Beutel" baseAuthorityYear="1997" box="[218,323,302,324]" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Noteridae</taxonomicName>
|
||
, the smooth, streamlined body form and modified legs are highly specialized (except in
|
||
<taxonomicName authority="Ueno" authorityName="Ueno" authorityYear="1957" class="Insecta" family="Noteridae" genus="Phreatodytes" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[636,767,331,352]" italics="true" pageId="15" pageNumber="354">Phreatodytes</emphasis>
|
||
Ueńo
|
||
</taxonomicName>
|
||
) for swimming and burrowing. Other derived features of the clade (
|
||
<bibRefCitation author="Beutel, R. G. & Roughley, R. E." box="[296,559,390,412]" journalOrPublisher="Canadian Journal of Zoology" pageId="15" pageNumber="354" pagination="1898 - 1905" part="65" refId="ref9941" refString="Beutel, R. G. & Roughley, R. E. (1987) On the systematic position of the genus Notomicrus Sharp (Hydradephaga, Coleoptera). Canadian Journal of Zoology, 65, 1898 - 1905." title="On the systematic position of the genus Notomicrus Sharp (Hydradephaga, Coleoptera)" type="journal article" year="1987">Beutel & Roughley, 1987</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Beutel, R. G." box="[572,700,390,412]" journalOrPublisher="Muzeum i Instytut Zoologii PAN, Warszawa" pageId="15" pageNumber="354" refId="ref9728" refString="Beutel, R. G. (1995) The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski), pp. 503 - 539. Muzeum i Instytut Zoologii PAN, Warszawa." title="The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski)" type="book" year="1995">Beutel, 1995</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Beutel, R." box="[712,762,390,412]" journalOrPublisher="Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF)" pageId="15" pageNumber="354" pagination="1 - 164" part="31" refId="ref9797" refString="Beutel, R. (1997) Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga. Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF), 31, 1 - 164." title="Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga" type="journal article" year="1997">1997</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Roughley, R. E." box="[161,331,419,441]" journalOrPublisher="American Beetles, Vol. 1 (ed. by R. H. Arnett, Jr and M. C. Thomas), pp. 147 - 152. CRC Press, Boca Raton, Florida" pageId="15" pageNumber="354" refId="ref10654" refString="Roughley, R. E. (2001 c) Noteridae C. G. Thompson, 1857. American Beetles, Vol. 1 (ed. by R. H. Arnett, Jr and M. C. Thomas), pp. 147 - 152. CRC Press, Boca Raton, Florida." title="Noteridae C. G. Thompson" type="book" year="2001">Roughley, 2001c</bibRefCitation>
|
||
) include the form of the scape, with a short globular base and more slender distal part, dilation of antennomeres
|
||
<collectionCode box="[315,335,477,499]" country="Canada" lsid="urn:lsid:biocol.org:col:13946" name="Royal British Columbia Museum - Herbarium" pageId="15" pageNumber="354" type="Museum">V</collectionCode>
|
||
,
|
||
<collectionCode box="[351,387,477,499]" pageId="15" pageNumber="354">VII</collectionCode>
|
||
and
|
||
<collectionCode box="[449,479,477,499]" pageId="15" pageNumber="354">IX</collectionCode>
|
||
, presence of a profemoral excavation and antenna cleaning organ, metacoxal platform and anterior paramedian angles.
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[495,550,536,557]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[495,550,536,557]" italics="true" pageId="15" pageNumber="354">Meru</emphasis>
|
||
</taxonomicName>
|
||
shares the characters of the antennal flagellum only, but does share a derived feature with
|
||
<taxonomicName authorityName="Sharp" authorityYear="1882" box="[299,415,594,615]" class="Insecta" family="Noteridae" genus="Notomicrus" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[299,415,594,615]" italics="true" pageId="15" pageNumber="354">Notomicrus</emphasis>
|
||
</taxonomicName>
|
||
, considered to be sister to most remaining noterids: complete fusion of the metacoxae with the metasternum (
|
||
<bibRefCitation author="Beutel, R. G. & Roughley, R. E." box="[419,703,653,675]" journalOrPublisher="Canadian Journal of Zoology" pageId="15" pageNumber="354" pagination="1898 - 1905" part="65" refId="ref9941" refString="Beutel, R. G. & Roughley, R. E. (1987) On the systematic position of the genus Notomicrus Sharp (Hydradephaga, Coleoptera). Canadian Journal of Zoology, 65, 1898 - 1905." title="On the systematic position of the genus Notomicrus Sharp (Hydradephaga, Coleoptera)" type="journal article" year="1987">Beutel & Roughley, 1987</bibRefCitation>
|
||
). The absence of a sutural stria is also shared between
|
||
<taxonomicName baseAuthorityName="Beutel" baseAuthorityYear="1997" box="[664,767,682,704]" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Noteridae</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[207,262,711,732]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[207,262,711,732]" italics="true" pageId="15" pageNumber="354">Meru</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="15.[161,767,156,1464]" pageId="15" pageNumber="354">
|
||
Although the body form of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[473,528,740,761]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[473,528,740,761]" italics="true" pageId="15" pageNumber="354">Meru</emphasis>
|
||
</taxonomicName>
|
||
is somewhat similar to that of an
|
||
<taxonomicName authorityName="LeConte" authorityYear="1853" box="[264,364,770,791]" class="Insecta" family="Amphizoidae" genus="Amphizoa" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[264,364,770,791]" italics="true" pageId="15" pageNumber="354">Amphizoa</emphasis>
|
||
</taxonomicName>
|
||
, the latter are very much larger, and the apomorphic features of
|
||
<taxonomicName authorityName="LeConte" authorityYear="1853" box="[419,556,799,821]" class="Insecta" family="Amphizoidae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Amphizoidae</taxonomicName>
|
||
(absence of mental suture, one-segmented galea, and strongly reduced metafurca) preclude any close relationship. In
|
||
<taxonomicName authorityName="LeConte" authorityYear="1853" box="[623,762,857,879]" class="Insecta" family="Amphizoidae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Amphizoidae</taxonomicName>
|
||
, legs are relatively unmodified for swimming and natatory abilities are poor (
|
||
<bibRefCitation author="Kavanaugh, D. H." box="[344,526,915,937]" journalOrPublisher="Proceedings of the California Academy of Sciences" pageId="15" pageNumber="354" pagination="67 - 109" part="44" refId="ref10137" refString="Kavanaugh, D. H. (1986) A systematic review of amphizoid beetles (Amphizoidae: Coleoptera) and their phylogenetic relationships to other Adephaga. Proceedings of the California Academy of Sciences, 44, 67 - 109." title="A systematic review of amphizoid beetles (Amphizoidae: Coleoptera) and their phylogenetic relationships to other Adephaga" type="journal article" year="1986">Kavanaugh, 1986</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Beutel, R. G." box="[536,663,915,937]" journalOrPublisher="Muzeum i Instytut Zoologii PAN, Warszawa" pageId="15" pageNumber="354" refId="ref9728" refString="Beutel, R. G. (1995) The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski), pp. 503 - 539. Muzeum i Instytut Zoologii PAN, Warszawa." title="The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski)" type="book" year="1995">Beutel, 1995</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Philips, T. K. & Weiping Xie" journalOrPublisher="American Beetles, Vol. 1 (ed. by R. H. Arnett, Jr and M. C. Thomas), pp. 153 - 155. CRC Press, Boca Raton, Florida" pageId="15" pageNumber="354" refId="ref10363" refString="Philips, T. K. & Weiping Xie (2001) Amphizoidae LeConte, 1853. American Beetles, Vol. 1 (ed. by R. H. Arnett, Jr and M. C. Thomas), pp. 153 - 155. CRC Press, Boca Raton, Florida." title="Amphizoidae LeConte" type="book" year="2001">Philips & Weiping Xie, 2001</bibRefCitation>
|
||
) but vestigial swimming hairs are present.
|
||
<taxonomicName box="[214,342,974,996]" class="Insecta" family="Hygrobiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Hygrobiidae</taxonomicName>
|
||
are also much larger, have long fringes of swimming hairs on the legs, and several unique autapomorphies as listed by
|
||
<bibRefCitation author="Beutel, R. G." box="[410,546,1032,1054]" journalOrPublisher="Muzeum i Instytut Zoologii PAN, Warszawa" pageId="15" pageNumber="354" refId="ref9728" refString="Beutel, R. G. (1995) The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski), pp. 503 - 539. Muzeum i Instytut Zoologii PAN, Warszawa." title="The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski)" type="book" year="1995">Beutel (1995</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Beutel, R." box="[563,612,1032,1054]" journalOrPublisher="Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF)" pageId="15" pageNumber="354" pagination="1 - 164" part="31" refId="ref9797" refString="Beutel, R. (1997) Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga. Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF), 31, 1 - 164." title="Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga" type="journal article" year="1997">1997</bibRefCitation>
|
||
) that are not found in
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[262,316,1062,1083]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[262,316,1062,1083]" italics="true" pageId="15" pageNumber="354">Meru</emphasis>
|
||
</taxonomicName>
|
||
. Members of the recently described family
|
||
<taxonomicName box="[161,281,1091,1113]" class="Insecta" family="Aspidytidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Aspidytidae</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Ribera, I. & Beutel, R. G. & Balke, M. & Vogler, A. P." box="[297,507,1091,1113]" journalOrPublisher="Proceedings of the Royal Society of London (B" pageId="15" pageNumber="354" pagination="2351 - 2356" part="269" refId="ref10419" refString="Ribera, I., Beutel, R. G., Balke, M. & Vogler, A. P. (2002 b) Discovery of Aspidytidae, a new family of aquatic Coleoptera. Proceedings of the Royal Society of London (B), 269, 2351 - 2356." title="Discovery of Aspidytidae, a new family of aquatic Coleoptera" type="journal article" year="2002">
|
||
Ribera
|
||
<emphasis box="[376,422,1091,1112]" italics="true" pageId="15" pageNumber="354">et al</emphasis>
|
||
., 2002b
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Balke, M. & Ribera, I. & Beutel, R. G." box="[520,707,1091,1113]" journalOrPublisher="Zoologisch-Botanische Gesellschaft in Osterreich and Wiener Coleopterologenverein, Vienna" pageId="15" pageNumber="354" refId="ref9521" refString="Balke, M., Ribera, I. & Beutel, R. G. (2003) Aspidytidae: on the discovery of a new beetle family: detailed morphological analysis, description of a second species, and a key to fossil and extant adephagan families. Water Beetles of China, III (ed. by M. A. Jach and L. Ji), pp. 53 - 66. Zoologisch-Botanische Gesellschaft in Osterreich and Wiener Coleopterologenverein, Vienna." title="Aspidytidae: on the discovery of a new beetle family: detailed morphological analysis, description of a second species, and a key to fossil and extant adephagan families. Water Beetles of China, III (ed. by M. A. Jach and L. Ji)" type="book" year="2003">
|
||
Balke
|
||
<emphasis box="[587,633,1091,1112]" italics="true" pageId="15" pageNumber="354">et al</emphasis>
|
||
., 2003
|
||
</bibRefCitation>
|
||
), also lacking swimming hairs (considered to be derived through loss) on the legs, have a smooth, noteridlike body form and unusual scapus and pedicellus very different from those of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[161,216,1208,1229]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[161,216,1208,1229]" italics="true" pageId="15" pageNumber="354">Meru</emphasis>
|
||
</taxonomicName>
|
||
. The carabidlike terrestrial
|
||
<taxonomicName authority="(Ball, 2001)" baseAuthorityName="Ball" baseAuthorityYear="2001" class="Insecta" family="Trachypachidae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">
|
||
Trachypachidae (
|
||
<bibRefCitation author="Ball, G. E." journalOrPublisher="American Beetles, Vol. 1 (ed. by R. H. Arnett, Jr and M. C. Thomas), pp. 144 - 146. CRC Press, Boca Raton, Florida" pageId="15" pageNumber="354" refId="ref9608" refString="Ball, G. E. (2001) Trachypachidae C. G. Thompson, 1857. American Beetles, Vol. 1 (ed. by R. H. Arnett, Jr and M. C. Thomas), pp. 144 - 146. CRC Press, Boca Raton, Florida." title="Trachypachidae C. G. Thompson" type="book" year="2001">Ball, 2001</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
have few similarities to
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[500,599,1237,1259]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
|
||
except for the groundplan adephagan features.
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[509,564,1267,1288]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[509,564,1267,1288]" italics="true" pageId="15" pageNumber="354">Meru</emphasis>
|
||
</taxonomicName>
|
||
lacks the protibial antennal cleaner of
|
||
<taxonomicName baseAuthorityName="Ball" baseAuthorityYear="2001" box="[359,526,1295,1317]" class="Insecta" family="Trachypachidae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Trachypachidae</taxonomicName>
|
||
. Relationships with this group and aquatic
|
||
<taxonomicName box="[353,457,1325,1347]" pageId="15" pageNumber="357" rank="subOrder" subOrder="Adephaga">Adephaga</taxonomicName>
|
||
have been the subject of many studies (
|
||
<bibRefCitation author="Bell, R. T." box="[248,354,1354,1376]" journalOrPublisher="Coleopterists Bulletin" pageId="15" pageNumber="354" pagination="107 - 112" part="20" refId="ref9665" refString="Bell, R. T. (1966) Trachypachus and the origin of the Hydradephaga (Coleoptera). Coleopterists Bulletin, 20, 107 - 112." title="Trachypachus and the origin of the Hydradephaga (Coleoptera)" type="journal article" year="1966">Bell, 1966</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Roughley, R. E." box="[369,534,1354,1376]" journalOrPublisher="Pan-Pacific Entomologist" pageId="15" pageNumber="354" pagination="272 - 285" part="57" refId="ref10519" refString="Roughley, R. E. (1981) TrachypachidaeandHydradephaga (Coleoptera): a monophyletic unit? Pan-Pacific Entomologist, 57, 272 - 285." title="TrachypachidaeandHydradephaga (Coleoptera): a monophyletic unit?" type="journal article" year="1981">Roughley, 1981</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Beutel, R. & Belkaceme, T." journalOrPublisher="Entomologica Basiliensia" pageId="15" pageNumber="354" pagination="221 - 229" part="11" refId="ref9834" refString="Beutel, R. & Belkaceme, T. (1986) Comparative studies on the metathorax of Hydradephaga and Trachypachidae. Entomologica Basiliensia, 11, 221 - 229." title="Comparative studies on the metathorax of Hydradephaga and Trachypachidae" type="journal article" year="1986">Beutel & Belkaceme, 1986</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Shull, V. L. & Vogler, A. P. & Baker, M. D. & Maddison, D. R. & Hammond, P. M." box="[225,410,1383,1405]" journalOrPublisher="Systematic Biology" pageId="15" pageNumber="354" pagination="945 - 969" part="50" refId="ref10808" refString="Shull, V. L., Vogler, A. P., Baker, M. D., Maddison, D. R. & Hammond, P. M. (2001) Sequence alignment of 18 S ribosomal RNA and the basal relationships of Adephagan beetles: evidence for monophyly of aquatic families and the placement of Trachypachidae. Systematic Biology, 50, 945 - 969." title="Sequence alignment of 18 S ribosomal RNA and the basal relationships of Adephagan beetles: evidence for monophyly of aquatic families and the placement of Trachypachidae" type="journal article" year="2001">
|
||
Shull
|
||
<emphasis box="[288,334,1383,1404]" italics="true" pageId="15" pageNumber="354">et al</emphasis>
|
||
., 2001
|
||
</bibRefCitation>
|
||
). Lastly, the characters of known fossil taxa, as discussed by
|
||
<bibRefCitation author="Balke, M. & Ribera, I. & Beutel, R. G." box="[439,628,1412,1434]" journalOrPublisher="Zoologisch-Botanische Gesellschaft in Osterreich and Wiener Coleopterologenverein, Vienna" pageId="15" pageNumber="354" refId="ref9521" refString="Balke, M., Ribera, I. & Beutel, R. G. (2003) Aspidytidae: on the discovery of a new beetle family: detailed morphological analysis, description of a second species, and a key to fossil and extant adephagan families. Water Beetles of China, III (ed. by M. A. Jach and L. Ji), pp. 53 - 66. Zoologisch-Botanische Gesellschaft in Osterreich and Wiener Coleopterologenverein, Vienna." title="Aspidytidae: on the discovery of a new beetle family: detailed morphological analysis, description of a second species, and a key to fossil and extant adephagan families. Water Beetles of China, III (ed. by M. A. Jach and L. Ji)" type="book" year="2003">
|
||
Balke
|
||
<emphasis box="[504,547,1413,1434]" italics="true" pageId="15" pageNumber="354">et al</emphasis>
|
||
. (2003)
|
||
</bibRefCitation>
|
||
, also rule out inclusion of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[288,343,1442,1463]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[288,343,1442,1463]" italics="true" pageId="15" pageNumber="354">Meru</emphasis>
|
||
</taxonomicName>
|
||
in any of these families.
|
||
</paragraph>
|
||
<paragraph blockId="15.[161,483,1528,1550]" box="[161,483,1528,1550]" pageId="15" pageNumber="354">
|
||
<heading bold="true" box="[161,483,1528,1550]" fontSize="9" level="1" pageId="15" pageNumber="354" reason="2">
|
||
<emphasis bold="true" box="[161,483,1528,1550]" pageId="15" pageNumber="354">Comments and conclusions</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="15.[161,767,1588,1843]" lastBlockId="15.[830,1437,156,1843]" pageId="15" pageNumber="354">
|
||
The overall body form of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[434,489,1588,1609]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[434,489,1588,1609]" italics="true" pageId="15" pageNumber="354">Meru</emphasis>
|
||
</taxonomicName>
|
||
, with rough surfaces, protruding eyes, and unmodified legs, is typical of aquatic beetles that are weaker swimmers of lentic water, or shallow marginal stream habitats, e.g.
|
||
<taxonomicName box="[467,573,1675,1697]" class="Insecta" family="Haliplidae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Haliplidae</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="LeConte" authorityYear="1853" box="[623,762,1675,1697]" class="Insecta" family="Amphizoidae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Amphizoidae</taxonomicName>
|
||
, and several nonadephagan groups that have acquired aquatic habits. For aquatic
|
||
<taxonomicName box="[447,554,1734,1756]" pageId="15" pageNumber="357" rank="subOrder" subOrder="Adephaga">Adephaga</taxonomicName>
|
||
, these characters are considered to be primitive, relative to the streamlined body and specialized swimming legs and methods of
|
||
<taxonomicName box="[652,763,1792,1814]" class="Insecta" family="Dytiscidae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Dytiscidae</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Beutel" baseAuthorityYear="1997" box="[161,266,1821,1843]" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Noteridae</taxonomicName>
|
||
, and
|
||
<taxonomicName authority="(Ribera et al., 2002 b)" baseAuthorityName="Ribera" baseAuthorityYear="2002" box="[324,658,1821,1843]" class="Insecta" family="Gyrinidae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">
|
||
Gyrinidae (
|
||
<bibRefCitation author="Ribera, I. & Beutel, R. G. & Balke, M. & Vogler, A. P." box="[441,652,1821,1843]" journalOrPublisher="Proceedings of the Royal Society of London (B" pageId="15" pageNumber="354" pagination="2351 - 2356" part="269" refId="ref10419" refString="Ribera, I., Beutel, R. G., Balke, M. & Vogler, A. P. (2002 b) Discovery of Aspidytidae, a new family of aquatic Coleoptera. Proceedings of the Royal Society of London (B), 269, 2351 - 2356." title="Discovery of Aspidytidae, a new family of aquatic Coleoptera" type="journal article" year="2002">
|
||
Ribera
|
||
<emphasis box="[520,565,1822,1843]" italics="true" pageId="15" pageNumber="354">et al</emphasis>
|
||
., 2002b
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
.
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[669,724,1822,1843]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[669,724,1822,1843]" italics="true" pageId="15" pageNumber="354">Meru</emphasis>
|
||
</taxonomicName>
|
||
has features of a beetle that has perhaps recently invaded the aquatic realm and/or has retained these primitive states, and this is in keeping with the postulated habitats of the earliest
|
||
<taxonomicName box="[915,1019,243,265]" pageId="15" pageNumber="357" rank="subOrder" subOrder="Adephaga">Adephaga</taxonomicName>
|
||
(Beutel, 1995,1997) at the margins of ponds or rivers, from which multiple invasions of aquatic and terrestrial to arboreal habitats may have occurred.
|
||
</paragraph>
|
||
<paragraph blockId="15.[830,1437,156,1843]" pageId="15" pageNumber="354">
|
||
The observed wing dimorphism, with atrophied wings in the majority of specimens, is not a common condition in aquatic beetles (
|
||
<bibRefCitation author="Spangler, P. J." box="[1005,1161,389,411]" journalOrPublisher="Proceedings of the Biology Society of Washington" pageId="15" pageNumber="354" pagination="697 - 718" part="92" refId="ref10911" refString="Spangler, P. J. (1979) A new genus of water beetle from austral South America (Coleoptera: Hydrophilidae). Proceedings of the Biology Society of Washington, 92, 697 - 718." title="A new genus of water beetle from austral South America (Coleoptera: Hydrophilidae)" type="journal article" year="1979">Spangler, 1979</bibRefCitation>
|
||
) and also not typical of obligate streamside lowland species of
|
||
<taxonomicName authorityName="Latreille" authorityYear="1802" box="[1205,1308,419,441]" class="Insecta" family="Carabidae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Carabidae</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Darlington, P. J." journalOrPublisher="Annals of the Entomological Society of America" pageId="15" pageNumber="354" pagination="136 - 179" part="29" refId="ref10065" refString="Darlington, P. J. (1936) Variation of flying wings of carabid beetles (Coleoptera). Annals of the Entomological Society of America, 29, 136 - 179." title="Variation of flying wings of carabid beetles (Coleoptera)" type="journal article" year="1936">Darlington, 1936</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Darlington, P. J." box="[889,940,448,470]" journalOrPublisher="Ecological Monographs" pageId="15" pageNumber="354" pagination="37 - 61" part="13" refId="ref10099" refString="Darlington, P. J. (1943) Carabidae of mountains and islands: data on the evolution of isolated faunas, and on atrophy of wings. Ecological Monographs, 13, 37 - 61." title="Carabidae of mountains and islands: data on the evolution of isolated faunas, and on atrophy of wings" type="journal article" year="1943">1943</bibRefCitation>
|
||
). Maintenance of both wing forms in a population is probably advantageous for a small aquatic beetle in a habitat subject to fluctuation or sudden changes, such as flooding or drying of the stream margin habitat.
|
||
</paragraph>
|
||
<paragraph blockId="15.[830,1437,156,1843]" pageId="15" pageNumber="354">
|
||
The unique apomorphic features of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[1255,1354,565,587]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
|
||
are an enigmatic assemblage: the pectinate tarsal claws, the possibly respiratory structures covering the body, the odd leaflike setae of the labrum, and the trifid tibial spurs. The very small size of these beetles, also considered to be derived (
|
||
<bibRefCitation author="Beutel, R. G. & Haas, A." box="[837,1054,711,733]" journalOrPublisher="Zoomorphology" pageId="15" pageNumber="354" pagination="103 - 116" part="18" refId="ref9865" refString="Beutel, R. G. & Haas, A. (1998) Larval head of Hydroscapha natans LeConte, 1874 (Coleoptera, Myxophaga, Hydroscaphidae) with special reference to miniaturization. Zoomorphology, 18, 103 - 116." title="Larval head of Hydroscapha natans LeConte, 1874 (Coleoptera, Myxophaga, Hydroscaphidae) with special reference to miniaturization" type="journal article" year="1998">Beutel & Haas, 1998</bibRefCitation>
|
||
) may be a contributing factor to the assemblage of other unusual characters of this taxon. Miniaturization in
|
||
<taxonomicName box="[1027,1140,769,791]" class="Magnoliopsida" higherTaxonomySource="GBIF" kingdom="Plantae" order="Coleoptera" pageId="15" pageNumber="354" phylum="Tracheophyta" rank="order">Coleoptera</taxonomicName>
|
||
results in alteration of other characters via fusion, reduction and loss. In
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[1278,1333,799,820]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[1278,1333,799,820]" italics="true" pageId="15" pageNumber="354">Meru</emphasis>
|
||
</taxonomicName>
|
||
, fusion of thoracic sclerites, loss of the oblongum cell and binding patch in the hind wing, reduced and membranous ovipositor, and probable loss of pygidial and prothracic defence glands, etc., can probably be attributed to miniaturization; similar conclusions have been drawn for other small-sized members of other beetle taxa, e.g.
|
||
<taxonomicName authorityName="Folkerts" authorityYear="1979" box="[1208,1363,974,995]" class="Insecta" family="Gyrinidae" genus="Spanglerogyrus" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[1208,1363,974,995]" italics="true" pageId="15" pageNumber="354">Spanglerogyrus</emphasis>
|
||
</taxonomicName>
|
||
in the
|
||
<taxonomicName authority="(Steiner & Anderson, 1981)" baseAuthorityName="Steiner & Anderson" baseAuthorityYear="1981" box="[830,1224,1003,1025]" class="Insecta" family="Gyrinidae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">
|
||
Gyrinidae (
|
||
<bibRefCitation author="Steiner, W. E. Jr & Anderson, J. J." box="[947,1218,1003,1025]" journalOrPublisher="Pan-Pacific Entomologist" pageId="15" pageNumber="354" pagination="124 - 132" part="57" refId="ref11013" refString="Steiner, W. E. Jr & Anderson, J. J. (1981) Notes on the natural history of Spanglerogyrus albiventris Folkerts, with a new distribution record (Coleoptera: Gyrinidae). Pan-Pacific Entomologist, 57, 124 - 132." title="Notes on the natural history of Spanglerogyrus albiventris Folkerts, with a new distribution record (Coleoptera: Gyrinidae)" type="journal article" year="1981">Steiner & Anderson, 1981</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="15.[830,1437,156,1843]" pageId="15" pageNumber="354">
|
||
The folding pattern of the fully developed wing of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[830,954,1062,1083]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="species" species="phyllisae">
|
||
<emphasis box="[830,954,1062,1083]" italics="true" pageId="15" pageNumber="354">M. phyllisae</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation box="[967,1037,1062,1084]" captionStart="Fig" captionStartId="10.[151,185,1224,1244]" captionTargetBox="[167,742,769,1196]" captionTargetId="figure-29@10.[167,742,767,1196]" captionTargetPageId="10" captionText="Fig. 12. Meru phyllisae, hind wing, macropterous condition, in folded position." figureDoi="http://doi.org/10.5281/zenodo.4311878" httpUri="https://zenodo.org/record/4311878/files/figure.png" pageId="15" pageNumber="354">Fig. 12</figureCitation>
|
||
) is of the adephagan
|
||
<typeStatus box="[1253,1297,1062,1084]" pageId="15" pageNumber="354">type</typeStatus>
|
||
(
|
||
<bibRefCitation author="Kavanaugh, D. H." journalOrPublisher="Proceedings of the California Academy of Sciences" pageId="15" pageNumber="354" pagination="67 - 109" part="44" refId="ref10137" refString="Kavanaugh, D. H. (1986) A systematic review of amphizoid beetles (Amphizoidae: Coleoptera) and their phylogenetic relationships to other Adephaga. Proceedings of the California Academy of Sciences, 44, 67 - 109." title="A systematic review of amphizoid beetles (Amphizoidae: Coleoptera) and their phylogenetic relationships to other Adephaga" type="journal article" year="1986">Kavanaugh, 1986</bibRefCitation>
|
||
; Kukalova´-Peck & Lawrence, 1993;
|
||
<bibRefCitation author="Lawrence, J. F. & Britton, E. B." journalOrPublisher="Melbourne University Press, Melbourne" pageId="15" pageNumber="354" refId="ref10210" refString="Lawrence, J. F. & Britton, E. B. (1994) Australian Beetles. Melbourne University Press, Melbourne." title="Australian Beetles" type="book" year="1994">Lawrence & Britton, 1994</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Beutel, R. G. & Haas, A." box="[990,1221,1120,1142]" journalOrPublisher="Cladistics" pageId="15" pageNumber="354" pagination="103 - 141" part="16" refId="ref9908" refString="Beutel, R. G. & Haas, A. (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 & Haas, 2000</bibRefCitation>
|
||
), although with the reduced venation and marginal fringe, the wing is more similar to that seen in members of Myxophaga. Similarities in the wing to that of myxophagan beetles, all of which are relatively small, may be the result of convergence, but a sister-group relationship between Myxophaga and
|
||
<taxonomicName box="[1334,1437,1266,1288]" pageId="15" pageNumber="357" rank="subOrder" subOrder="Adephaga">Adephaga</taxonomicName>
|
||
has been suggested by wing characters (
|
||
<bibRefCitation author="Kukalova-Peck, J. & Lawrence, J. F." journalOrPublisher="Canadian Entomologist" pageId="15" pageNumber="354" pagination="181 - 258" part="125" refId="ref10179" refString="Kukalova-Peck, J. & Lawrence, J. F. (1993) Evolution of the hind wing in Coleoptera. Canadian Entomologist, 125, 181 - 258." title="Evolution of the hind wing in Coleoptera" type="journal article" year="1993">Kukalová-Peck & Lawrence, 1993</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="15.[830,1437,156,1843]" pageId="15" pageNumber="354">
|
||
Serrate tibial spurs are found in most
|
||
<taxonomicName baseAuthorityName="Beutel" baseAuthorityYear="1997" box="[1327,1432,1354,1376]" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Noteridae</taxonomicName>
|
||
, especially the larger (inner) metatibial spur, which is also serrate in
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[930,1053,1413,1434]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="species" species="phyllisae">
|
||
<emphasis box="[930,1053,1413,1434]" italics="true" pageId="15" pageNumber="354">M. phyllisae</emphasis>
|
||
</taxonomicName>
|
||
. Some
|
||
<taxonomicName box="[1127,1233,1412,1434]" class="Insecta" family="Haliplidae" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="family">Haliplidae</taxonomicName>
|
||
also bear serrations on this spur. Whether this represents any synapomorphy among these families should be a subject of future study.
|
||
</paragraph>
|
||
<paragraph blockId="15.[830,1437,156,1843]" lastBlockId="16.[151,758,156,1843]" lastPageId="16" lastPageNumber="355" pageId="15" pageNumber="354">
|
||
The leaflike setae on the labrum, which oppose each other and overlap above a median emargination, along with the toothlike setae medial to these, may be related to a feeding specialization. The diet of these beetles is uncertain; observations on captive
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[1102,1234,1617,1638]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="354" phylum="Arthropoda" rank="species" species="phyllisae">
|
||
<emphasis box="[1102,1234,1617,1638]" italics="true" pageId="15" pageNumber="354">M. phyllisae</emphasis>
|
||
</taxonomicName>
|
||
suggest that they may be eating decaying leaf tissue or perhaps more likely, algal filaments and/or fungal hyphae growing on submerged leaves. The fact that captive beetles survived for nearly 200 days on decaying leaves, without any predatory or cannibalistic behaviour being observed during that time, supports this idea. Ancestral
|
||
<taxonomicName box="[1272,1376,1792,1814]" pageId="15" pageNumber="357" rank="subOrder" subOrder="Adephaga">Adephaga</taxonomicName>
|
||
were probably predators (
|
||
<bibRefCitation author="Beutel, R. G." box="[1060,1198,1821,1843]" journalOrPublisher="Muzeum i Instytut Zoologii PAN, Warszawa" pageId="15" pageNumber="354" refId="ref9728" refString="Beutel, R. G. (1995) The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski), pp. 503 - 539. Muzeum i Instytut Zoologii PAN, Warszawa." title="The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski)" type="book" year="1995">Beutel, 1995</bibRefCitation>
|
||
) as are most extant members, but predatory habits in
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[495,550,156,177]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[495,550,156,177]" italics="true" pageId="16" pageNumber="355">Meru</emphasis>
|
||
</taxonomicName>
|
||
are doubtful, as it is not an agile or rapid swimmer. Although the head is prognathous, the mandibles are rather stout, less suitable for predation, but the terebral ridges and teeth in
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[628,683,244,265]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[628,683,244,265]" italics="true" pageId="16" pageNumber="355">Meru</emphasis>
|
||
</taxonomicName>
|
||
mandibles would serve for cutting (
|
||
<bibRefCitation author="Acorn, J. H. & Ball, G. E." box="[453,656,273,295]" journalOrPublisher="Canadian Journal of Zoology" pageId="16" pageNumber="355" pagination="638 - 650" part="69" refId="ref9435" refString="Acorn, J. H. & Ball, G. E. (1991) The mandibles of some ground beetles: structure, function, and the evolution of herbivory (Coleoptera: Carabidae). Canadian Journal of Zoology, 69, 638 - 650." title="The mandibles of some ground beetles: structure, function, and the evolution of herbivory (Coleoptera: Carabidae)" type="journal article" year="1991">Acorn & Ball, 1991</bibRefCitation>
|
||
). Haliplid adults have stout mandibles and are known to be more omnivorous (
|
||
<bibRefCitation author="Seeger, W." box="[293,429,331,353]" journalOrPublisher="Archiv fur Hydrobiologie" pageId="16" pageNumber="355" pagination="400 - 435" part="68" refId="ref10772" refString="Seeger, W. (1971) Morphologie, Bionomie und Ethologie von Halipliden, unter besonderer Berucksichtigung funktionsmorphologischer Gesichtspunkte (Haliplidae; Coleoptera). Archiv fur Hydrobiologie, 68, 400 - 435." title="Morphologie, Bionomie und Ethologie von Halipliden, unter besonderer Berucksichtigung funktionsmorphologischer Gesichtspunkte (Haliplidae; Coleoptera)" type="journal article" year="1971">Seeger, 1971</bibRefCitation>
|
||
), whereas the larvae are algal specialists.
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[273,429,361,382]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="species" species="phyllisae">
|
||
<emphasis box="[273,429,361,382]" italics="true" pageId="16" pageNumber="355">Meru phyllisae</emphasis>
|
||
</taxonomicName>
|
||
could also be categorized as omnivorous, and if also an occasional predator, this would probably involve only small, perhaps sessile, prey species. Further comparisons of mandibular morphology, with scanning electron microscopy of the surfaces, will probably provide more information.
|
||
</paragraph>
|
||
<paragraph blockId="16.[151,758,156,1843]" pageId="16" pageNumber="355">
|
||
The labrum in adult haliplids (
|
||
<bibRefCitation author="Beutel, R. G. & Ruhnau, S." box="[486,728,536,558]" journalOrPublisher="Aquatic Insects" pageId="16" pageNumber="355" pagination="1 - 17" part="12" refId="ref10026" refString="Beutel, R. G. & Ruhnau, S. (1990) Phylogenetic analysis of the genera of Haliplidae (Coleoptera) based on characters of adults. Aquatic Insects, 12, 1 - 17." title="Phylogenetic analysis of the genera of Haliplidae (Coleoptera) based on characters of adults" type="journal article" year="1990">Beutel & Ruhnau, 1990</bibRefCitation>
|
||
) is also distinctly emarginate and with a fringe of flattened, blunt to tapered setae directed anteromedially. There are also similarities in the labrum of some Myxophaga, which are also small animals that feed on filaments of algae, e.g. the larva of
|
||
<taxonomicName authorityName="LeConte" authorityYear="1874" box="[356,488,682,703]" class="Insecta" family="Hydroscaphidae" genus="Hydroscapha" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[356,488,682,703]" italics="true" pageId="16" pageNumber="355">Hydroscapha</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Beutel, R. G. & Haas, A." box="[511,754,682,704]" journalOrPublisher="Zoomorphology" pageId="16" pageNumber="355" pagination="103 - 116" part="18" refId="ref9865" refString="Beutel, R. G. & Haas, A. (1998) Larval head of Hydroscapha natans LeConte, 1874 (Coleoptera, Myxophaga, Hydroscaphidae) with special reference to miniaturization. Zoomorphology, 18, 103 - 116." title="Larval head of Hydroscapha natans LeConte, 1874 (Coleoptera, Myxophaga, Hydroscaphidae) with special reference to miniaturization" type="journal article" year="1998">Beutel & Haas, 1998</bibRefCitation>
|
||
: fig. 2A) and adult
|
||
<taxonomicName authorityName="Steffan" authorityYear="1964" box="[334,493,711,733]" class="Insecta" family="Torridincolidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Torridincolidae</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Spangler, P. J." box="[505,658,711,733]" journalOrPublisher="Coleopterists Bulletin" pageId="16" pageNumber="355" pagination="145 - 158" part="34" refId="ref10950" refString="Spangler, P. J. (1980) A new species of Ytu from Brazil (Coleoptera: Torridincolidae). Coleopterists Bulletin, 34, 145 - 158." title="A new species of Ytu from Brazil (Coleoptera: Torridincolidae)" type="journal article" year="1980">Spangler, 1980</bibRefCitation>
|
||
: figs 4, 5).
|
||
</paragraph>
|
||
<paragraph blockId="16.[151,758,156,1843]" pageId="16" pageNumber="355">
|
||
Antennal characters of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[431,486,740,761]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[431,486,740,761]" italics="true" pageId="16" pageNumber="355">Meru</emphasis>
|
||
</taxonomicName>
|
||
are intriguing and show similarities to both
|
||
<taxonomicName box="[348,454,769,791]" class="Insecta" family="Haliplidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Haliplidae</taxonomicName>
|
||
and
|
||
<taxonomicName baseAuthorityName="Beutel" baseAuthorityYear="1997" box="[504,610,769,791]" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Noteridae</taxonomicName>
|
||
. The globular, short scape has been considered a derived feature of haliplids, and the condition in
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[490,618,828,849]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="species" species="phyllisae">
|
||
<emphasis box="[490,618,828,849]" italics="true" pageId="16" pageNumber="355">M. phyllisae</emphasis>
|
||
</taxonomicName>
|
||
seems to be even more reduced (derived) in that there is no separate distal portion; in haliplids, there is a slight constriction between basal and distal halves. The pseudo-two-segmented state in
|
||
<taxonomicName baseAuthorityName="Beutel" baseAuthorityYear="1997" box="[242,345,945,967]" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Noteridae</taxonomicName>
|
||
and
|
||
<taxonomicName box="[407,529,945,967]" class="Insecta" family="Aspidytidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Aspidytidae</taxonomicName>
|
||
appears to be intermediate in form and reduction between the scape in
|
||
<taxonomicName box="[151,257,1003,1025]" class="Insecta" family="Haliplidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Haliplidae</taxonomicName>
|
||
and that of other
|
||
<taxonomicName box="[471,578,1003,1025]" pageId="16" pageNumber="357" rank="subOrder" subOrder="Adephaga">Adephaga</taxonomicName>
|
||
, which have the distal portion generally longer than wide.
|
||
</paragraph>
|
||
<paragraph blockId="16.[151,758,156,1843]" pageId="16" pageNumber="355">
|
||
Modification of antennomeres
|
||
<collectionCode box="[502,522,1062,1084]" country="Canada" lsid="urn:lsid:biocol.org:col:13946" name="Royal British Columbia Museum - Herbarium" pageId="16" pageNumber="355" type="Museum">V</collectionCode>
|
||
,
|
||
<collectionCode box="[535,571,1062,1084]" pageId="16" pageNumber="355">VII</collectionCode>
|
||
and
|
||
<collectionCode box="[628,656,1062,1084]" pageId="16" pageNumber="355">IX</collectionCode>
|
||
has been considered unique to
|
||
<taxonomicName authority="(Beutel, 1997)" baseAuthorityName="Beutel" baseAuthorityYear="1997" box="[383,645,1091,1113]" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">
|
||
Noteridae (
|
||
<bibRefCitation author="Beutel, R." box="[504,639,1091,1113]" journalOrPublisher="Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF)" pageId="16" pageNumber="355" pagination="1 - 164" part="31" refId="ref9797" refString="Beutel, R. (1997) Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga. Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF), 31, 1 - 164." title="Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga" type="journal article" year="1997">Beutel, 1997</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. Increased size and sensory fields of the same alternating antennomeres among the two taxa would have a very remote chance of being a result of convergence, so this appears to be a synapomorphy with
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[318,420,1208,1230]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
|
||
. Perhaps the miniaturization (possibly an adaptation to the stream margin habitat) of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[179,302,1267,1288]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="species" species="phyllisae">
|
||
<emphasis box="[179,302,1267,1288]" italics="true" pageId="16" pageNumber="355">M. phyllisae</emphasis>
|
||
</taxonomicName>
|
||
has resulted in the loss of sensoria on antennomeres and reduction of the number to single organs on antennomeres
|
||
<collectionCode box="[299,335,1325,1347]" pageId="16" pageNumber="355">VII</collectionCode>
|
||
and
|
||
<collectionCode box="[385,416,1325,1347]" pageId="16" pageNumber="355">IX</collectionCode>
|
||
. Some
|
||
<taxonomicName box="[487,593,1325,1347]" class="Insecta" family="Haliplidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Haliplidae</taxonomicName>
|
||
possess vestiges of this pattern, but this character system needs more comparative studies among many taxa in order to draw conclusions on possible relationships.
|
||
</paragraph>
|
||
<paragraph blockId="16.[151,758,156,1843]" pageId="16" pageNumber="355">
|
||
The two brush-tipped setae of the anterior margin of the prosternum deserve mention, as we find similar setae in a number of
|
||
<taxonomicName baseAuthorityName="Beutel" baseAuthorityYear="1997" box="[314,419,1500,1522]" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Noteridae</taxonomicName>
|
||
, greater in number, but with the same general form. In
|
||
<taxonomicName authority="Sharp" authorityName="Sharp" authorityYear="1882" box="[415,673,1529,1551]" class="Insecta" family="Noteridae" genus="Notomicrus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="species" species="traili">
|
||
<emphasis box="[415,595,1530,1551]" italics="true" pageId="16" pageNumber="355">Notomicrus traili</emphasis>
|
||
Sharp
|
||
</taxonomicName>
|
||
, twelve such brush-tipped setae occur in a row along the margin; fewer are seen in the related
|
||
<taxonomicName authorityName="Spangler" authorityYear="1996" box="[458,579,1588,1609]" class="Insecta" family="Noteridae" genus="Speonoterus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[458,579,1588,1609]" italics="true" pageId="16" pageNumber="355">Speonoterus</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Spangler, P. J." box="[596,754,1588,1610]" journalOrPublisher="Insecta Mundi" pageId="16" pageNumber="355" pagination="241 - 259" part="10" refId="ref10981" refString="Spangler, P. J. (1996) Four new stygobiontic beetles (Coleoptera: Dytiscidae; Noteridae; Elmidae). Insecta Mundi, 10, 241 - 259." title="Four new stygobiontic beetles (Coleoptera: Dytiscidae; Noteridae; Elmidae)" type="journal article" year="1996">Spangler, 1996</bibRefCitation>
|
||
: fig. 58). Other larger noterids, e.g.
|
||
<taxonomicName authority="(Say)" baseAuthorityName="Say" class="Insecta" family="Noteridae" genus="Hydrocanthus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="species" species="iricolor">
|
||
<emphasis box="[532,757,1617,1638]" italics="true" pageId="16" pageNumber="355">Hydrocanthus iricolor</emphasis>
|
||
(Say)
|
||
</taxonomicName>
|
||
generally have a fringe of these setae. Their function is unknown, but the distinctive form and placement are similar in
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[267,366,1705,1727]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
|
||
and
|
||
<taxonomicName baseAuthorityName="Beutel" baseAuthorityYear="1997" box="[431,537,1705,1727]" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Noteridae</taxonomicName>
|
||
. Setae that line the prosternal margin in the haliplids examined appear to be of the simple, tapered form.
|
||
</paragraph>
|
||
<paragraph blockId="16.[151,758,156,1843]" lastBlockId="16.[821,1427,156,1843]" pageId="16" pageNumber="355">
|
||
The hidden scutellum is a feature common to all known
|
||
<taxonomicName box="[151,257,1821,1843]" class="Insecta" family="Haliplidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Haliplidae</taxonomicName>
|
||
and
|
||
<taxonomicName baseAuthorityName="Beutel" baseAuthorityYear="1997" box="[329,432,1821,1843]" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Noteridae</taxonomicName>
|
||
except
|
||
<taxonomicName authority="(Beutel, 1997)" baseAuthorityName="Beutel" baseAuthorityYear="1997" box="[530,879,156,1843]" class="Insecta" family="Noteridae" genus="Phreatodytes" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[530,661,1822,1843]" italics="true" pageId="16" pageNumber="355">Phreatodytes</emphasis>
|
||
(
|
||
<bibRefCitation author="Beutel, R." box="[685,872,156,1843]" journalOrPublisher="Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF)" pageId="16" pageNumber="355" pagination="1 - 164" part="31" refId="ref9797" refString="Beutel, R. (1997) Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga. Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF), 31, 1 - 164." title="Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga" type="journal article" year="1997">Beutel, 1997</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authority="Spangler (1996)" authorityName="Spangler" authorityYear="1996" box="[889,1179,156,178]" class="Insecta" family="Noteridae" genus="Speonoterus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[889,1010,156,177]" italics="true" pageId="16" pageNumber="355">Speonoterus</emphasis>
|
||
<bibRefCitation author="Spangler, P. J." box="[1017,1179,156,178]" journalOrPublisher="Insecta Mundi" pageId="16" pageNumber="355" pagination="241 - 259" part="10" refId="ref10981" refString="Spangler, P. J. (1996) Four new stygobiontic beetles (Coleoptera: Dytiscidae; Noteridae; Elmidae). Insecta Mundi, 10, 241 - 259." title="Four new stygobiontic beetles (Coleoptera: Dytiscidae; Noteridae; Elmidae)" type="journal article" year="1996">Spangler (1996)</bibRefCitation>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName authorityName="Sharp" authorityYear="1882" box="[1235,1351,156,177]" class="Insecta" family="Noteridae" genus="Notomicrus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[1235,1351,156,177]" italics="true" pageId="16" pageNumber="355">Notomicrus</emphasis>
|
||
</taxonomicName>
|
||
species (
|
||
<bibRefCitation author="Spangler, P. J." box="[828,990,185,207]" journalOrPublisher="Insecta Mundi" pageId="16" pageNumber="355" pagination="241 - 259" part="10" refId="ref10981" refString="Spangler, P. J. (1996) Four new stygobiontic beetles (Coleoptera: Dytiscidae; Noteridae; Elmidae). Insecta Mundi, 10, 241 - 259." title="Four new stygobiontic beetles (Coleoptera: Dytiscidae; Noteridae; Elmidae)" type="journal article" year="1996">Spangler, 1996</bibRefCitation>
|
||
); the latter have an exposed, but very small, scutellum, but according to
|
||
<bibRefCitation author="Beutel, R." box="[1172,1310,214,236]" journalOrPublisher="Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF)" pageId="16" pageNumber="355" pagination="1 - 164" part="31" refId="ref9797" refString="Beutel, R. (1997) Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga. Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF), 31, 1 - 164." title="Uber Phylogenese und Evolution der Coleoptera (Insecta), insbesondere der Adephaga" type="journal article" year="1997">Beutel (1997)</bibRefCitation>
|
||
, concealed. Several unrelated tribes of smaller-sized
|
||
<taxonomicName box="[1265,1373,243,265]" class="Insecta" family="Dytiscidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Dytiscidae</taxonomicName>
|
||
also have the scutellum concealed; the character may have arisen independently in some cases and may be another feature of convergence, related to the reduction in body size.
|
||
</paragraph>
|
||
<paragraph blockId="16.[821,1427,156,1843]" pageId="16" pageNumber="355">
|
||
The form of the proventriculus, with its eight lobes of alternating large and small size, is typical for all
|
||
<taxonomicName box="[1322,1426,389,411]" pageId="16" pageNumber="357" rank="subOrder" subOrder="Adephaga">Adephaga</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Balfour-Browne, F." box="[827,1066,419,441]" journalOrPublisher="Journal of the Royal Microscopical Society" pageId="16" pageNumber="355" pagination="68 - 117" part="64" refId="ref9484" refString="Balfour-Browne, F. (1944) The proventriculus of the Coleoptera (Adephaga) and other insects, a study in evolution. Journal of the Royal Microscopical Society, 64, 68 - 117." title="The proventriculus of the Coleoptera (Adephaga) and other insects, a study in evolution" type="journal article" year="1944">Balfour-Browne, 1944</bibRefCitation>
|
||
; Smrzˇ, 1982) but the backward pointing fringe of hairs on the smaller lobes is considered to be derived. If a proventriculus with lobes bearing sclerotized teeth is primitive compared with the purely hair filter
|
||
<typeStatus box="[876,920,536,558]" pageId="16" pageNumber="355">type</typeStatus>
|
||
(
|
||
<bibRefCitation author="Balfour-Browne, F." box="[933,1163,536,558]" journalOrPublisher="Journal of the Royal Microscopical Society" pageId="16" pageNumber="355" pagination="68 - 117" part="64" refId="ref9484" refString="Balfour-Browne, F. (1944) The proventriculus of the Coleoptera (Adephaga) and other insects, a study in evolution. Journal of the Royal Microscopical Society, 64, 68 - 117." title="The proventriculus of the Coleoptera (Adephaga) and other insects, a study in evolution" type="journal article" year="1944">Balfour-Browne, 1944</bibRefCitation>
|
||
),
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[1181,1236,536,557]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[1181,1236,536,557]" italics="true" pageId="16" pageNumber="355">Meru</emphasis>
|
||
</taxonomicName>
|
||
shares the derived state with
|
||
<taxonomicName baseAuthorityName="Steiner & Anderson" baseAuthorityYear="1981" box="[944,1049,565,587]" class="Insecta" family="Gyrinidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Gyrinidae</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Beutel" baseAuthorityYear="1997" box="[1068,1173,565,587]" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Noteridae</taxonomicName>
|
||
,
|
||
<taxonomicName box="[1192,1301,565,587]" class="Insecta" family="Haliplidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Haliplidae</taxonomicName>
|
||
, and some hydroporine
|
||
<taxonomicName box="[954,1065,594,616]" class="Insecta" family="Dytiscidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Dytiscidae</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="16.[821,1427,156,1843]" pageId="16" pageNumber="355">
|
||
The simple, membranous ovipositor with reduced gonocoxae in
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[969,1096,653,674]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="species" species="phyllisae">
|
||
<emphasis box="[969,1096,653,674]" italics="true" pageId="16" pageNumber="355">M. phyllisae</emphasis>
|
||
</taxonomicName>
|
||
indicates that these beetles are probably depositing eggs on surfaces rather than inserting them in plant tissue, as is the case for some
|
||
<taxonomicName box="[1320,1426,711,733]" class="Insecta" family="Haliplidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Haliplidae</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Beutel, R. G. & Ruhnau, S." box="[827,1076,740,762]" journalOrPublisher="Aquatic Insects" pageId="16" pageNumber="355" pagination="1 - 17" part="12" refId="ref10026" refString="Beutel, R. G. & Ruhnau, S. (1990) Phylogenetic analysis of the genera of Haliplidae (Coleoptera) based on characters of adults. Aquatic Insects, 12, 1 - 17." title="Phylogenetic analysis of the genera of Haliplidae (Coleoptera) based on characters of adults" type="journal article" year="1990">Beutel & Ruhnau, 1990</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Roughley, R. E." box="[1087,1262,740,762]" journalOrPublisher="American Beetles, Vol. 1 (ed. by R. H. Arnett, Jr and M. C. Thomas), pp. 138 - 143. CRC Press, Boca Raton, Florida" pageId="16" pageNumber="355" refId="ref10600" refString="Roughley, R. E. (2001 b) Haliplidae Aube', 1836. American Beetles, Vol. 1 (ed. by R. H. Arnett, Jr and M. C. Thomas), pp. 138 - 143. CRC Press, Boca Raton, Florida." title="Haliplidae Aube'" type="book" year="2001">Roughley, 2001b</bibRefCitation>
|
||
) and
|
||
<taxonomicName authority="(Beutel, 1995)" baseAuthorityName="Beutel" baseAuthorityYear="1995" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">
|
||
Noteridae (
|
||
<bibRefCitation author="Beutel, R. G." box="[828,957,769,791]" journalOrPublisher="Muzeum i Instytut Zoologii PAN, Warszawa" pageId="16" pageNumber="355" refId="ref9728" refString="Beutel, R. G. (1995) The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski), pp. 503 - 539. Muzeum i Instytut Zoologii PAN, Warszawa." title="The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski)" type="book" year="1995">Beutel, 1995</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. This appears to be the result of reductions of the sclerotized gonocoxae and setose appendages seen in most hydradephagan taxa.
|
||
</paragraph>
|
||
<paragraph blockId="16.[821,1427,156,1843]" pageId="16" pageNumber="355">
|
||
We believe that the dytiscoid
|
||
<typeStatus box="[1137,1180,857,879]" pageId="16" pageNumber="355">type</typeStatus>
|
||
of metacoxal fusion and metafurca, along with the other synapomorphies with
|
||
<taxonomicName baseAuthorityName="Beutel" baseAuthorityYear="1995" box="[821,924,915,937]" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Noteridae</taxonomicName>
|
||
, suggest a sister-group relationship between
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[821,918,945,967]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
|
||
<emphasis bold="true" box="[928,946,944,967]" italics="true" pageId="16" pageNumber="355">þ</emphasis>
|
||
<taxonomicName baseAuthorityName="Beutel" baseAuthorityYear="1995" box="[958,1059,945,967]" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Noteridae</taxonomicName>
|
||
<emphasis bold="true" box="[1069,1087,944,967]" italics="true" pageId="16" pageNumber="355">þ</emphasis>
|
||
the remaining Dytiscoidea. The globular antennal scape in
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[1100,1197,974,996]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
|
||
and
|
||
<taxonomicName box="[1252,1359,974,996]" class="Insecta" family="Haliplidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Haliplidae</taxonomicName>
|
||
, along with the nonpredatory feeding and associated features, could also indicate some relationship among these two taxa. A working hypothesis of a relationship among
|
||
<taxonomicName box="[1316,1423,1062,1084]" class="Insecta" family="Haliplidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Haliplidae</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Beutel" baseAuthorityYear="1995" box="[821,922,1091,1113]" class="Insecta" family="Noteridae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Noteridae</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[977,1074,1091,1113]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
|
||
needs consideration. The position of
|
||
<taxonomicName box="[847,951,1120,1142]" class="Insecta" family="Haliplidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Haliplidae</taxonomicName>
|
||
among the other families of
|
||
<taxonomicName box="[1235,1338,1120,1142]" pageId="16" pageNumber="357" rank="subOrder" subOrder="Adephaga">Adephaga</taxonomicName>
|
||
has long been controversial and we hope that the discovery of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[821,918,1178,1200]" class="Insecta" family="Meruidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Meruidae</taxonomicName>
|
||
will lead to clarification of this. The results of molecular analyses in progress, using fresh material of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[821,942,1237,1258]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="species" species="phyllisae">
|
||
<emphasis box="[821,942,1237,1258]" italics="true" pageId="16" pageNumber="355">M. phyllisae</emphasis>
|
||
</taxonomicName>
|
||
(M. Balke, pers. comm.), are eagerly awaited.
|
||
</paragraph>
|
||
<paragraph blockId="16.[821,1427,156,1843]" pageId="16" pageNumber="355">
|
||
Early aquatic
|
||
<taxonomicName box="[1002,1105,1266,1288]" pageId="16" pageNumber="357" rank="subOrder" subOrder="Adephaga">Adephaga</taxonomicName>
|
||
were probably nonswimming forms that lived at the edges of rivers or ponds (
|
||
<bibRefCitation author="Beutel, R. G." journalOrPublisher="Muzeum i Instytut Zoologii PAN, Warszawa" pageId="16" pageNumber="355" refId="ref9728" refString="Beutel, R. G. (1995) The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski), pp. 503 - 539. Muzeum i Instytut Zoologii PAN, Warszawa." title="The Adephaga (Coleoptera): phylogeny and evolutionary history. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80 th Birthday of Roy A. Crowson (ed. by J. Pakaluk and S. A. Slipinski)" type="book" year="1995">Beutel, 1995</bibRefCitation>
|
||
). Hygropetric habitats have existed as ‘edges’ for millennia, as the exposed bedrock has never been under forest canopy, perpetuating the sunlit microhabitats for surface algal growth and the assemblage of often relictual insects associated with them. The recently discovered ‘cliff water beetles’,
|
||
<taxonomicName box="[908,1031,1471,1493]" class="Insecta" family="Aspidytidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Aspidytidae</taxonomicName>
|
||
, demonstrate other unusual taxa to be found at these sites (
|
||
<bibRefCitation author="Ribera, I. & Beutel, R. G. & Balke, M. & Vogler, A. P." box="[1019,1214,1500,1522]" journalOrPublisher="Proceedings of the Royal Society of London (B" pageId="16" pageNumber="355" pagination="2351 - 2356" part="269" refId="ref10419" refString="Ribera, I., Beutel, R. G., Balke, M. & Vogler, A. P. (2002 b) Discovery of Aspidytidae, a new family of aquatic Coleoptera. Proceedings of the Royal Society of London (B), 269, 2351 - 2356." title="Discovery of Aspidytidae, a new family of aquatic Coleoptera" type="journal article" year="2002">
|
||
Ribera
|
||
<emphasis box="[1093,1134,1500,1521]" italics="true" pageId="16" pageNumber="355">et al</emphasis>
|
||
., 2002b
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Balke, M. & Ribera, I. & Beutel, R. G." box="[1223,1395,1500,1522]" journalOrPublisher="Zoologisch-Botanische Gesellschaft in Osterreich and Wiener Coleopterologenverein, Vienna" pageId="16" pageNumber="355" refId="ref9521" refString="Balke, M., Ribera, I. & Beutel, R. G. (2003) Aspidytidae: on the discovery of a new beetle family: detailed morphological analysis, description of a second species, and a key to fossil and extant adephagan families. Water Beetles of China, III (ed. by M. A. Jach and L. Ji), pp. 53 - 66. Zoologisch-Botanische Gesellschaft in Osterreich and Wiener Coleopterologenverein, Vienna." title="Aspidytidae: on the discovery of a new beetle family: detailed morphological analysis, description of a second species, and a key to fossil and extant adephagan families. Water Beetles of China, III (ed. by M. A. Jach and L. Ji)" type="book" year="2003">
|
||
Balke
|
||
<emphasis box="[1285,1326,1500,1521]" italics="true" pageId="16" pageNumber="355">et al</emphasis>
|
||
., 2003
|
||
</bibRefCitation>
|
||
). If there was a common ancestor to the modern aquatic
|
||
<taxonomicName box="[821,924,1558,1580]" pageId="16" pageNumber="357" rank="subOrder" subOrder="Adephaga">Adephaga</taxonomicName>
|
||
(excluding
|
||
<taxonomicName baseAuthorityName="Steiner & Anderson" baseAuthorityYear="1981" box="[1038,1142,1558,1580]" class="Insecta" family="Gyrinidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="family">Gyrinidae</taxonomicName>
|
||
), it probably had the general appearance of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[976,1030,1588,1609]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[976,1030,1588,1609]" italics="true" pageId="16" pageNumber="355">Meru</emphasis>
|
||
</taxonomicName>
|
||
, lacking modifications for swimming, but was probably larger, perhaps resembling an
|
||
<taxonomicName authorityName="LeConte" authorityYear="1853" box="[1321,1419,1617,1638]" class="Insecta" family="Amphizoidae" genus="Amphizoa" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[1321,1419,1617,1638]" italics="true" pageId="16" pageNumber="355">Amphizoa</emphasis>
|
||
</taxonomicName>
|
||
. From this form radiated the diverse and independent adaptations for different swimming, feeding and oviposition strategies. Future studies on additional characters of
|
||
<taxonomicName authorityName="Spangler & Steiner" authorityYear="2005" box="[821,951,1734,1755]" class="Insecta" family="Meruidae" genus="Meru" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="355" phylum="Arthropoda" rank="species" species="phyllisae">
|
||
<emphasis box="[821,951,1734,1755]" italics="true" pageId="16" pageNumber="355">M. phyllisae</emphasis>
|
||
</taxonomicName>
|
||
(parsimony analysis including musculature, presence of defence glands via thin sectioning, molecular data, etc.) and the discovery of the elusive larval stages will further clarify these notions.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |