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<document id="2206B3D94F475501C5F8BCB6B057C299" ID-DOI="10.11646/zootaxa.5458.2.5" ID-ISSN="1175-5326" ID-Zenodo-Dep="11369762" ID-ZooBank="BDE6853F-645E-4F45-88CF-F21938B6E6CE" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1716897820744" checkinUser="plazi" docAuthor="Prokin, Alexander A., Hájek, Jiří, Vasilenko, Dmitry V. &amp; Perkovsky, Evgeny E." docDate="2024" docId="03F7879FFF92FF8E1FB37C21FB33FCF9" docLanguage="en" docName="zootaxa.5458.2.5.pdf" docOrigin="Zootaxa 5458 (2)" docSource="http://dx.doi.org/10.11646/zootaxa.5360.3.8" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Rhantus villumi Prokin &amp; Hájek &amp; Vasilenko &amp; Perkovsky 2024, sp. nov." docType="treatment" docVersion="2" lastPageNumber="268" masterDocId="FFCEFFE7FF93FF8B1F247A39FFE6FFEC" masterDocTitle="The oldest Rhantus (Coleoptera, Dytiscidae) from the earliest Eocene Fur Formation, Denmark" masterLastPageNumber="274" masterPageNumber="263" pageNumber="264" updateTime="1716898317524" updateUser="ExternalLinkService" zenodo-license-document="CLOSED">
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<mods:titleInfo id="09894E87A2E0F0BEA9421F5113E467DB">
<mods:title id="81DA376C26C82201EADEDB693FD50736">The oldest Rhantus (Coleoptera, Dytiscidae) from the earliest Eocene Fur Formation, Denmark</mods:title>
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<mods:name id="A87920F0562618627295CFED9107C03A" type="personal">
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<mods:namePart id="AF61DA1BFBE784A8517DAC8AC13A9E53">Prokin, Alexander A.</mods:namePart>
<mods:nameIdentifier id="24E737665D78A3DB132FCF6971036CAE" type="ORCID">0000-0002-9345-5607</mods:nameIdentifier>
<mods:nameIdentifier id="C259ACD411031A43F74AC8EA7EF2C047" type="email">prokina@mail.ru</mods:nameIdentifier>
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<mods:namePart id="2B250E05907DE11875BC90ADBF42F01E">Hájek, Jiří</mods:namePart>
<mods:affiliation id="4A0E5A93441AA50E5FA19F886F816E9F">Department of Entomology, National Museum, Cirkusová 1740, CZ- 193 00 Prague 9 - Horní Počernice, Czech Republic.</mods:affiliation>
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<mods:namePart id="0589C7BE9CE9DDE68FFB397F988C14CA">Vasilenko, Dmitry V.</mods:namePart>
<mods:affiliation id="0DD559C74C2B197D53ED4B1546B56300">Laboratory of Arthropods, A. A. Borissiak Paleontological Institute, Russian Academy of Sciences, 117647 Moscow, Russia; &amp; Paleontological Laboratory, Cherepovets State University, 162600 Cherepovets, Russia;</mods:affiliation>
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<mods:namePart id="1722B70E59411F94CE66A5546D1E6A99">Perkovsky, Evgeny E.</mods:namePart>
<mods:affiliation id="9111B84FAE7E28131A5C9AF4DF062FD6">Natural History Museum of Denmark, Universitetsparken 15, Copenhagen 2100, Copenhagen, Denmark;</mods:affiliation>
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<mods:typeOfResource id="E7622EE874606E975B4353245390FE44">text</mods:typeOfResource>
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<mods:title id="1247B3CA13849A1EE910F364075019DF">Zootaxa</mods:title>
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<mods:part id="A105DEDA401B9007146F13FFE6A2B9A4">
<mods:date id="F4E305572FA1E63CCB2315205EB45A9F">2024</mods:date>
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<mods:number id="E288282878F74674CB5CD3F975CEBC1D">2024-05-28</mods:number>
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<mods:number id="00F2AFAE7D042FABCB446ADE7DA1E9A5">5458</mods:number>
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<mods:number id="54DB98467387188DFF0B5152B6D58917">2</mods:number>
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<mods:identifier id="982D4304278420B6A07E8FF5ED9DF8C0" type="ISSN">1175-5326</mods:identifier>
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<treatment id="03F7879FFF92FF8E1FB37C21FB33FCF9" LSID="urn:lsid:plazi:treatment:03F7879FFF92FF8E1FB37C21FB33FCF9" httpUri="http://treatment.plazi.org/id/03F7879FFF92FF8E1FB37C21FB33FCF9" lastPageId="5" lastPageNumber="268" pageId="1" pageNumber="264">
<subSubSection id="C3446502FF92FF8A1FB37C21FE28F9DE" box="[151,462,1559,1586]" pageId="1" pageNumber="264" type="nomenclature">
<paragraph id="8BE13689FF92FF8A1FB37C21FE28F9DE" blockId="1.[151,462,1559,1622]" box="[151,462,1559,1586]" pageId="1" pageNumber="264">
<heading id="D0A981E5FF92FF8A1FB37C21FE28F9DE" bold="true" box="[151,462,1559,1586]" fontSize="11" level="1" pageId="1" pageNumber="264" reason="1">
<emphasis id="B92AEA9BFF92FF8A1FB37C21FE28F9DE" bold="true" box="[151,462,1559,1586]" pageId="1" pageNumber="264">
†
<taxonomicName id="4C5E4D0AFF92FF8A1F827C2EFE8FF9DE" authority="Prokin &amp; Hájek &amp; Vasilenko &amp; Perkovsky, 2024" authorityName="Prokin &amp; Hájek &amp; Vasilenko &amp; Perkovsky" authorityYear="2024" box="[166,361,1559,1586]" class="Insecta" family="Dytiscidae" genus="Rhantus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="264" phylum="Arthropoda" rank="species" species="villumi" status="sp. nov.">
<emphasis id="B92AEA9BFF92FF8A1F827C2EFE8FF9DE" bold="true" box="[166,361,1559,1586]" italics="true" pageId="1" pageNumber="264">Rhantus villumi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A21957E0FF92FF8A1E547C21FE28F9DE" box="[368,462,1560,1586]" pageId="1" pageNumber="264" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
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<subSubSection id="C3446502FF92FF8A1FB37C05FEF4F9BA" box="[151,274,1596,1622]" pageId="1" pageNumber="264" type="description">
<paragraph id="8BE13689FF92FF8A1FB37C05FEF4F9BA" blockId="1.[151,462,1559,1622]" box="[151,274,1596,1622]" pageId="1" pageNumber="264">
(
<figureCitation id="13652A0CFF92FF8A1FBB7C05FEEFF9BA" box="[159,265,1596,1622]" captionStart-0="FIGURES 1–2" captionStart-1="FIGURES 3–4" captionStart-2="FIGURES 5–6" captionStart-3="FIGURES 7–8" captionStartId-0="2.[151,264,1526,1550]" captionStartId-1="3.[151,264,1096,1120]" captionStartId-2="3.[151,264,1734,1758]" captionStartId-3="4.[151,264,1943,1967]" captionTargetBox-0="[151,1436,489,1503]" captionTargetBox-1="[156,1429,192,1064]" captionTargetBox-2="[151,1435,1217,1710]" captionTargetBox-3="[195,1392,181,1918]" captionTargetId-0="figure-198@2.[151,1436,489,1503]" captionTargetId-1="figure-17@3.[151,1435,181,1072]" captionTargetId-2="figure-38@3.[151,1435,1217,1710]" captionTargetId-3="figure-33@4.[195,1392,181,1918]" captionTargetPageId-0="2" captionTargetPageId-1="3" captionTargetPageId-2="3" captionTargetPageId-3="4" captionText-0="FIGURES 1–2. †Rhantus villumi sp. nov., photographs of holotype: 1. direct impression; 2. counter impression. Scale bars, 4 mm." captionText-1="FIGURES 3–4. †Rhantus villumi sp. nov., line drawings of holotype: 3. dorsal view; 4. ventral view." captionText-2="FIGURES 5–6. †Rhantus villumi sp. nov., details of counter impression of holotype: 5. part of head, prosternum and mesoventrite, scale bar 1 mm; 6. anterior margin of head with maxillary palpi, base of antenna and distal portion of protarsus, scale bar 0.5 mm." captionText-3="FIGURES 7–8. †Rhantus villumi sp. nov., details of holotype: 7. left hind leg in direct impression, dorsal view; 8. right hind leg in counter impression, ventral view. Scale bars 1mm." figureDoi-0="http://doi.org/10.5281/zenodo.11369768" figureDoi-1="http://doi.org/10.5281/zenodo.11369772" figureDoi-2="http://doi.org/10.5281/zenodo.11369774" figureDoi-3="http://doi.org/10.5281/zenodo.11369776" httpUri-0="https://zenodo.org/record/11369768/files/figure.png" httpUri-1="https://zenodo.org/record/11369772/files/figure.png" httpUri-2="https://zenodo.org/record/11369774/files/figure.png" httpUri-3="https://zenodo.org/record/11369776/files/figure.png" pageId="1" pageNumber="264">Figs. 1–7</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="C3446502FF92FF8A1FB37CBDFB16F92F" pageId="1" pageNumber="264" type="materials_examined">
<paragraph id="8BE13689FF92FF8A1FB37CBDFB16F92F" blockId="1.[151,1436,1668,2019]" pageId="1" pageNumber="264">
<emphasis id="B92AEA9BFF92FF8A1FB37CBDFEA0F972" bold="true" box="[151,326,1668,1694]" pageId="1" pageNumber="264">Type material.</emphasis>
<typeStatus id="54E5882BFF92FF8A1E6A7CBDFE5DF972" box="[334,443,1668,1694]" pageId="1" pageNumber="264" type="holotype">Holotype</typeStatus>
: FUM-N 11573, complete (direct and counter impressions) female imprint. Oval beetle, with genitalia, partly preserved antennae, maxillary palpi, hind legs, and distal portion of protarsus.
</paragraph>
</subSubSection>
<subSubSection id="C3446502FF92FF891FE37CF5FABFFE3D" lastPageId="2" lastPageNumber="265" pageId="1" pageNumber="264" type="description">
<paragraph id="8BE13689FF92FF891FE37CF5FABFFE3D" blockId="1.[151,1436,1668,2019]" lastBlockId="2.[151,1436,151,465]" lastPageId="2" lastPageNumber="265" pageId="1" pageNumber="264">
<emphasis id="B92AEA9BFF92FF8A1FE37CF5FEBFF90A" bold="true" box="[199,345,1740,1766]" pageId="1" pageNumber="264">Description.</emphasis>
A large
<taxonomicName id="4C5E4D0AFF92FF8A1E917CF5FD8DF90A" box="[437,619,1740,1766]" pageId="1" pageNumber="264">
<emphasis id="B92AEA9BFF92FF8A1E917CF5FDF7F90A" box="[437,529,1740,1766]" italics="true" pageId="1" pageNumber="264">Rhantus</emphasis>
species
</taxonomicName>
, total length
<quantity id="4CA69B6CFF92FF8A1DDD7CF5FCB8F90B" box="[761,862,1740,1767]" metricMagnitude="-2" metricUnit="m" metricValue="2.1399999999999997" pageId="1" pageNumber="264" unit="mm" value="21.4">21.4 mm</quantity>
, length without head
<quantity id="4CA69B6CFF92FF8A1B687CF5FB57F90B" box="[1100,1201,1740,1767]" metricMagnitude="-2" metricUnit="m" metricValue="1.86" pageId="1" pageNumber="264" unit="mm" value="18.6">18.6 mm</quantity>
, length of elytra
<quantity id="4CA69B6CFF92FF8A1A4F7CF5FF23F8E7" metricMagnitude="-2" metricUnit="m" metricValue="1.52" pageId="1" pageNumber="264" unit="mm" value="15.2">15.2 mm</quantity>
, maximum width
<quantity id="4CA69B6CFF92FF8A1EB07CC9FE1BF8E7" box="[404,509,1776,1803]" metricMagnitude="-2" metricUnit="m" metricValue="1.09" pageId="1" pageNumber="264" unit="mm" value="10.9">10.9 mm</quantity>
. Head 1.85 times as wide as long, pale with darkened clypeus and subtrapezoidal transverse dark spot medially (
<figureCitation id="13652A0CFF92FF8A1EDC7D2DFDA1F8C2" box="[504,583,1812,1838]" captionStart="FIGURES 1–2" captionStartId="2.[151,264,1526,1550]" captionTargetBox="[151,1436,489,1503]" captionTargetId="figure-198@2.[151,1436,489,1503]" captionTargetPageId="2" captionText="FIGURES 1–2. †Rhantus villumi sp. nov., photographs of holotype: 1. direct impression; 2. counter impression. Scale bars, 4 mm." figureDoi="http://doi.org/10.5281/zenodo.11369768" httpUri="https://zenodo.org/record/11369768/files/figure.png" pageId="1" pageNumber="264">Figs. 1</figureCitation>
,
<figureCitation id="13652A0CFF92FF8A1D737D2DFD81F8C2" box="[599,615,1812,1838]" captionStart="FIGURES 3–4" captionStartId="3.[151,264,1096,1120]" captionTargetBox="[156,1429,192,1064]" captionTargetId="figure-17@3.[151,1435,181,1072]" captionTargetPageId="3" captionText="FIGURES 3–4. †Rhantus villumi sp. nov., line drawings of holotype: 3. dorsal view; 4. ventral view." figureDoi="http://doi.org/10.5281/zenodo.11369772" httpUri="https://zenodo.org/record/11369772/files/figure.png" pageId="1" pageNumber="264">3</figureCitation>
). Antenna filiform with antennomeres I–IV longer than others, ultimate maxillary palpomere longer than penultimate (
<figureCitation id="13652A0CFF92FF8A1D8E7D01FD17F8BE" box="[682,753,1848,1874]" captionStart="FIGURES 5–6" captionStartId="3.[151,264,1734,1758]" captionTargetBox="[151,1435,1217,1710]" captionTargetId="figure-38@3.[151,1435,1217,1710]" captionTargetPageId="3" captionText="FIGURES 5–6. †Rhantus villumi sp. nov., details of counter impression of holotype: 5. part of head, prosternum and mesoventrite, scale bar 1 mm; 6. anterior margin of head with maxillary palpi, base of antenna and distal portion of protarsus, scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.11369774" httpUri="https://zenodo.org/record/11369774/files/figure.png" pageId="1" pageNumber="264">Fig. 6</figureCitation>
). Eye 0.24 as wide as head. Pronotum 3.4 times as wide as long, 1.5 times as wide basally as at its anterior margin; pale medially, with broadly darkened anterior part, lateral parts, and narrowly darkened posterior part projecting anteridad submedially as two spots; lateral bead not evident (
<figureCitation id="13652A0CFF92FF8A1FBB7D9DFF0BF852" box="[159,237,1956,1982]" captionStart="FIGURES 1–2" captionStartId="2.[151,264,1526,1550]" captionTargetBox="[151,1436,489,1503]" captionTargetId="figure-198@2.[151,1436,489,1503]" captionTargetPageId="2" captionText="FIGURES 1–2. †Rhantus villumi sp. nov., photographs of holotype: 1. direct impression; 2. counter impression. Scale bars, 4 mm." figureDoi="http://doi.org/10.5281/zenodo.11369768" httpUri="https://zenodo.org/record/11369768/files/figure.png" pageId="1" pageNumber="264">Figs. 1</figureCitation>
,
<figureCitation id="13652A0CFF92FF8A1FDE7D9DFEECF852" box="[250,266,1956,1982]" captionStart="FIGURES 3–4" captionStartId="3.[151,264,1096,1120]" captionTargetBox="[156,1429,192,1064]" captionTargetId="figure-17@3.[151,1435,181,1072]" captionTargetPageId="3" captionText="FIGURES 3–4. †Rhantus villumi sp. nov., line drawings of holotype: 3. dorsal view; 4. ventral view." figureDoi="http://doi.org/10.5281/zenodo.11369772" httpUri="https://zenodo.org/record/11369772/files/figure.png" pageId="1" pageNumber="264">3</figureCitation>
). Scutellum broadly triangular (
<figureCitation id="13652A0CFF92FF8A1D547D9DFD59F852" box="[624,703,1956,1982]" captionStart="FIGURES 1–2" captionStartId="2.[151,264,1526,1550]" captionTargetBox="[151,1436,489,1503]" captionTargetId="figure-198@2.[151,1436,489,1503]" captionTargetPageId="2" captionText="FIGURES 1–2. †Rhantus villumi sp. nov., photographs of holotype: 1. direct impression; 2. counter impression. Scale bars, 4 mm." figureDoi="http://doi.org/10.5281/zenodo.11369768" httpUri="https://zenodo.org/record/11369768/files/figure.png" pageId="1" pageNumber="264">Figs. 2</figureCitation>
,
<figureCitation id="13652A0CFF92FF8A1DEF7D9DFD3DF853" box="[715,731,1956,1983]" captionStart="FIGURES 5–6" captionStartId="3.[151,264,1734,1758]" captionTargetBox="[151,1435,1217,1710]" captionTargetId="figure-38@3.[151,1435,1217,1710]" captionTargetPageId="3" captionText="FIGURES 5–6. †Rhantus villumi sp. nov., details of counter impression of holotype: 5. part of head, prosternum and mesoventrite, scale bar 1 mm; 6. anterior margin of head with maxillary palpi, base of antenna and distal portion of protarsus, scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.11369774" httpUri="https://zenodo.org/record/11369774/files/figure.png" pageId="1" pageNumber="264">5</figureCitation>
). Elytron translucent with black irrorations around small spots, integrated in rows better visible near lateral margin; epipleuron broad basaly, continuously decreasing to level of posterior margin of abdominal ventrite IV (
<figureCitation id="13652A0CFF91FF891D887AAEFCFDFF5D" box="[684,795,151,177]" captionStart-0="FIGURES 1–2" captionStart-1="FIGURES 3–4" captionStartId-0="2.[151,264,1526,1550]" captionStartId-1="3.[151,264,1096,1120]" captionTargetBox-0="[151,1436,489,1503]" captionTargetBox-1="[156,1429,192,1064]" captionTargetId-0="figure-198@2.[151,1436,489,1503]" captionTargetId-1="figure-17@3.[151,1435,181,1072]" captionTargetPageId-0="2" captionTargetPageId-1="3" captionText-0="FIGURES 1–2. †Rhantus villumi sp. nov., photographs of holotype: 1. direct impression; 2. counter impression. Scale bars, 4 mm." captionText-1="FIGURES 3–4. †Rhantus villumi sp. nov., line drawings of holotype: 3. dorsal view; 4. ventral view." figureDoi-0="http://doi.org/10.5281/zenodo.11369768" figureDoi-1="http://doi.org/10.5281/zenodo.11369772" httpUri-0="https://zenodo.org/record/11369768/files/figure.png" httpUri-1="https://zenodo.org/record/11369772/files/figure.png" pageId="2" pageNumber="265">Figs. 1–4</figureCitation>
). Metacoxal plate about twice as long as lateral lobe of metaventrite (also termed “metasternal wing”) (
<figureCitation id="13652A0CFF91FF891D9C7A82FCEFFF39" box="[696,777,187,213]" captionStart="FIGURES 1–2" captionStartId="2.[151,264,1526,1550]" captionTargetBox="[151,1436,489,1503]" captionTargetId="figure-198@2.[151,1436,489,1503]" captionTargetPageId="2" captionText="FIGURES 1–2. †Rhantus villumi sp. nov., photographs of holotype: 1. direct impression; 2. counter impression. Scale bars, 4 mm." figureDoi="http://doi.org/10.5281/zenodo.11369768" httpUri="https://zenodo.org/record/11369768/files/figure.png" pageId="2" pageNumber="265">Figs. 2</figureCitation>
,
<figureCitation id="13652A0CFF91FF891C337A82FCCEFF39" box="[791,808,187,213]" captionStart="FIGURES 3–4" captionStartId="3.[151,264,1096,1120]" captionTargetBox="[156,1429,192,1064]" captionTargetId="figure-17@3.[151,1435,181,1072]" captionTargetPageId="3" captionText="FIGURES 3–4. †Rhantus villumi sp. nov., line drawings of holotype: 3. dorsal view; 4. ventral view." figureDoi="http://doi.org/10.5281/zenodo.11369772" httpUri="https://zenodo.org/record/11369772/files/figure.png" pageId="2" pageNumber="265">4</figureCitation>
). Abdominal ventrites II and III nearly fused, ventrites I–V subequal in length; ventrite VI about 1.5 times as long as previous ventrites, evenly rounded (
<figureCitation id="13652A0CFF91FF891A3F7AE6FA88FF15" box="[1307,1390,223,249]" captionStart="FIGURES 1–2" captionStartId="2.[151,264,1526,1550]" captionTargetBox="[151,1436,489,1503]" captionTargetId="figure-198@2.[151,1436,489,1503]" captionTargetPageId="2" captionText="FIGURES 1–2. †Rhantus villumi sp. nov., photographs of holotype: 1. direct impression; 2. counter impression. Scale bars, 4 mm." figureDoi="http://doi.org/10.5281/zenodo.11369768" httpUri="https://zenodo.org/record/11369768/files/figure.png" pageId="2" pageNumber="265">Figs. 2</figureCitation>
,
<figureCitation id="13652A0CFF91FF891A5A7AE6FA69FF15" box="[1406,1423,223,249]" captionStart="FIGURES 3–4" captionStartId="3.[151,264,1096,1120]" captionTargetBox="[156,1429,192,1064]" captionTargetId="figure-17@3.[151,1435,181,1072]" captionTargetPageId="3" captionText="FIGURES 3–4. †Rhantus villumi sp. nov., line drawings of holotype: 3. dorsal view; 4. ventral view." figureDoi="http://doi.org/10.5281/zenodo.11369772" httpUri="https://zenodo.org/record/11369772/files/figure.png" pageId="2" pageNumber="265">4</figureCitation>
). Ultimate protarsomere (protarsomere 5) 3 times as long as penultimate protarsomere (protarsomere 4); protarsal claws subequal in size and shape (
<figureCitation id="13652A0CFF91FF891D047B1EFD80FEAD" box="[544,614,295,321]" captionStart="FIGURES 5–6" captionStartId="3.[151,264,1734,1758]" captionTargetBox="[151,1435,1217,1710]" captionTargetId="figure-38@3.[151,1435,1217,1710]" captionTargetPageId="3" captionText="FIGURES 5–6. †Rhantus villumi sp. nov., details of counter impression of holotype: 5. part of head, prosternum and mesoventrite, scale bar 1 mm; 6. anterior margin of head with maxillary palpi, base of antenna and distal portion of protarsus, scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.11369774" httpUri="https://zenodo.org/record/11369774/files/figure.png" pageId="2" pageNumber="265">Fig. 6</figureCitation>
). Metafemur 1.25 times as long as metatibia, with large cavity in apical third for receiving tibia (
<figureCitation id="13652A0CFF91FF891E967B72FE1EFE89" box="[434,504,331,357]" captionStart="FIGURES 7–8" captionStartId="4.[151,264,1943,1967]" captionTargetBox="[195,1392,181,1918]" captionTargetId="figure-33@4.[195,1392,181,1918]" captionTargetPageId="4" captionText="FIGURES 7–8. †Rhantus villumi sp. nov., details of holotype: 7. left hind leg in direct impression, dorsal view; 8. right hind leg in counter impression, ventral view. Scale bars 1mm." figureDoi="http://doi.org/10.5281/zenodo.11369776" httpUri="https://zenodo.org/record/11369776/files/figure.png" pageId="2" pageNumber="265">Fig. 7</figureCitation>
). Metatibia slightly widened towards apex, with line of 8 setigerous punctures on anterior surface, strong sharp setae on dorsal and apical margins, and with long and slender setae on ventral margin (
<figureCitation id="13652A0CFF91FF891FBB7BAAFF03FE41" box="[159,229,403,429]" captionStart="FIGURES 7–8" captionStartId="4.[151,264,1943,1967]" captionTargetBox="[195,1392,181,1918]" captionTargetId="figure-33@4.[195,1392,181,1918]" captionTargetPageId="4" captionText="FIGURES 7–8. †Rhantus villumi sp. nov., details of holotype: 7. left hind leg in direct impression, dorsal view; 8. right hind leg in counter impression, ventral view. Scale bars 1mm." figureDoi="http://doi.org/10.5281/zenodo.11369776" httpUri="https://zenodo.org/record/11369776/files/figure.png" pageId="2" pageNumber="265">Fig. 7</figureCitation>
). Metatarsomere I twice as long as metatarsomere II, III nearly 3 times as long as IV and V; all tarsomeres with long natatorial setae on dorsal margin, and strong sharp setae on ventral margin; claws unequal (
<figureCitation id="13652A0CFF91FF891A237B8EFAADFE3D" box="[1287,1355,439,465]" captionStart="FIGURES 7–8" captionStartId="4.[151,264,1943,1967]" captionTargetBox="[195,1392,181,1918]" captionTargetId="figure-33@4.[195,1392,181,1918]" captionTargetPageId="4" captionText="FIGURES 7–8. †Rhantus villumi sp. nov., details of holotype: 7. left hind leg in direct impression, dorsal view; 8. right hind leg in counter impression, ventral view. Scale bars 1mm." figureDoi="http://doi.org/10.5281/zenodo.11369776" httpUri="https://zenodo.org/record/11369776/files/figure.png" pageId="2" pageNumber="265">Fig. 8</figureCitation>
).
</paragraph>
</subSubSection>
<caption id="DF216601FF91FF891FB37FCFFF20F9DE" ID-DOI="http://doi.org/10.5281/zenodo.11369768" ID-Zenodo-Dep="11369768" httpUri="https://zenodo.org/record/11369768/files/figure.png" pageId="2" pageNumber="265" startId="2.[151,264,1526,1550]" targetBox="[151,1436,489,1503]" targetPageId="2" targetType="figure">
<paragraph id="8BE13689FF91FF891FB37FCFFF20F9DE" blockId="2.[151,1437,1526,1586]" pageId="2" pageNumber="265">
<emphasis id="B92AEA9BFF91FF891FB37FCFFEDDF9E2" bold="true" box="[151,315,1526,1550]" pageId="2" pageNumber="265">FIGURES 1–2.</emphasis>
†
<taxonomicName id="4C5E4D0AFF91FF891E6A7FCFFE08F9E2" authority="Prokin &amp; Hájek &amp; Vasilenko &amp; Perkovsky, 2024" authorityName="Prokin &amp; Hájek &amp; Vasilenko &amp; Perkovsky" authorityYear="2024" box="[334,494,1526,1550]" class="Insecta" family="Dytiscidae" genus="Rhantus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="265" phylum="Arthropoda" rank="species" species="villumi" status="sp. nov.">
<emphasis id="B92AEA9BFF91FF891E6A7FCFFE08F9E2" box="[334,494,1526,1550]" italics="true" pageId="2" pageNumber="265">Rhantus villumi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A21957E0FF91FF891ED17FCFFDA1F9E2" box="[501,583,1526,1550]" pageId="2" pageNumber="265" rank="species">
<emphasis id="B92AEA9BFF91FF891ED17FCFFDA7F9E2" bold="true" box="[501,577,1526,1550]" pageId="2" pageNumber="265">sp. nov</emphasis>
.
</taxonomicNameLabel>
, photographs of holotype: 1. direct impression; 2. counter impression. Scale bars, 4 mm.
</paragraph>
</caption>
<subSubSection id="C3446502FF91FF8E1FE37C59FCA6FD21" lastPageId="5" lastPageNumber="268" pageId="2" pageNumber="265" type="discussion">
<paragraph id="8BE13689FF91FF891FE37C59FCADF876" blockId="2.[151,1437,1632,1946]" pageId="2" pageNumber="265">
<emphasis id="B92AEA9BFF91FF891FE37C59FDF3F996" bold="true" box="[199,533,1632,1658]" pageId="2" pageNumber="265">Comments on classification.</emphasis>
Within the extant subfamilies of
<taxonomicName id="4C5E4D0AFF91FF891CAE7C59FBE3F996" box="[906,1029,1632,1658]" class="Insecta" family="Dytiscidae" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="265" phylum="Arthropoda" rank="family">Dytiscidae</taxonomicName>
, we place the fossil in the subfamily
<taxonomicName id="4C5E4D0AFF91FF891FB37CBDFED1F972" authorityName="Erichson" authorityYear="1837" box="[151,311,1668,1694]" class="Insecta" family="Dytiscidae" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="265" phylum="Arthropoda" rank="subFamily" subFamily="Colymbetinae">Colymbetinae</taxonomicName>
, based on the following combination of characters: 1) eyes anterolaterally emarginate; 2) apical maxillary palpomere simple, not biramous; 3) mesoscutellum (scutellar shield) clearly visible with elytra closed; 4) prosternal process without longitudinal groove, obtusely lanceolate; 5) apex of elytron evenly rounded; 6) lateral parts of metaventrite broadly triangular; 7) metacoxal lines present, broadly separated; 8) metafemur without linear series of setae near anteroapical angle; 9) metatarsal claws unequal (see
<bibRefCitation id="EFCF4B78FF91FF891CF67D2DFB01F8C2" author="Miller, K. B. &amp; Bergsten, J." box="[978,1255,1812,1839]" pageId="2" pageNumber="265" refId="ref6933" refString="Miller, K. B. &amp; Bergsten, J. (2016) Diving beetles of the World. Systematics and biology of Dytiscidae. John Hopkins University Press, Baltimore, Maryland, 320 pp." type="book" year="2016">Miller &amp; Bergsten 2016</bibRefCitation>
). It differs from the only extinct subfamily †Liadytiscinae in the maximum length of the metacoxal plate greater than the maximum length of the lateral part of the metaventrite, metatarsomeres 1–4 with lobed apical margins, metatibia shorter than metafemur, unequal metatarsal claws (see
<bibRefCitation id="EFCF4B78FF91FF891D567DB9FCA6F876" author="Prokin, A. A. &amp; Petrov, P. N. &amp; Wang, B. &amp; Ponomarenko, A. G." box="[626,832,1920,1946]" pageId="2" pageNumber="265" pagination="137 - 159" refId="ref7619" refString="Prokin, A. A., Petrov, P. N., Wang, B. &amp; Ponomarenko, A. G. (2013) New fossil taxa and notes on the Mesozoic evolution of Liadytidae and Dytiscidae (Coleoptera). Zootaxa, 3666 (2), 137 - 159. https: // doi. org / 10.11646 / zootaxa. 3666.2.2" type="journal article" year="2013">
Prokin
<emphasis id="B92AEA9BFF91FF891DE07DB8FD1AF876" box="[708,764,1920,1946]" italics="true" pageId="2" pageNumber="265">et al.</emphasis>
2013
</bibRefCitation>
).
</paragraph>
<caption id="DF216601FF90FF881FB37E71FB7BFB8C" ID-DOI="http://doi.org/10.5281/zenodo.11369772" ID-Zenodo-Dep="11369772" box="[151,1181,1096,1121]" httpUri="https://zenodo.org/record/11369772/files/figure.png" pageId="3" pageNumber="266" startId="3.[151,264,1096,1120]" targetBox="[156,1429,192,1064]" targetPageId="3" targetType="figure">
<paragraph id="8BE13689FF90FF881FB37E71FB7BFB8C" blockId="3.[151,1181,1096,1121]" box="[151,1181,1096,1121]" pageId="3" pageNumber="266">
<emphasis id="B92AEA9BFF90FF881FB37E71FEDDFB8C" bold="true" box="[151,315,1096,1120]" pageId="3" pageNumber="266">FIGURES 3–4.</emphasis>
†
<taxonomicName id="4C5E4D0AFF90FF881E697E71FE0BFB8C" authority="Prokin &amp; Hájek &amp; Vasilenko &amp; Perkovsky, 2024" authorityName="Prokin &amp; Hájek &amp; Vasilenko &amp; Perkovsky" authorityYear="2024" box="[333,493,1096,1120]" class="Insecta" family="Dytiscidae" genus="Rhantus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="266" phylum="Arthropoda" rank="species" species="villumi" status="sp. nov.">
<emphasis id="B92AEA9BFF90FF881E697E71FE0BFB8C" box="[333,493,1096,1120]" italics="true" pageId="3" pageNumber="266">Rhantus villumi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A21957E0FF90FF881ED07E70FDA0FB8C" box="[500,582,1097,1121]" pageId="3" pageNumber="266" rank="species">
<emphasis id="B92AEA9BFF90FF881ED07E70FDD9FB8D" bold="true" box="[500,575,1097,1121]" pageId="3" pageNumber="266">sp. nov</emphasis>
.
</taxonomicNameLabel>
, line drawings of holotype: 3. dorsal view; 4. ventral view.
</paragraph>
</caption>
<caption id="DF216601FF90FF881FB37CFFFEAAF8CA" ID-DOI="http://doi.org/10.5281/zenodo.11369774" ID-Zenodo-Dep="11369774" httpUri="https://zenodo.org/record/11369774/files/figure.png" pageId="3" pageNumber="266" startId="3.[151,264,1734,1758]" targetBox="[151,1435,1217,1710]" targetPageId="3" targetType="figure">
<paragraph id="8BE13689FF90FF881FB37CFFFEAAF8CA" blockId="3.[151,1437,1734,1830]" pageId="3" pageNumber="266">
<emphasis id="B92AEA9BFF90FF881FB37CFFFEA6F932" bold="true" box="[151,320,1734,1758]" pageId="3" pageNumber="266">FIGURES 5–6.</emphasis>
†
<taxonomicName id="4C5E4D0AFF90FF881E7D7CFFFE18F932" authority="Prokin &amp; Hájek &amp; Vasilenko &amp; Perkovsky, 2024" authorityName="Prokin &amp; Hájek &amp; Vasilenko &amp; Perkovsky" authorityYear="2024" box="[345,510,1734,1758]" class="Insecta" family="Dytiscidae" genus="Rhantus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="266" phylum="Arthropoda" rank="species" species="villumi" status="sp. nov.">
<emphasis id="B92AEA9BFF90FF881E7D7CFFFE18F932" box="[345,510,1734,1758]" italics="true" pageId="3" pageNumber="266">Rhantus villumi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A21957E0FF90FF881D2E7CFEFD84F932" box="[522,610,1735,1759]" pageId="3" pageNumber="266" rank="species">
<emphasis id="B92AEA9BFF90FF881D2E7CFEFDBDF933" bold="true" box="[522,603,1735,1759]" pageId="3" pageNumber="266">sp. nov</emphasis>
.
</taxonomicNameLabel>
, details of counter impression of holotype: 5. part of head, prosternum and mesoventrite, scale bar 1 mm; 6. anterior margin of head with maxillary palpi, base of antenna and distal portion of protarsus, scale bar 0.5 mm.
</paragraph>
</caption>
<caption id="DF216601FF97FF8F1FB37DAEFD30F83F" ID-DOI="http://doi.org/10.5281/zenodo.11369776" ID-Zenodo-Dep="11369776" httpUri="https://zenodo.org/record/11369776/files/figure.png" pageId="4" pageNumber="267" startId="4.[151,264,1943,1967]" targetBox="[195,1392,181,1918]" targetPageId="4" targetType="figure">
<paragraph id="8BE13689FF97FF8F1FB37DAEFD30F83F" blockId="4.[151,1436,1943,2003]" pageId="4" pageNumber="267">
<emphasis id="B92AEA9BFF97FF8F1FB37DAEFEDAF843" bold="true" box="[151,316,1943,1967]" pageId="4" pageNumber="267">FIGURES 7–8.</emphasis>
†
<taxonomicName id="4C5E4D0AFF97FF8F1E6B7DAEFE09F843" authority="Prokin &amp; Hájek &amp; Vasilenko &amp; Perkovsky, 2024" authorityName="Prokin &amp; Hájek &amp; Vasilenko &amp; Perkovsky" authorityYear="2024" box="[335,495,1943,1967]" class="Insecta" family="Dytiscidae" genus="Rhantus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="267" phylum="Arthropoda" rank="species" species="villumi" status="sp. nov.">
<emphasis id="B92AEA9BFF97FF8F1E6B7DAEFE09F843" box="[335,495,1943,1967]" italics="true" pageId="4" pageNumber="267">Rhantus villumi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A21957E0FF97FF8F1ED37DAEFDACF843" box="[503,586,1943,1967]" pageId="4" pageNumber="267" rank="species">
<emphasis id="B92AEA9BFF97FF8F1ED37DAEFDA5F843" bold="true" box="[503,579,1943,1967]" pageId="4" pageNumber="267">sp. nov</emphasis>
.
</taxonomicNameLabel>
, details of holotype: 7. left hind leg in direct impression, dorsal view; 8. right hind leg in counter impression, ventral view. Scale bars 1mm.
</paragraph>
</caption>
<paragraph id="8BE13689FF96FF8E1FE37AAEFA89FE19" blockId="5.[151,1437,151,789]" pageId="5" pageNumber="268">
Within the subfamily
<taxonomicName id="4C5E4D0AFF96FF8E1EEF7AAEFD8DFF5D" authorityName="Erichson" authorityYear="1837" box="[459,619,151,177]" pageId="5" pageNumber="268" rank="subFamily" subFamily="Colymbetinae">Colymbetinae</taxonomicName>
, the fossil is tentatively assigned to the cosmopolitan genus
<taxonomicName id="4C5E4D0AFF96FF8E1A647AAEFEC9FF39" authority="Dejean, 1833" authorityName="Dejean" authorityYear="1833" class="Insecta" family="Dytiscidae" genus="Rhantus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="268" phylum="Arthropoda" rank="genus">
<emphasis id="B92AEA9BFF96FF8E1A647AAEFA7AFF5D" box="[1344,1436,151,177]" italics="true" pageId="5" pageNumber="268">Rhantus</emphasis>
Dejean, 1833
</taxonomicName>
, mainly based on the following characters (see
<bibRefCitation id="EFCF4B78FF96FF8E1C1A7A82FBE7FF39" author="Balke, M. &amp; HAjek, J. &amp; Hendrich, L." box="[830,1025,187,213]" pageId="5" pageNumber="268" pagination="91 - 100" refId="ref5132" refString="Balke, M., HAjek, J. &amp; Hendrich, L. (2017) Generic reclassification of species formerly included in Rhantus Dejean (Coleoptera, Dytiscidae, Colymbetinae). Zootaxa, 4258 (1), 91 - 100. https: // doi. org / 10.11646 / zootaxa. 4258.1.7" type="journal article" year="2017">
Balke
<emphasis id="B92AEA9BFF96FF8E1CA27A82FC58FF39" box="[902,958,187,213]" italics="true" pageId="5" pageNumber="268">et al.</emphasis>
2017
</bibRefCitation>
): 1) body elongated oval, not slender or torpedo-shaped; 2) sides of pronotum with indistinct\narrow lateral bead; 3) posterior pronotal angles weakly obtuse; 4) anterior margin of metaventrite deeply incised for receiving prosternal process; 5) anterior surface of metatibia with several setigerous punctures arranged in linear row; 6) metatarsomeres weakly lobed on posteroapical margin. The genus
<taxonomicName id="4C5E4D0AFF96FF8E1E577B72FE29FE89" authorityName="Dejean" authorityYear="1833" box="[371,463,331,357]" class="Insecta" family="Dytiscidae" genus="Rhantus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="268" phylum="Arthropoda" rank="genus">
<emphasis id="B92AEA9BFF96FF8E1E577B72FE29FE89" box="[371,463,331,357]" italics="true" pageId="5" pageNumber="268">Rhantus</emphasis>
</taxonomicName>
, in the current sense, is only molecularly defined, without any known morphological synapomorphy and without clear known differences, especially from the genus
<taxonomicName id="4C5E4D0AFF96FF8E1B117B56FEA0FE41" authority="Balke, Hendrich &amp; Wewalka, 1992" authorityName="Balke, Hendrich &amp; Wewalka" authorityYear="1992" class="Insecta" family="Dytiscidae" genus="Carabdytes" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="268" phylum="Arthropoda" rank="genus">
<emphasis id="B92AEA9BFF96FF8E1B117B56FB50FE65" box="[1077,1206,367,393]" italics="true" pageId="5" pageNumber="268">Carabdytes</emphasis>
Balke, Hendrich &amp; Wewalka, 1992
</taxonomicName>
(see
<bibRefCitation id="EFCF4B78FF96FF8E1EA77BAAFDACFE41" author="Balke, M. &amp; HAjek, J. &amp; Hendrich, L." box="[387,586,403,429]" pageId="5" pageNumber="268" pagination="91 - 100" refId="ref5132" refString="Balke, M., HAjek, J. &amp; Hendrich, L. (2017) Generic reclassification of species formerly included in Rhantus Dejean (Coleoptera, Dytiscidae, Colymbetinae). Zootaxa, 4258 (1), 91 - 100. https: // doi. org / 10.11646 / zootaxa. 4258.1.7" type="journal article" year="2017">
Balke
<emphasis id="B92AEA9BFF96FF8E1EE87BAAFDE3FE41" box="[460,517,403,429]" italics="true" pageId="5" pageNumber="268">et al.</emphasis>
2017
</bibRefCitation>
). However, we consider the presence of the latter genus, which is currently distributed exclusively on Pacific Islands, in Eocene of Europe to be rather unlikely, and therefore prefer to classify the fossil as
<emphasis id="B92AEA9BFF96FF8E1E067BE2FE63FE19" box="[290,389,475,501]" italics="true" pageId="5" pageNumber="268">
<taxonomicName id="4C5E4D0AFF96FF8E1E067BE2FE67FE19" authorityName="Dejean" authorityYear="1833" box="[290,385,475,501]" class="Insecta" family="Dytiscidae" genus="Rhantus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="268" phylum="Arthropoda" rank="genus">Rhantus</taxonomicName>
,
</emphasis>
of which 11 species are currently known to occur in Europe (see
<bibRefCitation id="EFCF4B78FF96FF8E1B477BE2FA85FE19" author="Nilsson, A. N. &amp; HAjek, J." box="[1123,1379,475,503]" pageId="5" pageNumber="268" refId="ref7058" refString="Nilsson, A. N. &amp; HAjek, J. (2024) A World Catalogue of the family Dytiscidae, or the diving beetles (Coleoptera, Adephaga). Version 1 January 2024. Distributed as a PDF file via Internet. Available from: http: // www. waterbeetles. eu (accessed 20 January 2024)" type="url" year="2024">Nilsson &amp; HÁjek 2024</bibRefCitation>
).
</paragraph>
<paragraph id="8BE13689FF96FF8E1FE37BC6FCA6FD21" blockId="5.[151,1437,151,789]" pageId="5" pageNumber="268">
The female
<typeStatus id="54E5882BFF96FF8E1E6E7BC6FE4AFDF5" box="[330,428,511,537]" pageId="5" pageNumber="268" type="holotype">holotype</typeStatus>
imprint lacks the usual characters used to identify recent
<taxonomicName id="4C5E4D0AFF96FF8E1B057BC6FB35FDF5" box="[1057,1235,511,537]" pageId="5" pageNumber="268">
<emphasis id="B92AEA9BFF96FF8E1B057BC6FB9BFDF5" box="[1057,1149,511,537]" italics="true" pageId="5" pageNumber="268">Rhantus</emphasis>
species
</taxonomicName>
(elytral sculpture, shape of male claws, shape of male genitalia; see, e.g.
<bibRefCitation id="EFCF4B78FF96FF8E1C33781AFBD0FDD1" author="Nilsson, A. N. &amp; Holmen, M." box="[791,1078,546,573]" pageId="5" pageNumber="268" pagination="1 - 192" refId="ref7120" refString="Nilsson, A. N. &amp; Holmen, M. (1995) The aquatic Adephaga (Coleoptera) of Fennoscandia and Denmark. II. Dytiscidae. Fauna Entomologica Scandinavica, 32, 1 - 192. https: // doi. org / 10.1163 / 9789004273603 _ 001" type="journal article" year="1995">Nilsson &amp; Holmen 1995</bibRefCitation>
). Therefore, the new species is mainly distinguished on the basis of its large body size, which is significantly larger than in all known
<taxonomicName id="4C5E4D0AFF96FF8E1A64787EFF0EFD69" pageId="5" pageNumber="268">
<emphasis id="B92AEA9BFF96FF8E1A64787EFA7AFD8D" box="[1344,1436,583,609]" italics="true" pageId="5" pageNumber="268">Rhantus</emphasis>
species
</taxonomicName>
(body size
<quantity id="4CA69B6CFF96FF8E1E4C7853FE19FD69" box="[360,511,618,645]" metricMagnitude="-2" metricUnit="m" metricValue="1.175" metricValueMax="1.78" metricValueMin="0.57" pageId="5" pageNumber="268" unit="mm" value="11.75" valueMax="17.8" valueMin="5.7">5.7–17.8 mm</quantity>
, according to
<bibRefCitation id="EFCF4B78FF96FF8E1DBF7852FC4CFD69" author="Miller, K. B. &amp; Bergsten, J." box="[667,938,619,645]" pageId="5" pageNumber="268" refId="ref6933" refString="Miller, K. B. &amp; Bergsten, J. (2016) Diving beetles of the World. Systematics and biology of Dytiscidae. John Hopkins University Press, Baltimore, Maryland, 320 pp." type="book" year="2016">Miller &amp; Bergsten 2016</bibRefCitation>
). In addition, the colouration of the head and pronotum may also be of limited use for alpha taxonomy, although it varies in many extant species (see, e.g.
<bibRefCitation id="EFCF4B78FF96FF8E1A6378B6FEB7FD21" author="Nilsson, A. N. &amp; Holmen, M." pageId="5" pageNumber="268" pagination="1 - 192" refId="ref7120" refString="Nilsson, A. N. &amp; Holmen, M. (1995) The aquatic Adephaga (Coleoptera) of Fennoscandia and Denmark. II. Dytiscidae. Fauna Entomologica Scandinavica, 32, 1 - 192. https: // doi. org / 10.1163 / 9789004273603 _ 001" type="journal article" year="1995">Nilsson &amp; Holmen 1995</bibRefCitation>
: figs. 507–516;
<bibRefCitation id="EFCF4B78FF96FF8E1D21788DFD6DFD21" author="HAjek, J." box="[517,651,691,719]" pageId="5" pageNumber="268" pagination="1 - 32" refId="ref5967" refString="HAjek, J. (2009) Icones Insectorum Europae Centralis. Coleoptera: Dytiscidae. Folia Heyrovskyana, Series B, 13, 1 - 32." type="journal article" year="2009">HÁjek 2009</bibRefCitation>
: figs. 138–146).
</paragraph>
</subSubSection>
<subSubSection id="C3446502FF96FF8E1FE378EEFB33FCF9" pageId="5" pageNumber="268" type="etymology">
<paragraph id="8BE13689FF96FF8E1FE378EEFB33FCF9" blockId="5.[151,1437,151,789]" pageId="5" pageNumber="268">
<emphasis id="B92AEA9BFF96FF8E1FE378EEFEA8FD1D" bold="true" box="[199,334,727,753]" pageId="5" pageNumber="268">Etymology.</emphasis>
This species is dedicated to the Villum Foundation, named after Villum Kann Rasmussen (1909– 1993), whose financial support enables the EEP to continue the study of the Eocene insect fauna.
</paragraph>
</subSubSection>
</treatment>
</document>
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