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<mods:title id="ABDBDD06FAFA87FD06751446D63FCB6D">Reclassification of Cybistrinae Sharp, 1880 in the Neotropical Region (Coleoptera, Adephaga, Dytiscidae), with description of new taxa</mods:title>
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<mods:affiliation id="517FC6EEA52B91C0E349C7161F6D2A4B">Department of Biology and Museum of Southwestern Biology, University of New Mexico, Albuquerque, NM 87131 - 0001, USA</mods:affiliation>
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<mods:affiliation id="23A90A31EB3B7DAEFDEDD597A080349D">Instituto de Biodiversidad y Biologia Experimental y Aplicada, CONICET-Universidad de Buenos Aires, Buenos Aires, Argentina</mods:affiliation>
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<paragraph id="96AFFA6B355D3F8F93C586648948DC6C" pageId="0" pageNumber="125">
<taxonomicName id="E53581AF3FC0E27891425A858058DC17" LSID="B39BAFF7-362D-5968-AA6A-C3F82FC6FB23" authority="Brinck, 1945" authorityName="Brinck" authorityYear="1945" class="Insecta" family="Dytiscidae" genus="Bifurcitus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Bifurcitus" order="Coleoptera" pageId="0" pageNumber="125" phylum="Arthropoda" rank="genus" status="stat. nov.">Bifurcitus Brinck, 1945</taxonomicName>
<taxonomicNameLabel id="F39689758CF3506637A1F90E601C1FBD" pageId="0" pageNumber="125">stat. nov.</taxonomicNameLabel>
</paragraph>
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<subSubSection id="4DB2F3AFD6BAF70231BF60EF3BF2FC04" pageId="0" pageNumber="125" type="description">
<paragraph id="E3F934C4736C743122CD28AE8B54DDD5" pageId="0" pageNumber="125">
<figureCitation id="DD5AAAE4B1C87F04D73E33A9E7FF40A4" captionStart="Figures 18" captionStartId="F1" captionText="Figures 1 - 8. Cybistrinae morphological features 1 Bifurcitus lherminieri, right metatibia, posterior surface 2 Bifurcitus lherminieri, left metaleg, anterior surface 3 Trifurcitus robustus, left metaleg, anterior surface 4 Metaxydytes fraternus, left mesotarsomeres III-V, posterior surfaces 5 Cybister tripunctatus, left mesotarsomeres III-V, posterior surfaces 6 Megadytes latus, left half of metaventrite and left metacoxa (WV = metaventrite wing width, WC = metacoxal width) 7 Cybister tripunctatus, right metathoracic wing 8 Trifurcitus robustus, right metatrochanter, ventral surface. Scale bars: 1.0 mm." figureDoi="10.3897/zookeys.1188.110081.figures1-8" httpUri="https://binary.pensoft.net/fig/963583" pageId="0" pageNumber="125">Figs 1</figureCitation>
<figureCitation id="9ABA86E2BD690EBCF4FD502D9BF09E6C" captionStart="Figures 18" captionStartId="F1" captionText="Figures 1 - 8. Cybistrinae morphological features 1 Bifurcitus lherminieri, right metatibia, posterior surface 2 Bifurcitus lherminieri, left metaleg, anterior surface 3 Trifurcitus robustus, left metaleg, anterior surface 4 Metaxydytes fraternus, left mesotarsomeres III-V, posterior surfaces 5 Cybister tripunctatus, left mesotarsomeres III-V, posterior surfaces 6 Megadytes latus, left half of metaventrite and left metacoxa (WV = metaventrite wing width, WC = metacoxal width) 7 Cybister tripunctatus, right metathoracic wing 8 Trifurcitus robustus, right metatrochanter, ventral surface. Scale bars: 1.0 mm." figureDoi="10.3897/zookeys.1188.110081.figures1-8" httpUri="https://binary.pensoft.net/fig/963583" pageId="0" pageNumber="125">, 2</figureCitation>
<figureCitation id="7A5B75BD85BE23B58165AED585669D6A" captionStart="Figures 6266" captionStartId="F7" captionText="Figures 62 - 66. Neotropical Cybistrinae species, female genitalia including left gonocoxosternite, ventral aspect 62 Metaxydytes fraternus 63 Paramegadytes glaucus 64 Bifurcitus lherminieri 65 Trifurcitus robustus 66 Cybister fimbriolatus. Scale bars: 1.0 mm." figureDoi="10.3897/zookeys.1188.110081.figures62-66" httpUri="https://binary.pensoft.net/fig/963589" pageId="0" pageNumber="125">, 64</figureCitation>
<figureCitation id="7255DA0B9CA9F24A21EC5C25C1509954" captionStart="Figures 6771" captionStartId="F8" captionText="Figures 67 - 71. Third instar larva, anterior clypeal margin 67 Bifurcitus magnus 68 Megadytes latus 69 Metaxydytes carcharias 70 Paramegadytes glaucus 71 Trifurcitus fallax." figureDoi="10.3897/zookeys.1188.110081.figures67-71" httpUri="https://binary.pensoft.net/fig/963590" pageId="0" pageNumber="125">, 67</figureCitation>
</paragraph>
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<subSubSection id="B54A276FD6259688AABA41C97B15862B" pageId="0" pageNumber="125" type="reference_group">
<paragraph id="F875D696B3F966043FDF0E85ACCDDD64" pageId="0" pageNumber="125">
<taxonomicName id="404512F9BD353D9489AD2B116FFE1314" authorityName="Miller &amp; Michat &amp; Ferreira Jr" authorityYear="2024" class="Insecta" family="Dytiscidae" genus="Bifurcitus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Bifurcitus" order="Coleoptera" pageId="0" pageNumber="125" phylum="Arthropoda" rank="genus" status="stat. nov.">Bifurcitus</taxonomicName>
Brinck, 1945: 8.
</paragraph>
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<subSubSection id="F9B93C6AEC3C1E7F2B93B6ECE78ACC82" pageId="0" pageNumber="125" type="type species">
<paragraph id="54D463ECD72D49769AB34F5DF90C07A1" pageId="0" pageNumber="125">Type species.</paragraph>
<paragraph id="926708AC15890E69391A6B3291D9E4DC" pageId="0" pageNumber="125">
<taxonomicName id="0596BEA9EF2005570B2E45D77D13D920" authorityName="Laporte de Castelnau" authorityYear="1835" class="Insecta" family="Dytiscidae" genus="Cybister" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cybister giganteus" order="Coleoptera" pageId="0" pageNumber="125" phylum="Arthropoda" rank="species" species="giganteus">
<emphasis id="CF60AAE1F178213722A7F9F3A540C66D" italics="true" pageId="0" pageNumber="125">Cybister giganteus</emphasis>
</taxonomicName>
Laporte, 1835: 99 by original designation (=
<taxonomicName id="351F5596E353EB68C9AB5FA0FBEA388D" authorityName="Guérin-Méneville" authorityYear="1829" class="Insecta" family="Dytiscidae" genus="Dytiscus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Dytiscus lherminieri" order="Coleoptera" pageId="0" pageNumber="125" phylum="Arthropoda" rank="species" species="lherminieri">
<emphasis id="27BE49CC9B702F7119B138F52E6BEB3F" italics="true" pageId="0" pageNumber="125">Dytiscus lherminieri</emphasis>
</taxonomicName>
<normalizedToken id="B9C09F767508F595D71148C7D64985C4" originalValue="Guérin-Méneville">Guerin-Meneville</normalizedToken>
, 1829).
</paragraph>
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<subSubSection id="22644497003936A28BAF8B126FF4056A" pageId="0" pageNumber="125" type="diagnosis">
<paragraph id="A21EEF49A1DAC5A3B4428F6BBF0021A3" pageId="0" pageNumber="125">Diagnosis.</paragraph>
<paragraph id="A0D6440CEB9B5EDFBF31CAE82C9BB005" pageId="0" pageNumber="125">
Within
<taxonomicName id="FE8B2467C833089F824BDCB7BAA993FC" authorityName="Sharp" authorityYear="1880" lsidName="" pageId="0" pageNumber="125" rank="subFamily" subFamily="Cybistrinae">Cybistrinae</taxonomicName>
<taxonomicName id="0C73DBD5DDB7EC13A6833295FD950FCF" authorityName="Miller &amp; Michat &amp; Ferreira Jr" authorityYear="2024" class="Insecta" family="Dytiscidae" genus="Bifurcitus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Bifurcitus" order="Coleoptera" pageId="0" pageNumber="125" phylum="Arthropoda" rank="genus">
<emphasis id="293049A394531D3ABCA53B21E732C5DF" italics="true" pageId="0" pageNumber="125">Bifurcitus</emphasis>
</taxonomicName>
have (1) the lateral margins of the pronotum and elytra margined with yellow, (2) males and females each with two equal-length metatarsal claws, and (3) the posterior metatibial spur bifid (Figs
<figureCitation id="D0A39CAE7462AA01A0C21B797FF02243" captionStart="Figures 18" captionStartId="F1" captionText="Figures 1 - 8. Cybistrinae morphological features 1 Bifurcitus lherminieri, right metatibia, posterior surface 2 Bifurcitus lherminieri, left metaleg, anterior surface 3 Trifurcitus robustus, left metaleg, anterior surface 4 Metaxydytes fraternus, left mesotarsomeres III-V, posterior surfaces 5 Cybister tripunctatus, left mesotarsomeres III-V, posterior surfaces 6 Megadytes latus, left half of metaventrite and left metacoxa (WV = metaventrite wing width, WC = metacoxal width) 7 Cybister tripunctatus, right metathoracic wing 8 Trifurcitus robustus, right metatrochanter, ventral surface. Scale bars: 1.0 mm." figureDoi="10.3897/zookeys.1188.110081.figures1-8" httpUri="https://binary.pensoft.net/fig/963583" pageId="0" pageNumber="125">1</figureCitation>
,
<figureCitation id="3804106BE1FF6C49A963492B07768EA9" captionStart="Figures 18" captionStartId="F1" captionText="Figures 1 - 8. Cybistrinae morphological features 1 Bifurcitus lherminieri, right metatibia, posterior surface 2 Bifurcitus lherminieri, left metaleg, anterior surface 3 Trifurcitus robustus, left metaleg, anterior surface 4 Metaxydytes fraternus, left mesotarsomeres III-V, posterior surfaces 5 Cybister tripunctatus, left mesotarsomeres III-V, posterior surfaces 6 Megadytes latus, left half of metaventrite and left metacoxa (WV = metaventrite wing width, WC = metacoxal width) 7 Cybister tripunctatus, right metathoracic wing 8 Trifurcitus robustus, right metatrochanter, ventral surface. Scale bars: 1.0 mm." figureDoi="10.3897/zookeys.1188.110081.figures1-8" httpUri="https://binary.pensoft.net/fig/963583" pageId="0" pageNumber="125">2</figureCitation>
). These are the largest of all diving beetles with adult specimens 36-47 mm in total length (
<bibRefCitation id="AEF4E90A0321DF371239E6EAB04617E9" DOI="https://doi.org/10.11646/zootaxa.4586.3.8" author="Hendrich, L" journalOrPublisher="Zootaxa" pageId="0" pageNumber="125" pagination="517 - 535" refId="B16" refString="Hendrich, L, Manuel, M, Balke, M, 2019. The return of the Duke-locality data for Megadytes ducalis Sharp, 1882, the world's largest diving beetle, with notes on related species (Coleoptera: Dytiscidae). Zootaxa 4586 (3): 517 - 535, DOI: https://doi.org/10.11646/zootaxa.4586.3.8" title="The return of the Duke-locality data for Megadytes ducalis Sharp, 1882, the world's largest diving beetle, with notes on related species (Coleoptera: Dytiscidae)." url="https://doi.org/10.11646/zootaxa.4586.3.8" volume="4586" year="2019">Hendrich et al. 2019</bibRefCitation>
). Third instar larvae have (1) the median lobe of the frontoclypeus truncate apically with a tuft of setae (Fig.
<figureCitation id="0673FD46AC4F20D073B388B6314A635A" captionStart="Figures 6771" captionStartId="F8" captionText="Figures 67 - 71. Third instar larva, anterior clypeal margin 67 Bifurcitus magnus 68 Megadytes latus 69 Metaxydytes carcharias 70 Paramegadytes glaucus 71 Trifurcitus fallax." figureDoi="10.3897/zookeys.1188.110081.figures67-71" httpUri="https://binary.pensoft.net/fig/963590" pageId="0" pageNumber="125">67</figureCitation>
), (2) the median and lateral lobes of the frontoclypeus separated by a wide emargination (Fig.
<figureCitation id="B774B0BF24A233BADAAA052B1F27D2AB" captionStart="Figures 6771" captionStartId="F8" captionText="Figures 67 - 71. Third instar larva, anterior clypeal margin 67 Bifurcitus magnus 68 Megadytes latus 69 Metaxydytes carcharias 70 Paramegadytes glaucus 71 Trifurcitus fallax." figureDoi="10.3897/zookeys.1188.110081.figures67-71" httpUri="https://binary.pensoft.net/fig/963590" pageId="0" pageNumber="125">67</figureCitation>
), (3) the lateral lobes of the frontoclypeus apically simple (Fig.
<figureCitation id="E49E10618DDC17831F4D2B2ED212C79C" captionStart="Figures 6771" captionStartId="F8" captionText="Figures 67 - 71. Third instar larva, anterior clypeal margin 67 Bifurcitus magnus 68 Megadytes latus 69 Metaxydytes carcharias 70 Paramegadytes glaucus 71 Trifurcitus fallax." figureDoi="10.3897/zookeys.1188.110081.figures67-71" httpUri="https://binary.pensoft.net/fig/963590" pageId="0" pageNumber="125">67</figureCitation>
), (4) the lateral lobes of the frontoclypeus obtusely angulate (Fig.
<figureCitation id="93D1A08F03647FFA1DC73E4F73A457CD" captionStart="Figures 6771" captionStartId="F8" captionText="Figures 67 - 71. Third instar larva, anterior clypeal margin 67 Bifurcitus magnus 68 Megadytes latus 69 Metaxydytes carcharias 70 Paramegadytes glaucus 71 Trifurcitus fallax." figureDoi="10.3897/zookeys.1188.110081.figures67-71" httpUri="https://binary.pensoft.net/fig/963590" pageId="0" pageNumber="125">67</figureCitation>
), and (5) the cephalic capsule relatively short (head length / head width &lt;1.20).
</paragraph>
</subSubSection>
<subSubSection id="79FF3270A375B178114B4AFE0B2E2B2F" pageId="0" pageNumber="125" type="phylogenetic relationships">
<paragraph id="C203A6F00E181B54046CC8BDFB2946F0" pageId="0" pageNumber="125">Phylogenetic relationships.</paragraph>
<paragraph id="CD99FF5F05B384DA03D2CDC246C935D2" pageId="0" pageNumber="125">
<taxonomicName id="B4FCD7BC50A184312B087348C95615F9" authorityName="Miller &amp; Michat &amp; Ferreira Jr" authorityYear="2024" class="Insecta" family="Dytiscidae" genus="Bifurcitus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Bifurcitus" order="Coleoptera" pageId="0" pageNumber="125" phylum="Arthropoda" rank="genus">
<emphasis id="0D967CACC545047E82C14CD222DA564D" italics="true" pageId="0" pageNumber="125">Bifurcitus</emphasis>
</taxonomicName>
is sister group to the similar
<taxonomicName id="055BE36A1F1250411E83562968B81520" authorityName="Brinck" authorityYear="1945" class="Insecta" family="Dytiscidae" genus="Trifurcitus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Trifurcitus" order="Coleoptera" pageId="0" pageNumber="125" phylum="Arthropoda" rank="genus">
<emphasis id="4368596EA229CF27B865ED2EE2141C2E" italics="true" pageId="0" pageNumber="125">Trifurcitus</emphasis>
</taxonomicName>
(Figs
<figureCitation id="1152E0B36C73F0672C40C151FD4A2CA9" captionStart="Figure 75" captionStartId="F12" captionText="Figure 75. One of seven cladograms derived from parsimony analysis of 57 morphological characters from Cybistrinae and other Dytiscidae (len = 105, CI = 68, RI = 92) with characters mapped using ' fast' optimization in WinClada. Black hash marks indicate unambiguous changes, white hash marks indicate homoplasious changes or reversals. Numbers above hash marks are character numbers, those below hash marks are derived state numbers." figureDoi="10.3897/zookeys.1188.110081.figure75" httpUri="https://binary.pensoft.net/fig/963593" pageId="0" pageNumber="125">75</figureCitation>
,
<figureCitation id="3A9BD2E397D396D8AEE04F204D4B2CA4" captionStart="Figure 76" captionStartId="F13" captionText="Figure 76. Consensus cladogram of seven equally parsimonious cladograms from parsimony analysis of 57 morphological characters from Cybistrinae and other Dytiscidae morphology." figureDoi="10.3897/zookeys.1188.110081.figure76" httpUri="https://binary.pensoft.net/fig/963594" pageId="0" pageNumber="125">76</figureCitation>
;
<bibRefCitation id="334E01EA17F47006F0CAC4A984272169" DOI="https://doi.org/10.1111/j.1463-6409.2006.00254.x" author="Miller, KB" journalOrPublisher="Zoologica Scripta" pageId="0" pageNumber="125" pagination="41 - 59" refId="B29" refString="Miller, KB, Bergsten, J, Whiting, MF, 2007. Phylogeny and classification of diving beetles in the tribe Cybistrini (Coleoptera, Dytiscidae, Dytiscinae). Zoologica Scripta 36 (1): 41 - 59, DOI: https://doi.org/10.1111/j.1463-6409.2006.00254.x" title="Phylogeny and classification of diving beetles in the tribe Cybistrini (Coleoptera, Dytiscidae, Dytiscinae)." url="https://doi.org/10.1111/j.1463-6409.2006.00254.x" volume="36" year="2007">Miller et al. 2007</bibRefCitation>
). Both have males and females with equal-length metatarsal claws and the anterior metatibial spur either bifid or trifid (although these two conditions may not be homologous) (Figs
<figureCitation id="FC8CA264A58FF394890B63D24F188ECC" captionStart="Figures 18" captionStartId="F1" captionText="Figures 1 - 8. Cybistrinae morphological features 1 Bifurcitus lherminieri, right metatibia, posterior surface 2 Bifurcitus lherminieri, left metaleg, anterior surface 3 Trifurcitus robustus, left metaleg, anterior surface 4 Metaxydytes fraternus, left mesotarsomeres III-V, posterior surfaces 5 Cybister tripunctatus, left mesotarsomeres III-V, posterior surfaces 6 Megadytes latus, left half of metaventrite and left metacoxa (WV = metaventrite wing width, WC = metacoxal width) 7 Cybister tripunctatus, right metathoracic wing 8 Trifurcitus robustus, right metatrochanter, ventral surface. Scale bars: 1.0 mm." figureDoi="10.3897/zookeys.1188.110081.figures1-8" httpUri="https://binary.pensoft.net/fig/963583" pageId="0" pageNumber="125">1-3</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="4CCE689ED240DBA0E5B8AFFAC89795C5" pageId="0" pageNumber="125" type="discussion">
<paragraph id="871DDF79D9486F99D6E1616A0FE0E685" pageId="0" pageNumber="125">Discussion.</paragraph>
<paragraph id="C0335CC13A4552BF135D253E2D9CB142" pageId="0" pageNumber="125">
Although previous evidence suggested that the several subgenera of &quot;
<taxonomicName id="FC841C1117B83211151C83C405F08D24" authorityName="Sharp" authorityYear="1882" class="Insecta" family="Dytiscidae" genus="Megadytes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Megadytes" order="Coleoptera" pageId="0" pageNumber="125" phylum="Arthropoda" rank="genus">
<emphasis id="D006BC3C95435040DC294EA4A01F844A" italics="true" pageId="0" pageNumber="125">Megadytes</emphasis>
</taxonomicName>
&quot; are monophyletic, monophyly of this group is not supported here based in part on the discovery of undescribed cybistrine species with unique combinations of character states (Figs
<figureCitation id="CFE3955476DA3F7A2AADA8681D844E46" captionStart="Figure 75" captionStartId="F12" captionText="Figure 75. One of seven cladograms derived from parsimony analysis of 57 morphological characters from Cybistrinae and other Dytiscidae (len = 105, CI = 68, RI = 92) with characters mapped using ' fast' optimization in WinClada. Black hash marks indicate unambiguous changes, white hash marks indicate homoplasious changes or reversals. Numbers above hash marks are character numbers, those below hash marks are derived state numbers." figureDoi="10.3897/zookeys.1188.110081.figure75" httpUri="https://binary.pensoft.net/fig/963593" pageId="0" pageNumber="125">75</figureCitation>
,
<figureCitation id="214507609104245075DF8C0D6CA7A23B" captionStart="Figure 76" captionStartId="F13" captionText="Figure 76. Consensus cladogram of seven equally parsimonious cladograms from parsimony analysis of 57 morphological characters from Cybistrinae and other Dytiscidae morphology." figureDoi="10.3897/zookeys.1188.110081.figure76" httpUri="https://binary.pensoft.net/fig/963594" pageId="0" pageNumber="125">76</figureCitation>
). Given the situation, it seems appropriate to recognize these subgenera at the genus rank including
<taxonomicName id="2F19FB84A36440246E2412543D79875B" authorityName="Miller &amp; Michat &amp; Ferreira Jr" authorityYear="2024" class="Insecta" family="Dytiscidae" genus="Bifurcitus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Bifurcitus" order="Coleoptera" pageId="0" pageNumber="125" phylum="Arthropoda" rank="genus">
<emphasis id="719BAC687E813D1A573D6FE6DD7EA125" italics="true" pageId="0" pageNumber="125">Bifurcitus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="87EDF5E9CC0DC8A0E1DA12729022F02A" pageId="0" pageNumber="125">
There are three currently valid species in
<taxonomicName id="A28C9552E7E1906FE7FCEFCC568EB659" authorityName="Miller &amp; Michat &amp; Ferreira Jr" authorityYear="2024" class="Insecta" family="Dytiscidae" genus="Bifurcitus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Bifurcitus" order="Coleoptera" pageId="0" pageNumber="125" phylum="Arthropoda" rank="genus">
<emphasis id="14F48B33CB9EA974376A5E01A6F0D3D6" italics="true" pageId="0" pageNumber="125">Bifurcitus</emphasis>
</taxonomicName>
which were differentiated and characterized recently by
<bibRefCitation id="4147EF8F50B8157DAD48FF45F0257F65" DOI="https://doi.org/10.11646/zootaxa.4586.3.8" author="Hendrich, L" journalOrPublisher="Zootaxa" pageId="0" pageNumber="125" pagination="517 - 535" refId="B16" refString="Hendrich, L, Manuel, M, Balke, M, 2019. The return of the Duke-locality data for Megadytes ducalis Sharp, 1882, the world's largest diving beetle, with notes on related species (Coleoptera: Dytiscidae). Zootaxa 4586 (3): 517 - 535, DOI: https://doi.org/10.11646/zootaxa.4586.3.8" title="The return of the Duke-locality data for Megadytes ducalis Sharp, 1882, the world's largest diving beetle, with notes on related species (Coleoptera: Dytiscidae)." url="https://doi.org/10.11646/zootaxa.4586.3.8" volume="4586" year="2019">Hendrich et al. (2019)</bibRefCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>