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<mods:title id="7949B379F6C8AF084E3DD00CEB55CE75">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="8B1CA6DF6EFC947027C50751A3C4F9C1">Department of Biology and Museum of Southwestern Biology, University of New Mexico, Albuquerque, NM 87131 - 0001, USA</mods:affiliation>
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<mods:namePart id="B4F63F596829A2B5479A80B95ED39457">Michat, Mariano C.</mods:namePart>
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<mods:affiliation id="EFD1B2BD352F48ECBC97D957005A9FDD">Instituto de Biodiversidad y Biologia Experimental y Aplicada, CONICET-Universidad de Buenos Aires, Buenos Aires, Argentina</mods:affiliation>
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<mods:affiliation id="F95A8BC0F27CE9CA88741A0276173F91">Laboratorio de Entomologia, Departamento de Zoologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro, Rio de Janeiro, RJ, Brasi</mods:affiliation>
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<paragraph id="9BAF78237F00C92ABAB91878DEC0B1EC" pageId="0" pageNumber="125">
<taxonomicName id="444961D31B59A02EB9189C5796283903" LSID="EB5DE962-DDCF-57FC-88E9-060CCBFC845D" authority="Tremouilles &amp; Bachmann, 1980" authorityName="Tremouilles &amp; Bachmann" authorityYear="1980" class="Insecta" family="Dytiscidae" genus="Paramegadytes" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Paramegadytes" order="Coleoptera" pageId="0" pageNumber="125" phylum="Arthropoda" rank="genus" status="stat. nov.">
Paramegadytes
<normalizedToken id="D05B44261A3936C3E5C5FBA3DAFBA571" originalValue="Trémouilles">Tremouilles</normalizedToken>
&amp; Bachmann, 1980
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<taxonomicNameLabel id="310A57CEBC3F6B371FD31FB01CD65453" pageId="0" pageNumber="125">stat. nov.</taxonomicNameLabel>
</paragraph>
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<subSubSection id="E45AF83264B6350D36489937465352A2" pageId="0" pageNumber="125" type="description">
<paragraph id="2ECE8BE3DE40B0D732B1D24E8DFFDB7B" pageId="0" pageNumber="125">
<figureCitation id="3899BB645F94CF77AE28BD13AB5EBC2F" captionStart="Figures 1529" captionStartId="F3" captionText="Figures 15 - 29. Neotropical Cybistrinae species, metatarsal claws and tarsomere VI of males and females 15, 16 Cybister festae 17, 18 Cybister puncticollis 19, 20 Nilssondytes diversus 21, 22 Megadytes latus 23, 24 Megadytes parvus 25 species &quot; IR 57 &quot; (Ribera et al. 2008) 26, 27 Paramegadytes glaucus 28, 29 Metaxydytes fraternus. Scale bars: 1.0 mm." figureDoi="10.3897/zookeys.1188.110081.figures15-29" httpUri="https://binary.pensoft.net/fig/963585" pageId="0" pageNumber="125">Figs 26</figureCitation>
<figureCitation id="92EBF1274C55DAFF1F0EFDBF80FFC707" captionStart="Figures 1529" captionStartId="F3" captionText="Figures 15 - 29. Neotropical Cybistrinae species, metatarsal claws and tarsomere VI of males and females 15, 16 Cybister festae 17, 18 Cybister puncticollis 19, 20 Nilssondytes diversus 21, 22 Megadytes latus 23, 24 Megadytes parvus 25 species &quot; IR 57 &quot; (Ribera et al. 2008) 26, 27 Paramegadytes glaucus 28, 29 Metaxydytes fraternus. Scale bars: 1.0 mm." figureDoi="10.3897/zookeys.1188.110081.figures15-29" httpUri="https://binary.pensoft.net/fig/963585" pageId="0" pageNumber="125">, 27</figureCitation>
<figureCitation id="6281C811E46CF4AB67FF3439D71B2F2F" captionStart="Figures 5157" captionStartId="F5" captionText="Figures 51 - 57. Neotropical Cybistrinae species, male sternite IX, ventral aspect 51 Cybister festae 52 Cybister puncticollis 53 Nilssondytes diversus 54 Megadytes latus 55 Megadytes parvus 56 Paramegadytes glaucus 57 Metaxydytes fraternus. Scale bars: 1.0 mm." figureDoi="10.3897/zookeys.1188.110081.figures51-57" httpUri="https://binary.pensoft.net/fig/963587" pageId="0" pageNumber="125">, 56</figureCitation>
<figureCitation id="F478CCF12288A712F5415D13FAB31AB0" 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">, 63</figureCitation>
<figureCitation id="DAD8DCD44C681B03F1A15C0A713A581C" 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">, 70</figureCitation>
</paragraph>
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<paragraph id="44E4431FAFE1AB01783A14B928B27FA7" pageId="0" pageNumber="125">
<taxonomicName id="CB7A29A55478AF266F0EF792F9F43F68" authorityName="Miller &amp; Michat &amp; Ferreira Jr" authorityYear="2024" class="Insecta" family="Dytiscidae" genus="Paramegadytes" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Paramegadytes" order="Coleoptera" pageId="0" pageNumber="125" phylum="Arthropoda" rank="genus" status="stat. nov.">Paramegadytes</taxonomicName>
<normalizedToken id="D9661D0A539BF6E7A6A543025808CE05" originalValue="Trémouilles">Tremouilles</normalizedToken>
&amp; Bachmann, 1980: 101.
</paragraph>
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<subSubSection id="C0304D61690FF74D95DD5B3A5C4FB3A4" pageId="0" pageNumber="125" type="type species">
<paragraph id="A11D650C387FE492F6D7990241A906F2" pageId="0" pageNumber="125">Type species.</paragraph>
<paragraph id="AD353CF00A8D2786203122397CA1E6EA" pageId="0" pageNumber="125">
<taxonomicName id="832279BDCE2BC5A3DA6750C7E39AABC6" baseAuthorityName="Brullé" baseAuthorityYear="1837" class="Insecta" family="Dytiscidae" genus="Dyticus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Dyticus glaucus" order="Coleoptera" pageId="0" pageNumber="125" phylum="Arthropoda" rank="species" species="glaucus">
<emphasis id="75BCFE0A3A77F9CDBA95F5A68CCCDEA0" italics="true" pageId="0" pageNumber="125">Dyticus glaucus</emphasis>
</taxonomicName>
<normalizedToken id="94196FDA4F4614F11D7FBC57F62D5DDB" originalValue="Brullé">Brulle</normalizedToken>
, 1837: 46 by original designation.
</paragraph>
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<subSubSection id="FFDDA14CDE46A2D9599AC9083C472283" pageId="0" pageNumber="125" type="diagnosis">
<paragraph id="BA9A5D0272F1269C519A08E6CE3765C9" pageId="0" pageNumber="125">Diagnosis.</paragraph>
<paragraph id="E0C3D7BFDABD071F1D6F786EB7B6CECD" pageId="0" pageNumber="125">
Like
<taxonomicName id="8D0B0F28387E0FAE706E64A9DACF8952" authorityName="Miller &amp; Michat &amp; Ferreira Jr" authorityYear="2024" family="Dytiscidae" genus="Metaxydytes" kingdom="Animalia" lsidName="Metaxydytes" order="Coleoptera" pageId="0" pageNumber="125" rank="genus">
<emphasis id="76EB4A23AAC84C50814B86B3DB15C5E5" italics="true" pageId="0" pageNumber="125">Metaxydytes</emphasis>
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these species have both metatibial spurs apically simple, the medial margins of male abdominal sternite IX straight, and both males and females with two metatarsal claws, males with equal-length claws and females with the posterior claw reduced (Figs
<figureCitation id="6457646BCAE02A9FEDA55818E8C13D25" captionStart="Figures 1529" captionStartId="F3" captionText="Figures 15 - 29. Neotropical Cybistrinae species, metatarsal claws and tarsomere VI of males and females 15, 16 Cybister festae 17, 18 Cybister puncticollis 19, 20 Nilssondytes diversus 21, 22 Megadytes latus 23, 24 Megadytes parvus 25 species &quot; IR 57 &quot; (Ribera et al. 2008) 26, 27 Paramegadytes glaucus 28, 29 Metaxydytes fraternus. Scale bars: 1.0 mm." figureDoi="10.3897/zookeys.1188.110081.figures15-29" httpUri="https://binary.pensoft.net/fig/963585" pageId="0" pageNumber="125">26</figureCitation>
,
<figureCitation id="585D0F34EDFB5AA3167C1CA0C77021DC" captionStart="Figures 1529" captionStartId="F3" captionText="Figures 15 - 29. Neotropical Cybistrinae species, metatarsal claws and tarsomere VI of males and females 15, 16 Cybister festae 17, 18 Cybister puncticollis 19, 20 Nilssondytes diversus 21, 22 Megadytes latus 23, 24 Megadytes parvus 25 species &quot; IR 57 &quot; (Ribera et al. 2008) 26, 27 Paramegadytes glaucus 28, 29 Metaxydytes fraternus. Scale bars: 1.0 mm." figureDoi="10.3897/zookeys.1188.110081.figures15-29" httpUri="https://binary.pensoft.net/fig/963585" pageId="0" pageNumber="125">27</figureCitation>
). Females also share the characteristic of the medial margins of the gonocoxosternite with a series of spinous setae (Fig.
<figureCitation id="9BD3C2457A42FE691E101C8CFCB5E817" 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">63</figureCitation>
). The lateral pronotal margin has a diffuse, but distinctive lateral pale band. From
<taxonomicName id="9C9251F60D5475B51B76960D4F7B8E70" authorityName="Miller &amp; Michat &amp; Ferreira Jr" authorityYear="2024" family="Dytiscidae" genus="Metaxydytes" kingdom="Animalia" lsidName="Metaxydytes" order="Coleoptera" pageId="0" pageNumber="125" rank="genus">
<emphasis id="2528D1D4CE83F1113C2ACCDFC68966F2" italics="true" pageId="0" pageNumber="125">Metaxydytes</emphasis>
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these specimens are larger with the metaventrite wings relatively broader (see above under
<taxonomicName id="7DE1968545D0A871069D48B6B4734E1E" authorityName="Miller &amp; Michat &amp; Ferreira Jr" authorityYear="2024" family="Dytiscidae" genus="Metaxydytes" kingdom="Animalia" lsidName="Metaxydytes" order="Coleoptera" pageId="0" pageNumber="125" rank="genus">
<emphasis id="0C23B542232FA6A4DBA768427631CD59" italics="true" pageId="0" pageNumber="125">Metaxydytes</emphasis>
</taxonomicName>
for details of diagnostic comparisons). Third instar larvae have (1) the median lobe of the frontoclypeus truncate apically with a tuft of setae (Fig.
<figureCitation id="73EC18D414013E43E28D4BE47F4915EC" 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">70</figureCitation>
), (2) the median and lateral lobes of the frontoclypeus separated by a wide emargination (Fig.
<figureCitation id="9E90DDC5C7693B1C6DEDB5AA309B3361" 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">70</figureCitation>
), and (3) the lateral lobes of the frontoclypeus bilobed (Fig.
<figureCitation id="F86ECC1AD96EEB0DB02BC20DB7FE0D4A" 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">70</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="E9C5D1DC775AEA1BA65F5EA3025CC8E4" pageId="0" pageNumber="125" type="phylogenetics">
<paragraph id="D86440D648F94A4C1A78C1BEF6DD9742" pageId="0" pageNumber="125">Phylogenetics.</paragraph>
<paragraph id="3C933CA046B388602F8C20A5809B1F86" pageId="0" pageNumber="125">
This may be the sister genus to
<taxonomicName id="5402351FF67271703FA30E35BB9FD5CF" authorityName="Miller &amp; Michat &amp; Ferreira Jr" authorityYear="2024" family="Dytiscidae" genus="Metaxydytes" kingdom="Animalia" lsidName="Metaxydytes" order="Coleoptera" pageId="0" pageNumber="125" rank="genus">
<emphasis id="928BB398AC8DC106F93670427EEBA92C" italics="true" pageId="0" pageNumber="125">Metaxydytes</emphasis>
</taxonomicName>
(Figs
<figureCitation id="89D002E3F47A307634808FD37D5AAE7A" 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="A2D715D2A4BA826C5D360767B671DCEB" 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="5FF68541E70BF3BA159F3F6A532CC5C7" 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>
) although here it is in an unresolved position relative to
<taxonomicName id="3E564BFB30ED0388C67CB64CD8B40D8F" authorityName="Miller &amp; Michat &amp; Ferreira Jr" authorityYear="2024" family="Dytiscidae" genus="Nilssondytes" kingdom="Animalia" lsidName="Nilssondytes" order="Coleoptera" pageId="0" pageNumber="125" rank="genus">
<emphasis id="3116B627EFF7104C9285B828D23C81F4" italics="true" pageId="0" pageNumber="125">Nilssondytes</emphasis>
</taxonomicName>
,
<taxonomicName id="25B4A787D0FD284442E7EF3A99A31EFE" authorityName="Miller &amp; Michat &amp; Ferreira Jr" authorityYear="2024" family="Dytiscidae" genus="Metaxydytes" kingdom="Animalia" lsidName="Metaxydytes" order="Coleoptera" pageId="0" pageNumber="125" rank="genus">
<emphasis id="338C5E36276A0186CA541ACA8C9B24FC" italics="true" pageId="0" pageNumber="125">Metaxydytes</emphasis>
</taxonomicName>
and
<taxonomicName id="415F6CD4540E6CE1CEEB3D5034671A8F" authorityName="Miller &amp; Michat &amp; Ferreira Jr" authorityYear="2024" class="Insecta" family="Dytiscidae" genus="Megadytes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Megadytes" order="Coleoptera" pageId="0" pageNumber="125" phylum="Arthropoda" rank="genus">
<emphasis id="68E59DF0EF110BC56C1EAF30D524291F" italics="true" pageId="0" pageNumber="125">Megadytes</emphasis>
</taxonomicName>
+
<taxonomicName id="B9DC43B3E0F5E7A92E08E74A8910E5FD" authorityName="Curtis" authorityYear="1827" class="Insecta" family="Dytiscidae" genus="Cybister" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cybister" order="Coleoptera" pageId="0" pageNumber="125" phylum="Arthropoda" rank="genus">
<emphasis id="C017D7BCC3B530F21E284134105E9751" italics="true" pageId="0" pageNumber="125">Cybister</emphasis>
</taxonomicName>
(Figs
<figureCitation id="F11E31555BDAE9A494D7070D7A5B53DD" 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="8DD795F0A9C6C649C1399FB7FDAC296A" 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>
). See under
<taxonomicName id="E3153F6621B12FFEF27DCC3B2E7773C9" authorityName="Miller &amp; Michat &amp; Ferreira Jr" authorityYear="2024" family="Dytiscidae" genus="Metaxydytes" kingdom="Animalia" lsidName="Metaxydytes" order="Coleoptera" pageId="0" pageNumber="125" rank="genus">
<emphasis id="057033A35EFD0FABE32FEE36C4FAF59F" italics="true" pageId="0" pageNumber="125">Metaxydytes</emphasis>
</taxonomicName>
for further discussion.
</paragraph>
</subSubSection>
<subSubSection id="F76E553F871E78EB13D5F021A2F8C811" pageId="0" pageNumber="125" type="discussion">
<paragraph id="7A4293E99A43EA2A25EBFC3622179782" pageId="0" pageNumber="125">Discussion.</paragraph>
<paragraph id="0F666DD927F4DEE15E8691CAAB489584" pageId="0" pageNumber="125">
There are currently two valid species in this genus,
<taxonomicName id="DFDF0B6466B628A9E0F64CCA058EF61B" lsidName="P. australis" pageId="0" pageNumber="125" rank="species" species="australis">
<emphasis id="E50D40DF7C016C4CF2AAA34C1AA14054" italics="true" pageId="0" pageNumber="125">P. australis</emphasis>
</taxonomicName>
(Germain) and
<taxonomicName id="D9222E103BB3587160976FDD6B9D8FBC" lsidName="P. glaucus" pageId="0" pageNumber="125" rank="species" species="glaucus">
<emphasis id="4BC00E5B0C99C814DA677577428C5371" italics="true" pageId="0" pageNumber="125">P. glaucus</emphasis>
</taxonomicName>
(
<normalizedToken id="91D8771D9263359C89BEC07B51DA0934" originalValue="Brullé">Brulle</normalizedToken>
).
<bibRefCitation id="E98B1AC94A6D0428CBFCE1707C82F161" author="Tremouilles, ER" journalOrPublisher="Revista de la Sociedad Entomologica Argentina" pageId="0" pageNumber="125" pagination="101 - 125" refId="B40" refString="Tremouilles, ER, Bachmann, AO, 1980. La tribu Cybisterini en la Argentina (Coleoptera, Dytiscidae). Revista de la Sociedad Entomologica Argentina 39: 101 - 125" title="La tribu Cybisterini en la Argentina (Coleoptera, Dytiscidae)." volume="39" year="1980">
<normalizedToken id="35CE5E27D5732F43D40BC74EFE0FE93D" originalValue="Trémouilles">Tremouilles</normalizedToken>
and Bachmann (1980)
</bibRefCitation>
characterized and differentiated them.
</paragraph>
</subSubSection>
</treatment>
</document>