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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.869.33997" ID-GBIF-Dataset="5b500434-09e3-4370-82a1-1eb789d9d58a" ID-PMC="PMC6690892" ID-Pensoft-Pub="1313-2970-869-19" ID-Pensoft-UUID="C44C602AB0235FB2922DC2A4915C3288" ID-PubMed="31413659" ID-ZooBank="B7C88A64C06E4B67A352F2F9C8FB0D1C" ModsDocID="1313-2970-869-19" checkinTime="1565056339550" checkinUser="pensoft" docAuthor="Ranarilalatiana, Tolotra, Raveloson Ravaomanarivo, Lala Harivelo &amp; Bergsten, Johannes" docDate="2019" docId="585E67A8A2805A19B8996B9FBA19B8AA" docLanguage="en" docName="ZooKeys 869: 19-90" docOrigin="ZooKeys 869" docSource="http://dx.doi.org/10.3897/zookeys.869.33997" docTitle="Copelatus vokoka Ranarilalatiana &amp; Bergsten, sp. nov." docType="treatment" docUuid="8FF486E9-5A55-4581-9CE8-5990C65FAF6D" docUuidSource="ZooBank" docVersion="5" lastPageNumber="19" masterDocId="5E52FF84FFE5FF9193131A454057613C" masterDocTitle="Taxonomic revision of the genus Copelatus of Madagascar (Coleoptera, Dytiscidae, Copelatinae): the non- erichsonii group species" masterLastPageNumber="90" masterPageNumber="19" pageNumber="19" updateTime="1668167671827" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>Taxonomic revision of the genus Copelatus of Madagascar (Coleoptera, Dytiscidae, Copelatinae): the non- erichsonii group species</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ranarilalatiana, Tolotra</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Raveloson Ravaomanarivo, Lala Harivelo</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bergsten, Johannes</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
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<mods:part>
<mods:date>2019</mods:date>
<mods:detail type="volume">
<mods:number>869</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>19</mods:start>
<mods:end>90</mods:end>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.869.33997</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.869.33997</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-869-19</mods:identifier>
<mods:identifier type="ZooBank">B7C88A64C06E4B67A352F2F9C8FB0D1C</mods:identifier>
<mods:identifier type="Pensoft-UUID">C44C602AB0235FB2922DC2A4915C3288</mods:identifier>
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<treatment ID-GBIF-Taxon="158995551" LSID="urn:lsid:zoobank.org:act:8FF486E9-5A55-4581-9CE8-5990C65FAF6D" httpUri="http://treatment.plazi.org/id/585E67A8A2805A19B8996B9FBA19B8AA" lastPageNumber="19" pageId="0" pageNumber="19">
<subSubSection pageId="0" pageNumber="19" type="nomenclature">
<paragraph pageId="0" pageNumber="19">
<taxonomicName LSID="585e67a8-a280-5a19-b899-6b9fba19b8aa" authority="Ranarilalatiana &amp; Bergsten" class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus vokoka" order="Coleoptera" pageId="0" pageNumber="19" phylum="Arthropoda" rank="species" species="vokoka">Copelatus vokoka Ranarilalatiana &amp; Bergsten</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="19">sp. nov.</taxonomicNameLabel>
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Male genitalia. From left to right, aedeagus in right lateral, ventral, left lateral views and left paramere. A Copelatus insuetus B Copelatus kely sp. nov. C Copelatus vokoka sp. nov. D Copelatus ankaratra sp. nov. E Copelatus safiotra sp. nov." figureDoi="10.3897/zookeys.869.33997.figure6" httpUri="https://binary.pensoft.net/fig/322931" pageId="0" pageNumber="19">Figs 6C</figureCitation>
,
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Habitus, dorsal view. A Female: Copelatus insuetus B Male: Copelatus vokoka sp. nov. C Female: Copelatus kely sp. nov. D Female: Copelatus sp. 3 (Ivohibe)." figureDoi="10.3897/zookeys.869.33997.figure9" httpUri="https://binary.pensoft.net/fig/322934" pageId="0" pageNumber="19">9B</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="19" type="type locality">
<paragraph pageId="0" pageNumber="19">Type locality.</paragraph>
<paragraph pageId="0" pageNumber="19">Ivohibe Special Reserve [22.456683S, 46.956283E] [Madagascar, Ihorombe region, Ivohibe district]</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="19" type="type material">
<paragraph pageId="0" pageNumber="19">Type material.</paragraph>
<paragraph pageId="0" pageNumber="19">
<emphasis bold="true" pageId="0" pageNumber="19">Fianarantsoa. Ihorombe: Ivohibe</emphasis>
: -HT♂ (GP) (NHRS): // NHRS-JLKB | 000010849 // Madagascar: Fianarantsoa: Ihorombe: R.S. | Pic
<normalizedToken originalValue="dIvohibe">d'Ivohibe</normalizedToken>
Corridor: at the confluence of | two rivers Inganga and Anefitany: | S22.456683; E046.956283; 874 m; 09.XII.2013 | GB Nets &amp; sieves: forest pools with dead | leaves, Leg. J.H. Randriamihaja &amp; | T. Ranarialalatiana: Field# MAD13-55 // Holotype |
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus vokoka</emphasis>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 //
<emphasis bold="true" pageId="0" pageNumber="19">Paratypes</emphasis>
: -6♂ (GP), 2♀, 16 ex. (6♂, 10♀) (Alc.) (NHRS, NHMUK, DEUA &amp; PBZT/MBC): // NHRS-JLKB | 000010850-4, 10857, 65735-6, 10855(Alc.) // Madagascar: Fianarantsoa: Ihorombe: R.S. | Pic
<normalizedToken originalValue="dIvohibe">d'Ivohibe</normalizedToken>
Corridor: at the confluence of | two rivers Inganga and Anefitany: | S22.456683; E046.956283; 874 m; 09.XII.2013 | GB Nets &amp; sieves: forest pools with dead | leaves, Leg. J.H. Randriamihaja &amp; | T. Ranarialalatiana: Field# MAD13-55 // Paratype |
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus vokoka</emphasis>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 // -1♂ (GP), 1♀, 3♀ (Alc.) (NHRS): // NHRS-JLKB | 000010512-3, 10782(Alc.) // Madagascar: Fianarantsoa: Ihorombe: | R.S. Pic
<normalizedToken originalValue="dIvohibe">d'Ivohibe</normalizedToken>
Corridor: The | confluence of rivers Inganga and | Anefitany: S22.457283; E046.95535; 870 m, | 09.XII.2013, GB Nets &amp; sieves: big | muddy pool, Leg. J.H. Randriamihaja &amp; | T. Ranarialalatiana: Field# MAD13-57 // Paratype |
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus vokoka</emphasis>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 //
<emphasis bold="true" pageId="0" pageNumber="19">Fianarantsoa. Vatovavy Fitovinany: Ifanadiana</emphasis>
: -1♂ (GP), 4♂, 11♀ (coll.
<normalizedToken originalValue="Michaël">Michael</normalizedToken>
Manuel, Paris, NHRS): // Madagascar. Ex-prov. Fiana- | rantsoa. ca. 3.3 km WSW | Ranomafana. 28 XII 2017. | Ramahandrison &amp; Manuel leg. //
<geoCoordinate degrees="21" direction="south" minutes="16" orientation="latitude" precision="15" seconds="05" value="-21.268055">21°16'05&quot;S</geoCoordinate>
,
<geoCoordinate degrees="47" direction="east" minutes="25" orientation="longitude" precision="15" seconds="28" value="47.424446">47°25'28&quot;E</geoCoordinate>
Alt. 993 m. | Shallow shaded pond with large | quantity of dead tree leaves and | some Poaceae, in forest. | Ranomafana NP. // Coll. | M. Manuel | Paris // Paratype |
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus vokoka</emphasis>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 //
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="19" type="diagnosis">
<paragraph pageId="0" pageNumber="19">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="19">
Body shape slightly shorter, less elongate and slightly more oval than
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
, and eyes smaller. Elytral striae more deeply impressed and intervals therefore slightly more convex (
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Habitus, dorsal view. A Female: Copelatus insuetus B Male: Copelatus vokoka sp. nov. C Female: Copelatus kely sp. nov. D Female: Copelatus sp. 3 (Ivohibe)." figureDoi="10.3897/zookeys.869.33997.figure9" httpUri="https://binary.pensoft.net/fig/322934" pageId="0" pageNumber="19">Fig. 9B</figureCitation>
). Penis shape in lateral view gently curved from base to apex (
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Male genitalia. From left to right, aedeagus in right lateral, ventral, left lateral views and left paramere. A Copelatus insuetus B Copelatus kely sp. nov. C Copelatus vokoka sp. nov. D Copelatus ankaratra sp. nov. E Copelatus safiotra sp. nov." figureDoi="10.3897/zookeys.869.33997.figure6" httpUri="https://binary.pensoft.net/fig/322931" pageId="0" pageNumber="19">Fig. 6C</figureCitation>
), lacking the defined
<normalizedToken originalValue="“shoulder”">&quot;shoulder&quot;</normalizedToken>
of
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
and
<taxonomicName lsidName="C. kely" pageId="0" pageNumber="19" rank="species" species="kely">
<emphasis italics="true" pageId="0" pageNumber="19">C. kely</emphasis>
</taxonomicName>
. Penis more or less straight in ventral view, which separates the species from all other in the
<taxonomicName lsidName="insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">insuetus</emphasis>
</taxonomicName>
complex.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="19" type="description">
<paragraph pageId="0" pageNumber="19">Description.</paragraph>
<paragraph pageId="0" pageNumber="19">
Body length 3.9-4.5 mm. Body shape elongate oval, but slightly less elongate compared with
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
. Head rufotestaceous with only a faint suggesstion of an M-shaped infuscation between eyes. Pronotum rufotestaceous as the head, with weak medial infuscation Elytra dark brown, with testaceous band at base. Testaceous band generally broader and more diffusely transitioning into the darker elytral colour posteriorly (
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Habitus, dorsal view. A Female: Copelatus insuetus B Male: Copelatus vokoka sp. nov. C Female: Copelatus kely sp. nov. D Female: Copelatus sp. 3 (Ivohibe)." figureDoi="10.3897/zookeys.869.33997.figure9" httpUri="https://binary.pensoft.net/fig/322934" pageId="0" pageNumber="19">Fig. 9B</figureCitation>
) compared with
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
. Antennae, palps and legs testaceous.
</paragraph>
<paragraph pageId="0" pageNumber="19">
Elytra with six discal and one submarginal striae. First to fourth more or less full length, fifth and sixth slightly abbreviated anteriorly; submarginal stria starting approximately one half to one third from base. Striae more deeply impressed and intervals therefore slightly more convex compared to
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
. Head, pronotum, and elytra microreticulate and finely micropunctate. Pronotum extensively and coarsely striated on posterior surfaces, although reduced posteromedially. On some specimens striolation covers most of pronotum, also on anterior surfaces, but is reduced medially.
</paragraph>
<paragraph pageId="0" pageNumber="19">
Ventral side entirely testaceous except metacoxal plate which is variably infuscated, especially laterally in some individuals; metacoxa and abdominal sternites
<normalizedToken originalValue="IIIV">II-IV</normalizedToken>
striolate. Prosternal process with a slightly more pointed apex than in
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
and
<taxonomicName lsidName="C. kely" pageId="0" pageNumber="19" rank="species" species="kely">
<emphasis italics="true" pageId="0" pageNumber="19">C. kely</emphasis>
</taxonomicName>
, and lateral parts of metaventrite not as broad. Metacoxal lines long and not strongly diverging, as in
<taxonomicName lsidName="C. kely" pageId="0" pageNumber="19" rank="species" species="kely">
<emphasis italics="true" pageId="0" pageNumber="19">C. kely</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph pageId="0" pageNumber="19">
Male: first three pro- and mesotarsomeres widened, ventrally equipped with suction cups (same constellation as in
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
). Protibia modified, bisinuate and angled basally and broadened distally. Pro- and mesotarsal claws unmodified. Penis long, thin and simple; apex in ventral view straight and not leftturned; in lateral view rather evenly arched, lacking the distinct
<normalizedToken originalValue="“shoulder”">&quot;shoulder&quot;</normalizedToken>
characteristic of
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
but with a different type of postmedial and preapical suggested humps in the curvature. Apex in lateral view also broder closer to apex. Apex in left lateral view with a dorsal ridge crossing posterior dorsal margin but at a more acute angle (
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Male genitalia. From left to right, aedeagus in right lateral, ventral, left lateral views and left paramere. A Copelatus insuetus B Copelatus kely sp. nov. C Copelatus vokoka sp. nov. D Copelatus ankaratra sp. nov. E Copelatus safiotra sp. nov." figureDoi="10.3897/zookeys.869.33997.figure6" httpUri="https://binary.pensoft.net/fig/322931" pageId="0" pageNumber="19">Fig. 6C</figureCitation>
) compared with
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
. Preapically left side with fine transverse ridges. Right lateral and ventral side with fine longitudinal microsculpture but not forming coarse sulci as in
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
. Parameres as in
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Male genitalia. From left to right, aedeagus in right lateral, ventral, left lateral views and left paramere. A Copelatus insuetus B Copelatus kely sp. nov. C Copelatus vokoka sp. nov. D Copelatus ankaratra sp. nov. E Copelatus safiotra sp. nov." figureDoi="10.3897/zookeys.869.33997.figure6" httpUri="https://binary.pensoft.net/fig/322931" pageId="0" pageNumber="19">Figure 6C</figureCitation>
.
</paragraph>
<paragraph pageId="0" pageNumber="19">Female: very faint to no striolation on outer elytral intervals in the series from Ivohibe. All females from Ranomafana NP were densely striolated over the entire elytral surface except at the apical part, and entire pronotum. Density of striation approx. 5-8 striae in breadth across each elytral interval.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="19" type="etymology">
<paragraph pageId="0" pageNumber="19">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="19">
Vokoka is a Malagasy adjective for curvature, also used for an old person with a hunched back. Here it refers to the even curvature of the male aedeagus in lateral view where the even curvature sets it apart from
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
. It is a non-latinised adjective.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="19" type="distribution">
<paragraph pageId="0" pageNumber="19">Distribution.</paragraph>
<paragraph pageId="0" pageNumber="19">
Known from the mountainous escarpment of southeastern Madagascar at Ivohibe Special Reserve and Ranomafana NP (
<figureCitation captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Distribution maps of Copelatus species. A C. insuetus (circle), C. pseudostriatus sp. nov. (square) B C. kely sp. nov. (circle), C. vokoka sp. nov. (square) C C. ankaratra sp. nov. (circle), C. sp. 3 (square) D C. safiotra sp. nov. (circle), C. befasicus (square)." figureDoi="10.3897/zookeys.869.33997.figure12" httpUri="https://binary.pensoft.net/fig/322937" pageId="0" pageNumber="19">Fig. 12B</figureCitation>
). Our sampling is rather scant in the humid forests south of Ivohibe, for instance down to Andohahela NP so it is possible the species distribution continues further south.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="19" type="habitat">
<paragraph pageId="0" pageNumber="19">Habitat and ecology.</paragraph>
<paragraph pageId="0" pageNumber="19">
We collected this species in 2013 from forest pools with dead leaves next to streams at the Ivohibe reserve in pristine humid forest at an altitude of 870 m. The reserve of Pic
<normalizedToken originalValue="dIvohibe">d'Ivohibe</normalizedToken>
was established in 1964 and is managed through collaboration between Madagascar National Parks, local people associations, and other partners. During our visit in 2013, there were little signs of degradation except at the entrance and at the west edge of the reserve. Local people sometimes take zebu cattle with them on a path through the forest. At a larger scale zebu excrement could influence freshwater quality and species assemblages through eutrophication, but there were no signs of this inside the intact pristine forest. In Ranomafana NP it was collected in a shallow clear-water shaded forest pond with large quantity of dead tree leaves and some
<taxonomicName family="Poaceae" lsidName="" pageId="0" pageNumber="19" rank="family">Poaceae</taxonomicName>
(M. Manuel pers. comm.).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="19" type="comments">
<paragraph pageId="0" pageNumber="19">Comments.</paragraph>
<paragraph pageId="0" pageNumber="19">
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus vokoka" order="Coleoptera" pageId="0" pageNumber="19" phylum="Arthropoda" rank="species" species="vokoka">
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus vokoka</emphasis>
</taxonomicName>
sp. nov. falls in the
<taxonomicName lsidName="irinus" pageId="0" pageNumber="19" rank="species" species="irinus">irinus</taxonomicName>
group, based on the number of elytral striae. The new species is most closely related to
<taxonomicName lsidName="C. ankaratra" pageId="0" pageNumber="19" rank="species" species="ankaratra">
<emphasis italics="true" pageId="0" pageNumber="19">C. ankaratra</emphasis>
</taxonomicName>
according to the mitochondrial gene CO1 and belongs to the
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
species complex on Madagascar.
</paragraph>
</subSubSection>
</treatment>
</document>