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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="70C03B1EE8D25A8F8F488FFE0EB3CEB6" docLanguage="en" docName="ZooKeys 869: 19-90" docOrigin="ZooKeys 869" docSource="http://dx.doi.org/10.3897/zookeys.869.33997" docTitle="Copelatus : sp. 3" docType="treatment" docVersion="4" 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">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<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>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<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>
</mods:mods>
<treatment ID-GBIF-Taxon="158995542" LSID="urn:lsid:plazi:treatment:70C03B1EE8D25A8F8F488FFE0EB3CEB6" httpUri="http://treatment.plazi.org/id/70C03B1EE8D25A8F8F488FFE0EB3CEB6" lastPageNumber="19" pageId="0" pageNumber="19">
<subSubSection pageId="0" pageNumber="19" type="nomenclature">
<paragraph pageId="0" pageNumber="19">
<taxonomicName LSID="70c03b1e-e8d2-5a8f-8f48-8ffe0eb3ceb6" authority="(Ivohibe and North of Toamasina): sp. 3" class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus" order="Coleoptera" pageId="0" pageNumber="19" phylum="Arthropoda" rank="genus">Copelatus (Ivohibe and North of Toamasina): sp. 3</taxonomicName>
<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. 9D</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="19" type="materials_examined">
<paragraph pageId="0" pageNumber="19">Material studied.</paragraph>
<paragraph pageId="0" pageNumber="19">
<emphasis bold="true" pageId="0" pageNumber="19">Fianarantsoa. Ihorombe: Ivohibe</emphasis>
: -3♀ (NHRS): // NHRS-JLKB | 000010856, 65699, 65734 // Madagascar: Fianarantsoa: Ihorombe: R.S. Pic |
<normalizedToken originalValue="dIvohibe">d'Ivohibe</normalizedToken>
: Andaranovory: close to botanical | transect R.S. Pic
<normalizedToken originalValue="dIvohibe">d'Ivohibe</normalizedToken>
: S22.47511667 | E046.9559, 1106 m, 10.XII.2013, GB Nets and | sieves: small lake with dead leaves and | vegetation, Leg. J.H. Randriamihaja &amp; | T. Ranarialalatiana: Field# MAD13-61 //
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus</emphasis>
sp. nov. |
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
complex | Det. Ranarilalatiana | &amp; Bergsten, 2019 //
<emphasis bold="true" pageId="0" pageNumber="19">Toamasina. Atsinanana: Toamasina, Toamasina II</emphasis>
: -1♀ (NHRS): // NHRS-JLKB | 000010811 // Madagascar: Toamasina II: Analalava | reserve: MAD17-12: S of nursery plants: | S17.71055; E49.45002; 39 m: Forest | stream with side pools: 09/03/2017; | Leg. T. Ranarilalatiana //
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus</emphasis>
sp. nov. | C. insuetus complex | Det. Ranarilalatiana | &amp; Bergsten, 2019 // -1♀ (NHRS): // NHRS-JLKB | 000010779 // Madagascar: Toamasina: | Antsinanana: RN2, 6Km N | Toamasina by bridge: S18.06493 | E049.37856, 0 m. 15.XI.2011, | GB Nets and sieves: river and | sidepool: Field# MAD11-52 // Leg. J. Bergsten, R. | Bukontaite, T | Ranarilalatiana &amp; | J.H. Randriamihaja //
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus</emphasis>
sp. nov. |
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
complex | Det. Ranarilalatiana | &amp; Bergsten, 2019 //
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="19" type="comments">
<paragraph pageId="0" pageNumber="19">Comments.</paragraph>
<paragraph pageId="0" pageNumber="19">
The DNA data revealed that these females represent one or possibly two additional new species in the
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
complex (
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Majority-rule consensus tree from Bayesian analysis of CO 1. Values next to nodes indicate posterior probabilities." figureDoi="10.3897/zookeys.869.33997.figure2" httpUri="https://binary.pensoft.net/fig/322927" pageId="0" pageNumber="19">Figs 2</figureCitation>
,
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Result of the single-locus GMYC species delimitation using an ultrametric CO 1 genetree from BEAST. Black branches indicate interspecific divergences, red branches represent intraspecific coalescence events. Values above interspecific nodes indicate posterior probability from the Beast analysis under a strict clock model. The * indicates two nodes of further splitting from the GMYC analysis that is within the confidence interval of 2 Log likelihood units from the optimal solution." figureDoi="10.3897/zookeys.869.33997.figure3" httpUri="https://binary.pensoft.net/fig/322928" pageId="0" pageNumber="19">3</figureCitation>
). In fact, the CO1 data reveals that they are the most divergent in that group and are sister to a clade with all the other species:
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="C. vokoka" pageId="0" pageNumber="19" rank="species" species="vokoka">
<emphasis italics="true" pageId="0" pageNumber="19">C. vokoka</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="C. kely" pageId="0" pageNumber="19" rank="species" species="kely">
<emphasis italics="true" pageId="0" pageNumber="19">C. kely</emphasis>
</taxonomicName>
, and
<taxonomicName lsidName="C. ankaratra" pageId="0" pageNumber="19" rank="species" species="ankaratra">
<emphasis italics="true" pageId="0" pageNumber="19">C. ankaratra</emphasis>
</taxonomicName>
. The genetic distance between members of these two clades ranges from a minimum of 4.5% to a maximum of 7.1%, strongly indicating a separately evolving unit. The genetic distance between the specimens from Ivohibe and those from north of Toamasina was 2.3-2.4% (K2P), a distance that does not rule out conspecificity as the geographic and altitudinal distance are substantial between these localities (for distribution see
<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. 12C</figureCitation>
). It is also on the same level as the intraspecific distance found within
<taxonomicName lsidName="C. ankaratra" pageId="0" pageNumber="19" rank="species" species="ankaratra">
<emphasis italics="true" pageId="0" pageNumber="19">C. ankaratra</emphasis>
</taxonomicName>
between a peak population and a population at lower altitude of the Ankaratra Massif between which we do not find any morphological character differences to justify further separation. On the other hand,
<taxonomicName lsidName="C. kely" pageId="0" pageNumber="19" rank="species" species="kely">
<emphasis italics="true" pageId="0" pageNumber="19">C. kely</emphasis>
</taxonomicName>
and
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
are indistinguishable based on CO1 squences alone (
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Majority-rule consensus tree from Bayesian analysis of CO 1. Values next to nodes indicate posterior probabilities." figureDoi="10.3897/zookeys.869.33997.figure2" httpUri="https://binary.pensoft.net/fig/322927" pageId="0" pageNumber="19">Figs 2</figureCitation>
,
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Result of the single-locus GMYC species delimitation using an ultrametric CO 1 genetree from BEAST. Black branches indicate interspecific divergences, red branches represent intraspecific coalescence events. Values above interspecific nodes indicate posterior probability from the Beast analysis under a strict clock model. The * indicates two nodes of further splitting from the GMYC analysis that is within the confidence interval of 2 Log likelihood units from the optimal solution." figureDoi="10.3897/zookeys.869.33997.figure3" httpUri="https://binary.pensoft.net/fig/322928" pageId="0" pageNumber="19">3</figureCitation>
). We refrain from describing these as a new species since the shape of male genitalia is very important for identification in this group. Morphologically we note the following based on the females: in body size this species is similar to
<taxonomicName lsidName="C. vokoka" pageId="0" pageNumber="19" rank="species" species="vokoka">
<emphasis italics="true" pageId="0" pageNumber="19">C. vokoka</emphasis>
</taxonomicName>
, slightly larger than
<taxonomicName lsidName="C. kely" pageId="0" pageNumber="19" rank="species" species="kely">
<emphasis italics="true" pageId="0" pageNumber="19">C. kely</emphasis>
</taxonomicName>
but slightly smaller 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
<taxonomicName lsidName="C. ankaratra" pageId="0" pageNumber="19" rank="species" species="ankaratra">
<emphasis italics="true" pageId="0" pageNumber="19">C. ankaratra</emphasis>
</taxonomicName>
. Compared with females of
<taxonomicName lsidName="C. vokoka" pageId="0" pageNumber="19" rank="species" species="vokoka">
<emphasis italics="true" pageId="0" pageNumber="19">C. vokoka</emphasis>
</taxonomicName>
,
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus" order="Coleoptera" pageId="0" pageNumber="19" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus</emphasis>
</taxonomicName>
sp. 3 has a narrower testaceous basal band, flatter elytral intervals between striae, and more limited striolation on pronotum, restricted to posterolateral corners.
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus ankaratra" order="Coleoptera" pageId="0" pageNumber="19" phylum="Arthropoda" rank="species" species="ankaratra">
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus ankaratra</emphasis>
</taxonomicName>
can be distinguished based on its dark colour and extremely elongate body shape. Small
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
females can often be distinguished on the posteriorly extended testaceous basal medial band. It is most difficult to distinguish
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus" order="Coleoptera" pageId="0" pageNumber="19" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus</emphasis>
</taxonomicName>
sp. 3 females from large female specimens of
<taxonomicName lsidName="C. kely" pageId="0" pageNumber="19" rank="species" species="kely">
<emphasis italics="true" pageId="0" pageNumber="19">C. kely</emphasis>
</taxonomicName>
.
</paragraph>
<caption doi="10.3897/zookeys.869.33997.figure12" httpUri="https://binary.pensoft.net/fig/322937" pageId="0" pageNumber="19" start="Figure 12" startId="F12">
<paragraph pageId="0" pageNumber="19">
Figure 12. Distribution maps of
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus" order="Coleoptera" pageId="0" pageNumber="19" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus</emphasis>
</taxonomicName>
species.
<emphasis bold="true" pageId="0" pageNumber="19">A</emphasis>
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
(circle),
<taxonomicName lsidName="C. pseudostriatus" pageId="0" pageNumber="19" rank="species" species="pseudostriatus">
<emphasis italics="true" pageId="0" pageNumber="19">C. pseudostriatus</emphasis>
</taxonomicName>
sp. nov. (square)
<emphasis bold="true" pageId="0" pageNumber="19">B</emphasis>
<taxonomicName lsidName="C. kely" pageId="0" pageNumber="19" rank="species" species="kely">
<emphasis italics="true" pageId="0" pageNumber="19">C. kely</emphasis>
</taxonomicName>
sp. nov. (circle),
<taxonomicName lsidName="C. vokoka" pageId="0" pageNumber="19" rank="species" species="vokoka">
<emphasis italics="true" pageId="0" pageNumber="19">C. vokoka</emphasis>
</taxonomicName>
sp. nov. (square)
<emphasis bold="true" pageId="0" pageNumber="19">C</emphasis>
<taxonomicName lsidName="C. ankaratra" pageId="0" pageNumber="19" rank="species" species="ankaratra">
<emphasis italics="true" pageId="0" pageNumber="19">C. ankaratra</emphasis>
</taxonomicName>
sp. nov. (circle),
<taxonomicName genus="C." lsidName="C." pageId="0" pageNumber="19" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="19">C.</emphasis>
</taxonomicName>
sp. 3 (square)
<emphasis bold="true" pageId="0" pageNumber="19">D</emphasis>
<taxonomicName lsidName="C. safiotra" pageId="0" pageNumber="19" rank="species" species="safiotra">
<emphasis italics="true" pageId="0" pageNumber="19">C. safiotra</emphasis>
</taxonomicName>
sp. nov. (circle),
<taxonomicName lsidName="C. befasicus" pageId="0" pageNumber="19" rank="species" species="befasicus">
<emphasis italics="true" pageId="0" pageNumber="19">C. befasicus</emphasis>
</taxonomicName>
(square).
</paragraph>
</caption>
</subSubSection>
</treatment>
</document>