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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="851D9EEE861E5E29812C70504B087522" docLanguage="en" docName="ZooKeys 869: 19-90" docOrigin="ZooKeys 869" docSource="http://dx.doi.org/10.3897/zookeys.869.33997" docTitle="Copelatus safiotra Ranarilalatiana &amp; Bergsten, sp. nov." docType="treatment" docUuid="137C62A2-22DD-4550-8697-9E81CED4ED45" 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">
<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>
</mods:extent>
</mods:part>
</mods:relatedItem>
<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="158995550" LSID="urn:lsid:zoobank.org:act:137C62A2-22DD-4550-8697-9E81CED4ED45" httpUri="http://treatment.plazi.org/id/851D9EEE861E5E29812C70504B087522" lastPageNumber="19" pageId="0" pageNumber="19">
<subSubSection pageId="0" pageNumber="19" type="nomenclature">
<paragraph pageId="0" pageNumber="19">
<taxonomicName LSID="851d9eee-861e-5e29-812c-70504b087522" authority="Ranarilalatiana &amp; Bergsten" class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus safiotra" order="Coleoptera" pageId="0" pageNumber="19" phylum="Arthropoda" rank="species" species="safiotra">Copelatus safiotra 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 6E</figureCitation>
,
<figureCitation captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Habitus, dorsal view. A Female: Copelatus ankaratra sp. nov. B Male: Copelatus pseudostriatus sp. nov. C Male: Copelatus safiotra sp. nov. D Female: Copelatus befasicus." figureDoi="10.3897/zookeys.869.33997.figure10" httpUri="https://binary.pensoft.net/fig/322935" pageId="0" pageNumber="19">10C</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="19" type="type locality">
<paragraph pageId="0" pageNumber="19">Type locality.</paragraph>
<paragraph pageId="0" pageNumber="19">Anjanaharibe Sud reserve, [14.7414S, 049.4975E] [Madagascar, Sava region, Andapa 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">Antsiranana. Sava: Andapa</emphasis>
: -HT♂ (GP) (NHRS): // NHRS-JLKB | 000065415 // MAD: ANTS: Sava: Anjanaharibe | Sud NP: stream next to Camp | site: Mad14-62: medium size | sandy forest stream: 14.7414S | 49.4975E; 910 m: 14.XI.2014 // Leg. J. Bergsten, R. | Bukontaite, S. Holmgren, | J.H. Randriamihaja | &amp; T. Ranarilalatiana // Holotype |
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus safiotra</emphasis>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 //
<emphasis bold="true" pageId="0" pageNumber="19">Paratypes</emphasis>
: -2♀ (Alc.) (NHRS): // NHRS-JLKB | 000010785 // MAD: ANTS: Sava: Anjanaharibe | Sud NP: stream next to Camp | site: Mad14-62: medium size | sandy forest stream: 14.7414S | 49.4975E; 910 m: 14.XI.2014 // Leg. J. Bergsten, R. | Bukontaite, S. Holmgren, | J.H. Randriamihaja | &amp; T. Ranarilalatiana // Paratype |
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus safiotra</emphasis>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 // -4♂ (GP), 1♀, 12 ex. (Alc.), 3 ex. (Alc.): (NHRS, NHMUK, DEUA &amp; PBZT/MBC): // NHRS-JLKB | 000010506-7, 10566, 10595, 65414, 10775(Alc.), 10845(Alc.) // MAD: ANTS: Sava: Anjanaharibe | Sud NP: Camp site: Mad14-70: | forest stream: 14.7414S | 49.4975E; 910 m: 16.XI.2014 // Leg. J. Bergsten, R. | Bukontaite, S. Holmgren, | J.H. Randriamihaja | &amp; T. Ranarilalatiana // Paratype |
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus safiotra</emphasis>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 //
<emphasis bold="true" pageId="0" pageNumber="19">Toamasina. Alaotra Mangoro: Ambatondrazaka, Moramanga</emphasis>
: -4♂ (GP), 29 ex. (Alc.), 6 ex. (3♂, 3♀) (Alc.) (NHRS, NHMUK, DEUA &amp; PBZT/MBC): // NHRS-JLKB | 000010818, 10836-7, 65413, 10838(Alc.), 65752(Alc.) // Madagascar: Toamasina: Alaotra | Mangoro: Zahamena NP: Sect. | Antanandava: close to Camp Bemoara | S17.5108; E048.7287; 1060 m. 07.III.2018 | GB Nets, white pan and sieves: Waterfilled | goldigging holes: Field# MAD18-87 // Leg. J. Bergsten, &amp; | T. Ranarilalatiana // Paratype |
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus safiotra</emphasis>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 // -1♂ (teneral) (NHRS): // NHRS-JLKB | 000010871 // Madagascar: Toamasina: Alaotra | Mangoro: Zahamena NP: Sect. | Antanandava: Manambato stream by Camp | Cascade: S17.545; E048.7237; 1290 m. 09.III.2018 | GB Nets, white pan and sieves: large | foreststream: Field# MAD18-100 // Leg. J. Bergsten, &amp; | T. Ranarilalatiana // Paratype |
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus safiotra</emphasis>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 // -4♂ GP, 2♀, 6 ex. (1♂, 5♀) (Alc.) (NHRS, DEUA &amp; PBZT/MBC): // NHRS-JLKB | 000010589-94, 10774(Alc.) // Madagascar: Toamasina: Alaotra | Mangoro: Mantadia NP: Waterfall | 6km from park entrance: S18.83396 | E048.43777, 1000 m, 11.XI.2011 GB | Nets and sieves: forest stream in | rainforest: Field# MAD11-37 // Leg. J. Bergsten, R. | Bukontaite, T | Ranarilalatiana &amp; | J.H. Randriamihaja // Paratype |
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus safiotra</emphasis>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 // -4♀ (Alc.) (NHRS): // NHRS-JLKB | 000010784 // Madagascar: Toamasina: Alaotra | Mangoro: Mantadia NP: River | Sahanody 9 km from park entrance: S | 18.80973 E 048.42861, 930 m. 11.XI.2011 | GB Nets and sieves: forest stream in | rainforest: Field# MAD11-34 // Leg. J. Bergsten, R. | Bukontaite, T | Ranarilalatiana &amp; | J.H. Randriamihaja // Paratype |
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus safiotra</emphasis>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 //
<emphasis bold="true" pageId="0" pageNumber="19">Fianarantsoa. Matsiatra Ambony: Lalangina</emphasis>
: -1♂ (GP) (NHRS): // NHRS-JLKB | 000010846 // MAD: FIAN: Matsiatra Ambony | Ranomafana NP: next to Sahamalaotra | trail entrance: Mad14-04: forest stream: | with sandy bottom:
<geoCoordinate degrees="21.2395" direction="south" orientation="latitude" precision="5" value="-21.2395">21.2395S</geoCoordinate>
<geoCoordinate degrees="49.3947" direction="east" orientation="longitude" precision="5" value="49.3947">49.3947E</geoCoordinate>
: 1130 m: 02.XI.2014 // Leg. J. Bergsten, | T. Ranarilalatiana | &amp; S. Holmgren // Paratype |
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus safiotra</emphasis>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 //
<emphasis bold="true" pageId="0" pageNumber="19">Fianarantsoa. Ihorombe: Ihosy</emphasis>
: -2♂ (GP) (NHRS): // NHRS-JLKB | 000010847-8 // Madagascar: Fianarantsoa: Ihorombe: | Isalo NP: 300m into the canyon de | Makis: S
<geoCoordinate degrees="22.48665" direction="east" orientation="longitude" precision="1" value="22.48665">22.48665 E</geoCoordinate>
45.37966, 700 m | 13.XI.2012, GB nets and sieves: canyon | river with side pools: Leg. R. Bukontaite | &amp; J.H. Randriamihaja: Field# MAD12-03 // Paratype |
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus safiotra</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">
Somewhat similar to
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
on habitus appearance, but sturdier, broader pronotum and head with a much greater interocular distance compared to width of eyes, and subparalell along a longer distance of body with more rapidly attenuating anterior and posterior ends; easily distinguished from all other
<taxonomicName lsidName="irinus" pageId="0" pageNumber="19" rank="species" species="irinus">
<emphasis italics="true" pageId="0" pageNumber="19">irinus</emphasis>
</taxonomicName>
group species of Madagascar by the penis shape, which is of a type otherwise found in species closely related to
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus owas" order="Coleoptera" pageId="0" pageNumber="19" phylum="Arthropoda" rank="species" species="owas">
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus owas</emphasis>
</taxonomicName>
; penis has a large medial expansion in lateral view followed by an apical blade (
<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. 6E</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="19" type="description">
<paragraph pageId="0" pageNumber="19">Description.</paragraph>
<paragraph pageId="0" pageNumber="19">
Body length 4.3-5.2 mm. Body shape elongate and subparallel along a very long part of the body. Pronotum and head broad and eyes small creating a very wide interocular distance. Maximum width of pronotum clearly in front of hind corners. Head rufotestaceous with weak or absent infuscation in between and posterior of eyes. Pronotum infuscated medially with broadly testaceous lateral sides. Elytra brown, with a broad testaceous transverse band basally (
<figureCitation captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Habitus, dorsal view. A Female: Copelatus ankaratra sp. nov. B Male: Copelatus pseudostriatus sp. nov. C Male: Copelatus safiotra sp. nov. D Female: Copelatus befasicus." figureDoi="10.3897/zookeys.869.33997.figure10" httpUri="https://binary.pensoft.net/fig/322935" pageId="0" pageNumber="19">Fig. 10C</figureCitation>
). Testaceous band broader and transitioning posteriorly more diffusely into the brown colour 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 stria. Fifth stria abbreviated anteriorly, variably also first and third striae. In some individuals especially fifth but also first and sixth striae are rudimentary or with very shallow impressions. Interval between fifth and sixth striae narrow, approx. half interval between first and second striae. Submarginal stria short, starting around middle. Head, pronotum, and elytra microreticulate and finely micropunctate. Pronotum not striolate.</paragraph>
<paragraph pageId="0" pageNumber="19">
Ventral side entirely testaceous except last three abdominal ventrites may be vaguely infuscated and have lighter lateral spots. Metacoxa and abdominal sternite
<normalizedToken originalValue="II–IV">II-IV</normalizedToken>
striolate, but strioles shallower and finer 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>
. 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>
, the metacoxal lines are shorter, and the anterior traces suggest an inward curve towards the posterior metaventral margin. The lateral part of metaventrite is narrower 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>
at level of mesocoxa, equal to the width of mesofemur at middle. Posterior metaventral margin not straight but slightly angular at level of apex of mesotrochanter. Prosternal process is slightly more elongate.
</paragraph>
<paragraph pageId="0" pageNumber="19">
Male: first three pro- and mesotarsomeres widened, ventrally equipped with suction cups. Pattern of suction cups same as for
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
but tarsomeres not as wide and less developed as an integrated protarsal palett. Protibia modified, bisinuate, angled basally, and broadened distally. Pro- and mesotarsal claws unmodified. Penis very characteristic, rather broad and short and in lateral view with a medial expansion followed by a sharp constriction before the narrow blade-like 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. 6E</figureCitation>
) which is characteristic of several species related to
<taxonomicName lsidName="C. owas" pageId="0" pageNumber="19" rank="species" species="owas">
<emphasis italics="true" pageId="0" pageNumber="19">C. owas</emphasis>
</taxonomicName>
<normalizedToken originalValue="Régimbart">Regimbart</normalizedToken>
, 1895; apical blade curved leftwards in ventral view and serrated by transverse ridges on the convex right side. 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 6E</figureCitation>
, rather long and thin with a medial emargination on the concave side.
</paragraph>
<paragraph pageId="0" pageNumber="19">Female: on average smaller than males (Table 3). At least some females with elytral microreticulation slightly more strongly impressed than in males and therefore appearing more matt.</paragraph>
<caption ID-Table-UUID="F19D865534B08E4398BA0A0C892ECF52" httpUri="http://table.plazi.org/id/F19D865534B08E4398BA0A0C892ECF52" pageId="0" pageNumber="19">
<paragraph pageId="0" pageNumber="19">Table 3. Measurements of body length summarised as Min, Max, Mean, and Standard Deviation (SD) for each species, separated by sex. N = number of measured individuals, F = females, M = males.</paragraph>
</caption>
<paragraph pageId="0" pageNumber="19">
<table pageId="0" pageNumber="19">
<tr pageId="0" pageNumber="19">
<th colspan="1" pageId="0" pageNumber="19" rowspan="1">Species</th>
<th colspan="1" pageId="0" pageNumber="19" rowspan="1">Sex</th>
<th colspan="1" pageId="0" pageNumber="19" rowspan="1">N</th>
<th colspan="1" pageId="0" pageNumber="19" rowspan="1">Min</th>
<th colspan="1" pageId="0" pageNumber="19" rowspan="1">Max</th>
<th colspan="1" pageId="0" pageNumber="19" rowspan="1">Mean</th>
<th colspan="1" pageId="0" pageNumber="19" rowspan="1">SD</th>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="2">
<taxonomicName lsidName="C. befasicus" pageId="0" pageNumber="19" rank="species" species="befasicus">
<emphasis italics="true" pageId="0" pageNumber="19">C. befasicus</emphasis>
</taxonomicName>
</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">F</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.13</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.19</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.15</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.03</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">M</td>
<td colspan="1" isEmpty="true" pageId="0" pageNumber="19" rowspan="1">-</td>
<td colspan="1" isEmpty="true" pageId="0" pageNumber="19" rowspan="1">-</td>
<td colspan="1" isEmpty="true" pageId="0" pageNumber="19" rowspan="1">-</td>
<td colspan="1" isEmpty="true" pageId="0" pageNumber="19" rowspan="1">-</td>
<td colspan="1" isEmpty="true" pageId="0" pageNumber="19" rowspan="1">-</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="2">
<taxonomicName lsidName="C. marginipennis" pageId="0" pageNumber="19" rank="species" species="marginipennis">
<emphasis italics="true" pageId="0" pageNumber="19">C. marginipennis</emphasis>
</taxonomicName>
</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">F</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">26</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.16</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">6.32</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.75</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.30</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">M</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">45</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.48</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">6.58</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">6.02</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.26</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="2">
<taxonomicName lsidName="C. mahajanga" pageId="0" pageNumber="19" rank="species" species="mahajanga">
<emphasis italics="true" pageId="0" pageNumber="19">C. mahajanga</emphasis>
</taxonomicName>
</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">F</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">12</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.16</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">6.13</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.63</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.24</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">M</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">16</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.42</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">6.19</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.88</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.20</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="2">
<taxonomicName lsidName="C. pulchellus" pageId="0" pageNumber="19" rank="species" species="pulchellus">
<emphasis italics="true" pageId="0" pageNumber="19">C. pulchellus</emphasis>
</taxonomicName>
</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">F</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">1</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.74</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.74</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.74</td>
<td colspan="1" isEmpty="true" pageId="0" pageNumber="19" rowspan="1">-</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">M</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">3</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.48</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">6.06</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.83</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.30</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="2">
<taxonomicName lsidName="C. distinguendus" pageId="0" pageNumber="19" rank="species" species="distinguendus">
<emphasis italics="true" pageId="0" pageNumber="19">C. distinguendus</emphasis>
</taxonomicName>
</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">F</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">44</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.29</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">6.32</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.75</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.21</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">M</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">26</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.48</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">6.26</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.92</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.18</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="2">
<taxonomicName lsidName="C. peridinus" pageId="0" pageNumber="19" rank="species" species="peridinus">
<emphasis italics="true" pageId="0" pageNumber="19">C. peridinus</emphasis>
</taxonomicName>
</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">F</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">7</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.74</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">6.58</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">6.05</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.27</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">M</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">7</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.81</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">6.26</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">6.10</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.16</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="2">
<taxonomicName lsidName="C. baculiformis" pageId="0" pageNumber="19" rank="species" species="baculiformis">
<emphasis italics="true" pageId="0" pageNumber="19">C. baculiformis</emphasis>
</taxonomicName>
</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">F</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">1</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.00</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.00</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.00</td>
<td colspan="1" isEmpty="true" pageId="0" pageNumber="19" rowspan="1">-</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">M</td>
<td colspan="1" isEmpty="true" pageId="0" pageNumber="19" rowspan="1">-</td>
<td colspan="1" isEmpty="true" pageId="0" pageNumber="19" rowspan="1">-</td>
<td colspan="1" isEmpty="true" pageId="0" pageNumber="19" rowspan="1">-</td>
<td colspan="1" isEmpty="true" pageId="0" pageNumber="19" rowspan="1">-</td>
<td colspan="1" isEmpty="true" pageId="0" pageNumber="19" rowspan="1">-</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="2">
<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.
</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">F</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">7</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">3.81</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.26</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.01</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.16</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">M</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">3.87</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.32</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.03</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.17</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="2">
<taxonomicName lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
</taxonomicName>
</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">F</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">27</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.26</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.03</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.60</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.20</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">M</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">24</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.26</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.90</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.61</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.17</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="2">
<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.
</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">F</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">3</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.32</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.71</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.58</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.22</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">M</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">13</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.52</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.16</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.86</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.19</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="2">
<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.
</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">F</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">6</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">3.94</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.32</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.18</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.13</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">M</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">8</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.06</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.45</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.29</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.12</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="2">
<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.
</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">F</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">19</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.39</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.03</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.70</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.17</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">M</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">30</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.65</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.16</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">4.87</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.16</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="2">
<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.
</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">F</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">2</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.42</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.61</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.52</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">0.14</td>
</tr>
<tr pageId="0" pageNumber="19">
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">M</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">1</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.29</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.29</td>
<td colspan="1" pageId="0" pageNumber="19" rowspan="1">5.29</td>
<td colspan="1" isEmpty="true" pageId="0" pageNumber="19" rowspan="1">-</td>
</tr>
</table>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="19" type="etymology">
<paragraph pageId="0" pageNumber="19">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="19">
The species name
<taxonomicName lsidName="safiotra" pageId="0" pageNumber="19" rank="species" species="safiotra">
<emphasis italics="true" pageId="0" pageNumber="19">safiotra</emphasis>
</taxonomicName>
is a Malagasy noun for hybrid, here referring to the unusual combination of a male genitalia type, typical of the
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus owas" order="Coleoptera" pageId="0" pageNumber="19" phylum="Arthropoda" rank="species" species="owas">
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus owas</emphasis>
</taxonomicName>
species complex, in a body with a 6+1 striated elytra very much resembling 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 of species. It is a non-latinised noun in apposition.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="19" type="distribution">
<paragraph pageId="0" pageNumber="19">Distribution.</paragraph>
<paragraph pageId="0" pageNumber="19">
Endemic to Madagascar. This species has a rather large distribution in the eastern humid forest from Anjanaharibe Sud reserve in the NE, all along the eastern escarpment including Zahamena NP, Mantadia NP, and Ranomafana NP, and even extending to the rather isolated western patch of subhumid forest at Isalo 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. 12D</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="19" type="habitat">
<paragraph pageId="0" pageNumber="19">Habitat and ecology.</paragraph>
<paragraph pageId="0" pageNumber="19">
This species seems to be strongly associated with clean streams having sandy substrate in humid forests. At these localities, individuals can be found in sidepools, at margins or sites protected from waterflow (e.g., by fallen logs) where dead leaves and debris accumulate. The species has been found at altitudes between 700 and 1300 m but most numerous at elevations above 900 m in primary humid forest. The discovery in Isalo NP, in a sandy river running through a Canyon, indicates that subhumid forests may also be part of the
<normalizedToken originalValue="species’">species'</normalizedToken>
niche.
</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 safiotra" order="Coleoptera" pageId="0" pageNumber="19" phylum="Arthropoda" rank="species" species="safiotra">
<emphasis italics="true" pageId="0" pageNumber="19">Copelatus safiotra</emphasis>
</taxonomicName>
sp. nov. falls in the
<taxonomicName lsidName="irinus" pageId="0" pageNumber="19" rank="species" species="irinus">
<emphasis italics="true" pageId="0" pageNumber="19">irinus</emphasis>
</taxonomicName>
group, based on the number of elytral striae (six discal and one submarginal striae). However, the genitalia is of a very different type and characteristic of the complex of species close to
<taxonomicName lsidName="C. owas" pageId="0" pageNumber="19" rank="species" species="owas">
<emphasis italics="true" pageId="0" pageNumber="19">C. owas</emphasis>
</taxonomicName>
in the
<emphasis italics="true" pageId="0" pageNumber="19">erichsonii</emphasis>
group with ten discal and one submarginal striae. We hypothesize that this species, despite the body shape and number of elytral striae, is not related to the other species treated here, but belongs to the radiation of species around
<taxonomicName lsidName="C. owas" pageId="0" pageNumber="19" rank="species" species="owas">
<emphasis italics="true" pageId="0" pageNumber="19">C. owas</emphasis>
</taxonomicName>
. This would reinforce the idea that the number of elytral striae is a very homoplastic character and not reliable to create phylogenetically sound groups (
<bibRefCitation DOI="https://doi.org/10.1016/j.ympev.2004.03.014" author="Balke, M" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="19" pagination="866 - 880" refId="B5" refString="Balke, M, Ribera, I, Vogler, AP, 2004. MtDNA phylogeny and biogeography of Copelatinae, a highly diverse group of tropical diving beetles (Dytiscidae). . Molecular Phylogenetics and Evolution 32: 866 - 880" title="MtDNA phylogeny and biogeography of Copelatinae, a highly diverse group of tropical diving beetles (Dytiscidae)." url="https://doi.org/10.1016/j.ympev.2004.03.014" volume="32" year="2004">Balke et al. 2004</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>
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