<document id="43CA916973DA4FC75A9C507848A000D1" ID-CLB-Dataset="26103" 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="246290F17F4059918CAB1EF8A036B9D5" docLanguage="en" docName="ZooKeys 869: 19-90" docOrigin="ZooKeys 869" docSource="http://dx.doi.org/10.3897/zookeys.869.33997" docTitle="Copelatus befasicus Guignot 1956" docType="treatment" docVersion="6" 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="1732686602329" updateUser="ExternalLinkService">
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<mods:title id="231325F3D2BCC453D5B8EB87586AF180">Taxonomic revision of the genus Copelatus of Madagascar (Coleoptera, Dytiscidae, Copelatinae): the non- erichsonii group species</mods:title>
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<mods:namePart id="AB23292BEABBC7FE886D5E4C214FF1A5">Ranarilalatiana, Tolotra</mods:namePart>
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<mods:namePart id="F442D22F5F50F566376FA0FE3817D4B5">Raveloson Ravaomanarivo, Lala Harivelo</mods:namePart>
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<mods:namePart id="03A80E59DC4EF8E9C6F9D2545C69D232">Bergsten, Johannes</mods:namePart>
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<treatment id="246290F17F4059918CAB1EF8A036B9D5" ID-GBIF-Taxon="158995557" LSID="urn:lsid:plazi:treatment:246290F17F4059918CAB1EF8A036B9D5" httpUri="http://treatment.plazi.org/id/246290F17F4059918CAB1EF8A036B9D5" lastPageNumber="19" pageId="0" pageNumber="19" scope_family="Dytiscidae" scope_order="Coleoptera" scope_subFamily="Copelatinae">
<subSubSection id="2AE16679299A03A0A6A0C376A77F49F4" pageId="0" pageNumber="19" type="nomenclature">
<paragraph id="42AD66F862C09BB7C1E2C462E498B7D1" pageId="0" pageNumber="19">
<taxonomicName id="8BAA35D97BDDB389B5EFBAEFA39B4F5E" ID-CoL="Y34D" LSID="246290f1-7f40-5991-8cab-1ef8a036b9d5" authority="Guignot, 1956" authorityName="Guignot" authorityYear="1956" class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus befasicus" order="Coleoptera" pageId="0" pageNumber="19" phylum="Arthropoda" rank="species" species="befasicus">Copelatus befasicus Guignot, 1956</taxonomicName>
<figureCitation id="B5CFE0962A829D991512878D76145A5A" 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. 10D</figureCitation>
</paragraph>
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<subSubSection id="0C01BE6D0090DBDBC35E7BE840C6E349" pageId="0" pageNumber="19" type="reference_group">
<paragraph id="694FDA7F7F281BA5CBA6A021EB017B2C" pageId="0" pageNumber="19">
<taxonomicName id="80CA223DE03330768030F05A3982C58A" class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus befasicus" order="Coleoptera" pageId="0" pageNumber="19" phylum="Arthropoda" rank="species" species="befasicus">Copelatus befasicus</taxonomicName>
Guignot, 1956: 79.
</paragraph>
</subSubSection>
<subSubSection id="D71572E285E364C5DEEEEB50C58A8232" pageId="0" pageNumber="19" type="type locality">
<paragraph id="519DBD3E6C33A10E182600510645118F" pageId="0" pageNumber="19">Type locality.</paragraph>
<paragraph id="8D620EBB290B8A1E8D9FA5C2420E514D" pageId="0" pageNumber="19">Madagascar, Morondava, forest south of Befasy.</paragraph>
</subSubSection>
<subSubSection id="9F3890C86ED3EB25081CF4045BF84EF9" pageId="0" pageNumber="19" type="description">
<paragraph id="8C270E4056D7515440CF133F312E1BBC" pageId="0" pageNumber="19">Type information from original description.</paragraph>
<paragraph id="5A9192B5526D42421905B9F156B44536" pageId="0" pageNumber="19">based on an unknown number of female type specimens but holotype and paratypes are distinguished in introduction. Collected in January 1956 by R. Paulian.</paragraph>
</subSubSection>
<subSubSection id="FEA3A08FE79613705192BAEA0F2E6FF4" pageId="0" pageNumber="19" type="type material">
<paragraph id="7B047F1B2227F3FA3B0643AD81ACC48F" pageId="0" pageNumber="19">Type material studied.</paragraph>
<paragraph id="CEC56179E27BF96317A2AC2B7A7C206F" pageId="0" pageNumber="19">
<emphasis id="89A1677272A3408D2813F8EA3EFA3B31" bold="true" pageId="0" pageNumber="19">Toliara. Menabe: Morondava</emphasis>
: -HT♀ (MNHN, &quot;coll. IRSM&quot;): // Data in NHRS | JLKB | 000030021 // Morondava | 
<normalizedToken id="FE2B2F862B3F3B3F65B5817F5879243B" originalValue="fôret">foret</normalizedToken>
sud | de Befasy | I-56 R.P // Type [red label] // INSTITUT | SCIENTIFIQUE | MADAGASCAR // Guignot det., 1956 | 
<emphasis id="1F76B9290B1873CE9BF370903D6931AE" italics="true" pageId="0" pageNumber="19">Copelatus</emphasis>
| 
<emphasis id="54033EBD2FFA472DDF07E5CEA1CE27AF" italics="true" pageId="0" pageNumber="19">befasicus</emphasis>
n. sp. | Type // -2PT ♀ (MNHN, &quot;coll. Guignot&quot;): // Data in NHRS | JLKB | 000030300-1// Morondava | 
<normalizedToken id="A16BBC825C09CABFF2AB04040CEE689F" originalValue="fôret">foret</normalizedToken>
sud | de Befasy // I-56 R.P // Paratype [red label] // [female symbol] //
</paragraph>
</subSubSection>
<subSubSection id="57F1B4C30FC9BBC2FC4896B318D1F139" pageId="0" pageNumber="19" type="materials_examined">
<paragraph id="701A1EAB4CAA3B3A0FAFC25E49A1F487" pageId="0" pageNumber="19">Additional material studied.</paragraph>
<paragraph id="4013D807D928DC7FFFC0561BCFD8B5A2" pageId="0" pageNumber="19">
<emphasis id="1102C5B5D1856287F7E152DBBE6ABCA1" bold="true" pageId="0" pageNumber="19">Mahajanga. Melaky: Morafenobe</emphasis>
: -1♀ (NHRS): // NHRS-JLKB | 000010860 (JB204) // Madagascar: Mahajanga: Melaky | Btw. 
<normalizedToken id="2595351A7A63F70ECF078614504163E9" originalValue="Morafenobe–Ambohijanahary">Morafenobe-Ambohijanahary</normalizedToken>
| S18.19091; E045.19986, 290 m.a.o | 19.XII.2009 Water Net, Field# MAD09-74 | Leg: J. Bergsten, N. 
<normalizedToken id="960483D82BFAC909D821785A83329278" originalValue="Jönsson">Joensson</normalizedToken>
, T. | Ranarilalatiana, H.J. Randriamihaja //
</paragraph>
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<subSubSection id="ACD5CE45CEAB5C9E318EA123CDCD78D5" pageId="0" pageNumber="19" type="diagnosis">
<paragraph id="42D5AE5F433FE9E92DE051B6F5C2F39D" pageId="0" pageNumber="19">Diagnosis.</paragraph>
<paragraph id="F37D1ED3AD7432FFF1D08F18694D1668" pageId="0" pageNumber="19">
Similar to 
<taxonomicName id="9681130C6366B26138FF41FF857FF691" lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis id="BD3AB6B07164DDE41CC26A84ED78459B" italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
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and related species in habitus by being small, elongate, and subparalell, but 
<taxonomicName id="25D9A4F7406CC5B60D294FEB63D62345" lsidName="C. befasicus" pageId="0" pageNumber="19" rank="species" species="befasicus">
<emphasis id="B55CBCC6347B673E7B420138A0BF46AC" italics="true" pageId="0" pageNumber="19">C. befasicus</emphasis>
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is distinguished from 
<taxonomicName id="1FE92B68F46573A0034B9052AB4ADBD0" lsidName="C. insuetus" pageId="0" pageNumber="19" rank="species" species="insuetus">
<emphasis id="2094BA547351AE9631711CDE6F75EF42" italics="true" pageId="0" pageNumber="19">C. insuetus</emphasis>
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and from all other Malagasy 
<taxonomicName id="804E516C354B052F50FC3F5243F204AB" class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus" order="Coleoptera" pageId="0" pageNumber="19" phylum="Arthropoda" rank="genus">
<emphasis id="3F6598150B367032D69A13C2C4B4FD66" italics="true" pageId="0" pageNumber="19">Copelatus</emphasis>
</taxonomicName>
species by the presence of only five elytral striae, and without submarginal striae. In addition, the first stria is shortened, present only in the posterior third (
<figureCitation id="31962BAE972F265B1CAB3C752E6EF873" 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. 10D</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="A291ABA0057EA097821012109BDF58ED" pageId="0" pageNumber="19" type="description">
<paragraph id="237619F7EEAD87C2BE6CABFB51C0AA21" pageId="0" pageNumber="19">Description.</paragraph>
<paragraph id="235FC94E617DCF58E7D27F3DB5ACEE02" pageId="0" pageNumber="19">
Body length 4.1-4.2 mm. Body shape elongate and subparallel, dorsal surface reddish brown with a lighter elytral base. Head and pronotum uniformly rufus brown. Head, pronotum and elytra with dense punctation. Elytra and pronotum covered with dense punctures and the whole dorsal surface with a microsculpture. Lateral sides of pronotum striolate with the widest striolate area in the posterior corners. Elytra light brown with a distinct testaceous band basally (
<figureCitation id="9B44A1FE11D34D63F2862E44C0348EF3" 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. 10D</figureCitation>
). First elytral stria shortened and present only in posterior third. Second to fifth elytral stria starting more or less at base and all striae approaching the apex of elytron, but the second and fourth a little shorter. Submarginal striae absent. Appendages testaceous.
</paragraph>
<paragraph id="055EBBA3D5C274CAAD6F661846E9E3AC" pageId="0" pageNumber="19">Ventral side light brown. Metacoxa and abdominal ventrites punctate and striolate. Prosternal process rather short and medially raised, triangular in cross-section. Lateral parts of metaventrite medium broad. Metacoxal lines anteriorly diverging but rather weakly so, abbreviated well before metaventral margin. Antennae, palps and legs testaceous.</paragraph>
<paragraph id="B0DE742A8FEE654698A2148F0AC16097" pageId="0" pageNumber="19">Male: unknown.</paragraph>
</subSubSection>
<subSubSection id="26E92E524012CB7BFD1322C42E607FE2" pageId="0" pageNumber="19" type="distribution">
<paragraph id="E25CFA9CFB105EF3D2FB8751A078FB67" pageId="0" pageNumber="19">Distribution.</paragraph>
<paragraph id="67CD4429D81EAA8B56DA0C8D748E89C2" pageId="0" pageNumber="19">
Known only from two localities in the western part of Madagascar, the deciduous forest south of Befasy, Morondava, and at one locality between Morafenobe and Beravina village (
<figureCitation id="335F9BCFC01D9750AE3AC0BA6C1A0028" 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 id="D594F8B28EF31F69355BA9CA67519996" pageId="0" pageNumber="19" type="habitat">
<paragraph id="9F1B705B852D2DE28ACD3305CBC25364" pageId="0" pageNumber="19">Habitat and ecology.</paragraph>
<paragraph id="68E2FAD60F72109B633F7CC1769A91D3" pageId="0" pageNumber="19">
Paulian collected the species in 1956 in the western dry deciduous forest south of Befasy, Morondava. We rediscovered the species in 2009, when we found one female specimen of 
<taxonomicName id="D3BC807ADFBC697981D1E98A5225C11E" lsidName="C. befasicus" pageId="0" pageNumber="19" rank="species" species="befasicus">
<emphasis id="5D31705E7044F91DCD3B67FC76A9FBFE" italics="true" pageId="0" pageNumber="19">C. befasicus</emphasis>
</taxonomicName>
also in the western part but a bit further north than the type locality, along the road between Morafenobe and Beravina village at an altitude of 290 m. This locality consisted of dry savannah with mixed wood and grassland ecosystem after deforestation. The habitat consisted of muddy/sandy residual pools with some dead leaves in a temporary stream after the rainy season. The dry deciduous forest ecosystem in western Madagascar has suffered immensely from deforestation and very little of this habitat remains (
<bibRefCitation id="7EAA43B361832C515E27B993F8799465" DOI="https://doi.org/10.1046/j.1365-3008.2001.00201.x" author="Ganzhorn, JU" journalOrPublisher="Oryx" pageId="0" pageNumber="19" pagination="346 - 348" refId="B19" refString="Ganzhorn, JU, Lowry, PP, Schatz, GE, Sommer, A, 2001. The biodiversity of Madagascar: one of the world's hottest hotspots on its way out. . Oryx 35: 346 - 348" title="The biodiversity of Madagascar: one of the world's hottest hotspots on its way out." url="https://doi.org/10.1046/j.1365-3008.2001.00201.x" volume="35" year="2001">Ganzhorn et al. 2001</bibRefCitation>
; 
<bibRefCitation id="E41C55AC802ADD61C5C59C44E4F08760" DOI="https://doi.org/10.1017/S0030605309001756" author="Whitehurst, AS" journalOrPublisher="Oryx" pageId="0" pageNumber="19" pagination="275 - 283" refId="B75" refString="Whitehurst, AS, Sexton, JO, Dollar, L, 2009. Land cover change in western Madagascar's dry deciduous forests: a comparison of forest changes in and around Kirindy Mite National Park. . Oryx 43: 275 - 283" title="Land cover change in western Madagascar's dry deciduous forests: a comparison of forest changes in and around Kirindy Mite National Park." url="https://doi.org/10.1017/S0030605309001756" volume="43" year="2009">Whitehurst et al. 2009</bibRefCitation>
). The species is rare and likely threatened due to the disappearance of western deciduous forests in Madagascar. That all four known specimens are females may suggest an uneven sex ratio as a test of equal sex ratio is marginally non-significant (p = 0.0625) if considered randomly picked from the population.
</paragraph>
</subSubSection>
<subSubSection id="D7F2CA4540C19C930D959B278E49CB80" pageId="0" pageNumber="19" type="comments">
<paragraph id="BB90C2B89FDFEF1EC80D8666D3C94899" pageId="0" pageNumber="19">Comments.</paragraph>
<paragraph id="23F9F3D8FB7BA4B96D6B4D152FD12537" pageId="0" pageNumber="19">
This is the only species in the 
<taxonomicName id="D317F81430308FB2D2EE89BB65134227" class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus longicornis" order="Coleoptera" pageId="0" pageNumber="19" phylum="Arthropoda" rank="species" species="longicornis">
<emphasis id="E2116487D8626C74D8F2CD7B00747867" italics="true" pageId="0" pageNumber="19">Copelatus longicornis</emphasis>
</taxonomicName>
group from Madagascar. The 
<taxonomicName id="D8FB6A9C26E027F4B25D923477E7A12A" lsidName="longicornis" pageId="0" pageNumber="19" rank="species" species="longicornis">
<emphasis id="FBD759C2092C2DE4C9B78631965D4BA6" italics="true" pageId="0" pageNumber="19">longicornis</emphasis>
</taxonomicName>
species group currently contains 38 species distributed mainly in the Neotropical and Afrotropical regions but also with few species present in Japan, New Guinea, and Fiji Islands (
<bibRefCitation id="E4E87F0924092505D558DA7E9026BBFE" author="Nilsson, AN" journalOrPublisher="Tropical Zoology" pageId="0" pageNumber="19" publicationUrl="http://www.waterbeetles.eu" refId="B57" refString="Nilsson, AN, Hajek, J, 2018. . http://www.waterbeetles.eu" url="http://www.waterbeetles.eu" year="2018">
Nilsson and 
<normalizedToken id="C8D29D23014B68D5E96F0DE41DD49AC7" originalValue="Hájek">Hajek</normalizedToken>
2018
</bibRefCitation>
). The group as currently defined is certainly artificial from a phylogenetic perspective and the only character they have in common is the low number of elytral striae.
</paragraph>
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