treatments-xml/data/8D/D5/86/8DD58697A6F15D9EA211ED0F0F19FBB4.xml

<document ID-DOI="http://dx.doi.org/10.3897/zookeys.871.36337" ID-GBIF-Dataset="09f4db46-2943-4141-909b-8fe8401d88bf" ID-PMC="PMC6700060" ID-Pensoft-Pub="1313-2970-871-89" ID-Pensoft-UUID="256119825E9A58B5A3D98E60E176273E" ID-PubMed="31435188" ID-ZooBank="56C3FB7DAF714C1B8BD108417883D624" ModsDocID="1313-2970-871-89" checkinTime="1565637245384" checkinUser="pensoft" docAuthor="Ranarilalatiana, Tolotra &amp; Bergsten, Johannes" docDate="2019" docId="8DD58697A6F15D9EA211ED0F0F19FBB4" docLanguage="en" docName="ZooKeys 871: 89-118" docOrigin="ZooKeys 871" docSource="http://dx.doi.org/10.3897/zookeys.871.36337" docTitle="Copelatus amphibius Ranarilalatiana &amp; Bergsten, 2019, sp. nov." docType="treatment" docUuid="440DF910-AAFD-4430-BDF8-EE5E59B1907C" docUuidSource="ZooBank" docVersion="5" id="256119825E9A58B5A3D98E60E176273E" lastPageNumber="94" masterDocId="256119825E9A58B5A3D98E60E176273E" masterDocTitle="Discovery of a specialist Copelatinae fauna on Madagascar: highly ephemeral tropical forest floor depressions as an overlooked habitat for diving beetles (Coleoptera, Dytiscidae)" masterLastPageNumber="118" masterPageNumber="89" pageNumber="92" updateTime="1668167722462" updateUser="ExternalLinkService">
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<mods:title>Discovery of a specialist Copelatinae fauna on Madagascar: highly ephemeral tropical forest floor depressions as an overlooked habitat for diving beetles (Coleoptera, Dytiscidae)</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Ranarilalatiana, Tolotra</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Bergsten, Johannes</mods:namePart>
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<mods:title>ZooKeys</mods:title>
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<mods:date>2019</mods:date>
<mods:detail type="volume">
<mods:number>871</mods:number>
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<mods:start>89</mods:start>
<mods:end>118</mods:end>
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<mods:url>http://dx.doi.org/10.3897/zookeys.871.36337</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.871.36337</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-871-89</mods:identifier>
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<treatment ID-GBIF-Taxon="159120444" LSID="urn:lsid:zoobank.org:act:440DF910-AAFD-4430-BDF8-EE5E59B1907C" httpUri="http://treatment.plazi.org/id/8DD58697A6F15D9EA211ED0F0F19FBB4" lastPageId="5" lastPageNumber="94" pageId="3" pageNumber="92">
<subSubSection pageId="3" pageNumber="92" type="nomenclature">
<paragraph pageId="3" pageNumber="92">
<taxonomicName LSID="8dd58697-a6f1-5d9e-a211-ed0f0f19fbb4" class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus amphibius" order="Coleoptera" pageId="3" pageNumber="92" phylum="Arthropoda" rank="species" species="amphibius">Copelatus amphibius</taxonomicName>
<taxonomicNameLabel pageId="3" pageNumber="92">sp. nov.</taxonomicNameLabel>
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Habitus, dorsal view. A Copelatus amphibius sp. nov. (female) B Copelatus zanatanensis sp. nov. (female) C Copelatus betampona sp. nov. (female) D Madaglymbus semifactus sp. nov. (female) E Madaglymbus kelimaso sp. nov. (male) F Madaglymbus menalamba sp. nov. (female)." figureDoi="10.3897/zookeys.871.36337.figure1" httpUri="https://binary.pensoft.net/fig/325108" pageId="3" pageNumber="92">Figs 1A</figureCitation>
, 
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Male genitalia. From left to right aedeagus in right lateral, ventral, left lateral views, and left paramere. A Copelatus amphibius sp. nov. B Copelatus zanatanensis sp. nov. C Copelatus betampona sp. nov." figureDoi="10.3897/zookeys.871.36337.figure2" httpUri="https://binary.pensoft.net/fig/325109" pageId="3" pageNumber="92">2A</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="92" type="type locality">
<paragraph pageId="3" pageNumber="92">Type locality.</paragraph>
<paragraph pageId="3" pageNumber="92">Masoala National Park [15.6713S; 49.9672E] (Madagascar, Analanjirofo region, Maroantsetra)</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="92" type="type material">
<paragraph pageId="3" pageNumber="92">Type material.</paragraph>
<paragraph pageId="3" pageNumber="92">
<emphasis bold="true" italics="true" pageId="3" pageNumber="92">Holotype</emphasis>
♂ GP (NHRS): // NHRS-JLKB | 000066350 // Madagascar: Toamasina: Analanjirofo: | Masoala NP: lowalt. rainforest | MAD18-53: depression on forest floor | on path ~1.6 km NE of Andranobe camp | 
<geoCoordinate degrees="15.6713" direction="south" orientation="latitude" precision="5" value="-15.6713">15.6713S</geoCoordinate>
, 
<geoCoordinate degrees="49.9672" direction="east" orientation="longitude" precision="5" value="49.9672">49.9672E</geoCoordinate>
, 220 m, 18.II.2018 | Leg. J. Bergsten &amp; T. Ranarilalatiana // Holotype | 
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus amphibius" order="Coleoptera" pageId="3" pageNumber="92" phylum="Arthropoda" rank="species" species="amphibius">
<emphasis italics="true" pageId="3" pageNumber="92">Copelatus amphibius</emphasis>
</taxonomicName>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 //
</paragraph>
<paragraph pageId="3" pageNumber="92">
<emphasis bold="true" italics="true" pageId="3" pageNumber="92">Paratypes</emphasis>
: -7♂ GP, 6♀, 45 ex. (Alc.) (NHRS, DEUA &amp; PBZT/MBC): // NHRS-JLKB | 000011230-1, 65651-2, 66016, 66347-9, 66351-5, 11232(Alc.) // Madagascar: Toamasina: Analanjirofo: | Masoala NP: lowalt. rainforest | MAD18-53: depression on forest floor | on path ~1.6 km NE of Andranobe camp | 
<geoCoordinate degrees="15.6713" direction="south" orientation="latitude" precision="5" value="-15.6713">15.6713S</geoCoordinate>
, 
<geoCoordinate degrees="49.9672" direction="east" orientation="longitude" precision="5" value="49.9672">49.9672E</geoCoordinate>
, 220 m, 18.II.2018 | Leg. J. Bergsten &amp; T. Ranarilalatiana // Paratype | 
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus amphibius" order="Coleoptera" pageId="3" pageNumber="92" phylum="Arthropoda" rank="species" species="amphibius">
<emphasis italics="true" pageId="3" pageNumber="92">Copelatus amphibius</emphasis>
</taxonomicName>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 //
</paragraph>
<paragraph pageId="3" pageNumber="92">
-3♂ GP, 5♀, 42 ex. (Alc.) (NHRS, DEUA &amp; PBZT/MBC): // NHRS-JLKB | 000011234, 65649-50, 65788, 65790, 65795, 65435-6, 11233(Alc.) // Madagascar: Toamasina: Analanjirofo: | Nosy Mangabe, Masoala NP: MAD18-58 | rainfallpool with dead leaves nr path | after lighthouse, lowalt. rainforest | 
<geoCoordinate degrees="15.5079" direction="south" orientation="latitude" precision="5" value="-15.5079">15.5079S</geoCoordinate>
, 
<geoCoordinate degrees="49.7641" direction="east" orientation="longitude" precision="5" value="49.7641">49.7641E</geoCoordinate>
, 195 m, 19.II.2018 | Leg. J. Bergsten &amp; T. Ranarilalatiana // Paratype | 
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus amphibius" order="Coleoptera" pageId="3" pageNumber="92" phylum="Arthropoda" rank="species" species="amphibius">
<emphasis italics="true" pageId="3" pageNumber="92">Copelatus amphibius</emphasis>
</taxonomicName>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 //
</paragraph>
<paragraph pageId="3" pageNumber="92">
-1♂ GP, 2♀ (NHRS): // NHRS-JLKB | 000065642-4 // Madagascar: Antsiranana: Sava: | Marojejy NP: midalt. rainforest: | small stream above camp II on | trail towards Taktajania, MAD18-23 | 
<geoCoordinate degrees="14.4375" direction="south" orientation="latitude" precision="5" value="-14.4375">14.4375S</geoCoordinate>
, 
<geoCoordinate degrees="49.7612" direction="east" orientation="longitude" precision="5" value="49.7612">49.7612E</geoCoordinate>
, 860 m, 09.II.2018 | Leg. J. Bergsten &amp; T. Ranarilalatiana // Paratype | 
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus amphibius" order="Coleoptera" pageId="3" pageNumber="92" phylum="Arthropoda" rank="species" species="amphibius">
<emphasis italics="true" pageId="3" pageNumber="92">Copelatus amphibius</emphasis>
</taxonomicName>
sp. nov. | Det. Ranarilalatiana | &amp; Bergsten, 2019 //
</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="92" type="diagnosis">
<paragraph pageId="3" pageNumber="92">Diagnosis.</paragraph>
<paragraph pageId="3" pageNumber="92">
A small species with medially infuscated testaceous elytra and oblong-oval body shape. Penis in lateral view with low ventral 
<normalizedToken originalValue="”hump”">&quot;hump&quot;</normalizedToken>
, apical blade with acute apex and somewhat curved non-straight ventral margin, in ventral view apical blade is left-angled (
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Male genitalia. From left to right aedeagus in right lateral, ventral, left lateral views, and left paramere. A Copelatus amphibius sp. nov. B Copelatus zanatanensis sp. nov. C Copelatus betampona sp. nov." figureDoi="10.3897/zookeys.871.36337.figure2" httpUri="https://binary.pensoft.net/fig/325109" pageId="3" pageNumber="92">Fig. 2A</figureCitation>
). Significantly smaller than described species from Madagascar with similar type of genitalia such as 
<taxonomicName lsidName="C. owas" pageId="3" pageNumber="92" rank="species" species="owas">
<emphasis italics="true" pageId="3" pageNumber="92">C. owas</emphasis>
</taxonomicName>
<normalizedToken originalValue="Régimbart">Regimbart</normalizedToken>
, 1895 and 
<taxonomicName lsidName="C. acamas" pageId="3" pageNumber="92" rank="species" species="acamas">
<emphasis italics="true" pageId="3" pageNumber="92">C. acamas</emphasis>
</taxonomicName>
Guignot, 1955, and genitalia details also clearly different.
</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="92" type="description">
<paragraph pageId="3" pageNumber="92">Description.</paragraph>
<paragraph pageId="3" pageNumber="92">Body length: 4.6-5.4 mm (♀: 4.6-5 mm, ♂: 4.8-5.4 mm).</paragraph>
<paragraph pageId="3" pageNumber="92">
Body shape oval (
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Habitus, dorsal view. A Copelatus amphibius sp. nov. (female) B Copelatus zanatanensis sp. nov. (female) C Copelatus betampona sp. nov. (female) D Madaglymbus semifactus sp. nov. (female) E Madaglymbus kelimaso sp. nov. (male) F Madaglymbus menalamba sp. nov. (female)." figureDoi="10.3897/zookeys.871.36337.figure1" httpUri="https://binary.pensoft.net/fig/325108" pageId="3" pageNumber="92">Fig. 1A</figureCitation>
).
</paragraph>
<caption doi="10.3897/zookeys.871.36337.figure1" httpUri="https://binary.pensoft.net/fig/325108" pageId="3" pageNumber="92" start="Figure 1" startId="F1">
<paragraph pageId="3" pageNumber="92">
Figure 1. Habitus, dorsal view. 
<emphasis bold="true" pageId="3" pageNumber="92">A</emphasis>
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus amphibius" order="Coleoptera" pageId="3" pageNumber="92" phylum="Arthropoda" rank="species" species="amphibius">
<emphasis italics="true" pageId="3" pageNumber="92">Copelatus amphibius</emphasis>
</taxonomicName>
sp. nov. (female) 
<emphasis bold="true" pageId="3" pageNumber="92">B</emphasis>
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus zanatanensis" order="Coleoptera" pageId="3" pageNumber="92" phylum="Arthropoda" rank="species" species="zanatanensis">
<emphasis italics="true" pageId="3" pageNumber="92">Copelatus zanatanensis</emphasis>
</taxonomicName>
sp. nov. (female) 
<emphasis bold="true" pageId="3" pageNumber="92">C</emphasis>
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus betampona" order="Coleoptera" pageId="3" pageNumber="92" phylum="Arthropoda" rank="species" species="betampona">
<emphasis italics="true" pageId="3" pageNumber="92">Copelatus betampona</emphasis>
</taxonomicName>
sp. nov. (female) 
<emphasis bold="true" pageId="3" pageNumber="92">D</emphasis>
<taxonomicName class="Insecta" family="Dytiscidae" genus="Madaglymbus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Madaglymbus semifactus" order="Coleoptera" pageId="3" pageNumber="92" phylum="Arthropoda" rank="species" species="semifactus">
<emphasis italics="true" pageId="3" pageNumber="92">Madaglymbus semifactus</emphasis>
</taxonomicName>
sp. nov. (female) 
<emphasis bold="true" pageId="3" pageNumber="92">E</emphasis>
<taxonomicName class="Insecta" family="Dytiscidae" genus="Madaglymbus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Madaglymbus kelimaso" order="Coleoptera" pageId="3" pageNumber="92" phylum="Arthropoda" rank="species" species="kelimaso">
<emphasis italics="true" pageId="3" pageNumber="92">Madaglymbus kelimaso</emphasis>
</taxonomicName>
sp. nov. (male) 
<emphasis bold="true" pageId="3" pageNumber="92">F</emphasis>
<taxonomicName class="Insecta" family="Dytiscidae" genus="Madaglymbus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Madaglymbus menalamba" order="Coleoptera" pageId="3" pageNumber="92" phylum="Arthropoda" rank="species" species="menalamba">
<emphasis italics="true" pageId="3" pageNumber="92">Madaglymbus menalamba</emphasis>
</taxonomicName>
sp. nov. (female).
</paragraph>
</caption>
<paragraph pageId="3" pageNumber="92">
Head rufotestaceous with a rather weak v-shaped infuscation between eyes. Pronotum dark brown medially and testaceous laterally. Elytra testaceous brown, variably with darker infuscation medially especially along the striae (
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Habitus, dorsal view. A Copelatus amphibius sp. nov. (female) B Copelatus zanatanensis sp. nov. (female) C Copelatus betampona sp. nov. (female) D Madaglymbus semifactus sp. nov. (female) E Madaglymbus kelimaso sp. nov. (male) F Madaglymbus menalamba sp. nov. (female)." figureDoi="10.3897/zookeys.871.36337.figure1" httpUri="https://binary.pensoft.net/fig/325108" pageId="3" pageNumber="92">Fig. 1A</figureCitation>
). Basally, laterally and apically lighter testaceous. Appendages testaceous except metatarsus somewhat rufotestaceous.
</paragraph>
<paragraph pageId="3" pageNumber="92">Elytra with ten discal and one submarginal striae. Ninth striae avbreviated anteriorly. Submarginal striae present posteriorly only, starting at about middle. Posteriorly every second striae abbreviated (2nd, 4th, 6th, 8th, and 10th). Pronotum striolated laterally and basally. Lateral margin of pronotum with a narrow bead, not reaching anterior corner. Head, pronotum and elytra with same type of microreticulation and micropunctures.</paragraph>
<paragraph pageId="3" pageNumber="92">
Ventral side rufotestaceous except metacoxal plate infuscated brown. Abdominal sternites with vague testaceous spots laterally. Metacoxal plate with coarse strioles, abdominal sternites 
<normalizedToken originalValue="II–IV">II-IV</normalizedToken>
with finer strioles. Metacoxal lines anteriorly diverging and ending well before metaventral suture. Prosternal process lanceolate, short, and anterior metaventral process rather broad.
</paragraph>
<paragraph pageId="3" pageNumber="92">
Male protibia modified, angled at base and expanding distally. Pro and mesotarsal segments 
<normalizedToken originalValue="I–III">I-III</normalizedToken>
dilated and ventrally equipped with adhesive discs (constellation I:3 (row 1), 4 (row 2), II:4, III:4). Longer metatibial spur apically slightly more curved than in female.
</paragraph>
<paragraph pageId="3" pageNumber="92">
Male genitalia as in 
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Male genitalia. From left to right aedeagus in right lateral, ventral, left lateral views, and left paramere. A Copelatus amphibius sp. nov. B Copelatus zanatanensis sp. nov. C Copelatus betampona sp. nov." figureDoi="10.3897/zookeys.871.36337.figure2" httpUri="https://binary.pensoft.net/fig/325109" pageId="3" pageNumber="92">Figure 2A</figureCitation>
. Penis curved and robust in lateral view with a comparatively low ventral hump and sinuate before the apical blade. Apical blade left-turned in ventral view. Right lateral side with strong rugosity or transverse ridges apically. Parameres as in 
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Male genitalia. From left to right aedeagus in right lateral, ventral, left lateral views, and left paramere. A Copelatus amphibius sp. nov. B Copelatus zanatanensis sp. nov. C Copelatus betampona sp. nov." figureDoi="10.3897/zookeys.871.36337.figure2" httpUri="https://binary.pensoft.net/fig/325109" pageId="3" pageNumber="92">Figure 2A</figureCitation>
.
</paragraph>
<caption doi="10.3897/zookeys.871.36337.figure2" httpUri="https://binary.pensoft.net/fig/325109" pageId="3" pageNumber="92" start="Figure 2" startId="F2">
<paragraph pageId="3" pageNumber="92">
Figure 2. Male genitalia. From left to right aedeagus in right lateral, ventral, left lateral views, and left paramere. 
<emphasis bold="true" pageId="3" pageNumber="92">A</emphasis>
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus amphibius" order="Coleoptera" pageId="3" pageNumber="92" phylum="Arthropoda" rank="species" species="amphibius">
<emphasis italics="true" pageId="3" pageNumber="92">Copelatus amphibius</emphasis>
</taxonomicName>
sp. nov. 
<emphasis bold="true" pageId="3" pageNumber="92">B</emphasis>
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus zanatanensis" order="Coleoptera" pageId="3" pageNumber="92" phylum="Arthropoda" rank="species" species="zanatanensis">
<emphasis italics="true" pageId="3" pageNumber="92">Copelatus zanatanensis</emphasis>
</taxonomicName>
sp. nov. 
<emphasis bold="true" pageId="3" pageNumber="92">C</emphasis>
<taxonomicName class="Insecta" family="Dytiscidae" genus="Copelatus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copelatus betampona" order="Coleoptera" pageId="3" pageNumber="92" phylum="Arthropoda" rank="species" species="betampona">
<emphasis italics="true" pageId="3" pageNumber="92">Copelatus betampona</emphasis>
</taxonomicName>
sp. nov.
</paragraph>
</caption>
<paragraph pageId="3" pageNumber="92">Dorsal structures of females not significantly different from male, but body size on average smaller.</paragraph>
</subSubSection>
<subSubSection pageId="4" pageNumber="93" type="distribution">
<paragraph pageId="4" pageNumber="93">
<pageBreakToken pageId="4" pageNumber="93" start="start">Distribution</pageBreakToken>
.
</paragraph>
<paragraph pageId="4" pageNumber="93">
The northeastern humid forest from Marojejy NP to Masoala NP including the island of Nosy Mangabe (
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Maps of Madagascar with species distributions and administrative divisions. A Copelatus amphibius sp. nov. (circle), Copelatus betampona sp. nov. (square) B Copelatus zanatanensis sp. nov. (circle), Madaglymbus semifactus sp. nov. (square) C Madaglymbus kelimaso sp. nov. (circle), Madaglymbus menalamba sp. nov. (square) D current 22 regions and six former provinces of Madagascar." figureDoi="10.3897/zookeys.871.36337.figure4" httpUri="https://binary.pensoft.net/fig/325111" pageId="4" pageNumber="93">Fig. 4A</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection lastPageId="5" lastPageNumber="94" pageId="4" pageNumber="93" type="ecology">
<paragraph pageId="4" pageNumber="93">Ecology and conservation.</paragraph>
<paragraph lastPageId="5" lastPageNumber="94" pageId="4" pageNumber="93">
<taxonomicName lsidName="C. amphibius" pageId="4" pageNumber="93" rank="species" species="amphibius">
<emphasis italics="true" pageId="4" pageNumber="93">C. amphibius</emphasis>
</taxonomicName>
sp. nov. was found in primary humid forests in dry shallow forest floor depression with dead leaves and soil at Masoala and in 
<pageBreakToken pageId="5" pageNumber="94" start="start">a</pageBreakToken>
rainwater-filled pool on Nosy Mangabe (
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Habitat photo of locality MAD 18 - 58, Nosy Mangabe, where the new species Copelatus amphibius sp. nov. and Madaglymbus menalamba sp. nov. were found. Of five different localities recorded for M. menalamba sp. nov. this was the only one where the species was found in water." figureDoi="10.3897/zookeys.871.36337.figure8" httpUri="https://binary.pensoft.net/fig/325115" pageId="5" pageNumber="94">Fig. 8</figureCitation>
). Both localities were at low altitudes but we also collected the species from Marojejy NP, in residual pools of a small stream at mid-altitude (860 m). Marojejy NP is unique in having continuous humid forest stretching from lowaltitude to the alpine zone and hence it is not surprising that the species can reach into the lower mid-altitude zone here. Both Marojejy and Masoala are since 2007 part of the UNESCO World Heritage Site Rainforests of the Atsinanana. Unfortunately, since 2010 Atsinanana is also on the list of World Heritage in Danger following a surge in illegal logging and hunting threatening its outstanding biodiversity values.
</paragraph>
</subSubSection>
<subSubSection pageId="5" pageNumber="94" type="etymology">
<paragraph pageId="5" pageNumber="94">Etymology.</paragraph>
<paragraph pageId="5" pageNumber="94">
The Latin adjective amphibius comes from the ancient Greek word 
<normalizedToken originalValue="“amphibios”">&quot;amphibios&quot;</normalizedToken>
and means capable of living both in water and on land.
</paragraph>
</subSubSection>
</treatment>
</document>