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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.994.57012" ID-GBIF-Dataset="8345955a-c4e9-4b46-9252-2219a5257656" ID-PMC="PMC7686220" ID-Pensoft-Pub="1313-2970-994-149" ID-Pensoft-UUID="D35BA35FB0BB5A1FB90EF99300F210A9" ID-PubMed="33273885" ID-ZooBank="D54ACBDAEB1944B3A8386274DD8BDC03" ModsDocID="1313-2970-994-149" checkinTime="1605693956575" checkinUser="pensoft" docAuthor="Vieites, David R., Nieto-Roman, Sandra, Fernandez, Marcos Peso &amp; Santos-Santos, Javier H." docDate="2020" docId="91B4828C26115107BE98F2D0ED179A84" docLanguage="en" docName="ZooKeys 994: 149-166" docOrigin="ZooKeys 994" docSource="http://dx.doi.org/10.3897/zookeys.994.57012" docTitle="Blommersia nataliae Vieites &amp; Nieto-Román &amp; Fernández &amp; Santos-Santos 2020, sp. nov." docType="treatment" docUuid="8D15676C-402C-417E-B185-5551C356CA44" docUuidSource="ZooBank" docVersion="4" id="D35BA35FB0BB5A1FB90EF99300F210A9" lastPageNumber="149" masterDocId="D35BA35FB0BB5A1FB90EF99300F210A9" masterDocTitle="Hidden in plain sight: a new frog species of the genus Blommersia from the oceanic island of Mayotte, Comoros archipelago" masterLastPageNumber="166" masterPageNumber="149" pageNumber="149" updateTime="1668169923665" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Hidden in plain sight: a new frog species of the genus Blommersia from the oceanic island of Mayotte, Comoros archipelago</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Vieites, David R.</mods:namePart>
<mods:affiliation>Integrative Biogeography and Global Change Group, Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Cientificas (MNCN-CSIC), 28006 Madrid, Spain</mods:affiliation>
<mods:nameIdentifier type="orcid">https://orcid.org/0000-0001-5551-7419</mods:nameIdentifier>
<mods:nameIdentifier type="email">vieites@mncn.csic.es</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Nieto-Roman, Sandra</mods:namePart>
<mods:affiliation>Integrative Biogeography and Global Change Group, Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Cientificas (MNCN-CSIC), 28006 Madrid, Spain</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Fernandez, Marcos Peso</mods:namePart>
<mods:affiliation>Integrative Biogeography and Global Change Group, Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Cientificas (MNCN-CSIC), 28006 Madrid, Spain &amp; Department of Animal Biology, University of Barcelona, Avenida Diagonal 645, 08028, Barcelona, Spain</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Santos-Santos, Javier H.</mods:namePart>
<mods:affiliation>Integrative Biogeography and Global Change Group, Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Cientificas (MNCN-CSIC), 28006 Madrid, Spain &amp; Science and Business Ltd. Calle Peru, 6, 28290 Las Rozas de Madrid, Madrid, Spain</mods:affiliation>
<mods:nameIdentifier type="orcid">https://orcid.org/0000-0002-4622-9682</mods:nameIdentifier>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2020</mods:date>
<mods:detail type="volume">
<mods:number>994</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>149</mods:start>
<mods:end>166</mods:end>
</mods:extent>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.994.57012</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.994.57012</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-994-149</mods:identifier>
<mods:identifier type="ZooBank">D54ACBDAEB1944B3A8386274DD8BDC03</mods:identifier>
<mods:identifier type="Pensoft-UUID">D35BA35FB0BB5A1FB90EF99300F210A9</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="169094412" LSID="urn:lsid:zoobank.org:act:8D15676C-402C-417E-B185-5551C356CA44" httpUri="http://treatment.plazi.org/id/91B4828C26115107BE98F2D0ED179A84" lastPageNumber="149" pageId="0" pageNumber="149">
<subSubSection pageId="0" pageNumber="149" type="nomenclature">
<paragraph pageId="0" pageNumber="149">
<taxonomicName LSID="91B4828C-2611-5107-BE98-F2D0ED179A84" authority="Vieites &amp; Nieto-Román &amp; Fernández &amp; Santos-Santos, 2020" authorityName="Vieites &amp; Nieto-Román &amp; Fernández &amp; Santos-Santos" authorityYear="2020" class="Amphibia" family="Mantellidae" genus="Blommersia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blommersia nataliae" order="Anura" pageId="0" pageNumber="149" phylum="Chordata" rank="species" species="nataliae" status="sp. nov.">Blommersia nataliae</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="149">sp. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="149" type="holotype">
<paragraph pageId="0" pageNumber="149">Holotype.</paragraph>
<paragraph pageId="0" pageNumber="149">
An adult male, left thigh muscle removed for genetic analyses. Original field number: &quot;DRV6867, David R. Vieites collection&quot;. Museo Nacional de Ciencias Naturales catalog number: MNCN50456. Collected in a degraded forest with giant bamboo, in forest leaf litter at the Mont
<normalizedToken originalValue="MSapere">M'Sapere</normalizedToken>
, island of Mayotte (French Overseas Department), Comoros archipelago, -
<geoCoordinate degrees="12.7656" direction="south" orientation="latitude" precision="5" value="-12.7656">12.7656°S</geoCoordinate>
,
<geoCoordinate degrees="45.1852" direction="east" orientation="longitude" precision="5" value="45.1852">45.1852°E</geoCoordinate>
500 m a.s.l. the 25th November 2012 by D. Vieites and M. Peso
<normalizedToken originalValue="Fernández">Fernandez</normalizedToken>
.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="149" type="paratypes">
<paragraph pageId="0" pageNumber="149">Paratypes.</paragraph>
<paragraph pageId="0" pageNumber="149">
Females DRV6808 (MNCN50451), DRV6854 (MNCN50447), DRV6855 (MNCN50448), DRV6868 (MNCN50450), DRV6869 (MNCN50449); males DRV6857 (MNCN50453), DRV6859 (MNCN50455), DRV6860 (MNCN50458), DRV6861 (MNCN50454), DRV6862 (MNCN50452), DRV6863 (MNCN50457), collected at the type locality at the Mont
<normalizedToken originalValue="MSapere">M'Sapere</normalizedToken>
in 2012 by D. Vieites and M. Peso
<normalizedToken originalValue="Fernández">Fernandez</normalizedToken>
.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="149" type="etymology">
<paragraph pageId="0" pageNumber="149">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="149">Noun in the genitive case. D. Vieites and S. Nieto dedicate this species to their daughter Natalia Vieites Nieto, who has a birthmark resembling the beautiful conspicuous round moon-like brown spot characteristic of the species.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="149" type="diagnosis">
<paragraph pageId="0" pageNumber="149">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="149">
Assigned to the genus
<taxonomicName authorityName="Dubois" authorityYear="1992" class="Amphibia" family="Mantellidae" genus="Blommersia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blommersia" order="Anura" pageId="0" pageNumber="149" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="149">Blommersia</emphasis>
</taxonomicName>
in the family
<taxonomicName authorityName="Laurent" authorityYear="1946" family="Mantellidae" lsidName="" pageId="0" pageNumber="149" rank="family">Mantellidae</taxonomicName>
and subfamily
<taxonomicName authorityName="Laurent" authorityYear="1946" lsidName="" pageId="0" pageNumber="149" rank="subFamily" subFamily="Mantellinae">Mantellinae</taxonomicName>
by a combination of (1) presence of femoral glands and absence of nuptial pads in males, (2) presence of intercalary elements between ultimate and penultimate phalanges of fingers and toes (verified by external examination and microCT scanning), (3) presence of a single subgular vocal sac in males, (4) small size (adult SVL &lt;30 mm), and (5) molecular data.
</paragraph>
<paragraph pageId="0" pageNumber="149">
Within the genus
<taxonomicName authorityName="Dubois" authorityYear="1992" class="Amphibia" family="Mantellidae" genus="Blommersia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blommersia" order="Anura" pageId="0" pageNumber="149" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="149">Blommersia</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="B. nataliae" pageId="0" pageNumber="149" rank="species" species="nataliae">
<emphasis italics="true" pageId="0" pageNumber="149">B. nataliae</emphasis>
</taxonomicName>
sp. nov. is characterized by the following unique suite of morphological characters: (1) small adult body size (SVL 18-23 mm), (2) round femoral glands that are distantly separated in males, (3) inconspicuous vomerine teeth, (4) ovoid tongue, (5) tibiotarsal articulation reaching between the eye and the nostril when adpressed along the body. Furthermore, the new species is differentiated from all other species of
<taxonomicName authorityName="Dubois" authorityYear="1992" class="Amphibia" family="Mantellidae" genus="Blommersia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blommersia" order="Anura" pageId="0" pageNumber="149" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="149">Blommersia</emphasis>
</taxonomicName>
by a significant molecular genetic differentiation (≥ 4.3% uncorrected pairwise-distance in 16S).
</paragraph>
<paragraph pageId="0" pageNumber="149">
<taxonomicName authorityName="Vieites &amp; Nieto-Román &amp; Fernández &amp; Santos-Santos" authorityYear="2020" class="Amphibia" family="Mantellidae" genus="Blommersia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blommersia nataliae" order="Anura" pageId="0" pageNumber="149" phylum="Chordata" rank="species" species="nataliae">
<emphasis italics="true" pageId="0" pageNumber="149">Blommersia nataliae</emphasis>
</taxonomicName>
sp. nov. can be distinguished from all other described
<taxonomicName authorityName="Dubois" authorityYear="1992" class="Amphibia" family="Mantellidae" genus="Blommersia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blommersia" order="Anura" pageId="0" pageNumber="149" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="149">Blommersia</emphasis>
</taxonomicName>
species except
<taxonomicName lsidName="B. wittei" pageId="0" pageNumber="149" rank="species" species="wittei">
<emphasis italics="true" pageId="0" pageNumber="149">B. wittei</emphasis>
</taxonomicName>
and
<taxonomicName lsidName="B. transmarina" pageId="0" pageNumber="149" rank="species" species="transmarina">
<emphasis italics="true" pageId="0" pageNumber="149">B. transmarina</emphasis>
</taxonomicName>
by the presence of vomerine teeth (vs. absence) and having separated metatarsalia (vs. unseparated).
</paragraph>
<caption doi="10.3897/zookeys.994.57012.figure2" httpUri="https://binary.pensoft.net/fig/476542" pageId="0" pageNumber="149" start="Figure 2" startId="F2">
<paragraph pageId="0" pageNumber="149">
<emphasis bold="true" pageId="0" pageNumber="149">Figure 2.</emphasis>
Dorsolateral and ventral views of the holotype of
<taxonomicName authorityName="Vieites &amp; Nieto-Román &amp; Fernández &amp; Santos-Santos" authorityYear="2020" class="Amphibia" family="Mantellidae" genus="Blommersia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blommersia nataliae" order="Anura" pageId="0" pageNumber="149" phylum="Chordata" rank="species" species="nataliae">
<emphasis italics="true" pageId="0" pageNumber="149">Blommersia nataliae</emphasis>
</taxonomicName>
sp. nov. DRV6867 (MNCN50456). Note the color and shape of the femoral glands which are a diagnostic character to distinguish it from its sister taxon
<taxonomicName lsidName="B. transmarina" pageId="0" pageNumber="149" rank="species" species="transmarina">
<emphasis italics="true" pageId="0" pageNumber="149">B. transmarina</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="149">
<taxonomicName authorityName="Vieites &amp; Nieto-Román &amp; Fernández &amp; Santos-Santos" authorityYear="2020" class="Amphibia" family="Mantellidae" genus="Blommersia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blommersia nataliae" order="Anura" pageId="0" pageNumber="149" phylum="Chordata" rank="species" species="nataliae">
<emphasis italics="true" pageId="0" pageNumber="149">Blommersia nataliae</emphasis>
</taxonomicName>
sp. nov. can be distinguished from its syntopic sister taxon
<taxonomicName lsidName="B. transmarina" pageId="0" pageNumber="149" rank="species" species="transmarina">
<emphasis italics="true" pageId="0" pageNumber="149">B. transmarina</emphasis>
</taxonomicName>
by its rounded, distantly separated femoral glands (versus oblong, less separated glands; Figs
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Dorsolateral and ventral views of the holotype of Blommersia nataliae sp. nov. DRV 6867 (MNCN 50456). Note the color and shape of the femoral glands which are a diagnostic character to distinguish it from its sister taxon B. transmarina." figureDoi="10.3897/zookeys.994.57012.figure2" httpUri="https://binary.pensoft.net/fig/476542" pageId="0" pageNumber="149">2</figureCitation>
-
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Dorsolateral and ventral views of Blommersia transmarina DRV 6807 (MNCN 50446), adult male collected at Mont M'Sapere in 2012 by D. Vieites and M. Peso Fernandez. Note the shape and color of the femoral glands." figureDoi="10.3897/zookeys.994.57012.figure3" httpUri="https://binary.pensoft.net/fig/476543" pageId="0" pageNumber="149">3</figureCitation>
). Fig.
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Scatterplot of relative femoral gland length (FGL / SVL ratio) and the relative distance between the inner edges of the femoral glands (FGD / SVL) in the two new species from Mayotte and their sister taxon Blommersia wittei from Madagascar. Measurements are based on Table 1; and for B. wittei on Vences et al. (2010) and Pabijan et al. (2011). Note the intermediate position of B. wittei between B. transmarina and B. nataliae sp. nov." figureDoi="10.3897/zookeys.994.57012.figure4" httpUri="https://binary.pensoft.net/fig/476544" pageId="0" pageNumber="149">4</figureCitation>
represents the relative femoral gland length versus the relative distance between the femoral
<normalizedToken originalValue="glands">glands'</normalizedToken>
inner edges for both species and their sister taxon from Madagascar,
<taxonomicName lsidName="B. wittei" pageId="0" pageNumber="149" rank="species" species="wittei">
<emphasis italics="true" pageId="0" pageNumber="149">B. wittei</emphasis>
</taxonomicName>
.
<taxonomicName lsidName="B. wittei" pageId="0" pageNumber="149" rank="species" species="wittei">
<emphasis italics="true" pageId="0" pageNumber="149">B. wittei</emphasis>
</taxonomicName>
has an intermediate position between
<taxonomicName lsidName="B. nataliae" pageId="0" pageNumber="149" rank="species" species="nataliae">
<emphasis italics="true" pageId="0" pageNumber="149">B. nataliae</emphasis>
</taxonomicName>
sp. nov. and
<taxonomicName lsidName="B. transmarina" pageId="0" pageNumber="149" rank="species" species="transmarina">
<emphasis italics="true" pageId="0" pageNumber="149">B. transmarina</emphasis>
</taxonomicName>
; where
<taxonomicName lsidName="B. nataliae" pageId="0" pageNumber="149" rank="species" species="nataliae">
<emphasis italics="true" pageId="0" pageNumber="149">B. nataliae</emphasis>
</taxonomicName>
sp. nov. presents rounded and shorter glands that are ca. two times more separated between each other than in
<taxonomicName lsidName="B. transmarina" pageId="0" pageNumber="149" rank="species" species="transmarina">
<emphasis italics="true" pageId="0" pageNumber="149">B. transmarina</emphasis>
</taxonomicName>
(median FGD 2.6
<normalizedToken originalValue="±">+/-</normalizedToken>
0.2 mm vs. 1.3
<normalizedToken originalValue="±">+/-</normalizedToken>
0.3 mm, respectively) and ca. half shorter (median FGL 2.6
<normalizedToken originalValue="±">+/-</normalizedToken>
0.4 mm vs. 4.8
<normalizedToken originalValue="±">+/-</normalizedToken>
0.5 mm) (see also Figs
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Dorsolateral and ventral views of the holotype of Blommersia nataliae sp. nov. DRV 6867 (MNCN 50456). Note the color and shape of the femoral glands which are a diagnostic character to distinguish it from its sister taxon B. transmarina." figureDoi="10.3897/zookeys.994.57012.figure2" httpUri="https://binary.pensoft.net/fig/476542" pageId="0" pageNumber="149">2</figureCitation>
-
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Dorsolateral and ventral views of Blommersia transmarina DRV 6807 (MNCN 50446), adult male collected at Mont M'Sapere in 2012 by D. Vieites and M. Peso Fernandez. Note the shape and color of the femoral glands." figureDoi="10.3897/zookeys.994.57012.figure3" httpUri="https://binary.pensoft.net/fig/476543" pageId="0" pageNumber="149">3</figureCitation>
).
<taxonomicName authorityName="Vieites &amp; Nieto-Román &amp; Fernández &amp; Santos-Santos" authorityYear="2020" class="Amphibia" family="Mantellidae" genus="Blommersia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blommersia nataliae" order="Anura" pageId="0" pageNumber="149" phylum="Chordata" rank="species" species="nataliae">
<emphasis italics="true" pageId="0" pageNumber="149">Blommersia nataliae</emphasis>
</taxonomicName>
sp. nov. also differs from
<taxonomicName lsidName="B. transmarina" pageId="0" pageNumber="149" rank="species" species="transmarina">
<emphasis italics="true" pageId="0" pageNumber="149">B. transmarina</emphasis>
</taxonomicName>
in having inconspicuous vomerine teeth versus well developed and showing a V shape, ovoid tongue (vs. bifid), shorter hindlimbs with tibiotarsal articulation reaching between the eye and the nostril (vs. surpassing well the snout when appressed along the body), less distinct inner metatarsal tubercle, webbing formula [1(1), 2i(1.75), 2e(1), 3i(2.5), 3e(2), 4i/e(3), 5(1.5) versus 1(1), 2i(1-1.5), 2e(0.5), 3i(1.5), 3e(1), 4i(2-2.5), 4e(1.5-2), 5(0.5)], and by showing a brown facial mask from the snout, under the loreal region, to the tympanum, and by the presence of (usually) one very conspicuous moon-like spot on the back of each flank, close to the pelvic region and the hindlimbs. From
<taxonomicName lsidName="B. wittei" pageId="0" pageNumber="149" rank="species" species="wittei">
<emphasis italics="true" pageId="0" pageNumber="149">B. wittei</emphasis>
</taxonomicName>
, it differs in femoral gland dimensions and position (see Fig.
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Scatterplot of relative femoral gland length (FGL / SVL ratio) and the relative distance between the inner edges of the femoral glands (FGD / SVL) in the two new species from Mayotte and their sister taxon Blommersia wittei from Madagascar. Measurements are based on Table 1; and for B. wittei on Vences et al. (2010) and Pabijan et al. (2011). Note the intermediate position of B. wittei between B. transmarina and B. nataliae sp. nov." figureDoi="10.3897/zookeys.994.57012.figure4" httpUri="https://binary.pensoft.net/fig/476544" pageId="0" pageNumber="149">4</figureCitation>
), ovoid tongue (vs. bifid), slightly longer hindlimbs with tibiotarsal articulation reaching between the eye and the nostril (vs. reaching the anterior corner of the eye), less distinct inner metatarsal tubercle, webbing formula, and in coloration.
<taxonomicName baseAuthorityName="Guibé" baseAuthorityYear="1974" class="Amphibia" family="Mantellidae" genus="Blommersia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blommersia wittei" order="Anura" pageId="0" pageNumber="149" phylum="Chordata" rank="species" species="wittei">
<emphasis italics="true" pageId="0" pageNumber="149">Blommersia wittei</emphasis>
</taxonomicName>
has a proportionally smaller tympanum than both
<taxonomicName lsidName="B. nataliae" pageId="0" pageNumber="149" rank="species" species="nataliae">
<emphasis italics="true" pageId="0" pageNumber="149">B. nataliae</emphasis>
</taxonomicName>
sp. nov. and
<taxonomicName lsidName="B. transmarina" pageId="0" pageNumber="149" rank="species" species="transmarina">
<emphasis italics="true" pageId="0" pageNumber="149">B. transmarina</emphasis>
</taxonomicName>
(mean ratio TD/SVL 0.068
<normalizedToken originalValue="±">+/-</normalizedToken>
0.007, vs. 0.081
<normalizedToken originalValue="±">+/-</normalizedToken>
0.009 in
<taxonomicName lsidName="B. transmarina" pageId="0" pageNumber="149" rank="species" species="transmarina">
<emphasis italics="true" pageId="0" pageNumber="149">B. transmarina</emphasis>
</taxonomicName>
and 0.080
<normalizedToken originalValue="±">+/-</normalizedToken>
0.010 in
<taxonomicName lsidName="B. nataliae" pageId="0" pageNumber="149" rank="species" species="nataliae">
<emphasis italics="true" pageId="0" pageNumber="149">B. nataliae</emphasis>
</taxonomicName>
sp. nov.).
</paragraph>
<caption doi="10.3897/zookeys.994.57012.figure3" httpUri="https://binary.pensoft.net/fig/476543" pageId="0" pageNumber="149" start="Figure 3" startId="F3">
<paragraph pageId="0" pageNumber="149">
<emphasis bold="true" pageId="0" pageNumber="149">Figure 3.</emphasis>
Dorsolateral and ventral views of
<taxonomicName authorityName="Glaw, Hawlitschek, Glaw &amp; Vences" authorityYear="2019" class="Amphibia" family="Mantellidae" genus="Blommersia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blommersia transmarina" order="Anura" pageId="0" pageNumber="149" phylum="Chordata" rank="species" species="transmarina">
<emphasis italics="true" pageId="0" pageNumber="149">Blommersia transmarina</emphasis>
</taxonomicName>
DRV6807 (MNCN50446), adult male collected at Mont
<normalizedToken originalValue="MSapere">M'Sapere</normalizedToken>
in 2012 by D. Vieites and M. Peso
<normalizedToken originalValue="Fernández">Fernandez</normalizedToken>
. Note the shape and color of the femoral glands.
</paragraph>
</caption>
<caption doi="10.3897/zookeys.994.57012.figure4" httpUri="https://binary.pensoft.net/fig/476544" pageId="0" pageNumber="149" start="Figure 4" startId="F4">
<paragraph pageId="0" pageNumber="149">
<emphasis bold="true" pageId="0" pageNumber="149">Figure 4.</emphasis>
Scatterplot of relative femoral gland length (FGL/SVL ratio) and the relative distance between the inner edges of the femoral glands (FGD/SVL) in the two new species from Mayotte and their sister taxon
<taxonomicName baseAuthorityName="Guibé" baseAuthorityYear="1974" class="Amphibia" family="Mantellidae" genus="Blommersia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blommersia wittei" order="Anura" pageId="0" pageNumber="149" phylum="Chordata" rank="species" species="wittei">
<emphasis italics="true" pageId="0" pageNumber="149">Blommersia wittei</emphasis>
</taxonomicName>
from Madagascar. Measurements are based on Table
<tableCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Measurements of Blommersia nataliae sp. nov. and B. transmarina for comparison. See methods for abbreviations. The holotype of B. nataliae sp. nov. is shown in bold letters." httpUri="http://table.plazi.org/id/AE037202BE724C1E12E2F65537319F22" pageId="0" pageNumber="149" tableUuid="AE037202BE724C1E12E2F65537319F22">1</tableCitation>
; and for
<taxonomicName lsidName="B. wittei" pageId="0" pageNumber="149" rank="species" species="wittei">
<emphasis italics="true" pageId="0" pageNumber="149">B. wittei</emphasis>
</taxonomicName>
on
<bibRefCitation DOI="https://doi.org/10.1080/21564574.2010.512961" author="Vences, M" journalOrPublisher="African Journal of Herpetology" pageId="0" pageNumber="149" pagination="133 - 156" refId="B20" refString="Vences, M, Koehler, J, Pabijan, M, Glaw, F, 2010. Two syntopic and microendemic new frogs of the genus Blommersia from the East coast of Madagascar. African Journal of Herpetology 59: 133 - 156, DOI: https://doi.org/10.1080/21564574.2010.512961" title="Two syntopic and microendemic new frogs of the genus Blommersia from the East coast of Madagascar." url="https://doi.org/10.1080/21564574.2010.512961" volume="59" year="2010">Vences et al. (2010)</bibRefCitation>
and
<bibRefCitation DOI="https://doi.org/10.11646/zootaxa.2978.1.2" author="Pabijan, M" journalOrPublisher="Zootaxa" pageId="0" pageNumber="149" pagination="34 - 50" refId="B14" refString="Pabijan, M, Gehring, P-S, Koehler, J, Glaw, F, Vences, M, 2011. A new microendemic frog species of the genus Blommersia (Anura: Mantellidae) from the East coast of Madagascar. Zootaxa 2978: 34 - 50, DOI: https://doi.org/10.11646/zootaxa.2978.1.2" title="A new microendemic frog species of the genus Blommersia (Anura: Mantellidae) from the East coast of Madagascar." url="https://doi.org/10.11646/zootaxa.2978.1.2" volume="2978" year="2011">Pabijan et al. (2011)</bibRefCitation>
. Note the intermediate position of
<taxonomicName lsidName="B. wittei" pageId="0" pageNumber="149" rank="species" species="wittei">
<emphasis italics="true" pageId="0" pageNumber="149">B. wittei</emphasis>
</taxonomicName>
between
<taxonomicName lsidName="B. transmarina" pageId="0" pageNumber="149" rank="species" species="transmarina">
<emphasis italics="true" pageId="0" pageNumber="149">B. transmarina</emphasis>
</taxonomicName>
and
<taxonomicName lsidName="B. nataliae" pageId="0" pageNumber="149" rank="species" species="nataliae">
<emphasis italics="true" pageId="0" pageNumber="149">B. nataliae</emphasis>
</taxonomicName>
sp. nov.
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="149">
<emphasis bold="true" pageId="0" pageNumber="149">
Description of the holotype (Fig.
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Dorsolateral and ventral views of the holotype of Blommersia nataliae sp. nov. DRV 6867 (MNCN 50456). Note the color and shape of the femoral glands which are a diagnostic character to distinguish it from its sister taxon B. transmarina." figureDoi="10.3897/zookeys.994.57012.figure2" httpUri="https://binary.pensoft.net/fig/476542" pageId="0" pageNumber="149">2</figureCitation>
).
</emphasis>
Male specimen in good state of preservation. Part of the left thigh taken for genetic analyses. SVL = 18.6 mm. The body is slender; the head is slightly longer than wide but not wider than the body. Snout slightly pointed and rounded in lateral views with protuberant nostrils directed laterally, nearer to the tip of snout than to eye; canthus rostralis indistinct and straight; loreal region straight; tympanum distinct and rounded, with a diameter of 60% of the eye diameter; supratympanic fold present and slightly distinct behind the tympanum, but indistinct in its anterior part between the eye and the tympanum; tongue slender and ovoid, slightly notched posteriorly but not bifid; vomerine teeth present but very inconspicuous and very small, hard to see, and not grouped; maxillary teeth rudimentary; choanae rounded. The arms are slender with distinct, single subarticular tubercles, the inner and outer metacarpal tubercles distinct, the fingers without webbing, and the relative length of the fingers is 1&lt;2&lt;4&lt;3; terminal finger discs are enlarged and nuptial pads absent. Hindlimbs are relatively robust; the tibiotarsal articulation reaches between the eye and the nostril when the hindlimb is appressed along the body; the lateral metatarsalia are separated; the inner metatarsal tubercle is small and the outer distinct; toe discs are enlarged, and the webbing between toes weakly developed [1(1), 2i(1.75), 2e(1), 3i(2.5), 3e(2), 4i/e(3), 5(1.5)]. The skin on the dorsal surface is smooth without folds or ridges. The ventral skin is uniformly smooth. Femoral glands are very distinct in life, as well as after ethanol preservation, in external view.
</paragraph>
<paragraph pageId="0" pageNumber="149">
Coloration of the Holotype (Fig.
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Dorsolateral and ventral views of the holotype of Blommersia nataliae sp. nov. DRV 6867 (MNCN 50456). Note the color and shape of the femoral glands which are a diagnostic character to distinguish it from its sister taxon B. transmarina." figureDoi="10.3897/zookeys.994.57012.figure2" httpUri="https://binary.pensoft.net/fig/476542" pageId="0" pageNumber="149">2</figureCitation>
). In life, the overall color is creamy light brown with golden spots on the flanks, arms, and legs. It shows a thin yellowish line from the midpoint between the eyes to the vent. The legs are slightly darker brown and bands are visible. It shows a dark brown spot on the flanks and a characteristic larger moon-like spot on each
<normalizedToken originalValue="flanks">flank's</normalizedToken>
back close to the pelvic region and the hindlimbs. It presents a dark brown facial mask that covers from the snout, under the loreal region, to the tympanum (see Fig.
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Dorsolateral and ventral views of the holotype of Blommersia nataliae sp. nov. DRV 6867 (MNCN 50456). Note the color and shape of the femoral glands which are a diagnostic character to distinguish it from its sister taxon B. transmarina." figureDoi="10.3897/zookeys.994.57012.figure2" httpUri="https://binary.pensoft.net/fig/476542" pageId="0" pageNumber="149">2</figureCitation>
). The loreal region, as well as the outer iris periphery, shows a thin golden-colored line. The pupil is black and the inner iris area dark brown, while the outer iris area is golden with dark reticulations. The throat is brownish. The belly is light brown with some whitish, silver, and gold spots. The femoral glands are oval with a yellowish coloration and 9-10 circular internal rounded structures. After eight years in preservative, the back shows a creamy brown coloration that gets lighter towards the sides of the body, but the golden spots and dorsal line are lost. Ventral coloration is light brown without evident golden spots. The moon-like brown spot in the posterior part of the flanks is still evident, as well as the small ones on the flanks behind the arms. The femoral glands are whitish.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="149" type="variation">
<paragraph pageId="0" pageNumber="149">Variation.</paragraph>
<paragraph pageId="0" pageNumber="149">
The measurements of the holotype and paratypes are provided in Table
<tableCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Measurements of Blommersia nataliae sp. nov. and B. transmarina for comparison. See methods for abbreviations. The holotype of B. nataliae sp. nov. is shown in bold letters." httpUri="http://table.plazi.org/id/AE037202BE724C1E12E2F65537319F22" pageId="0" pageNumber="149" tableUuid="AE037202BE724C1E12E2F65537319F22">1</tableCitation>
. Sexual dimorphism is apparent in several characters: males present distinctive femoral glands, females are larger than males [males: median
<normalizedToken originalValue="±">+/-</normalizedToken>
SD SVL= 18.5
<normalizedToken originalValue="±">+/-</normalizedToken>
0.8 mm (min-max=17.9-20.5); females: 20
<normalizedToken originalValue="±">+/-</normalizedToken>
1.5 mm (min-max=19.6-23 mm)]. The color pattern is rather homogeneous, but females show an overall much creamier coloration than males, which are slightly darker. Both males and females show the characteristic brown rounded moon-like spot on the posterior flanks of the body, as well as some blotches on the lateral body sides behind the arms of variable size and shape. Vomerine teeth are more evident in specimen DRV6854 (MNCN50447), but only on one side of the vomer, and in DRV6855 (MNCN50448) and DRV6808 (MNCN50451) on both sides and more evident than in the holotype. Female DRV6855 (MNCN50448) lacks the moon-like blotch in the posterior side of the body, but shows a large circular one behind the arms. Female DRV6808 (MNCN50451) shows a similar pattern, but with a smaller blotch. Female DRV6868 (MNCN50450) shows a constellation of small rounded to irregular dark blotches from behind the arms to the inguinal region.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="149" type="natural history">
<paragraph pageId="0" pageNumber="149">
Natural history of
<taxonomicName authorityName="Vieites &amp; Nieto-Román &amp; Fernández &amp; Santos-Santos" authorityYear="2020" class="Amphibia" family="Mantellidae" genus="Blommersia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blommersia nataliae" order="Anura" pageId="0" pageNumber="149" phylum="Chordata" rank="species" species="nataliae">
<emphasis italics="true" pageId="0" pageNumber="149">Blommersia nataliae</emphasis>
</taxonomicName>
sp. nov.
</paragraph>
<paragraph pageId="0" pageNumber="149">
The species was found on the ground and in its breeding places: cut bamboo trunks filled with water (Fig.
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Photo of three specimens of Blommersia nataliae sp. nov. during reproduction in a cut bamboo trunk filled with water. On top of the picture, a male can be seen on top of a female, and another male is in the water on the left side of the trunk." figureDoi="10.3897/zookeys.994.57012.figure5" httpUri="https://binary.pensoft.net/fig/476545" pageId="0" pageNumber="149">5</figureCitation>
). There, we observed several males waiting for females to reproduce the night of 28th November 2012, with a temperature of 24.6 °C. No frogs were seen in the breeding places during the day. No call was ever heard during reproductive periods despite several attempts and leaving a digital recorder running for two hours at a breeding spot with active frogs at night, while
<taxonomicName lsidName="B. transmarina" pageId="0" pageNumber="149" rank="species" species="transmarina">
<emphasis italics="true" pageId="0" pageNumber="149">B. transmarina</emphasis>
</taxonomicName>
and
<taxonomicName lsidName="B. nauticus" pageId="0" pageNumber="149" rank="species" species="nauticus">
<emphasis italics="true" pageId="0" pageNumber="149">B. nauticus</emphasis>
</taxonomicName>
were calling. The clutches were placed on the
<normalizedToken originalValue="bamboos">bamboo's</normalizedToken>
inner walls above the water, but only a few eggs seemed to be fertilized and showed embryonic development. We counted three clutches of 42, 43, and 22 eggs on the walls of cut bamboo trunks in November 2012 at Mont
<normalizedToken originalValue="MSapere">M'Sapere</normalizedToken>
. Females seem to deposit several unfertilized eggs in the water that can serve as food for the tadpoles, but more research is needed to disentangle the
<normalizedToken originalValue="species">species'</normalizedToken>
reproductive strategy. Individuals were seen during the day on the ground in the forest leaf litter, mixed with
<taxonomicName lsidName="B. transmarina" pageId="0" pageNumber="149" rank="species" species="transmarina">
<emphasis italics="true" pageId="0" pageNumber="149">B. transmarina</emphasis>
</taxonomicName>
. No frogs were ever seen in other microhabitats like swamps, ponds, streams, or similar water bodies, where
<taxonomicName lsidName="B. transmarina" pageId="0" pageNumber="149" rank="species" species="transmarina">
<emphasis italics="true" pageId="0" pageNumber="149">B. transmarina</emphasis>
</taxonomicName>
reproduces. The species reproduces during the rainy season if the bamboo holes are filled with water. We observed clutches and tadpoles in November-December 2012 and April 2014. In some years with little rain (e.g., November 2019), we observed the frogs, but all usual reproduction sites were empty of water with no clutches or tadpoles. The scarcity of rain may strongly affect this species in the near future, limiting its possibility to reproduce.
</paragraph>
<caption doi="10.3897/zookeys.994.57012.figure5" httpUri="https://binary.pensoft.net/fig/476545" pageId="0" pageNumber="149" start="Figure 5" startId="F5">
<paragraph pageId="0" pageNumber="149">
<emphasis bold="true" pageId="0" pageNumber="149">Figure 5.</emphasis>
Photo of three specimens of
<taxonomicName authorityName="Vieites &amp; Nieto-Román &amp; Fernández &amp; Santos-Santos" authorityYear="2020" class="Amphibia" family="Mantellidae" genus="Blommersia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blommersia nataliae" order="Anura" pageId="0" pageNumber="149" phylum="Chordata" rank="species" species="nataliae">
<emphasis italics="true" pageId="0" pageNumber="149">Blommersia nataliae</emphasis>
</taxonomicName>
sp. nov. during reproduction in a cut bamboo trunk filled with water. On top of the picture, a male can be seen on top of a female, and another male is in the water on the left side of the trunk.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="149" type="distribution">
<paragraph pageId="0" pageNumber="149">Distribution.</paragraph>
<paragraph pageId="0" pageNumber="149">
Initially found on the slopes of Mont
<normalizedToken originalValue="MSapere">M'Sapere</normalizedToken>
only where there is still forest present and giant bamboos, between 235 m a.s.l. and 409 m a.s.l. (2012, 2014, and 2019). In the 2014 expedition, we also found the new species at Mont
<normalizedToken originalValue="Bénara">Benara</normalizedToken>
(
<geoCoordinate degrees="12.8712" direction="south" orientation="latitude" precision="5" value="-12.8712">12.8712°S</geoCoordinate>
,
<geoCoordinate degrees="45.15614" direction="east" orientation="longitude" precision="1" value="45.15614">45.15614°E</geoCoordinate>
, 317 m a.s.l.) in a forested place with fewer bamboo stands available, but few specimens. It is possible that the species breeds in other microhabitats (e.g., tree holes) as in the places where it occurs at Mont
<normalizedToken originalValue="Bénara">Benara</normalizedToken>
there are not many bamboo stands available, but this hypothesis needs to be confirmed. After several trips and visits around the whole entire main island of Mayotte and surrounding islets, with very intense fieldwork, we have not found it anywhere else, and the habitat appears to be degraded for the species.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="149" type="conservation">
<paragraph pageId="0" pageNumber="149">Conservation.</paragraph>
<paragraph pageId="0" pageNumber="149">
The new species is only known from two localities and seems restricted to mountain areas where the forest is still present with giant bamboo stands. The area of occupancy is estimated to be less than 10 km2 from the elevation range where it was observed (235 to 409 m a.sl.) and the remaining forest available at the Reserve
<normalizedToken originalValue="Forestière">Forestiere</normalizedToken>
de Majimbini (Mont
<normalizedToken originalValue="MSapere">M'Sapere</normalizedToken>
) and the Reserve
<normalizedToken originalValue="Forèstiere">Forestiere</normalizedToken>
du Mont Benara (
<bibRefCitation author="Dupuy, S" journalOrPublisher="Journal of Biogeography" pageId="0" pageNumber="149" refId="B5" refString="Dupuy, S, Laine, G, Tassin, J, Sarrailh, JM, 2019. &quot;Mayotte - Typologie des espaces boises - 2010 - (donnee brute)&quot;, CIRAD Dataverse, V2." title="&quot; Mayotte - Typologie des espaces boises - 2010 - (donnee brute) &quot;, CIRAD Dataverse, V 2." year="2019">Dupuy et al. 2019</bibRefCitation>
). There are bamboo stands also at lower elevations in degraded forests, but we never detected the species there. The distribution range is extremely small, the habitat is increasingly degraded, the breeding places (broken bamboo trunks) not frequent, and the fact that the observed densities seem to be very low suggest to consider the species as Critically Endangered according to IUCN criteria, in need of urgent conservation actions considering the ongoing degradation of these forest habitats. The impacts of local harvesting of bamboos or fires are not known, but it seems critical for the conservation of the species. The introduction of the chytrid fungus in Mayotte could decimate this species as well as
<taxonomicName lsidName="B. transmarina" pageId="0" pageNumber="149" rank="species" species="transmarina">
<emphasis italics="true" pageId="0" pageNumber="149">B. transmarina</emphasis>
</taxonomicName>
in a few years. The consequences of climate change, such as a reduction in rainfall that fills its reproductive sites, may strongly affect the species as well.
</paragraph>
</subSubSection>
</treatment>
</document>