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<document id="BF2CD73063DCCA4AB8F49421A9E43344" ID-DOI="10.11646/zootaxa.5397.4.1" ID-ISSN="1175-5326" ID-Zenodo-Dep="10469003" ID-ZooBank="2F745EBA-05B4-4F85-A0EE-A56A73683B7A" IM.bibliography_approvedBy="valdenar" IM.illustrations_approvedBy="valdenar" IM.materialsCitations_approvedBy="valdenar" IM.metadata_approvedBy="felipe" IM.tables_approvedBy="valdenar" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="valdenar" checkinTime="1704709229503" checkinUser="plazi" docAuthor="Gabriel, Hugh, Rothe, Laila-Denise, Köhler, Jörn, Rakotomanga, Sandratra, Edmonds, Devin, Galán, Pedro, Glaw, Frank, Lehtinen, Richard M., Rakotoarison, Andolalao &amp; Vences, Miguel" docDate="2024" docId="03B3990EC52F4641FF2CFA0DFC6CFAAC" docLanguage="en" docName="zootaxa.5397.4.1.pdf" docOrigin="Zootaxa 5397 (4)" docSource="http://dx.doi.org/10.11646/zootaxa.5397.4.1" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Guibemantis vakoa Gabriel &amp; Rothe &amp; Köhler &amp; Rakotomanga &amp; Edmonds &amp; Galán &amp; Glaw &amp; Lehtinen &amp; Rakotoarison &amp; Vences 2024, sp. nov." docType="treatment" docUuid="03D4783D-8313-4F15-8476-F73C046001D8" docUuidSource="ZooBank" docVersion="3" lastPageNumber="479" masterDocId="FF8AE176C537465DFFBBFFFAFFB6FFF9" masterDocTitle="Unexpected diversity and co-occurrence of phytotelmic frogs (Guibemantis) around Andasibe, one of the most intensively surveyed amphibian hotspots of Madagascar, and descriptions of three new species" masterLastPageNumber="485" masterPageNumber="451" pageNumber="475" updateTime="1704736861007" updateUser="valdenar" zenodo-license-document="CLOSED">
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<mods:title id="301221B4C09BBC7CB751ECD286870C5B">Unexpected diversity and co-occurrence of phytotelmic frogs (Guibemantis) around Andasibe, one of the most intensively surveyed amphibian hotspots of Madagascar, and descriptions of three new species</mods:title>
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<mods:namePart id="C2252B9F75D19DCAB816A8836D171891">Gabriel, Hugh</mods:namePart>
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<mods:namePart id="3F78A9C0E56B8A0236F4252618F98D24">Rakotomanga, Sandratra</mods:namePart>
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<mods:affiliation id="5CE3587EDA73E9F495B59A5557B72323">Universidad de A Coruña, Facultad de Ciencias, Campus de A Zapateira, s / n, (Grupo de Investigación en Biología Evolutiva, GIBE), E- 15071 A Coruña, Spain.</mods:affiliation>
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<mods:namePart id="15CF63F3737F8E68FFCC5E9681D10004">Glaw, Frank</mods:namePart>
<mods:affiliation id="C25B4D4BAADF4737A8D2EC8CAB3CBF6A">Zoologische Staatssammlung München (ZSM-SNSB), Münchhausenstr. 21, 81247 München, Germany.</mods:affiliation>
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<mods:namePart id="E94A08BBC53CA3BC5A67301108A851EC">Lehtinen, Richard M.</mods:namePart>
<mods:affiliation id="D3B24D6CCF45CBEE3C243FC0A29C932A">The College of Wooster, Department of Biology, 931 College Mall, Wooster, Ohio, 44691 USA.</mods:affiliation>
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<mods:namePart id="F3050F522D2A00609BDC7798E5640FEC">Rakotoarison, Andolalao</mods:namePart>
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<mods:affiliation id="DA68BF290FDB308618DFD018446A3A1B">Mention Agriculture, Universite de lItasy, Faliarivo Ambohidanerana, 118 Soavinandriana Itasy, Madagascar.</mods:affiliation>
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<mods:namePart id="47440F7638206A3CB80A9C1603A075D6">Vences, Miguel</mods:namePart>
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<mods:date id="088CA76725D48D7FF2F37EDE4F30F800">2024</mods:date>
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<heading id="D0ED9F74C52F4645FF2CFA0DFE50F9EB" bold="true" box="[151,486,1527,1554]" fontSize="11" level="1" pageId="24" pageNumber="475" reason="1">
<emphasis id="B96EF40AC52F4645FF2CFA0DFE50F9EB" bold="true" box="[151,486,1527,1554]" pageId="24" pageNumber="475">
<taxonomicName id="4C1A539BC52F4645FF2CFA0DFE37F9EB" authority="Gabriel &amp; Rothe &amp; Köhler &amp; Rakotomanga &amp; Edmonds &amp; Galán &amp; Glaw &amp; Lehtinen &amp; Rakotoarison &amp; Vences, 2024" authorityName="Gabriel &amp; Rothe &amp; Köhler &amp; Rakotomanga &amp; Edmonds &amp; Galán &amp; Glaw &amp; Lehtinen &amp; Rakotoarison &amp; Vences" authorityYear="2024" box="[151,385,1527,1554]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="species" species="vakoa" status="sp. nov.">
<emphasis id="B96EF40AC52F4645FF2CFA0DFE37F9EB" bold="true" box="[151,385,1527,1554]" italics="true" pageId="24" pageNumber="475">Guibemantis vakoa</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A25D4971C52F4645FE33FA02FE50F9EB" box="[392,486,1528,1554]" pageId="24" pageNumber="475" rank="species">sp. nov.</taxonomicNameLabel>
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<paragraph id="8BA52818C52F4645FF2CF9E6FE9EF9CF" blockId="24.[151,486,1527,1590]" box="[151,296,1564,1590]" pageId="24" pageNumber="475">
(
<figureCitation id="1321349DC52F4645FF24F9E6FF40F9CF" box="[159,246,1564,1590]" captionStart="FIGURE 10" captionStartId="25.[151,250,1915,1939]" captionTargetBox="[151,1436,273,1891]" captionTargetId="figure-55@25.[151,1436,273,1891]" captionTargetPageId="25" captionText="FIGURE 10. Male individuals of Guibemantis methueni from Andasibe and Besariaka. (AB) ZSM 3223/2012, (ZCMV 14089) from Andasibe. (CD) ZSM 391/2004 (ZCMV 933) from Besariaka. (EF) Specimen from Besariaka that cannot be reliably assigned to a voucher specimen and is assigned to G. methueni based on morphology." figureDoi="http://doi.org/10.5281/zenodo.10469027" httpUri="https://zenodo.org/record/10469027/files/figure.png" pageId="24" pageNumber="475">Figs 10</figureCitation>
, 12)
</paragraph>
<paragraph id="8BA52818C52F4645FF2CF99EFBE8F93F" blockId="24.[151,1437,1636,2022]" pageId="24" pageNumber="475">
<emphasis id="B96EF40AC52F4645FF2CF99EFF58F987" box="[151,238,1636,1662]" italics="true" pageId="24" pageNumber="475">Remark</emphasis>
. Individuals of this species have been reported as
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<emphasis id="B96EF40AC52F4645FCA7F99EFC1DF987" box="[796,939,1636,1662]" italics="true" pageId="24" pageNumber="475">Guibemantis</emphasis>
sp.
</taxonomicName>
<taxonomicName id="4C1A539BC52F4645FC63F99EFB26F987" authority="Moramanga" authorityName="Moramanga" baseAuthorityName="Boettger" baseAuthorityYear="1913" box="[984,1168,1636,1662]" class="Amphibia" family="Mantellidae" genus="Guibemantis" isUncertain="true" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="species" species="bicalcaratus">
aff.
<emphasis id="B96EF40AC52F4645FBBFF99EFB26F987" box="[1028,1168,1636,1662]" italics="true" pageId="24" pageNumber="475">bicalcaratus</emphasis>
</taxonomicName>
“Moramanga” by
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, and as
<emphasis id="B96EF40AC52F4645FE15F972FE7FF95B" box="[430,457,1672,1698]" italics="true" pageId="24" pageNumber="475">G.</emphasis>
sp.
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Vieites
<emphasis id="B96EF40AC52F4645FDC0F973FD05F95B" box="[635,691,1672,1698]" italics="true" pageId="24" pageNumber="475">et al.</emphasis>
(2009)
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and
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sp. Ca
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Perl
<emphasis id="B96EF40AC52F4645FBBDF973FB88F95B" box="[1030,1086,1672,1698]" italics="true" pageId="24" pageNumber="475">et al.</emphasis>
(2014)
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, but with incorrect field number information for the sample used therein (reported erroneously as ZCMV 466).
</paragraph>
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<materialsCitation id="3B722245C52F4645FF7CF92AFE27F8CB" collectingDate="2004-02-15" collectionCode="ZCMV" collectorName="M. Vences &amp; Vences, E &amp; C. Woodhead" country="Madagascar" elevation="976" latitude="-19.12863" location="Besariaka" longLatPrecision="1" longitude="48.28063" municipality="Moramanga" pageId="24" pageNumber="475" specimenCode="ZCMV 931" specimenCount="1" typeStatus="holotype">
<emphasis id="B96EF40AC52F4645FF7CF92AFE98F913" box="[199,302,1744,1770]" italics="true" pageId="24" pageNumber="475">
<typeStatus id="54A196BAC52F4645FF7CF92AFE98F913" box="[199,302,1744,1770]" pageId="24" pageNumber="475" type="holotype">Holotype</typeStatus>
</emphasis>
. ZSM 389/2004 (
<specimenCode id="DBBC8063C52F4645FE46F92AFD30F913" box="[509,646,1744,1770]" collectionCode="ZCMV" pageId="24" pageNumber="475">ZCMV 931</specimenCode>
), adult male, collected by
<collectorName id="26EF4DCEC52F4645FC01F92AFB83F913" box="[954,1077,1744,1770]" pageId="24" pageNumber="475">M. Vences</collectorName>
, E. Edwards and
<collectorName id="26EF4DCEC52F4645FAB9F92AFA2AF913" box="[1282,1436,1744,1770]" pageId="24" pageNumber="475">C. Woodhead</collectorName>
on
<date id="FFA40ED8C52F4645FF04F90EFE39F8F7" box="[191,399,1780,1807]" pageId="24" pageNumber="475" value="2004-02-15">
<collectingDate id="EFE0F730C52F4645FF04F90EFE39F8F7" box="[191,399,1780,1807]" pageId="24" pageNumber="475" value="2004-02-15">15 February 2004</collectingDate>
</date>
at
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(south of
<collectingMunicipality id="6BC1B262C52F4645FD13F90EFC82F8F7" box="[680,820,1780,1806]" pageId="24" pageNumber="475">Moramanga</collectingMunicipality>
), central eastern
<collectingCountry id="F30D6888C52F4645FC45F90EFB31F8F7" box="[1022,1159,1780,1806]" name="Madagascar" pageId="24" pageNumber="475">Madagascar</collectingCountry>
, coordinates
<geoCoordinate id="EE2E4EDFC52F4645FA98F90EFA21F8F7" box="[1315,1431,1780,1806]" degrees="19.12863" direction="south" orientation="latitude" pageId="24" pageNumber="475" precision="1" value="-19.12863">-19.12863</geoCoordinate>
,
<geoCoordinate id="EE2E4EDFC52F4645FF2CF8E2FEB5F8CB" box="[151,259,1816,1842]" degrees="48.28063" direction="east" orientation="longitude" pageId="24" pageNumber="475" precision="1" value="48.28063">48.28063</geoCoordinate>
,
<elevation id="0037CF2BC52F4645FEB5F8E2FE27F8CB" box="[270,401,1816,1842]" metricMagnitude="2" metricUnit="m" metricValue="9.76" pageId="24" pageNumber="475" unit="m" value="976.0">
<quantity id="4CE285FDC52F4645FEB5F8E2FEE3F8CB" box="[270,341,1816,1842]" metricMagnitude="2" metricUnit="m" metricValue="9.76" pageId="24" pageNumber="475" unit="m" value="976.0">976 m</quantity>
a.s.l.
</elevation>
</materialsCitation>
</paragraph>
<paragraph id="8BA52818C52F4645FF7CF8C7FAD3F8AF" blockId="24.[151,1437,1636,2022]" box="[199,1381,1852,1878]" pageId="24" pageNumber="475">
<materialsCitation id="3B722245C52F4645FF7CF8C7FAD4F8AF" box="[199,1378,1852,1878]" collectingDate="2004-02-15" collectionCode="ZCMV" collectorName="M. Vences &amp; Vences, E &amp; C. Woodhead" country="Madagascar" elevation="976" latitude="-19.12863" location="Besariaka" longLatPrecision="1" longitude="48.28063" municipality="Moramanga" pageId="24" pageNumber="475" specimenCode="ZCMV 930" specimenCount="1" typeStatus="paratype">
<emphasis id="B96EF40AC52F4645FF7CF8C7FE98F8AF" box="[199,302,1853,1878]" italics="true" pageId="24" pageNumber="475">
<typeStatus id="54A196BAC52F4645FF7CF8C7FE98F8AF" box="[199,302,1853,1878]" pageId="24" pageNumber="475" type="paratype">Paratype</typeStatus>
</emphasis>
.
<specimenCount id="9D1CE391C52F4645FE87F8C6FE68F8AF" box="[316,478,1852,1878]" count="1" pageId="24" pageNumber="475" type="generic">One specimen</specimenCount>
, ZSM 388/2004 (
<specimenCode id="DBBC8063C52F4645FD19F8C6FC9FF8AF" box="[674,809,1852,1878]" collectionCode="ZCMV" pageId="24" pageNumber="475">ZCMV 930</specimenCode>
), adult male with same collection data as holotype
</materialsCitation>
.
</paragraph>
<paragraph id="8BA52818C52F4644FF7CF89AFEB4FF00" blockId="24.[151,1437,1636,2022]" lastBlockId="25.[151,1437,151,249]" lastPageId="25" lastPageNumber="476" pageId="24" pageNumber="475">
<emphasis id="B96EF40AC52F4645FF7CF89AFE8FF883" box="[199,313,1888,1914]" italics="true" pageId="24" pageNumber="475">Definition</emphasis>
. Assigned to the subgenus
<taxonomicName id="4C1A539BC52F4645FDC9F89AFCACF883" box="[626,794,1888,1914]" pageId="24" pageNumber="475" rank="subGenus" subGenus="Pandanusicola">
<emphasis id="B96EF40AC52F4645FDC9F89AFCACF883" box="[626,794,1888,1914]" italics="true" pageId="24" pageNumber="475">Pandanusicola</emphasis>
</taxonomicName>
in the genus
<taxonomicName id="4C1A539BC52F4645FC0EF89AFBF2F883" authorityName="Dubois" authorityYear="1992" box="[949,1092,1888,1914]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="24" pageNumber="475" phylum="Chordata" rank="genus">
<emphasis id="B96EF40AC52F4645FC0EF89AFBF2F883" box="[949,1092,1888,1914]" italics="true" pageId="24" pageNumber="475">Guibemantis</emphasis>
</taxonomicName>
based on its small to medium body size, phytotelm-dwelling habitats (in
<taxonomicName id="4C1A539BC52F4645FDC1F87EFD5CF867" baseAuthorityName="Lehtinen" baseAuthorityYear="2002" box="[634,746,1924,1950]" class="Liliopsida" family="Pandanaceae" genus="Pandanus" kingdom="Plantae" order="Pandanales" pageId="24" pageNumber="475" phylum="Tracheophyta" rank="genus">
<emphasis id="B96EF40AC52F4645FDC1F87EFD5CF867" box="[634,746,1924,1950]" italics="true" pageId="24" pageNumber="475">Pandanus</emphasis>
</taxonomicName>
plants), near-absent webbing between toes, connected lateral metatarsalia, presence of both inner and outer metatarsal tubercles, intercalary elements between ultimate and penultimate phalanges of fingers and toes (verified by external examination), presence of well-delimited femoral macroglands consisting of discrete enlarged gland granules and without externally visible central depression and of whitish color laterally on throat (marking the vocal sac) in males (absence in females), and molecular phylogenetic affinities.
</paragraph>
<caption id="DF657890C52E4644FF2CF881FC4AF822" ID-DOI="http://doi.org/10.5281/zenodo.10469027" ID-Zenodo-Dep="10469027" httpUri="https://zenodo.org/record/10469027/files/figure.png" pageId="25" pageNumber="476" startId="25.[151,250,1915,1939]" targetBox="[151,1436,273,1891]" targetPageId="25" targetType="figure">
<paragraph id="8BA52818C52E4644FF2CF881FC4AF822" blockId="25.[151,1436,1915,2011]" pageId="25" pageNumber="476">
<emphasis id="B96EF40AC52E4644FF2CF881FEAEF86A" bold="true" box="[151,280,1915,1939]" pageId="25" pageNumber="476">FIGURE 10</emphasis>
. Male individuals of
<taxonomicName id="4C1A539BC52E4644FE57F881FD66F86A" box="[492,720,1915,1939]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="25" pageNumber="476" phylum="Chordata" rank="species" species="methueni">
<emphasis id="B96EF40AC52E4644FE57F881FD66F86A" box="[492,720,1915,1939]" italics="true" pageId="25" pageNumber="476">Guibemantis methueni</emphasis>
</taxonomicName>
from Andasibe and Besariaka. (
<emphasis id="B96EF40AC52E4644FBABF881FBF6F86A" bold="true" box="[1040,1088,1915,1939]" pageId="25" pageNumber="476">AB</emphasis>
) ZSM 3223/2012, (ZCMV 14089) from Andasibe. (
<emphasis id="B96EF40AC52E4644FEFEF865FEC3F84E" bold="true" box="[325,373,1951,1975]" pageId="25" pageNumber="476">CD</emphasis>
) ZSM 391/2004 (ZCMV 933) from Besariaka. (
<emphasis id="B96EF40AC52E4644FCD1F865FC21F84E" bold="true" box="[874,919,1951,1975]" pageId="25" pageNumber="476">EF</emphasis>
) Specimen from Besariaka that cannot be reliably assigned to a voucher specimen and is assigned to
<taxonomicName id="4C1A539BC52E4644FD23F839FCA2F822" box="[664,788,1987,2011]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="25" pageNumber="476" phylum="Chordata" rank="species" species="methueni">
<emphasis id="B96EF40AC52E4644FD23F839FCA2F822" box="[664,788,1987,2011]" italics="true" pageId="25" pageNumber="476">G. methueni</emphasis>
</taxonomicName>
based on morphology.
</paragraph>
</caption>
<caption id="DF657890C52D4647FF2CF86DFADDF801" ID-DOI="http://doi.org/10.5281/zenodo.10469029" ID-Zenodo-Dep="10469029" httpUri="https://zenodo.org/record/10469029/files/figure.png" pageId="26" pageNumber="477" startId="26.[151,250,1943,1967]" targetBox="[188,1402,181,1918]" targetPageId="26" targetType="figure">
<paragraph id="8BA52818C52D4647FF2CF86DFADDF801" blockId="26.[151,1437,1943,2040]" pageId="26" pageNumber="477">
<emphasis id="B96EF40AC52D4647FF2CF86DFEA1F856" bold="true" box="[151,279,1943,1967]" pageId="26" pageNumber="477">FIGURE 11</emphasis>
. Dorsal and ventral views of preserved holotypes of new
<taxonomicName id="4C1A539BC52D4647FCEBF862FB95F849" box="[848,1059,1944,1968]" pageId="26" pageNumber="477">
<emphasis id="B96EF40AC52D4647FCEBF862FC67F849" box="[848,977,1944,1968]" italics="true" pageId="26" pageNumber="477">Guibemantis</emphasis>
species
</taxonomicName>
:
<emphasis id="B96EF40AC52D4647FB90F86DFBD4F856" bold="true" box="[1067,1122,1943,1967]" pageId="26" pageNumber="477">AB.</emphasis>
<taxonomicName id="4C1A539BC52D4647FBDDF862FAF0F856" authority="Gabriel &amp; Rothe &amp; Köhler &amp; Rakotomanga &amp; Edmonds &amp; Galán &amp; Glaw &amp; Lehtinen &amp; Rakotoarison &amp; Vences, 2024" authorityName="Gabriel &amp; Rothe &amp; Köhler &amp; Rakotomanga &amp; Edmonds &amp; Galán &amp; Glaw &amp; Lehtinen &amp; Rakotoarison &amp; Vences" authorityYear="2024" box="[1126,1350,1944,1968]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="26" pageNumber="477" phylum="Chordata" rank="species" species="rianasoa" status="sp. nov.">
<emphasis id="B96EF40AC52D4647FBDDF862FAF0F856" box="[1126,1350,1944,1968]" italics="true" pageId="26" pageNumber="477">Guibemantis rianasoa</emphasis>
</taxonomicName>
<emphasis id="B96EF40AC52D4647FAF0F862FA2AF856" bold="true" box="[1355,1436,1944,1968]" pageId="26" pageNumber="477">
<taxonomicNameLabel id="A25D4971C52D4647FAF0F862FA2AF856" box="[1355,1436,1944,1968]" pageId="26" pageNumber="477" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
(sp. Ca14), male holotype ZSM 149/2022 (ZCMV 13705);
<emphasis id="B96EF40AC52D4647FD46F841FC83F82A" bold="true" box="[765,821,1979,2003]" pageId="26" pageNumber="477">CD.</emphasis>
<taxonomicName id="4C1A539BC52D4647FC87F846FB8AF82D" authority="Gabriel &amp; Rothe &amp; Köhler &amp; Rakotomanga &amp; Edmonds &amp; Galán &amp; Glaw &amp; Lehtinen &amp; Rakotoarison &amp; Vences, 2024" authorityName="Gabriel &amp; Rothe &amp; Köhler &amp; Rakotomanga &amp; Edmonds &amp; Galán &amp; Glaw &amp; Lehtinen &amp; Rakotoarison &amp; Vences" authorityYear="2024" box="[828,1084,1980,2004]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="26" pageNumber="477" phylum="Chordata" rank="species" species="ambakoana" status="sp. nov.">
<emphasis id="B96EF40AC52D4647FC87F846FB8AF82D" box="[828,1084,1980,2004]" italics="true" pageId="26" pageNumber="477">Guibemantis ambakoana</emphasis>
</taxonomicName>
<emphasis id="B96EF40AC52D4647FBFFF846FB20F82D" bold="true" box="[1092,1174,1980,2004]" pageId="26" pageNumber="477">
<taxonomicNameLabel id="A25D4971C52D4647FBFFF846FB20F82D" box="[1092,1174,1980,2004]" pageId="26" pageNumber="477" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
(sp. Ca3), male holotype ZSM 152/2022 (ZCMV 13707);
<emphasis id="B96EF40AC52D4647FE5FF81AFDA3F801" bold="true" box="[484,533,2016,2040]" pageId="26" pageNumber="477">E, F.</emphasis>
<taxonomicName id="4C1A539BC52D4647FDA7F81AFD69F801" authority="Gabriel &amp; Rothe &amp; Köhler &amp; Rakotomanga &amp; Edmonds &amp; Galán &amp; Glaw &amp; Lehtinen &amp; Rakotoarison &amp; Vences, 2024" authorityName="Gabriel &amp; Rothe &amp; Köhler &amp; Rakotomanga &amp; Edmonds &amp; Galán &amp; Glaw &amp; Lehtinen &amp; Rakotoarison &amp; Vences" authorityYear="2024" box="[540,735,2016,2040]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="26" pageNumber="477" phylum="Chordata" rank="species" species="vakoa" status="sp. nov.">
<emphasis id="B96EF40AC52D4647FDA7F81AFD69F801" box="[540,735,2016,2040]" italics="true" pageId="26" pageNumber="477">Guibemantis vakoa</emphasis>
</taxonomicName>
<emphasis id="B96EF40AC52D4647FD5DF81AFC81F801" bold="true" box="[742,823,2016,2040]" pageId="26" pageNumber="477">
<taxonomicNameLabel id="A25D4971C52D4647FD5DF81AFC81F801" box="[742,823,2016,2040]" pageId="26" pageNumber="477" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
(sp. Ca8), male holotype ZSM 389/2004 (ZCMV 931).
</paragraph>
</caption>
<paragraph id="8BA52818C52C4646FF7CFF6DFC6CFE9C" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="478">
The new species is characterized by the combination of the following characters: (1) small size with adult SVL up to
<quantity id="4CE285FDC52C4646FF62FF41FE9FFF2C" box="[217,297,187,213]" metricMagnitude="-2" metricUnit="m" metricValue="2.4" pageId="27" pageNumber="478" unit="mm" value="24.0">24 mm</quantity>
in males (females unknown), (2) light brownish dorsal color, often without distinct darker markings or spots, (3) distinct brown rostral stripe, (4) a few, small whitish spots on (posterior) flanks, (5) absence of distinct dorsolateral stripes, (6) absence of webbing on hand and only traces of web on foot, (7) vomerine teeth present, (8) femoral (macro)glands of males distinct, very clearly recognizable and of contrasting orange-brown color in life, single gland granules well recognizable externally in life and preservative.
</paragraph>
<paragraph id="8BA52818C52C4646FF7CFE95FD03FDC4" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="478">
Although this combination of character states will allow a distinction from most other species of
<taxonomicName id="4C1A539BC52C4646FABDFE95FA23FE70" authorityName="Dubois" authorityYear="1992" box="[1286,1429,367,393]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="genus">
<emphasis id="B96EF40AC52C4646FABDFE95FA23FE70" box="[1286,1429,367,393]" italics="true" pageId="27" pageNumber="478">Guibemantis</emphasis>
</taxonomicName>
, it is more difficult to diagnose the species against other brownish-colored species, especially
<emphasis id="B96EF40AC52C4646FB72FE69FE08FE28" italics="true" pageId="27" pageNumber="478">
<taxonomicName id="4C1A539BC52C4646FB72FE69FAC2FE54" authorityName="Gabriel &amp; Rothe &amp; Köhler &amp; Rakotomanga &amp; Edmonds &amp; Galán &amp; Glaw &amp; Lehtinen &amp; Rakotoarison &amp; Vences" authorityYear="2024" box="[1225,1396,403,429]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="ambakoana">G. ambakoana</taxonomicName>
,
<taxonomicName id="4C1A539BC52C4646FA3AFE69FE90FE28" baseAuthorityName="Boettger" baseAuthorityYear="1913" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="bicalcaratus">G. bicalcaratus</taxonomicName>
,
<taxonomicName id="4C1A539BC52C4646FE8BFE4DFE0CFE28" box="[304,442,439,465]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="methueni">G. methueni</taxonomicName>
,
</emphasis>
and
<emphasis id="B96EF40AC52C4646FE49FE4DFD27FE29" box="[498,657,439,465]" italics="true" pageId="27" pageNumber="478">
<taxonomicName id="4C1A539BC52C4646FE49FE4DFD3BFE29" authorityName="Lehtinen, Glaw &amp; Vences" authorityYear="2011" box="[498,653,439,465]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="wattersoni">G. wattersoni</taxonomicName>
.
</emphasis>
The new species can be reliably diagnosed from all other
<taxonomicName id="4C1A539BC52C4646FAB6FE4DFF5EFE0C" pageId="27" pageNumber="478">
<emphasis id="B96EF40AC52C4646FAB6FE4DFA2AFE28" box="[1293,1436,439,465]" italics="true" pageId="27" pageNumber="478">Guibemantis</emphasis>
species
</taxonomicName>
based on nucleotide positions in the mitochondrial 16S rRNA gene: MolD identified a robust diagnostic nucleotide combination of an 'G' in the site 998, 'A' in the site 1132, 'C' in the site 1251 (positions relative to the full 16S rRNA gene of
<taxonomicName id="4C1A539BC52C4646FED5FDD9FD12FDC4" baseAuthorityName="Grandidier" baseAuthorityYear="1872" box="[366,676,547,573]" class="Amphibia" family="Mantellidae" genus="Mantella" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="madagascariensis">
<emphasis id="B96EF40AC52C4646FED5FDD9FD12FDC4" box="[366,676,547,573]" italics="true" pageId="27" pageNumber="478">Mantella madagascariensis</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph id="8BA52818C52C4646FF7CFDBDFB50FD34" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="478">
<emphasis id="B96EF40AC52C4646FF7CFDBDFEF6FD99" box="[199,320,583,608]" italics="true" pageId="27" pageNumber="478">Diagnosis.</emphasis>
Distinction from
<emphasis id="B96EF40AC52C4646FDBDFDBDFBB4FD98" box="[518,1026,583,609]" italics="true" pageId="27" pageNumber="478">
<taxonomicName id="4C1A539BC52C4646FDBDFDBDFD2EFD98" authorityName="Koppetsch, Pabijan, Hutter, Köhler, Gehring, Rakotoarison, Ratsoavina, Scherz, Vieites, Glaw &amp; Vences" authorityYear="2023" box="[518,664,583,609]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="fotsitenda">G. fotsitenda</taxonomicName>
,
<taxonomicName id="4C1A539BC52C4646FD18FDBDFD4FFD98" baseAuthorityName="Peracca" baseAuthorityYear="1893" box="[675,761,583,609]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="liber">G. liber</taxonomicName>
,
<taxonomicName id="4C1A539BC52C4646FCB8FDBDFC3AFD98" authorityName="Koppetsch, Pabijan, Hutter, Köhler, Gehring, Rakotoarison, Ratsoavina, Scherz, Vieites, Glaw &amp; Vences" authorityYear="2023" box="[771,908,583,609]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="razandry">G. razandry</taxonomicName>
,
<taxonomicName id="4C1A539BC52C4646FC2DFDBDFBB4FD98" authorityName="Koppetsch, Pabijan, Hutter, Köhler, Gehring, Rakotoarison, Ratsoavina, Scherz, Vieites, Glaw &amp; Vences" authorityYear="2023" box="[918,1026,583,609]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="razoky">G. razoky</taxonomicName>
</emphasis>
and
<taxonomicName id="4C1A539BC52C4646FB8CFDBDFB00FD98" authorityName="Lehtinen, Glaw, Andreone, Pabijan &amp; Vences" authorityYear="2012" box="[1079,1206,583,609]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="tasifotsy">
<emphasis id="B96EF40AC52C4646FB8CFDBDFB00FD98" box="[1079,1206,583,609]" italics="true" pageId="27" pageNumber="478">G. tasifotsy</emphasis>
</taxonomicName>
: male femoral gland consisting of a well-delimited field of discrete enlarged single gland granules (vs. a diffuse field of small granules covering most of the ventral shank), occurrence limited to
<taxonomicName id="4C1A539BC52C4646FC82FD75FC1FFD50" baseAuthorityName="Lehtinen" baseAuthorityYear="2002" box="[825,937,655,681]" class="Liliopsida" family="Pandanaceae" genus="Pandanus" kingdom="Plantae" order="Pandanales" pageId="27" pageNumber="478" phylum="Tracheophyta" rank="genus">
<emphasis id="B96EF40AC52C4646FC82FD75FC1FFD50" box="[825,937,655,681]" italics="true" pageId="27" pageNumber="478">Pandanus</emphasis>
</taxonomicName>
leaf axils (vs. calling and breeding in open swamps), and calls emitted from
<taxonomicName id="4C1A539BC52C4646FDB0FD49FDCDFD34" baseAuthorityName="Lehtinen" baseAuthorityYear="2002" box="[523,635,691,717]" class="Liliopsida" family="Pandanaceae" genus="Pandanus" kingdom="Plantae" order="Pandanales" pageId="27" pageNumber="478" phylum="Tracheophyta" rank="genus">
<emphasis id="B96EF40AC52C4646FDB0FD49FDCDFD34" box="[523,635,691,717]" italics="true" pageId="27" pageNumber="478">Pandanus</emphasis>
</taxonomicName>
vs. calls emitted from exposed perches above swamps.
</paragraph>
<paragraph id="8BA52818C52C4646FF7CFD2DFD1FFCEC" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="478">
Distinction from
<taxonomicName id="4C1A539BC52C4646FE33FD2DFD83FD08" baseAuthorityName="Blommers-Schlosser &amp; Blanc" baseAuthorityYear="1991" box="[392,565,727,753]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="albolineatus">
<emphasis id="B96EF40AC52C4646FE33FD2DFD83FD08" box="[392,565,727,753]" italics="true" pageId="27" pageNumber="478">G. albolineatus</emphasis>
</taxonomicName>
: absence of well-delimited dorsolateral bands (vs. presence), and light brown ground color on dorsum (vs. chocolate-brown).
</paragraph>
<paragraph id="8BA52818C52C4646FF7CFCE5FEA5FC78" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="478">
Distinction from
<taxonomicName id="4C1A539BC52C4646FE3EFCE5FDFCFCC0" authorityName="Lehtinen, Glaw, Vences, Rakotoarison &amp; Scherz" authorityYear="2018" box="[389,586,799,825]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="albomaculatus">
<emphasis id="B96EF40AC52C4646FE3EFCE5FDFCFCC0" box="[389,586,799,825]" italics="true" pageId="27" pageNumber="478">G. albomaculatus</emphasis>
</taxonomicName>
: absence of fine white dotting on body (vs. presence, especially on hindlimbs and sometimes on dorsum), and males with white color only laterally on throat (vs. white color covering most of the throat).
</paragraph>
<paragraph id="8BA52818C52C4646FF7CFC71FBDDFC5C" blockId="27.[151,1437,151,2013]" box="[199,1131,907,933]" pageId="27" pageNumber="478">
Distinction from
<taxonomicName id="4C1A539BC52C4646FE33FC71FD9BFC5C" authorityName="Gabriel &amp; Rothe &amp; Köhler &amp; Rakotomanga &amp; Edmonds &amp; Galán &amp; Glaw &amp; Lehtinen &amp; Rakotoarison &amp; Vences" authorityYear="2024" box="[392,557,907,933]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="ambakoana">
<emphasis id="B96EF40AC52C4646FE33FC71FD9BFC5C" box="[392,557,907,933]" italics="true" pageId="27" pageNumber="478">G. ambakoana</emphasis>
</taxonomicName>
: absence of light dorsolateral bands (vs. presence).
</paragraph>
<paragraph id="8BA52818C52C4646FF7CFC55FB52FC14" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="478">
Distinction from
<taxonomicName id="4C1A539BC52C4646FE34FC55FD94FC30" authorityName="Lehtinen, Glaw &amp; Vences" authorityYear="2011" box="[399,546,943,969]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="annulatus">
<emphasis id="B96EF40AC52C4646FE34FC55FD94FC30" box="[399,546,943,969]" italics="true" pageId="27" pageNumber="478">G. annulatus</emphasis>
</taxonomicName>
: absence of a regular pattern of small dark dots dorsally (vs. presence), and absent or poorly contrasted light ring proximal to finger and toe discs (vs. distinct and contrasted).
</paragraph>
<paragraph id="8BA52818C52C4646FF7CFC0DFE76FBA0" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="478">
Distinction from
<taxonomicName id="4C1A539BC52C4646FE35FC0DFD89FBE8" baseAuthorityName="Boettger" baseAuthorityYear="1913" box="[398,575,1015,1041]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="bicalcaratus">
<emphasis id="B96EF40AC52C4646FE35FC0DFD89FBE8" box="[398,575,1015,1041]" italics="true" pageId="27" pageNumber="478">G. bicalcaratus</emphasis>
</taxonomicName>
: presence of a black vertical line in upper part of iris (vs. absence), and a weakly pigmented grayish flank with a few silvery white markings (vs. flanks of similar color as dorsum and without silvery markings).
</paragraph>
<paragraph id="8BA52818C52C4646FF7CFB99FC11FB58" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="478">
Distinction from
<taxonomicName id="4C1A539BC52C4646FE3CFB99FDFFFB84" baseAuthorityName="Blommers-Schlosser" baseAuthorityYear="1979" box="[391,585,1123,1149]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="flavobrunneus">
<emphasis id="B96EF40AC52C4646FE3CFB99FDFFFB84" box="[391,585,1123,1149]" italics="true" pageId="27" pageNumber="478">G. flavobrunneus</emphasis>
</taxonomicName>
: smaller body size (male SVL
<quantity id="4CE285FDC52C4646FC19FB99FBACFB84" box="[930,1050,1123,1149]" metricMagnitude="-2" metricUnit="m" metricValue="2.35" metricValueMax="2.4" metricValueMin="2.3" pageId="27" pageNumber="478" unit="mm" value="23.5" valueMax="24.0" valueMin="23.0">2324 mm</quantity>
vs. ≥
<quantity id="4CE285FDC52C4646FBEDFB99FB1FFB84" box="[1110,1193,1123,1149]" metricMagnitude="-2" metricUnit="m" metricValue="3.0" pageId="27" pageNumber="478" unit="mm" value="30.0">30 mm</quantity>
), and dorsum more or less uniformly brown (vs. brown with contrasted light brown pattern).
</paragraph>
<paragraph id="8BA52818C52C4646FF7CFB51FC1AFAF4" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="478">
Distinction from
<taxonomicName id="4C1A539BC52C4646FE3FFB51FDBAFB3C" box="[388,524,1195,1221]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="methueni">
<emphasis id="B96EF40AC52C4646FE3FFB51FDBAFB3C" box="[388,524,1195,1221]" italics="true" pageId="27" pageNumber="478">G. methueni</emphasis>
</taxonomicName>
: largely uniformly colored dorsum (vs. usually dark markings or spots on dorsum), femoral glands typically with contrasting yellow-orange color (vs. without contrasting color), and males with white color only laterally on throat (vs. white color covering most of throat).
</paragraph>
<paragraph id="8BA52818C52C4646FF7CFAEDFEA6FA80" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="478">
Distinction from
<taxonomicName id="4C1A539BC52C4646FE32FAEDFD85FAC8" authorityName="Bletz, Scherz, Rakotoarison, Lehtinen, Glaw &amp; Vences" authorityYear="2018" box="[393,563,1303,1329]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="milingilingy">
<emphasis id="B96EF40AC52C4646FE32FAEDFD85FAC8" box="[393,563,1303,1329]" italics="true" pageId="27" pageNumber="478">G. milingilingy</emphasis>
</taxonomicName>
: absence of dorsolateral bands (vs. broad and continuous dorsolateral bands), and dorsal color brown typically without greenish shade (vs. yellowish green dorsolateral bands and dorsal markings).
</paragraph>
<paragraph id="8BA52818C52C4646FF7CFA79FCD6FA38" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="478">
Distinction from
<taxonomicName id="4C1A539BC52C4646FE30FA79FDB3FA64" baseAuthorityName="Boulenger" baseAuthorityYear="1882" box="[395,517,1411,1437]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="pulcher">
<emphasis id="B96EF40AC52C4646FE30FA79FDB3FA64" box="[395,517,1411,1437]" italics="true" pageId="27" pageNumber="478">G. pulcher</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1A539BC52C4646FD85FA79FD41FA64" authorityName="Vences, Hutter, Glaw, Rakotoarison, Raselimanana &amp; Scherz" authorityYear="2023" box="[574,759,1411,1437]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="pulcherrimus">
<emphasis id="B96EF40AC52C4646FD85FA79FD41FA64" box="[574,759,1411,1437]" italics="true" pageId="27" pageNumber="478">G. pulcherrimus</emphasis>
</taxonomicName>
: brown dorsal color (vs. translucent green), and absence of purple-reddish dots and markings on the dorsum (vs. presence).
</paragraph>
<paragraph id="8BA52818C52C4646FF7CFA31FA81FA1C" blockId="27.[151,1437,151,2013]" box="[199,1335,1483,1509]" pageId="27" pageNumber="478">
Distinction from
<taxonomicName id="4C1A539BC52C4646FE33FA31FDA1FA1D" baseAuthorityName="Blommers-Schlosser" baseAuthorityYear="1979" box="[392,535,1483,1509]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="punctatus">
<emphasis id="B96EF40AC52C4646FE33FA31FDA1FA1D" box="[392,535,1483,1509]" italics="true" pageId="27" pageNumber="478">G. punctatus</emphasis>
</taxonomicName>
: absence of a regular pattern of small dark dots dorsally (vs. presence).
</paragraph>
<paragraph id="8BA52818C52C4646FF7CFA15FF4AF9A8" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="478">
Distinction from
<taxonomicName id="4C1A539BC52C4646FE37FA15FDA3F9F1" authorityName="Gabriel &amp; Rothe &amp; Köhler &amp; Rakotomanga &amp; Edmonds &amp; Galán &amp; Glaw &amp; Lehtinen &amp; Rakotoarison &amp; Vences" authorityYear="2024" box="[396,533,1519,1545]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="rianasoa">
<emphasis id="B96EF40AC52C4646FE37FA15FDA3F9F1" box="[396,533,1519,1545]" italics="true" pageId="27" pageNumber="478">G. rianasoa</emphasis>
</taxonomicName>
: larger body size (male SVL
<quantity id="4CE285FDC52C4646FCD0FA15FC50F9F0" box="[875,998,1519,1545]" metricMagnitude="-2" metricUnit="m" metricValue="2.35" metricValueMax="2.4" metricValueMin="2.3" pageId="27" pageNumber="478" unit="mm" value="23.5" valueMax="24.0" valueMin="23.0">2324 mm</quantity>
vs. ≤
<quantity id="4CE285FDC52C4646FB9CFA15FBCBF9F0" box="[1063,1149,1519,1545]" metricMagnitude="-2" metricUnit="m" metricValue="2.0" pageId="27" pageNumber="478" unit="mm" value="20.0">20 mm</quantity>
), brown dorsal color (vs. olive greenish), and large femoral glands with contrasting yellowish color in males (vs. small and inconspicuously colored).
</paragraph>
<paragraph id="8BA52818C52C4646FF7CF9A1FCCCF960" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="478">
Distinction from
<taxonomicName id="4C1A539BC52C4646FE3DF9A1FDABF98D" authorityName="Lehtinen, Glaw &amp; Vences" authorityYear="2011" box="[390,541,1627,1653]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="wattersoni">
<emphasis id="B96EF40AC52C4646FE3DF9A1FDABF98D" box="[390,541,1627,1653]" italics="true" pageId="27" pageNumber="478">G. wattersoni</emphasis>
</taxonomicName>
: absence of dorsolateral bands (vs. poorly contrasted bands usually present), and presence of a black vertical line in upper part of iris (vs. absence).
</paragraph>
<paragraph id="8BA52818C52C4646FF7CF959FADDF918" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="478">
Distinction from
<taxonomicName id="4C1A539BC52C4646FE33F959FDBDF945" authorityName="Lehtinen, Glaw, Vences, Rakotoarison &amp; Scherz" authorityYear="2018" box="[392,523,1699,1725]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="27" pageNumber="478" phylum="Chordata" rank="species" species="woosteri">
<emphasis id="B96EF40AC52C4646FE33F959FDBDF945" box="[392,523,1699,1725]" italics="true" pageId="27" pageNumber="478">G. woosteri</emphasis>
</taxonomicName>
: absence of dorsolateral bands (vs. poorly contrasted dorsolateral bands present), and a brown dorsal color without greenish elements (vs. often a green shade, especially of dorsolateral bands).
</paragraph>
<paragraph id="8BA52818C52C4641FF7CF911FAF1FEB8" blockId="27.[151,1437,151,2013]" lastBlockId="28.[151,1437,151,1365]" lastPageId="28" lastPageNumber="479" pageId="27" pageNumber="478">
<emphasis id="B96EF40AC52C4646FF7CF911FE42F8FC" box="[199,500,1771,1797]" italics="true" pageId="27" pageNumber="478">
Description of the
<typeStatus id="54A196BAC52C4646FE2FF911FE42F8FC" box="[404,500,1771,1797]" pageId="27" pageNumber="478" type="holotype">holotype</typeStatus>
</emphasis>
. Adult male in good state of preservation (
<figureCitation id="1321349DC52C4646FC73F911FBA1F8FC" box="[968,1047,1771,1797]" captionStart="FIGURE 11" captionStartId="26.[151,250,1943,1967]" captionTargetBox="[188,1402,181,1918]" captionTargetId="figure-36@26.[185,1403,181,1919]" captionTargetPageId="26" captionText="FIGURE 11. Dorsal and ventral views of preserved holotypes of new Guibemantis species: AB. Guibemantis rianasoa sp. nov. (sp. Ca14), male holotype ZSM 149/2022 (ZCMV 13705); CD. Guibemantis ambakoana sp. nov. (sp. Ca3), male holotype ZSM 152/2022 (ZCMV 13707); E, F. Guibemantis vakoa sp. nov. (sp. Ca8), male holotype ZSM 389/2004 (ZCMV 931)." figureDoi="http://doi.org/10.5281/zenodo.10469029" httpUri="https://zenodo.org/record/10469029/files/figure.png" pageId="27" pageNumber="478">Fig. 11</figureCitation>
). Tissue removed from ventral side of left thigh for DNA extraction. Head longer than wide and wider than body; snout pointed in dorsal and ventral views, more rounded in lateral view; canthus rostralis straight, loreal region flat; nostrils nearer to tip of snout than to eye, tympanum distinct, 43% of horizontal eye diameter; supratympanic fold distinct, curved; vomerine odontophores distinct, located medially between eye and choanae on either side of head; maxillary teeth present; tongue ovoid, and distinctly bifid at its tip. Arms slightly slender; relative finger length 1&lt;2&lt;4&lt;3, second finger very slightly shorter than fourth finger; finger discs moderately enlarged and squared off at tips in a rounded T shape, no webbing between fingers recognizable, subarticular tubercles distinct, unpaired. Hindlimbs moderately slender, foot length 91% of tibia length; lateral metatarsalia largely connected by muscular tissue; inner metatarsal tubercle oblong and recognizable, relatively small; outer metatarsal tubercle round, small and distinct; only traces of webbing recognizable between toes, relative length of toes 1&lt;2&lt;5&lt;3&lt;4, third toe slightly shorter than fifth; toe discs moderately enlarged. Femoral glands distinct and prominent. For morphometric measurements see
<tableCitation id="C6981DA3C52B4641FB56FEDDFAF7FEB8" box="[1261,1345,295,321]" captionStart="TABLE 1" captionStartId="18.[150,238,482,508]" captionTargetPageId="18" captionText="TABLE 1. Morphometric measurements (all in mm) of genotyped adults of species of Guibemantis targeted in this study. For abbreviations of measurements, see Materials and Methods; other abbreviations: HT, holotype; PT, paratype; M, male; F, female." pageId="28" pageNumber="479">Table 1</tableCitation>
.
</paragraph>
<paragraph id="8BA52818C52B4641FF7CFEB1FE2FFDC4" blockId="28.[151,1437,151,1365]" pageId="28" pageNumber="479">
After almost 20 years in preservative (
<figureCitation id="1321349DC52B4641FD36FEB1FD54FE9C" box="[653,738,331,357]" captionStart="FIGURE 11" captionStartId="26.[151,250,1943,1967]" captionTargetBox="[188,1402,181,1918]" captionTargetId="figure-36@26.[185,1403,181,1919]" captionTargetPageId="26" captionText="FIGURE 11. Dorsal and ventral views of preserved holotypes of new Guibemantis species: AB. Guibemantis rianasoa sp. nov. (sp. Ca14), male holotype ZSM 149/2022 (ZCMV 13705); CD. Guibemantis ambakoana sp. nov. (sp. Ca3), male holotype ZSM 152/2022 (ZCMV 13707); E, F. Guibemantis vakoa sp. nov. (sp. Ca8), male holotype ZSM 389/2004 (ZCMV 931)." figureDoi="http://doi.org/10.5281/zenodo.10469029" httpUri="https://zenodo.org/record/10469029/files/figure.png" pageId="28" pageNumber="479">Fig. 11</figureCitation>
), dorsal surface is beige with indistinct brownish spots and markings. The flanks are darker brown. No obvious light dorsolateral bands are recognizable. Forelimbs and hindlimbs beige-grayish, with brown crossbands. Ventral side beige. In life (
<figureCitation id="1321349DC52B4641FBABFE69FBE1FE54" box="[1040,1111,403,429]" captionStart="FIGURE 9" captionStartId="24.[151,250,1313,1337]" captionTargetBox="[151,1436,381,1290]" captionTargetId="figure-123@24.[151,1436,381,1290]" captionTargetPageId="24" captionText="FIGURE 9. Male individuals of Guibemantis vakoa sp. nov. from Besariaka (south of Moramanga), photographed in 2004. (AB) Holotype ZSM 389/2004 (ZCMV 931). (CD) Paratype ZSM 388/2004 (ZCMV 930) (assigned to species based on morphology only). Note the distinct orangish-brown femoral glands and only limited white throat pigmentation in comparison to syntopic specimens of G. methueni which have less distinctly pigmented glands and extended white throat color (Fig. 10)." figureDoi="http://doi.org/10.5281/zenodo.10469025" httpUri="https://zenodo.org/record/10469025/files/figure.png" pageId="28" pageNumber="479">Fig. 9</figureCitation>
), dorsal surface was golden brown with a few dark brown spots, a dark brown rostral stripe was present, and limbs were grayish brown with dark brown spots and crossbands. A few silvery white spots were visible on the posterior flanks. Ventrally, mostly unpigmented/translucent, with very little white pigment on throat. Femoral glands were very distinct and contrasted, of orange-brown color.
</paragraph>
<paragraph id="8BA52818C52B4641FF7CFDBDFC98FD7C" blockId="28.[151,1437,151,1365]" pageId="28" pageNumber="479">
<emphasis id="B96EF40AC52B4641FF7CFDBDFE99FD99" box="[199,303,583,608]" italics="true" pageId="28" pageNumber="479">Variation</emphasis>
. The known males all have distinct brown-orange femoral glands and only a little amount of white color laterally on the throat (
<figureCitation id="1321349DC52B4641FE6CFD91FDAAFD7C" box="[471,540,619,645]" captionStart="FIGURE 9" captionStartId="24.[151,250,1313,1337]" captionTargetBox="[151,1436,381,1290]" captionTargetId="figure-123@24.[151,1436,381,1290]" captionTargetPageId="24" captionText="FIGURE 9. Male individuals of Guibemantis vakoa sp. nov. from Besariaka (south of Moramanga), photographed in 2004. (AB) Holotype ZSM 389/2004 (ZCMV 931). (CD) Paratype ZSM 388/2004 (ZCMV 930) (assigned to species based on morphology only). Note the distinct orangish-brown femoral glands and only limited white throat pigmentation in comparison to syntopic specimens of G. methueni which have less distinctly pigmented glands and extended white throat color (Fig. 10)." figureDoi="http://doi.org/10.5281/zenodo.10469025" httpUri="https://zenodo.org/record/10469025/files/figure.png" pageId="28" pageNumber="479">Fig. 9</figureCitation>
). Females are unknown.
</paragraph>
<paragraph id="8BA52818C52B4641FF7CFD75FE6DFD08" blockId="28.[151,1437,151,1365]" pageId="28" pageNumber="479">
<emphasis id="B96EF40AC52B4641FF7CFD75FE89FD50" box="[199,319,655,681]" italics="true" pageId="28" pageNumber="479">Etymology</emphasis>
. The species name is derived from the Malagasy word
<taxonomicName id="4C1A539BC52B4641FC1CFD75FC4DFD50" authorityName="Gabriel &amp; Rothe &amp; Köhler &amp; Rakotomanga &amp; Edmonds &amp; Galán &amp; Glaw &amp; Lehtinen &amp; Rakotoarison &amp; Vences" authorityYear="2024" box="[935,1019,655,681]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="28" pageNumber="479" phylum="Chordata" rank="species" species="vakoa">“vakoa</taxonomicName>
”, used in eastern
<collectingCountry id="F30D6888C52B4641FB06FD75FAF2FD50" box="[1213,1348,655,681]" name="Madagascar" pageId="28" pageNumber="479">Madagascar</collectingCountry>
to refer to
<taxonomicName id="4C1A539BC52B4641FF0EFD49FE93FD34" baseAuthorityName="Lehtinen" baseAuthorityYear="2002" box="[181,293,691,717]" class="Liliopsida" family="Pandanaceae" genus="Pandanus" kingdom="Plantae" order="Pandanales" pageId="28" pageNumber="479" phylum="Tracheophyta" rank="genus">
<emphasis id="B96EF40AC52B4641FF0EFD49FE93FD34" box="[181,293,691,717]" italics="true" pageId="28" pageNumber="479">Pandanus</emphasis>
</taxonomicName>
screw pines and used to highlight the occurrence of this species in
<taxonomicName id="4C1A539BC52B4641FB9FFD49FB22FD34" baseAuthorityName="Lehtinen" baseAuthorityYear="2002" box="[1060,1172,691,717]" class="Liliopsida" family="Pandanaceae" genus="Pandanus" kingdom="Plantae" order="Pandanales" pageId="28" pageNumber="479" phylum="Tracheophyta" rank="genus">
<emphasis id="B96EF40AC52B4641FB9FFD49FB22FD34" box="[1060,1172,691,717]" italics="true" pageId="28" pageNumber="479">Pandanus</emphasis>
</taxonomicName>
leaf axils. The name is used as a noun in apposition.
</paragraph>
<paragraph id="8BA52818C52B4641FF7CFD01FC8BFCA4" blockId="28.[151,1437,151,1365]" pageId="28" pageNumber="479">
<emphasis id="B96EF40AC52B4641FF7CFD01FEC0FCEC" box="[199,374,763,789]" italics="true" pageId="28" pageNumber="479">Natural history</emphasis>
. Very poorly known. Specimens were found on
<date id="FFA40ED8C52B4641FC13FD00FBC2FCEC" box="[936,1140,762,789]" pageId="28" pageNumber="479" value="2004-02-15">
15 February
<quantity id="4CE285FDC52B4641FB87FD01FBC2FCEC" box="[1084,1140,763,789]" metricMagnitude="1" metricUnit="m" metricValue="5.090159999999999" pageId="28" pageNumber="479" unit="in" value="2004.0">2004</quantity>
</date>
in a forest fragment near Besariaka (destroyed by slash-and-burn agriculture two years later) in
<taxonomicName id="4C1A539BC52B4641FC0FFCE5FB92FCC0" baseAuthorityName="Lehtinen" baseAuthorityYear="2002" box="[948,1060,799,825]" class="Liliopsida" family="Pandanaceae" genus="Pandanus" kingdom="Plantae" order="Pandanales" pageId="28" pageNumber="479" phylum="Tracheophyta" rank="genus">
<emphasis id="B96EF40AC52B4641FC0FFCE5FB92FCC0" box="[948,1060,799,825]" italics="true" pageId="28" pageNumber="479">Pandanus</emphasis>
</taxonomicName>
plants, calling during the day. At Besariaka, specimens occurred in syntopy with
<taxonomicName id="4C1A539BC52B4641FD16FCB9FC81FCA4" box="[685,823,835,861]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="28" pageNumber="479" phylum="Chordata" rank="species" species="methueni">
<emphasis id="B96EF40AC52B4641FD16FCB9FC81FCA4" box="[685,823,835,861]" italics="true" pageId="28" pageNumber="479">G. methueni</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BA52818C52B4641FF7CFC9DFE6FFBA0" blockId="28.[151,1437,151,1365]" pageId="28" pageNumber="479">
<emphasis id="B96EF40AC52B4641FF7CFC9DFEE0FC79" box="[199,342,871,896]" italics="true" pageId="28" pageNumber="479">Bioacoustics</emphasis>
. Advertisement calls recorded on
<date id="FFA40ED8C52B4641FD63FC9CFC16FC78" box="[728,928,870,897]" pageId="28" pageNumber="479" value="2004-02-15">15 February 2004</date>
at Besariaka (air temperature 21°C;
<figureCitation id="1321349DC52B4641FA85FC9DFA22FC78" box="[1342,1428,871,897]" captionStart="FIGURE 4" captionStartId="11.[151,250,1715,1739]" captionTargetBox="[151,1435,181,1692]" captionTargetId="figure-22@11.[151,1435,181,1692]" captionTargetPageId="11" captionText="FIGURE 4. Audiospectrograms and oscillograms of advertisement calls of six species of Guibemantis of the subgenus Pandanusicola. A, call series (containing 3 calls) of Guibemantis rianasoa sp. nov., recorded on 19 November 2022 at Andasibe. Recording high-pass filtered at 2000 Hz. B, two calls of Guibemantis ambakoana sp. nov., recorded on 19 November 2022 at Andasibe. Recording high-pass filtered at 1500 Hz. C, three calls of Guibemantis vakoa sp. nov. recorded on 15 February 2004 at Besariaka. Recording band-pass filtered at 15009200 Hz. D, call series (containing 5 calls) of Guibemantis methueni recorded on 8 November 2011 at Andasibe. Recording high-pass filtered at 1000 Hz. EF, calls of G. annulatus (from Lehtinen et al. 2011) and G. pulcher (from Vences et al. 2011)." figureDoi="http://doi.org/10.5281/zenodo.10469015" httpUri="https://zenodo.org/record/10469015/files/figure.png" pageId="28" pageNumber="479">Fig. 4C</figureCitation>
) consist of a single very short click-note, emitted at long, somewhat irregular intervals. Amplitude modulation is evident in each call, with maximum call energy present at the beginning of the call, gradually decreasing towards the calls end. Numerical parameters of 28 analysed calls from at least two individuals are as follows: call duration (= note duration) 814 ms (10.2 ± 1.7 ms); inter-call intervals 6231286 ms (869.5 ± 214.3 ms); dominant frequency 35213972 Hz (3720 ± 206 Hz); prevalent bandwidth 20005500 Hz. Call repetition rate was approximately between 6585 calls/minute.
</paragraph>
<paragraph id="8BA52818C52B4641FF7CFB99FD87FAF4" blockId="28.[151,1437,151,1365]" pageId="28" pageNumber="479">
Calls recorded in 2007 at Betampona (
<bibRefCitation id="EF8B55E9C52B4641FDC3FB99FC86FB84" author="Rosa, G. M. &amp; Marquez, R. &amp; Andreone, F." box="[632,816,1123,1149]" pageId="28" pageNumber="479" refId="ref21688" refString="Rosa, G. M., Marquez, R. &amp; Andreone, F. (2011) The astonishing calls of the frogs of Betampona. Museo Regionale di Scienze Naturali di Torino &amp; Fonoteca Madrid, Torino. [CD and booklet]" type="book" year="2011">
Rosa
<emphasis id="B96EF40AC52B4641FD0CFB99FD59FB84" box="[695,751,1123,1149]" italics="true" pageId="28" pageNumber="479">et al.</emphasis>
2011
</bibRefCitation>
, track 27, as
<emphasis id="B96EF40AC52B4641FC78FB99FC61FB84" box="[963,983,1123,1149]" italics="true" pageId="28" pageNumber="479">G</emphasis>
. sp.
<taxonomicName id="4C1A539BC52B4641FBB7FB99FB73FB84" baseAuthorityName="Boettger" baseAuthorityYear="1913" box="[1036,1221,1123,1149]" class="Amphibia" family="Mantellidae" genus="Guibemantis" isUncertain="true" kingdom="Animalia" order="Anura" pageId="28" pageNumber="479" phylum="Chordata" rank="species" species="bicalcaratus">
aff.
<emphasis id="B96EF40AC52B4641FB82FB99FB73FB84" box="[1081,1221,1123,1149]" italics="true" pageId="28" pageNumber="479">bicalcaratus</emphasis>
</taxonomicName>
) agree in character with those reported from Besariaka in consisting of a single very short click-note emitted at variable, but long intervals. Numerical parameters of 32 analysed calls are as follows: call duration (= note duration) 39 ms (5.9 ± 1.9 ms); inter-call intervals 4711470 ms (767.9 ± 363.8 ms); dominant frequency 34233779 Hz (3576 ± 135 Hz); prevalent bandwidth 20005700 Hz.
</paragraph>
<paragraph id="8BA52818C52B4641FF7CFAEDFC6CFAAC" blockId="28.[151,1437,151,1365]" pageId="28" pageNumber="479">
<emphasis id="B96EF40AC52B4641FF7CFAEDFEE6FAC8" box="[199,336,1303,1329]" italics="true" pageId="28" pageNumber="479">Distribution</emphasis>
. Besides (1) the
<typeStatus id="54A196BAC52B4641FDA7FAEDFDFAFAC8" box="[540,588,1303,1329]" pageId="28" pageNumber="479">type</typeStatus>
locality Besariaka, the species has also been recorded, based on genetic evidence, from (2) Sahafina, (3) Betampona, and (4) Fierenana (
<figureCitation id="1321349DC52B4641FCDCFAC1FC7BFAAC" box="[871,973,1339,1365]" captionStart-0="FIGURE 1" captionStart-1="FIGURE 2" captionStartId-0="6.[151,250,1908,1932]" captionStartId-1="9.[151,250,1874,1898]" captionTargetBox-0="[370,1218,181,1885]" captionTargetBox-1="[227,1361,181,1850]" captionTargetId-0="figure-36@6.[370,1218,181,1885]" captionTargetId-1="figure-22@9.[226,1361,181,1850]" captionTargetPageId-0="6" captionTargetPageId-1="9" captionText-0="FIGURE 1. Maximum Likelihood tree based on a DNA sequences of a fragment of the mitochondrial 16S rRNA gene (alignment length 545 bp) for 368 specimens of all species and candidate species of Guibemantis. See Koppetsch et al. (2023) for a discussion of the mitochondrial non-monophyly of species in the G. liber complex (G. fotsitenda, G. liber, G. razandry, G. razoky)." captionText-1="FIGURE 2. Map of Madagascar showing genetically verified locality records of the target species of Guibemantis studied herein (colors as in Fig. 1). The base map shows vegetation across Madagascar from the Madagascar Vegetation Mapping Project (Moat &amp; Smith 2007; formerly available at www.vegmad.org). Vegetation is colored as follows: green, humid forest (rainforest); red, western dry deciduous forest; bluish, western subhumid forest; orange, south western dry spiny forest-thicket; yellow, tapia forest." figureDoi-0="http://doi.org/10.5281/zenodo.10469005" figureDoi-1="http://doi.org/10.5281/zenodo.10469009" httpUri-0="https://zenodo.org/record/10469005/files/figure.png" httpUri-1="https://zenodo.org/record/10469009/files/figure.png" pageId="28" pageNumber="479">Figs 12</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>