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<document id="7DC2228C0B84AB8A4F8A957AB2FC592A" ID-DOI="10.11646/zootaxa.4006.3.1" ID-GBIF-Dataset="9b8d3194-5435-46fb-92ee-301fd23b0782" ID-ISSN="1175-5326" ID-Zenodo-Dep="240836" ID-ZooBank="136F727A-1628-449F-B762-254371D27360" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1461211198470" checkinUser="plazi" docAuthor="Köhler, Jörn, Glaw, Frank, Pabijan, Maciej &amp; Vences, Miguel" docDate="2015" docId="039D87F4FFAFC80808C1FF4CFDCDE3A6" docLanguage="en" docName="zt04006p438.pdf" docOrigin="Zootaxa 4006 (3)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Aglyptodactylus inguinalis Gunther 1877" docType="treatment" docVersion="9" lastPageNumber="423" masterDocId="FFA4FF8CFFBBC81E0856FFDBFFFCE677" masterDocTitle="Integrative taxonomic revision of mantellid frogs of the genus Aglyptodactylus (Anura: Mantellidae)" masterLastPageNumber="438" masterPageNumber="401" pageNumber="421" updateTime="1698602080951" updateUser="plazi">
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<mods:title id="51795DCBA664F4CA3A7BCBB67712278A">Integrative taxonomic revision of mantellid frogs of the genus Aglyptodactylus (Anura: Mantellidae)</mods:title>
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<mods:namePart id="B3DCF9163438F64E00E5F9739CE97497">Köhler, Jörn</mods:namePart>
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<mods:namePart id="2DBEC850A2D11DFF810F4BAC0B251632">Glaw, Frank</mods:namePart>
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<mods:namePart id="758665BE394FB7B8A8D4E07C3CA3F0D3">Pabijan, Maciej</mods:namePart>
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<mods:namePart id="CFD535285AB936496139F4DFCECEC20F">Vences, Miguel</mods:namePart>
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<mods:title id="DCB17AB550356A2BC43447AE7570D06E">Zootaxa</mods:title>
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<mods:date id="59DBC16757328F2D369E7FD4B9B3B702">2015</mods:date>
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<mods:number id="41AD8AD4B849F935FF7C25A8E391EC3B">4006</mods:number>
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<mods:classification id="A9B47E69793ACB8D7E3CADA485549235">journal article</mods:classification>
<mods:identifier id="AF8135FFC737487414DFEC924C093542" type="DOI">10.11646/zootaxa.4006.3.1</mods:identifier>
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<treatment id="039D87F4FFAFC80808C1FF4CFDCDE3A6" ID-DOI="http://doi.org/10.5281/zenodo.5684810" ID-GBIF-Taxon="119630513" ID-Zenodo-Dep="5684810" LSID="urn:lsid:plazi:treatment:039D87F4FFAFC80808C1FF4CFDCDE3A6" httpUri="http://treatment.plazi.org/id/039D87F4FFAFC80808C1FF4CFDCDE3A6" lastPageId="22" lastPageNumber="423" pageId="20" pageNumber="421">
<subSubSection id="C32E6569FFAFC80A08C1FF4CFCA2E6C5" box="[151,862,151,178]" pageId="20" pageNumber="421" type="nomenclature">
<paragraph id="8B8B36E2FFAFC80A08C1FF4CFCA2E6C5" blockId="20.[151,862,151,178]" box="[151,862,151,178]" pageId="20" pageNumber="421">
<heading id="D0C3818EFFAFC80A08C1FF4CFCA2E6C5" bold="true" box="[151,862,151,178]" fontSize="11" level="1" pageId="20" pageNumber="421" reason="1">
<emphasis id="B940EAF0FFAFC80A08C1FF4CFCA2E6C5" bold="true" box="[151,862,151,178]" pageId="20" pageNumber="421">
<taxonomicName id="4C344D61FFAFC80A08C1FF4CFD5FE6C5" ID-CoL="663B7" authority="Gunther, 1877" authorityName="Gunther" authorityYear="1877" box="[151,675,151,178]" class="Amphibia" family="Mantellidae" genus="Aglyptodactylus" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="inguinalis">
<emphasis id="B940EAF0FFAFC80A08C1FF4CFE24E6C6" bold="true" box="[151,472,151,177]" italics="true" pageId="20" pageNumber="421">Aglyptodactylus inguinalis</emphasis>
(
<bibRefCitation id="EFA54B13FFAFC80A09BEFF43FD67E6C5" author="Gunther" box="[488,667,152,178]" pageId="20" pageNumber="421" refString="Gunther, A. (1877) Descriptions of some new species of reptiles from Madagascar. Annals and Magazine of Natural History, Series 4, 19, 313 - 317." type="journal article" year="1877">Günther, 1877</bibRefCitation>
)
</taxonomicName>
—bona species
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C32E6569FFAFC80A08C1FF0CFD69E74F" pageId="20" pageNumber="421" type="discussion">
<paragraph id="8B8B36E2FFAFC80A08C1FF0CFD69E74F" blockId="20.[151,1437,215,2027]" pageId="20" pageNumber="421">
<emphasis id="B940EAF0FFAFC80A08C1FF0CFEF7E687" bold="true" box="[151,267,215,240]" pageId="20" pageNumber="421">Remarks.</emphasis>
This species has been referred to as
<taxonomicName id="4C344D61FFAFC80A0AFDFF03FC9CE687" box="[683,864,216,240]" class="Amphibia" family="Mantellidae" genus="Aglyptodactylus" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="genus">
<emphasis id="B940EAF0FFAFC80A0AFDFF03FC9CE687" box="[683,864,216,240]" italics="true" pageId="20" pageNumber="421">Aglyptodactylus</emphasis>
</taxonomicName>
sp.
<taxonomicName id="4C344D61FFAFC80A0BD8FF0CFB7AE687" authority="Ranomafana" authorityName="Ranomafana" box="[910,1158,215,240]" class="Amphibia" family="Mantellidae" genus="Aglyptodactylus" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="madagascariensis">
aff.
<emphasis id="B940EAF0FFAFC80A0BEAFF03FB7AE687" box="[956,1158,216,240]" italics="true" pageId="20" pageNumber="421">madagascariensis</emphasis>
</taxonomicName>
”Ranomafana” by Glaw &amp; Vences (2007) and
<taxonomicName id="4C344D61FFAFC80A09F0FF27FDA7E763" box="[422,603,252,276]" class="Amphibia" family="Mantellidae" genus="Aglyptodactylus" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="genus">
<emphasis id="B940EAF0FFAFC80A09F0FF27FDA7E763" box="[422,603,252,276]" italics="true" pageId="20" pageNumber="421">Aglyptodactylus</emphasis>
</taxonomicName>
sp. 3 by
<bibRefCitation id="EFA54B13FFAFC80A0A8AFF20FC2CE763" box="[732,976,251,276]" pageId="20" pageNumber="421" refString="Vieites, D. R., Wollenberg, K. C., Andreone, F., Kohler, J., Glaw, F. &amp; Vences, M. (2009) Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences of the U. S. A., 106, 8267 - 8272. http: // dx. doi. org / 10.1073 / pnas. 0810821106" type="journal article">
Vieites
<emphasis id="B940EAF0FFAFC80A0B6EFF27FC8BE763" box="[824,887,252,276]" italics="true" pageId="20" pageNumber="421">et al.</emphasis>
(2009)
</bibRefCitation>
and
<bibRefCitation id="EFA54B13FFAFC80A0C42FF20FAC0E763" box="[1044,1340,251,276]" pageId="20" pageNumber="421" refString="Wollenberg, K. C. W., Vieites, D. R., Glaw, F. &amp; Vences, M. (2011) Speciation in little: the role of range and body size in the diversification of Malagasy mantellid frogs. BMC Evolutionary Biology, 11, 217." type="journal article">
Wollenberg
<emphasis id="B940EAF0FFAFC80A0CF5FF27FB1EE763" box="[1187,1250,252,276]" italics="true" pageId="20" pageNumber="421">et al.</emphasis>
(2011)
</bibRefCitation>
, and as
<taxonomicName id="4C344D61FFAFC80A08C1FEC4FEB0E740" box="[151,332,287,311]" class="Amphibia" family="Mantellidae" genus="Aglyptodactylus" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="genus">
<emphasis id="B940EAF0FFAFC80A08C1FEC4FEB0E740" box="[151,332,287,311]" italics="true" pageId="20" pageNumber="421">Aglyptodactylus</emphasis>
</taxonomicName>
sp. Ca3 by
<bibRefCitation id="EFA54B13FFAFC80A0986FEC4FD6DE74F" box="[464,657,287,312]" pageId="20" pageNumber="421" refString="Perl, R. G. B., Nagy, Z. T., Sonet, G., Glaw, F., Wollenberg, K. C. &amp; Vences, M. (2014) DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia, 35, 197 - 206. http: // dx. doi. org / 10.1163 / 15685381 - 00002942" type="journal article">
Perl
<emphasis id="B940EAF0FFAFC80A0A52FEFBFDC0E740" box="[516,572,287,311]" italics="true" pageId="20" pageNumber="421">et al.</emphasis>
(2014)
</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C32E6569FFAFC80B0891FE98FAF6E76B" lastPageId="21" lastPageNumber="422" pageId="20" pageNumber="421" type="materials_examined">
<paragraph id="8B8B36E2FFAFC80A0891FE98FED8E479" blockId="20.[151,1437,215,2027]" pageId="20" pageNumber="421">
<emphasis id="B940EAF0FFAFC80A0891FE98FE40E72B" bold="true" box="[199,444,323,348]" pageId="20" pageNumber="421">Material examined.</emphasis>
<collectionCode id="ED25AE27FFAFC80A0999FE9FFDDFE72C" box="[463,547,324,347]" pageId="20" pageNumber="421">BMNH</collectionCode>
1947.2.26.54,
<typeStatus id="548F8840FFAFC80A0AB2FE98FCBAE72B" box="[740,838,323,348]" pageId="20" pageNumber="421" type="holotype">holotype</typeStatus>
(probably a male, currently unavailable), from Anzahamaru, [central eastern]
<collectingCountry id="F3237672FFAFC80A0A58FEBDFD65E708" box="[526,665,358,383]" name="Madagascar" pageId="20" pageNumber="421">Madagascar</collectingCountry>
;
<collectionCode id="ED25AE27FFAFC80A0AF8FEBCFD14E708" box="[686,744,359,383]" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="20" pageNumber="421">ZSM</collectionCode>
5213/2005 (
<collectionCode id="ED25AE27FFAFC80A0BDEFEBCFC2DE708" box="[904,977,359,383]" pageId="20" pageNumber="421">FGZC</collectionCode>
2802), adult female, from Marojejy,
<geoCoordinate id="EE005025FFAFC80A08C1FE51FED6E7D4" box="[151,298,394,419]" direction="south" orientation="latitude" pageId="20" pageNumber="421" precision="1" value="-14.437667">14°26.260' S</geoCoordinate>
,
<geoCoordinate id="EE005025FFAFC80A096FFE51FE33E7D5" box="[313,463,394,419]" direction="east" orientation="longitude" pageId="20" pageNumber="421" precision="1" value="49.77555">49°46.533' E</geoCoordinate>
,
<quantity id="4CCC9B07FFAFC80A098BFE50FDDAE7D5" box="[477,550,395,419]" metricMagnitude="2" metricUnit="m" metricValue="4.81" pageId="20" pageNumber="421" unit="m" value="481.0">481 m</quantity>
a.s.l., northern
<collectingCountry id="F3237672FFAFC80A0A8DFE51FC9AE7D4" box="[731,870,394,419]" name="Madagascar" pageId="20" pageNumber="421">Madagascar</collectingCountry>
;
<collectionCode id="ED25AE27FFAFC80A0B22FE50FC52E7D4" box="[884,942,395,419]" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="20" pageNumber="421">ZSM</collectionCode>
183/2002 (
<collectionCode id="ED25AE27FFAFC80A0C67FE50FB78E7D4" box="[1073,1156,395,419]" pageId="20" pageNumber="421">FGMV</collectionCode>
2001.1482), adult male, from Tolagnaro, southeastern
<collectingCountry id="F3237672FFAFC80A09BFFE75FD88E7B0" box="[489,628,430,455]" name="Madagascar" pageId="20" pageNumber="421">Madagascar</collectingCountry>
;
<collectionCode id="ED25AE27FFAFC80A0AD6FE74FD45E7B0" box="[640,697,431,455]" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="20" pageNumber="421">ZSM</collectionCode>
2271/2007 (
<collectionCode id="ED25AE27FFAFC80A0B1FFE74FC60E7B0" box="[841,924,431,455]" pageId="20" pageNumber="421">ZCMV</collectionCode>
5429), adult female, from Manombo Special Reserve (precise coordinates not taken, but roughly at
<geoCoordinate id="EE005025FFAFC80A0B17FE09FC56E79D" box="[833,938,466,490]" direction="south" orientation="latitude" pageId="20" pageNumber="421" precision="925" value="-23.033333">23°02' S</geoCoordinate>
,
<geoCoordinate id="EE005025FFAFC80A0B96FE09FBD0E79D" box="[960,1068,466,490]" direction="east" orientation="longitude" pageId="20" pageNumber="421" precision="925" value="47.733334">47°44' E</geoCoordinate>
,
<quantity id="4CCC9B07FFAFC80A0C16FE09FB50E79D" box="[1088,1196,466,490]" metricMagnitude="1" metricUnit="m" metricValue="3.5" metricValueMax="4.0" metricValueMin="3.0" pageId="20" pageNumber="421" unit="m" value="35.0" valueMax="40.0" valueMin="30.0">30‒40 m</quantity>
a.s.l.), southeastern
<collectingCountry id="F3237672FFAFC80A08C1FE2EFEDDE479" box="[151,289,501,526]" name="Madagascar" pageId="20" pageNumber="421">Madagascar</collectingCountry>
.
</paragraph>
<paragraph id="8B8B36E2FFAFC80A0891FDC2FE93E55B" blockId="20.[151,1437,215,2027]" pageId="20" pageNumber="421">
<emphasis id="B940EAF0FFAFC80A0891FDC2FED7E445" bold="true" box="[199,299,537,562]" pageId="20" pageNumber="421">Identity.</emphasis>
Described as
<taxonomicName id="4C344D61FFAFC80A099DFDC1FD81E446" box="[459,637,537,561]" class="Amphibia" family="Ranidae" genus="Rana" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="inguinalis">
<emphasis id="B940EAF0FFAFC80A099DFDC1FD81E446" box="[459,637,537,561]" italics="true" pageId="20" pageNumber="421">Rana inguinalis</emphasis>
</taxonomicName>
from Anzahamaru by
<bibRefCitation id="EFA54B13FFAFC80A0B2BFDC2FBD7E445" author="Gunther" box="[893,1067,537,562]" pageId="20" pageNumber="421" refString="Gunther, A. (1877) Descriptions of some new species of reptiles from Madagascar. Annals and Magazine of Natural History, Series 4, 19, 313 - 317." type="journal article" year="1877">Günther (1877)</bibRefCitation>
and first considered to represent a junior synonym of
<taxonomicName id="4C344D61FFAFC80A09DAFDE5FD8BE422" box="[396,631,573,597]" class="Amphibia" family="Mantellidae" genus="Aglyptodactylus" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="madagascariensis">
<emphasis id="B940EAF0FFAFC80A09DAFDE5FD8BE422" box="[396,631,573,597]" italics="true" pageId="20" pageNumber="421">A. madagascariensis</emphasis>
</taxonomicName>
by
<bibRefCitation id="EFA54B13FFAFC80A0AF1FDE6FC88E421" author="Boulenger" box="[679,884,573,598]" pageId="20" pageNumber="421" refString="Boulenger, G. A. (1918) On the Madagascar frogs of the genus Mantidactylus Blgr. Proceedings of the Zoological Society London, 19, 257 - 261." type="journal article" year="1918">Boulenger (1918)</bibRefCitation>
. According to
<bibRefCitation id="EFA54B13FFAFC80A0C76FDE6FB2FE421" author="Gunther" box="[1056,1235,573,598]" pageId="20" pageNumber="421" refString="Gunther, A. (1877) Descriptions of some new species of reptiles from Madagascar. Annals and Magazine of Natural History, Series 4, 19, 313 - 317." type="journal article" year="1877">Günther (1877)</bibRefCitation>
, the
<typeStatus id="548F8840FFAFC80A0D58FDE5FAC2E421" box="[1294,1342,574,598]" pageId="20" pageNumber="421">type</typeStatus>
locality Anzahamaru most probably corresponds to a village close to Mahanoro, south of Tamatave (= Toamasina). This locality was later suggested to correspond to Anjahamaro, a locality close to Mahanoro by
<bibRefCitation id="EFA54B13FFAFC80A0CF3FD5EFAB3E4EA" author="Brygoo" box="[1189,1359,645,669]" pageId="20" pageNumber="421" refString="Brygoo, E. R. (1990) Toponymie herpetologique Malgache. Stations - Recoltes - Terra typica des Reptiles de Madagascar. Unpublished manuscript, MNHN, Paris. [unkown pagination]" type="book" year="1990">Brygoo (1990)</bibRefCitation>
. Later,
<bibRefCitation id="EFA54B13FFAFC80A08C1FD73FDADE4B6" author="Blommers-Schlosser" box="[151,593,680,705]" pageId="20" pageNumber="421" refString="Blommers-Schlosser, R. M. A. &amp; Blanc, C. P. (1991) Amphibiens (premiere partie). Faune de Madagascar, 75 (1), 1 - 379." type="journal article" year="1991">Blommers-Schlösser &amp; Blanc (1991)</bibRefCitation>
by mistake called this locality Anzahamani in their
<taxonomicName id="4C344D61FFAFC80A0CB1FD73FA60E4B7" box="[1255,1436,680,704]" class="Amphibia" family="Mantellidae" genus="Aglyptodactylus" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="genus">
<emphasis id="B940EAF0FFAFC80A0CB1FD73FA60E4B7" box="[1255,1436,680,704]" italics="true" pageId="20" pageNumber="421">Aglyptodactylus</emphasis>
</taxonomicName>
distribution map and placed it at the east coast, but considerably north of Toamasina. According to the information provided in the literature, we here consider Anzahamaru (= Anjahamaro) to most probably represent a locality very close to Mahanoro.
</paragraph>
<paragraph id="8B8B36E2FFAFC80A0891FCECFAA9E2C3" blockId="20.[151,1437,215,2027]" pageId="20" pageNumber="421">
All samples from this area of the east coast corresponded to the coastal clade, with two strongly divergent sublineages, one from the Betampona/Tampolo area and one from south-eastern localities. It might be speculated that the Anzahamaru population probably belongs to the Betampona/Tampolo sub-lineage since these three sites are located north of the Mangoro river estuary which is known to act as a barrier to dispersal for some coastal frog lineages (e.g.,
<bibRefCitation id="EFA54B13FFAFC80A096FFC1DFDEAE5A8" author="Gehring" box="[313,534,966,991]" pageId="20" pageNumber="421" refString="Gehring, P. S., Pabijan, M., Randrianirina, J. E., Glaw, F., &amp; Vences, M. (2012). The influence of riverine barriers on phylogeographic patterns of Malagasy reed frogs (Heterixalus). Molecular phylogenetics and evolution, 64 (3), 618 - 632." type="journal article" year="2012">
Gehring
<emphasis id="B940EAF0FFAFC80A09CDFC1CFE30E5A9" box="[411,460,966,990]" italics="true" pageId="20" pageNumber="421">et al</emphasis>
. 2012
</bibRefCitation>
). The comparatively small size of the
<typeStatus id="548F8840FFAFC80A0BE8FC1DFBDCE5A8" box="[958,1056,966,991]" pageId="20" pageNumber="421" type="holotype">holotype</typeStatus>
(probably a male; Table 3) agrees well with other specimens of the coastal clade (e.g.,
<collectionCode id="ED25AE27FFAFC80A0AA1FC31FCCDE275" box="[759,817,1002,1026]" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="20" pageNumber="421">ZSM</collectionCode>
183/2002 from Pic St. Louis near Tolagnaro), and its relative head length (0.398) fits well with the values of other coastal specimens (see section on morphological differentiation above). We therefore assign this nomen to the populations of the coastal clade. Due to the strong differentiation among the three sub-lineages here assigned to
<taxonomicName id="4C344D61FFAFC80A0B07FB8DFC1DE21A" box="[849,993,1109,1133]" class="Amphibia" family="Ranidae" genus="Rana" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="inguinalis">
<emphasis id="B940EAF0FFAFC80A0B07FB8DFC1DE21A" box="[849,993,1109,1133]" italics="true" pageId="20" pageNumber="421">A. inguinalis</emphasis>
</taxonomicName>
(Marojejy vs. Tampolo/Betampona vs. south-east) it cannot be excluded that this is indeed a species complex warranting further subdivision, in which case it would be necessary to ascertain the identity of the Anzahamaru population by new collections in this area.
</paragraph>
<paragraph id="8B8B36E2FFAFC80A0891FB1BFD91E086" blockId="20.[151,1437,215,2027]" pageId="20" pageNumber="421">
<emphasis id="B940EAF0FFAFC80A0891FB1BFD93E2AE" bold="true" box="[199,623,1216,1241]" pageId="20" pageNumber="421">Morphology of reference specimen.</emphasis>
<collectionCode id="ED25AE27FFAFC80A0A21FB1AFD4DE2AE" box="[631,689,1217,1241]" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="20" pageNumber="421">ZSM</collectionCode>
183/2002, adult male (
<figureCitation id="130F2A67FFAFC80A0BEFFB1BFBF6E2AE" box="[953,1034,1216,1241]" captionStart="FIGURE 11" captionStartId="21.[151,250,1167,1189]" captionTargetBox="[211,1376,326,1150]" captionTargetId="figure@21.[211,1377,326,1150]" captionTargetPageId="21" captionText="FIGURE 11. Photographs in life of Aglyptodactylus inguinalis: (a, b) dorsolateral and ventral views of a male specimen from the base of Pic St Louis near Tolagnaro (ZSM 183 / 2002); (c, d) dorsolateral and ventral views of a female specimen from Marojejy (ZSM 5213 / 2005)." httpUri="https://zenodo.org/record/240847/files/figure.png" pageId="20" pageNumber="421">Fig. 11</figureCitation>
a, b), as indicated by brown nuptial pads on the first finger. Intercalary element between ultimate and penultimate phalanges presumably present given the typical bent-down pattern of toe tips. Body slender; head slightly broader than body; snout pointed in dorsal view, nostrils directed dorsolaterally, distinctly nearer to snout tip than to eye; canthus rostralis concave, loreal region distinctly concave, tympanum distinct, tympanum round, its horizontal diameter 63% of eye diameter; distinct supratympanic fold; tongue oval bifid posteriorly. Vomerine teeth distinct in two well separated, round patches, the diameter of the group of vomerine teeth about the same diameter as choanae. Arms slender, fingers with distinct single subarticular tubercles, outer metacarpal tubercle oval, slightly elevated; inner metacarpal tubercle prominent and elliptical; fingers without webbing; comparative finger length 2=4&lt;1&lt;3, slightly developed terminal finger disks, without circummarginal grooves; limbs slender, tibiotarsal articulation reaches widely beyond snout tip if limb is adpressed along body; foot with an elongated inner metatarsal tubercle, no recognizable outer metatarsal tubercle, subarticular tubercle distinct, single, rounded; terminal toe disks slightly developed, lateral metatarsalia deeply separated, comparative toe length 1&lt;2&lt;3&lt;5&lt;4. Webbing formula 1(1),
<date id="FF8A1022FFAFC80A0C88F9B6FB08E0F1" box="[1246,1268,1645,1670]" pageId="20" pageNumber="421">2i</date>
(1.5), 2e (0.5),
<date id="FF8A1022FFAFC80A08C1F94AFF50E0DD" box="[151,172,1681,1706]" pageId="20" pageNumber="421">3i</date>
(2), 3e (1),
<date id="FF8A1022FFAFC80A0964F94AFEBBE0DD" box="[306,327,1681,1706]" pageId="20" pageNumber="421">4i</date>
(2), 4e (1.5), 5 (0.5). Skin on the upper surface smooth, throat smooth, chest and anterior throat tip with keratinized spicules. Venter slightly granular and covered with similar but smaller keratinized spicules. For morphological measurements see Table 3.
</paragraph>
<paragraph id="8B8B36E2FFAFC80B0891F927FB4DE6A4" blockId="20.[151,1437,215,2027]" lastBlockId="21.[151,1436,151,284]" lastPageId="21" lastPageNumber="422" pageId="20" pageNumber="421">
Colour of reference specimen (
<collectionCode id="ED25AE27FFAFC80A0A71F926FD9EE162" box="[551,610,1789,1813]" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="20" pageNumber="421">ZSM</collectionCode>
183/2002) in preservative (after 13 years in alcohol): head, dorsum, and dorsal surfaces of limbs brown. Inguinal region with a small and sharply bordered black marking on each side. Flanks greyish. Loreal and temporal region greyish, tympanum brown. A fine black line below the supratympanic fold and a triangular blotch posterior to tympanum. A blackish line running from the anterior margin of the eye to tip of snout, expanding to triangular shape at mid-level between eye and nostril. Upper lip whitish. Dorsal surface of hands whitish covered with minute brown spotting, dorsal surface of feet cream; thighs and shanks with brown crossbands. Posterior surfaces of thighs without a reticulated pattern. Cloacal region dark brown. Ventral surfaces of feet dark brown. Arms greyish-brown with some irregular brown markings. Iris blackish, pupil grey. Belly and throat cream except for the brown spicules, anterior venter and ventral surface of arms cream.
</paragraph>
<paragraph id="8B8B36E2FFAEC80B0891FF3BFAF6E76B" blockId="21.[151,1436,151,284]" pageId="21" pageNumber="422">
In life, the general pattern was the same as in preservative, except for brown dorsal surfaces exhibiting a yellowish tint (
<figureCitation id="130F2A67FFAEC80B0969FED8FE73E76B" box="[319,399,259,284]" captionStart="FIGURE 11" captionStartId="21.[151,250,1167,1189]" captionTargetBox="[211,1376,326,1150]" captionTargetId="figure@21.[211,1377,326,1150]" captionTargetPageId="21" captionText="FIGURE 11. Photographs in life of Aglyptodactylus inguinalis: (a, b) dorsolateral and ventral views of a male specimen from the base of Pic St Louis near Tolagnaro (ZSM 183 / 2002); (c, d) dorsolateral and ventral views of a female specimen from Marojejy (ZSM 5213 / 2005)." httpUri="https://zenodo.org/record/240847/files/figure.png" pageId="21" pageNumber="422">Fig. 11</figureCitation>
a) and the venter and throat being bright yellow with brown spicules (
<figureCitation id="130F2A67FFAEC80B0CCAFED8FB11E76B" box="[1180,1261,259,284]" captionStart="FIGURE 11" captionStartId="21.[151,250,1167,1189]" captionTargetBox="[211,1376,326,1150]" captionTargetId="figure@21.[211,1377,326,1150]" captionTargetPageId="21" captionText="FIGURE 11. Photographs in life of Aglyptodactylus inguinalis: (a, b) dorsolateral and ventral views of a male specimen from the base of Pic St Louis near Tolagnaro (ZSM 183 / 2002); (c, d) dorsolateral and ventral views of a female specimen from Marojejy (ZSM 5213 / 2005)." httpUri="https://zenodo.org/record/240847/files/figure.png" pageId="21" pageNumber="422">Fig. 11</figureCitation>
b).
</paragraph>
</subSubSection>
<caption id="DF4B666AFFAEC80B08C1FB54FE44E294" httpUri="https://zenodo.org/record/240847/files/figure.png" pageId="21" pageNumber="422" targetBox="[211,1376,326,1150]" targetPageId="21">
<paragraph id="8B8B36E2FFAEC80B08C1FB54FE44E294" blockId="21.[151,1435,1167,1251]" pageId="21" pageNumber="422">
<emphasis id="B940EAF0FFAEC80B08C1FB54FEE5E2D2" bold="true" box="[151,281,1167,1189]" pageId="21" pageNumber="422">FIGURE 11</emphasis>
. Photographs in life of
<taxonomicName id="4C344D61FFAEC80B0A5BFB54FCE0E2D2" box="[525,796,1167,1189]" class="Amphibia" family="Mantellidae" genus="Aglyptodactylus" kingdom="Animalia" order="Anura" pageId="21" pageNumber="422" phylum="Chordata" rank="species" species="inguinalis">
<emphasis id="B940EAF0FFAEC80B0A5BFB54FCE0E2D2" box="[525,796,1167,1189]" italics="true" pageId="21" pageNumber="422">Aglyptodactylus inguinalis</emphasis>
</taxonomicName>
: (a,b) dorsolateral and ventral views of a male specimen from the base of Pic St Louis near Tolagnaro (ZSM 183/2002); (c,d) dorsolateral and ventral views of a female specimen from Marojejy (ZSM 5213/2005).
</paragraph>
</caption>
<subSubSection id="C32E6569FFAEC8080891FAD4FEE3E36A" lastPageId="22" lastPageNumber="423" pageId="21" pageNumber="422" type="description">
<paragraph id="8B8B36E2FFAEC80B0891FAD4FD13E03F" blockId="21.[151,1437,1295,2004]" pageId="21" pageNumber="422">
<emphasis id="B940EAF0FFAEC80B0891FAD4FEC7E35F" bold="true" box="[199,315,1295,1320]" pageId="21" pageNumber="422">Variation.</emphasis>
For variation in morphological measurements see Table 3. There is a considerable amount of variation in colour pattern among specimens collected throughout the huge supposed range of this species. This mainly refers to the extent and distribution of dark markings, flecks and spots on flanks and blotches and stripes in the canthal and tympanal regions of the head. These vary from being very few, faint and indistinct to numerous and/or dark and sharply outlined. However, in life, brown dorsal markings seem to be rather indistinct in most specimens. In some specimens, the iris is golden with some dark brown reticulation in both, upper and lower iris area (
<figureCitation id="130F2A67FFAEC80B0883FA3CFED8E077" box="[213,292,1511,1536]" captionStart="FIGURE 11" captionStartId="21.[151,250,1167,1189]" captionTargetBox="[211,1376,326,1150]" captionTargetId="figure@21.[211,1377,326,1150]" captionTargetPageId="21" captionText="FIGURE 11. Photographs in life of Aglyptodactylus inguinalis: (a, b) dorsolateral and ventral views of a male specimen from the base of Pic St Louis near Tolagnaro (ZSM 183 / 2002); (c, d) dorsolateral and ventral views of a female specimen from Marojejy (ZSM 5213 / 2005)." httpUri="https://zenodo.org/record/240847/files/figure.png" pageId="21" pageNumber="422">Fig. 11</figureCitation>
a), whereas in other specimens the lower iris area can be distinctly darker (
<figureCitation id="130F2A67FFAEC80B0C33FA3CFB49E077" box="[1125,1205,1511,1536]" captionStart="FIGURE 11" captionStartId="21.[151,250,1167,1189]" captionTargetBox="[211,1376,326,1150]" captionTargetId="figure@21.[211,1377,326,1150]" captionTargetPageId="21" captionText="FIGURE 11. Photographs in life of Aglyptodactylus inguinalis: (a, b) dorsolateral and ventral views of a male specimen from the base of Pic St Louis near Tolagnaro (ZSM 183 / 2002); (c, d) dorsolateral and ventral views of a female specimen from Marojejy (ZSM 5213 / 2005)." httpUri="https://zenodo.org/record/240847/files/figure.png" pageId="21" pageNumber="422">Fig. 11</figureCitation>
c). Given the genetic differentiation within
<taxonomicName id="4C344D61FFAEC80B09C4F9D7FDD8E053" box="[402,548,1548,1572]" class="Amphibia" family="Mantellidae" genus="Aglyptodactylus" kingdom="Animalia" order="Anura" pageId="21" pageNumber="422" phylum="Chordata" rank="species" species="inguinalis">
<emphasis id="B940EAF0FFAEC80B09C4F9D7FDD8E053" box="[402,548,1548,1572]" italics="true" pageId="21" pageNumber="422">A. inguinalis</emphasis>
</taxonomicName>
and the small number of voucher specimens examined, more morphological variation than currently documented can be expected.
</paragraph>
<paragraph id="8B8B36E2FFAEC80B0891F988FB96E1FB" blockId="21.[151,1437,1295,2004]" pageId="21" pageNumber="422">
<emphasis id="B940EAF0FFAEC80B0891F988FEFCE01B" bold="true" box="[199,256,1619,1644]" pageId="21" pageNumber="422">Call.</emphasis>
The call of
<taxonomicName id="4C344D61FFAEC80B09DAF98FFDE0E01B" box="[396,540,1620,1644]" class="Amphibia" family="Mantellidae" genus="Aglyptodactylus" kingdom="Animalia" order="Anura" pageId="21" pageNumber="422" phylum="Chordata" rank="species" species="inguinalis">
<emphasis id="B940EAF0FFAEC80B09DAF98FFDE0E01B" box="[396,540,1620,1644]" italics="true" pageId="21" pageNumber="422">A. inguinalis</emphasis>
</taxonomicName>
, recorded at Tolagnaro, differs remarkably from calls of other
<taxonomicName id="4C344D61FFAEC80B0CB1F98FFA60E01B" box="[1255,1436,1620,1644]" class="Amphibia" family="Mantellidae" genus="Aglyptodactylus" kingdom="Animalia" order="Anura" pageId="21" pageNumber="422" phylum="Chordata" rank="genus">
<emphasis id="B940EAF0FFAEC80B0CB1F98FFA60E01B" box="[1255,1436,1620,1644]" italics="true" pageId="21" pageNumber="422">Aglyptodactylus</emphasis>
</taxonomicName>
species in being composed of short and longer notes combined in a somewhat irregular pattern. A typical call is shown in
<figureCitation id="130F2A67FFAEC80B0951F940FE8BE0C3" box="[263,375,1691,1716]" captionStart="FIGURE 12" captionStartId="22.[151,250,1156,1178]" captionTargetBox="[151,1436,193,1134]" captionTargetId="figure@22.[151,1436,193,1134]" captionTargetPageId="22" captionText="FIGURE 12. Audiospectrogram (above) and oscillogram (below) of an advertisement call of Aglyptodactylus inguinalis, recorded at the base of Pic St. Louis near Tolagnaro." httpUri="https://zenodo.org/record/240848/files/figure.png" pageId="21" pageNumber="422">Figure 12</figureCitation>
, showing the following alternation of notes: short, long, short, long, short, short, long, short, short, long, short, short, short, long, short, short, short, short. This note pattern is similar but not identical in all calls and may also vary slightly among calls of the same individual. All calls show overall amplitude modulation with almost constantly increasing call energy from the beginning to the end of the call. Both short and long notes are pulsed, with distinctly higher pulse rate in short notes. Number of pulses per note is very similar in short and long notes, and note duration seems to result from modulation of the pulse repetition rate. In some calls, pulses within short notes are indistinct or partly fused. For numerical call parameters refer to Table 2.
</paragraph>
<paragraph id="8B8B36E2FFAEC80B0891F84CFA61E1A3" blockId="21.[151,1437,1295,2004]" pageId="21" pageNumber="422">
The release call of
<taxonomicName id="4C344D61FFAEC80B09EEF843FCB8E1C7" box="[440,836,1944,1968]" class="Amphibia" family="Mantellidae" genus="Aglyptodactylus" kingdom="Animalia" order="Anura" pageId="21" pageNumber="422" phylum="Chordata" rank="species" species="madagascariensis">
<emphasis id="B940EAF0FFAEC80B09EEF843FCB8E1C7" box="[440,836,1944,1968]" italics="true" pageId="21" pageNumber="422">Aglyptodactylus madagascariensis</emphasis>
</taxonomicName>
from Betampona provided by Rosa
<emphasis id="B940EAF0FFAEC80B0D53F842FAB9E1C7" box="[1285,1349,1944,1968]" italics="true" pageId="21" pageNumber="422">et al.</emphasis>
(2011) probably corresponds to
<taxonomicName id="4C344D61FFAEC80B09FAF867FDC7E1A3" box="[428,571,1980,2004]" class="Amphibia" family="Mantellidae" genus="Aglyptodactylus" kingdom="Animalia" order="Anura" pageId="21" pageNumber="422" phylum="Chordata" rank="species" species="inguinalis">
<emphasis id="B940EAF0FFAEC80B09FAF867FDC7E1A3" box="[428,571,1980,2004]" italics="true" pageId="21" pageNumber="422">A. inguinalis</emphasis>
</taxonomicName>
, given the sequence data provided by
<bibRefCitation id="EFA54B13FFAEC80B0BBFF867FB4DE1A3" box="[1001,1201,1980,2004]" pageId="21" pageNumber="422" refString="Rosa, G. M., Andreone, F., Crottini, A., Hauswald, J. S., Noel, J., Rabibisoa, N. H., Randriambahiniarime, M. O., Rebelo, R. &amp; Raxworthy, C. J. (2012) The amphibians of the relict Betampona low-elevation rainforest, eastern Madagascar: an application of the integrative taxonomy approach to biodiversity assessments. Biodiversity and Conservation, 21, 1531 - 1559. http: // dx. doi. org / 10.1007 / s 10531 - 012 - 0262 - x" type="journal article">
Rosa
<emphasis id="B940EAF0FFAEC80B0C71F867FB9CE1A3" box="[1063,1120,1980,2004]" italics="true" pageId="21" pageNumber="422">et al.</emphasis>
(2012)
</bibRefCitation>
and included herein.
</paragraph>
<caption id="DF4B666AFFADC80808C1FB5FFD56E2CE" httpUri="https://zenodo.org/record/240848/files/figure.png" pageId="22" pageNumber="423" targetBox="[151,1436,193,1134]" targetPageId="22">
<paragraph id="8B8B36E2FFADC80808C1FB5FFD56E2CE" blockId="22.[151,1435,1156,1209]" pageId="22" pageNumber="423">
<emphasis id="B940EAF0FFADC80808C1FB5FFEE1E2ED" bold="true" box="[151,285,1156,1178]" pageId="22" pageNumber="423">FIGURE 12</emphasis>
. Audiospectrogram (above) and oscillogram (below) of an advertisement call of
<taxonomicName id="4C344D61FFADC8080CD5FB5FFA69E2ED" box="[1155,1429,1156,1178]" class="Amphibia" family="Mantellidae" genus="Aglyptodactylus" kingdom="Animalia" order="Anura" pageId="22" pageNumber="423" phylum="Chordata" rank="species" species="inguinalis">
<emphasis id="B940EAF0FFADC8080CD5FB5FFA69E2ED" box="[1155,1429,1156,1178]" italics="true" pageId="22" pageNumber="423">Aglyptodactylus inguinalis</emphasis>
</taxonomicName>
, recorded at the base of Pic St. Louis near Tolagnaro.
</paragraph>
</caption>
<paragraph id="8B8B36E2FFADC8080891FB3BFEE3E36A" blockId="22.[151,1437,1248,1489]" pageId="22" pageNumber="423">
<emphasis id="B940EAF0FFADC8080891FB3BFE78E28E" bold="true" box="[199,388,1248,1273]" pageId="22" pageNumber="423">Natural history.</emphasis>
To our knowledge, there are no available data particularly referring to populations assigned to this species.
</paragraph>
</subSubSection>
<subSubSection id="C32E6569FFADC8080891FAF3FDCDE3A6" pageId="22" pageNumber="423" type="distribution">
<paragraph id="8B8B36E2FFADC8080891FAF3FDCDE3A6" blockId="22.[151,1437,1248,1489]" pageId="22" pageNumber="423">
<emphasis id="B940EAF0FFADC8080891FAF3FE9CE336" bold="true" box="[199,352,1320,1345]" pageId="22" pageNumber="423">Distribution.</emphasis>
As defined here based on genetics, bioacoustics and
<typeStatus id="548F8840FFADC8080B91FAF2FC0BE336" box="[967,1015,1321,1345]" pageId="22" pageNumber="423">type</typeStatus>
information, we tentatively consider the following populations to represent
<taxonomicName id="4C344D61FFADC8080A2EFA97FC44E312" authority="Anzahamaru" authorityName="Anzahamaru" box="[632,952,1356,1381]" class="Amphibia" family="Mantellidae" genus="Aglyptodactylus" kingdom="Animalia" order="Anura" pageId="22" pageNumber="423" phylum="Chordata" rank="species" species="inguinalis">
<emphasis id="B940EAF0FFADC8080A2EFA97FCECE313" box="[632,784,1356,1380]" italics="true" pageId="22" pageNumber="423">A. inguinalis</emphasis>
: Anzahamaru
</taxonomicName>
(
<typeStatus id="548F8840FFADC8080B86FA97FBFEE313" box="[976,1026,1356,1380]" pageId="22" pageNumber="423">type</typeStatus>
locality), Betampona, Ifanadiana, Manombo, Marojejy, Ranomafana village, Tampolo, and Tolagnaro (
<figureCitation id="130F2A67FFADC8080BF4FAABFC15E3FF" box="[930,1001,1392,1417]" captionStart="FIGURE 2" captionStartId="5.[151,250,1820,1842]" captionTargetBox="[248,1326,193,1777]" captionTargetId="figure@5.[236,1326,193,1778]" captionTargetPageId="5" captionText="FIGURE 2. Map showing the reliably known distribution ranges of all six species of Aglyptodactylus. Note that many more records under the name A. madagascariensis are available mainly from eastern Madagascar (e. g., Blommers-Schlösser &amp; Blanc 1991); the map shown here only includes records verified by molecular data (except Anzahamaru, the type locality of A. inguinalis, and Kirindy, the type locality of A. securifer). The Marovato record of A. australis is based on sequence data alone and in need of confirmation. The different blue colours of A. inguinalis records symbolize their substantial genetic differentiation. The map shows the remaining primary vegetation of Madagascar (www. vegmad. org), green colours indicating rainforest, reddish colours deciduous dry forest, and orange colour arid spiny forest." httpUri="https://zenodo.org/record/240838/files/figure.png" pageId="22" pageNumber="423">Fig. 2</figureCitation>
). These localities are all located along the east coast of
<collectingCountry id="F3237672FFADC8080907FA4FFE24E3DA" box="[337,472,1428,1453]" name="Madagascar" pageId="22" pageNumber="423">Madagascar</collectingCountry>
at elevations from sea level up to
<quantity id="4CCC9B07FFADC8080B0EFA4FFC63E3DB" box="[856,927,1428,1452]" metricMagnitude="2" metricUnit="m" metricValue="4.81" pageId="22" pageNumber="423" unit="m" value="481.0">481 m</quantity>
a.s.l. (highest locality Marojejy at a campsite locally known as &quot;Camp
<taxonomicName id="4C344D61FFADC80809E7FA63FDE0E3A6" box="[433,540,1464,1489]" class="Amphibia" family="Mantellidae" genus="Mantella" kingdom="Animalia" order="Anura" pageId="22" pageNumber="433" phylum="Chordata" rank="genus">Mantella</taxonomicName>
&quot;).
</paragraph>
</subSubSection>
</treatment>
</document>
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