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<document id="ADA503D596A294CC0EA5C07CDD8BDAF4" ID="10.11646/zootaxa.4273.3.1SLASH11254" ID-CLB-Dataset="32906" ID-DOI="10.11646/zootaxa.4273.3.1" ID-GBIF-Dataset="c96e6bc4-729c-469c-9f1e-1f301d83357d" ID-ISSN="1175-5326" ID-Zenodo-Dep="803049" ID-ZooBank="4D576A21-DA71-41C2-9408-96BB24B9CFD" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1496734297629" checkinUser="plazi" docAuthor="Scherz, Mark D., Hawlitschek, Oliver, Andreone, Franco, Rakotoarison, Andolalao, Vences, Miguel &amp; Glaw, Frank" docDate="2017" docId="03DE3444FFFEFFF0D4D4FA1EFBF6F82C" docLanguage="en" docName="zootaxa.4273.3.1SLASH11254.pdf" docOrigin="Zootaxa 4273 (3)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Rhombophryne guentherpetersi Guibe 1974" docType="treatment" docVersion="7" lastPageNumber="312" masterDocId="FFE74C3CFFF8FFFBD443FFA7FF80FF92" masterDocTitle="A review of the taxonomy and osteology of the Rhombophryne serratopalpebrosa species group (Anura: Microhylidae) from Madagascar, with comments on the value of volume rendering of micro-CT data to taxonomists" masterLastPageNumber="340" masterPageNumber="301" pageNumber="307" updateTime="1698440179113" updateUser="ExternalLinkService">
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<mods:title id="0F17FA3DB505DF72C0E4A34735A76B40">A review of the taxonomy and osteology of the Rhombophryne serratopalpebrosa species group (Anura: Microhylidae) from Madagascar, with comments on the value of volume rendering of micro-CT data to taxonomists</mods:title>
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<treatment id="03DE3444FFFEFFF0D4D4FA1EFBF6F82C" ID-DOI="http://doi.org/10.5281/zenodo.6039695" ID-GBIF-Taxon="131812312" ID-Zenodo-Dep="6039695" LSID="urn:lsid:plazi:treatment:03DE3444FFFEFFF0D4D4FA1EFBF6F82C" httpUri="http://treatment.plazi.org/id/03DE3444FFFEFFF0D4D4FA1EFBF6F82C" lastPageId="11" lastPageNumber="312" pageId="6" pageNumber="307">
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<paragraph id="8BC88552FFFEFFFDD4D4FA1EFD47FA41" blockId="6.[151,1210,1465,1560]" box="[151,711,1465,1491]" pageId="6" pageNumber="307">
<heading id="D080323EFFFEFFFDD4D4FA1EFD47FA41" bold="true" box="[151,711,1465,1491]" fontSize="11" level="1" pageId="6" pageNumber="307" reason="1">
<taxonomicName id="4C77FED1FFFEFFFDD4D4FA1EFD47FA41" ID-CoL="4SK5L" authority="Guibe, 1974" authorityName="Guibe" authorityYear="1974" box="[151,711,1465,1491]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="6" pageNumber="307" phylum="Chordata" rank="species" species="guentherpetersi">
<emphasis id="B9035940FFFEFFFDD4D4FA1EFD47FA41" bold="true" box="[151,711,1465,1491]" pageId="6" pageNumber="307">
<emphasis id="B9035940FFFEFFFDD4D4FA1EFD96FA41" bold="true" box="[151,534,1465,1491]" italics="true" pageId="6" pageNumber="307">Rhombophryne guentherpetersi</emphasis>
(
<bibRefCitation id="EFE6F8A3FFFEFFFDD665FA1EFD3FFA41" author="Guibe" box="[550,703,1465,1491]" pageId="6" pageNumber="307" refString="Guibe, J. (1974) Batraciens nouveaux de Madagascar. Bulletin du Museum national d'Histoire Naturelle, 3, 1170 - 1192." type="journal article" year="1974">Guibé, 1974</bibRefCitation>
)
</emphasis>
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</heading>
</paragraph>
<paragraph id="8BC88552FFFEFFFDD4D4FA7BFEE2F98A" blockId="6.[151,1210,1465,1560]" pageId="6" pageNumber="307">
Common name: Tsaratanana saw-browed diamond frog (modified from
<bibRefCitation id="EFE6F8A3FFFEFFFDD7FCFA7BFB32FA67" author="Frank" box="[959,1202,1500,1525]" pageId="6" pageNumber="307" refString="Frank, N. &amp; Ramus, E. (1995) Complete Guide to Scientific and Common Names of Amphibians and Reptiles of the World. N. G. Publishing Inc., Pottsville, Pennsylvania, 377 pp." type="book" year="1995">Frank &amp; Ramus 1995</bibRefCitation>
) (
<figureCitation id="134C99D7FFFEFFFDD4DCFA58FE84F98A" box="[159,260,1535,1560]" captionStart-0="FIGURE 4" captionStart-1="FIGURE 5" captionStart-2="FIGURE 6" captionStart-3="FIGURE 7" captionStartId-0="7.[151,250,1697,1719]" captionStartId-1="8.[151,250,1879,1901]" captionStartId-2="9.[151,250,1925,1947]" captionStartId-3="10.[151,250,1056,1078]" captionTargetBox-0="[151,1436,414,1675]" captionTargetBox-1="[151,1436,949,1858]" captionTargetBox-2="[170,1414,193,1885]" captionTargetBox-3="[174,1403,201,1035]" captionTargetId-0="figure@7.[151,1436,414,1675]" captionTargetId-1="figure@8.[151,1436,949,1858]" captionTargetId-2="figure@9.[170,1414,193,1885]" captionTargetId-3="figure@10.[164,1422,193,1035]" captionTargetPageId-0="7" captionTargetPageId-1="8" captionTargetPageId-2="9" captionTargetPageId-3="10" captionText-0="FIGURE 4. Photographs of the holotypes of the species treated in this manuscript, in dorsal (top row) and ventral (bottom row) views." captionText-1="FIGURE 5. Comparative morphology of the heads, feet, and hands of members of the Rhombophryne serratopalpebrosa species group. All specimens are the holotypes of their respective species, except R. coronata, which is paratype ZSM 694 / 2001. Asterisks indicate mirrored images. Not to scale." captionText-2="FIGURE 6. Rhombophryne guentherpetersi in life. (a c) ZSM 607 / 2014 in (a) dorsolateral (with inset showing superciliary spines), (b) dorsal, and (c) ventral view; and (d, e) other specimens in dorsolateral view (assignment to field and collection numbers unknown)." captionText-3="FIGURE 7. Osteology of Rhombophryne guentherpetersi (MNHN 1953.165). Full skeleton in (a) dorsal, (b) ventral, and (c) lateral view; skull in (d) dorsal, (e) ventral, and (f) lateral view. Abbreviations: angspl = angulosplenial, col = columella, exoc = exoccipital, fpar = frontoparietal, max = maxilla, mmk = mentomeckelian, npl = neopalatine, pmx = premaxilla, povom = postchoanal vomer, proot = prootic, prvom = prechoanal vomer, prsph = parasphenoid, pter = pterygoid, qj = quadratojugal, smax = septomaxilla, spheth = sphenethmoid, sq = squamosal." httpUri-0="https://zenodo.org/record/803057/files/figure.png" httpUri-1="https://zenodo.org/record/803059/files/figure.png" httpUri-2="https://zenodo.org/record/803061/files/figure.png" httpUri-3="https://zenodo.org/record/803063/files/figure.png" pageId="6" pageNumber="307">Figs 47</figureCitation>
, S1S2)
</paragraph>
</subSubSection>
<subSubSection id="C36DD6D9FFFEFFFDD4D4F9EEFC9CF94A" pageId="6" pageNumber="307" type="reference_group">
<paragraph id="8BC88552FFFEFFFDD4D4F9EEFD1CF9CC" blockId="6.[151,1436,1608,1752]" box="[151,668,1608,1630]" pageId="6" pageNumber="307">
<treatmentCitationGroup id="AB67A27CFFFEFFFDD4D4F9EEFD1CF9CC" box="[151,668,1608,1630]" pageId="6" pageNumber="307">
<taxonomicName id="4C77FED1FFFEFFFDD4D4F9EEFDDCF9CC" ID-CoL="736MB" authority="Guibe, 1974: 1181" authorityName="Guibe" authorityPageNumber="1181" authorityYear="1974" box="[151,604,1608,1630]" class="Amphibia" family="Microhylidae" genus="Mantipus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="6" pageNumber="307" phylum="Chordata" rank="species" species="guentherpetersi">
<emphasis id="B9035940FFFEFFFDD4D4F9EEFE1CF9CC" box="[151,412,1608,1630]" italics="true" pageId="6" pageNumber="307">Mantipus guentherpetersi</emphasis>
<treatmentCitation id="0AD6A343FFFEFFFDD5E1F9EFFDDCF9CC" author="Guibe" box="[418,604,1608,1630]" page="1181" pageId="6" pageNumber="307" year="1974">
<bibRefCitation id="EFE6F8A3FFFEFFFDD5E1F9EFFDA2F9CC" author="Guibe" box="[418,546,1608,1630]" pageId="6" pageNumber="307" refString="Guibe, J. (1974) Batraciens nouveaux de Madagascar. Bulletin du Museum national d'Histoire Naturelle, 3, 1170 - 1192." type="journal article" year="1974">Guibé, 1974</bibRefCitation>
: 1181
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1182
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</paragraph>
<paragraph id="8BC88552FFFEFFFDD4D4F9C1FECAF909" blockId="6.[151,1436,1608,1752]" pageId="6" pageNumber="307">
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<taxonomicName id="4C77FED1FFFEFFFDD4D4F9C1FE64F9EE" ID-CoL="6VQ8D" box="[151,484,1638,1660]" class="Amphibia" family="Microhylidae" genus="Plethodontohyla" kingdom="Animalia" order="Anura" pageId="6" pageNumber="307" phylum="Chordata" rank="species" species="guentherpetersi">
<emphasis id="B9035940FFFEFFFDD4D4F9C1FE64F9EE" box="[151,484,1638,1660]" italics="true" pageId="6" pageNumber="307">Plethodontohyla guentherpetersi</emphasis>
</taxonomicName>
<treatmentCitation id="0AD6A343FFFEFFFDD640F9C1FC18F9EE" author="Blommers-Schlosser" box="[515,920,1638,1660]" page="56" pageId="6" pageNumber="307" year="1991">
<bibRefCitation id="EFE6F8A3FFFEFFFDD640F9C1FCF4F9EE" author="Blommers-Schlosser" box="[515,884,1638,1660]" pageId="6" pageNumber="307" refString="Blommers-Schlosser, R. M. A. &amp; Blanc, C. P. (1991) Amphibiens (premiere partie). Faune de Madagascar, 75, 1 - 397." type="journal article" year="1991">Blommers-Schlösser &amp; Blanc, 1991</bibRefCitation>
: 56
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57. First depicted in life under this name in
<bibRefCitation id="EFE6F8A3FFFEFFFDD123F9C1FEC6F909" pageId="6" pageNumber="307" refString="Stuart, S. N., Hoffmann, M., Chanson, J. S., Cox, N. A., Berridge, R. J., Ramani, P. &amp; Young, B. E. (2008) Threatened Amphibians of the World. Lynx Edicions, Barcelona; IUCN, Gland &amp; Conservation International, Arlington, Virginia, 758 pp." type="book">
Stuart
<emphasis id="B9035940FFFEFFFDD484F921FF7AF909" box="[199,250,1669,1691]" italics="true" pageId="6" pageNumber="307">et al.</emphasis>
(2008)
</bibRefCitation>
.
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</paragraph>
<paragraph id="8BC88552FFFEFFFDD4D4F903FC9CF94A" blockId="6.[151,1436,1608,1752]" pageId="6" pageNumber="307">
<treatmentCitationGroup id="AB67A27CFFFEFFFDD4D4F903FC9CF94A" pageId="6" pageNumber="307">
<taxonomicName id="4C77FED1FFFEFFFDD4D4F903FE56F928" ID-CoL="4SK5L" box="[151,470,1700,1722]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="6" pageNumber="307" phylum="Chordata" rank="species" species="guentherpetersi">
<emphasis id="B9035940FFFEFFFDD4D4F903FE56F928" box="[151,470,1700,1722]" italics="true" pageId="6" pageNumber="307">Rhombophryne guentherpetersi</emphasis>
</taxonomicName>
<treatmentCitation id="0AD6A343FFFEFFFDD5B3F903FC81F928" author="Glaw" box="[496,769,1700,1722]" page="118" pageId="6" pageNumber="307" year="2007">
<bibRefCitation id="EFE6F8A3FFFEFFFDD5B3F903FD50F928" author="Glaw" box="[496,720,1700,1722]" pageId="6" pageNumber="307" refString="Glaw, F. &amp; Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Vences &amp; Glaw Verlags GbR, Koln, 496 pp." type="book" year="2007">Glaw &amp; Vences, 2007</bibRefCitation>
: 118
</treatmentCitation>
. The photo provided on page 119 of this book is misattributed and depicts an undescribed species (Scherz
<emphasis id="B9035940FFFEFFFDD614F964FD0AF94A" box="[599,650,1730,1752]" italics="true" pageId="6" pageNumber="307">et al.</emphasis>
unpubl. data).
</treatmentCitationGroup>
</paragraph>
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<paragraph id="8BC88552FFFEFFFDD4D4F8A2FD71F8D0" blockId="6.[151,1436,1797,2038]" pageId="6" pageNumber="307">
<emphasis id="B9035940FFFEFFFDD4D4F8A2FE8BF88C" bold="true" box="[151,267,1797,1822]" pageId="6" pageNumber="307">Holotype.</emphasis>
MNHN 1953.165, an adult female specimen captured by an unknown collector on an unknown date on the Tsaratanana Massif at
<quantity id="4C8F28B7FFFEFFFDD5F8F88EFD90F8D3" box="[443,528,1833,1858]" metricMagnitude="3" metricUnit="m" metricValue="2.6" pageId="6" pageNumber="307" unit="m" value="2600.0">2600 m</quantity>
a.s.l. (
<figureCitation id="134C99D7FFFEFFFDD619F88EFD22F8D0" box="[602,674,1833,1858]" captionStart="FIGURE 4" captionStartId="7.[151,250,1697,1719]" captionTargetBox="[151,1436,414,1675]" captionTargetId="figure@7.[151,1436,414,1675]" captionTargetPageId="7" captionText="FIGURE 4. Photographs of the holotypes of the species treated in this manuscript, in dorsal (top row) and ventral (bottom row) views." httpUri="https://zenodo.org/record/803057/files/figure.png" pageId="6" pageNumber="307">Figs 4</figureCitation>
,
<figureCitation id="134C99D7FFFEFFFDD6EDF88EFD3EF8D0" box="[686,702,1833,1858]" captionStart="FIGURE 5" captionStartId="8.[151,250,1879,1901]" captionTargetBox="[151,1436,949,1858]" captionTargetId="figure@8.[151,1436,949,1858]" captionTargetPageId="8" captionText="FIGURE 5. Comparative morphology of the heads, feet, and hands of members of the Rhombophryne serratopalpebrosa species group. All specimens are the holotypes of their respective species, except R. coronata, which is paratype ZSM 694 / 2001. Asterisks indicate mirrored images. Not to scale." httpUri="https://zenodo.org/record/803059/files/figure.png" pageId="6" pageNumber="307">5</figureCitation>
, S2)
</paragraph>
<paragraph id="8BC88552FFFEFFFDD484F8EAFB8FF83C" blockId="6.[151,1436,1797,2038]" pageId="6" pageNumber="307">
<emphasis id="B9035940FFFEFFFDD484F8EAFEC6F8F4" bold="true" box="[199,326,1869,1894]" pageId="6" pageNumber="307">Paratypes.</emphasis>
MNHN 1953.165A and MNHN 1953.165B, a juvenile and adult female with the same collection data as the holotype.
<materialsCitation id="3B1F8F0FFFFEFFFDD5C5F8D5FB8BF83C" ID-GBIF-Occurrence="1563404814" collectionCode="MNHN, ORSTOM" collectorName="Charles P. &amp; Mission" pageId="6" pageNumber="307" specimenCode="MNHN 1973.592, MNHN 1973.593" specimenCount="1" typeStatus="holotype">
<specimenCode id="DBD12D29FFFEFFFDD5C5F8D5FDCCF818" box="[390,588,1905,1930]" collectionCode="MNHN" country="France" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34988" name="Museum National d'Histoire Naturelle" pageId="6" pageNumber="307">
<collectionCode id="ED661D97FFFEFFFDD5C5F8D5FE5BF81B" box="[390,475,1906,1929]" country="France" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34988" name="Museum National d'Histoire Naturelle" pageId="6" pageNumber="307">MNHN</collectionCode>
1973.592
</specimenCode>
and
<specimenCode id="DBD12D29FFFEFFFDD6C0F8D5FCCCF818" box="[643,844,1905,1930]" collectionCode="MNHN" country="France" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34988" name="Museum National d'Histoire Naturelle" pageId="6" pageNumber="307">
<collectionCode id="ED661D97FFFEFFFDD6C0F8D5FD58F81B" box="[643,728,1906,1929]" country="France" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34988" name="Museum National d'Histoire Naturelle" pageId="6" pageNumber="307">MNHN</collectionCode>
1973.593
</specimenCode>
, a subadult female and juvenile collected by
<collectorName id="2682E084FFFEFFFDD104F8D6FF2BF83F" pageId="6" pageNumber="307">Charles P.</collectorName>
Blanc in 1966 on
<collectorName id="2682E084FFFEFFFDD53BF832FE52F83C" box="[376,466,1941,1966]" pageId="6" pageNumber="307">Mission</collectorName>
<collectionCode id="ED661D97FFFEFFFDD599F831FDCEF83C" box="[474,590,1942,1966]" country="Bolivia" httpUri="http://grbio.org/cool/mdr4-z094" name="Office de la Recherche scientifique et Technique Outre-mer" pageId="6" pageNumber="307">ORSTOM</collectionCode>
from the same location as the
<typeStatus id="54CC3BF0FFFEFFFDD7E6F832FB8BF83C" box="[933,1035,1941,1966]" pageId="6" pageNumber="307" type="holotype">holotype</typeStatus>
</materialsCitation>
.
</paragraph>
<paragraph id="8BC88552FFFEFFFDD484F81EFA1CF867" blockId="6.[151,1436,1797,2038]" pageId="6" pageNumber="307">
<emphasis id="B9035940FFFEFFFDD484F81EFE3BF840" bold="true" box="[199,443,1977,2002]" pageId="6" pageNumber="307">Referred specimens.</emphasis>
ZSM 606/2014 (DRV 6220), a female, ZSM 607/2014 (DRV 6223), a male, and ZSM 608/2014 (DRV 6231), a female (Fig. S1), three adult specimens collected by D. Vieites, M. Vences, R.D.
</paragraph>
<paragraph id="8BC88552FFFFFFFCD4D4FF30FA04FEF8" blockId="7.[151,1437,151,362]" pageId="7" pageNumber="308">
<materialsCitation id="3B1F8F0FFFFFFFFCD4D4FF30FD59FF6C" ID-GBIF-Occurrence="1563404819" collectingDate="2010-06" collectionCode="ZSM" collectorName="Randrianiaina, F. M. &amp; Ratsoavina, S. &amp; Rasamison, A. &amp; Rakotoarison, E. &amp; Rajeriarison, F. &amp; Randrianasolo, F. &amp; T. Rajoafiarison" elevation="2503" latitude="-14.0801" location="Andranomadio" longLatPrecision="7" longitude="48.9854" municipality="Tsaratanana" pageId="7" pageNumber="308" specimenCode="ZSM 606" specimenCount="1">
<collectorName id="2682E084FFFFFFFCD4D4FF30FA1DFF3D" box="[151,1437,151,176]" pageId="7" pageNumber="308">Randrianiaina, F.M. Ratsoavina, S. Rasamison, A. Rakotoarison, E. Rajeriarison, F. Randrianasolo, F.</collectorName>
Randrianasolo and
<collectorName id="2682E084FFFFFFFCD52DFF67FDA4FF45" box="[366,548,190,215]" pageId="7" pageNumber="308">T. Rajoafiarison</collectorName>
,
<specimenCode id="DBD12D29FFFFFFFCD66CFF18FD18FF45" box="[559,664,190,215]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="7" pageNumber="308">
<collectionCode id="ED661D97FFFFFFFCD66CFF18FDE9FF45" box="[559,617,191,215]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="7" pageNumber="308">ZSM</collectionCode>
606
</specimenCode>
/2014 and 607/2014 on the 16th of
<collectingDate id="EF8D5A7AFFFFFFFCD056FF67FB06FF45" box="[1045,1158,191,215]" pageId="7" pageNumber="308" value="2010-06">
<date id="FFC9A392FFFFFFFCD056FF67FB06FF45" box="[1045,1158,191,215]" pageId="7" pageNumber="308" value="2010-06">June 2010</date>
</collectingDate>
at
<location id="8EA8D389FFFFFFFCD0EBFF19FAD0FF45" LSID="urn:lsid:plazi:treatment:03DE3444FFFEFFF0D4D4FA1EFBF6F82C:8EA8D389FFFFFFFCD0EBFF19FAD0FF45" box="[1192,1360,190,215]" latitude="-14.0801" longLatPrecision="7" longitude="48.9854" municipality="Tsaratanana" name="Andranomadio" pageId="7" pageNumber="308">Andranomadio</location>
(camp 4),
<collectingMunicipality id="6BAC1F28FFFFFFFCD4FCFF41FEC8FF6F" box="[191,328,230,253]" pageId="7" pageNumber="308">Tsaratanana</collectingMunicipality>
,
<geoCoordinate id="EE43E395FFFFFFFCD515FF41FE4EFF6C" box="[342,462,230,254]" direction="south" orientation="latitude" pageId="7" pageNumber="308" precision="5" value="-14.0801">14.0801°S</geoCoordinate>
,
<geoCoordinate id="EE43E395FFFFFFFCD59EFF42FDD8FF6C" box="[477,600,229,254]" direction="east" orientation="longitude" pageId="7" pageNumber="308" precision="5" value="48.9854">48.9854°E</geoCoordinate>
,
<elevation id="005A6261FFFFFFFCD625FF42FD3EFF6F" box="[614,702,229,254]" metricMagnitude="3" metricUnit="m" metricValue="2.503" pageId="7" pageNumber="308" unit="m" value="2503.0">
<quantity id="4C8F28B7FFFFFFFCD625FF42FD3EFF6F" box="[614,702,229,254]" metricMagnitude="3" metricUnit="m" metricValue="2.503" pageId="7" pageNumber="308" unit="m" value="2503.0">2503 m</quantity>
</elevation>
a
</materialsCitation>
.
<materialsCitation id="3B1F8F0FFFFFFFFCD69DFF42FC95FEB0" ID-GBIF-Occurrence="1563404815" collectingDate="2010-06" collectionCode="ZSM" elevation="2732" latitude="-14.0665" location="Tsaratanana mountain" longLatPrecision="7" longitude="48.9832" pageId="7" pageNumber="308" specimenCode="ZSM 608" specimenCount="1">
s.l.;
<specimenCode id="DBD12D29FFFFFFFCD74CFF41FCFBFF6C" box="[783,891,229,254]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="7" pageNumber="308">
<collectionCode id="ED661D97FFFFFFFCD74CFF41FCC8FF6C" box="[783,840,230,254]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="7" pageNumber="308">ZSM</collectionCode>
608
</specimenCode>
/2014 on the 18th of
<collectingDate id="EF8D5A7AFFFFFFFCD02EFF41FB63FF6C" box="[1133,1251,230,254]" pageId="7" pageNumber="308" value="2010-06">
<date id="FFC9A392FFFFFFFCD02EFF41FB63FF6C" box="[1133,1251,230,254]" pageId="7" pageNumber="308" value="2010-06">June 2010</date>
</collectingDate>
at a site on the
<location id="8EA8D389FFFFFFFCD4D4FEADFE0FFEB0" LSID="urn:lsid:plazi:treatment:03DE3444FFFEFFF0D4D4FA1EFBF6F82C:8EA8D389FFFFFFFCD4D4FEADFE0FFEB0" box="[151,399,265,290]" latitude="-14.0665" longLatPrecision="7" longitude="48.9832" name="Tsaratanana mountain" pageId="7" pageNumber="308">Tsaratanana mountain</location>
<geoCoordinate id="EE43E395FFFFFFFCD5D4FEAEFD90FEB0" box="[407,528,265,290]" direction="south" orientation="latitude" pageId="7" pageNumber="308" precision="5" value="-14.0665">14.0665°S</geoCoordinate>
,
<geoCoordinate id="EE43E395FFFFFFFCD65EFEADFD17FEB0" box="[541,663,266,290]" direction="east" orientation="longitude" pageId="7" pageNumber="308" precision="5" value="48.9832">48.9832°E</geoCoordinate>
,
<elevation id="005A6261FFFFFFFCD6E7FEADFD7BFEB3" box="[676,763,266,290]" metricMagnitude="3" metricUnit="m" metricValue="2.732" pageId="7" pageNumber="308" unit="m" value="2732.0">
<quantity id="4C8F28B7FFFFFFFCD6E7FEADFD7BFEB3" box="[676,763,266,290]" metricMagnitude="3" metricUnit="m" metricValue="2.732" pageId="7" pageNumber="308" unit="m" value="2732.0">2732 m</quantity>
</elevation>
a
</materialsCitation>
.s.l. A further three specimens from Andranomadio (with same collecting data as ZSM specimens from this site) were deposited in the UADBA collection: UADBA-A 60775 (DRV 6210) and UADBA-A 60776 (ZCMV 12401), adults; UADBA-A 60782 (ZCMV 12435), subadult.
</paragraph>
</subSubSection>
<caption id="DF08D5DAFFFFFFFCD4D4F906FF59F944" httpUri="https://zenodo.org/record/803057/files/figure.png" pageId="7" pageNumber="308" targetBox="[151,1436,414,1675]" targetPageId="7">
<paragraph id="8BC88552FFFFFFFCD4D4F906FF59F944" blockId="7.[151,1436,1697,1750]" pageId="7" pageNumber="308">
<emphasis id="B9035940FFFFFFFCD4D4F906FE92F924" bold="true" box="[151,274,1697,1719]" pageId="7" pageNumber="308">FIGURE 4.</emphasis>
Photographs of the holotypes of the species treated in this manuscript, in dorsal (top row) and ventral (bottom row) views.
</paragraph>
</caption>
<subSubSection id="C36DD6D9FFFFFFF3D484F95AFB8CFE1A" lastPageId="8" lastPageNumber="309" pageId="7" pageNumber="308" type="diagnosis">
<paragraph id="8BC88552FFFFFFFCD484F95AFCE6F858" blockId="7.[151,1436,1789,2030]" pageId="7" pageNumber="308">
<emphasis id="B9035940FFFFFFFCD484F95AFD93F884" bold="true" box="[199,531,1789,1814]" pageId="7" pageNumber="308">Diagnosis and comparisons.</emphasis>
<taxonomicName id="4C77FED1FFFFFFFCD659F95AFCFCF887" box="[538,892,1789,1813]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="7" pageNumber="308" phylum="Chordata" rank="species" species="guentherpetersi">
<emphasis id="B9035940FFFFFFFCD659F95AFCFCF887" box="[538,892,1789,1813]" italics="true" pageId="7" pageNumber="308">Rhombophryne guentherpetersi</emphasis>
</taxonomicName>
is a member of the genus
<taxonomicName id="4C77FED1FFFFFFFCD0E7F95AFACFF887" box="[1188,1359,1789,1813]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="7" pageNumber="308" phylum="Chordata" rank="genus">
<emphasis id="B9035940FFFFFFFCD0E7F95AFACFF887" box="[1188,1359,1789,1813]" italics="true" pageId="7" pageNumber="308">Rhombophryne</emphasis>
</taxonomicName>
on the basis of molecular phylogenetic affinities (
<figureCitation id="134C99D7FFFFFFFCD636F886FD37F8A8" box="[629,695,1825,1850]" captionStart="FIGURE 3" captionStartId="5.[151,250,1424,1446]" captionTargetBox="[168,1428,732,1392]" captionTargetId="figure@5.[151,1436,716,1405]" captionTargetPageId="5" captionText="FIGURE 3. Majority-rule consensus tree of the Rhombophryne serratopalpebrosa species group and its sister clade, obtained by partitioned BI analysis, based on 2491 nucleotide characters of two mitochondrial and one nuclear gene (16 S, cox 1, sacs). Asterisks indicate Bayesian posterior probability values (* 0.95 0.98, ** 0.99 1.0; not shown if &lt;0.95). Stumpffia psologlossa and other species of Rhombophryne were used as hierarchical outgroups and are here excluded from the figure as they are not the subject of this study. Members of the R. serratopalpebrosa species group are depicted in life, not to scale." httpUri="https://zenodo.org/record/803055/files/figure.png" pageId="7" pageNumber="308">Fig. 3</figureCitation>
) and the possession of a clavicle combined with the absence of T- or Y-shaped terminal phalanges (vs. either absence of a clavicle or possession of a clavicle combined with T- or Yshaped terminal phalanges in the morphologically similar
<taxonomicName id="4C77FED1FFFFFFFCD778F8CDFB40F810" authority="Boulenger, 1882" authorityName="Boulenger" authorityYear="1882" box="[827,1216,1897,1922]" class="Amphibia" family="Microhylidae" genus="Plethodontohyla" kingdom="Animalia" order="Anura" pageId="7" pageNumber="308" phylum="Chordata" rank="genus">
<emphasis id="B9035940FFFFFFFCD778F8CDFC74F810" box="[827,1012,1898,1922]" italics="true" pageId="7" pageNumber="308">Plethodontohyla</emphasis>
<bibRefCitation id="EFE6F8A3FFFFFFFCD7BDF8CEFB40F810" author="Boulenger" box="[1022,1216,1897,1922]" pageId="7" pageNumber="308" refString="Boulenger, G. A. (1882) Catalogue of the Batrachia Salientia s. Caudata in the collection of the British Museum. 2 nd Edition. Taylor and Francis, London, xvi + 503 pp." type="book" year="1882">Boulenger, 1882</bibRefCitation>
</taxonomicName>
). Within the genus
<taxonomicName id="4C77FED1FFFFFFFCD4D4F82AFEC2F837" box="[151,322,1933,1957]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="7" pageNumber="308" phylum="Chordata" rank="genus">
<emphasis id="B9035940FFFFFFFCD4D4F82AFEC2F837" box="[151,322,1933,1957]" italics="true" pageId="7" pageNumber="308">Rhombophryne</emphasis>
</taxonomicName>
, it is assigned to the
<taxonomicName id="4C77FED1FFFFFFFCD603F829FCB0F837" box="[576,816,1933,1957]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="7" pageNumber="308" phylum="Chordata" rank="species" species="serratopalpebrosa">
<emphasis id="B9035940FFFFFFFCD603F829FCB0F837" box="[576,816,1933,1957]" italics="true" pageId="7" pageNumber="308">R. serratopalpebrosa</emphasis>
</taxonomicName>
species group on the basis of possessing superciliary spines and of molecular phylogenetic affinities (
<figureCitation id="134C99D7FFFFFFFCD6F0F816FD77F858" box="[691,759,1969,1994]" captionStart="FIGURE 3" captionStartId="5.[151,250,1424,1446]" captionTargetBox="[168,1428,732,1392]" captionTargetId="figure@5.[151,1436,716,1405]" captionTargetPageId="5" captionText="FIGURE 3. Majority-rule consensus tree of the Rhombophryne serratopalpebrosa species group and its sister clade, obtained by partitioned BI analysis, based on 2491 nucleotide characters of two mitochondrial and one nuclear gene (16 S, cox 1, sacs). Asterisks indicate Bayesian posterior probability values (* 0.95 0.98, ** 0.99 1.0; not shown if &lt;0.95). Stumpffia psologlossa and other species of Rhombophryne were used as hierarchical outgroups and are here excluded from the figure as they are not the subject of this study. Members of the R. serratopalpebrosa species group are depicted in life, not to scale." httpUri="https://zenodo.org/record/803055/files/figure.png" pageId="7" pageNumber="308">Fig. 3</figureCitation>
,
<tableCitation id="C6F5B0E9FFFFFFFCD740F816FCD5F858" box="[771,853,1969,1994]" captionStart="TABLE 1" captionStartId="6.[151,239,836,859]" captionText="TABLE 1. Uncorrected p-distances (in percent) in the ca. 550 bp 3 - terminal segment of the 16 S gene (lower diagonal) and in a ca. 600 bp segment of the cox 1 gene (upper diagonal). Grey cells are within-group distances in 16 S. Note that not all available sequences were included in the comparisons because some sequences of much shorter length were excluded. NA, not applicable (due to missing sequences for distance calculation)." pageId="7" pageNumber="308">Table 1</tableCitation>
).
</paragraph>
<paragraph id="8BC88552FFFFFFF3D484F872FB8CFE1A" blockId="7.[151,1436,1789,2030]" lastBlockId="8.[151,1437,151,896]" lastPageId="8" lastPageNumber="309" pageId="7" pageNumber="308">
<taxonomicName id="4C77FED1FFFFFFFCD484F872FDABF87F" box="[199,555,2005,2029]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="7" pageNumber="308" phylum="Chordata" rank="species" species="guentherpetersi">
<emphasis id="B9035940FFFFFFFCD484F872FDABF87F" box="[199,555,2005,2029]" italics="true" pageId="7" pageNumber="308">Rhombophryne guentherpetersi</emphasis>
</taxonomicName>
is distinguished from all other described congeners by the following unique suite of characters: adult SVL
<quantity id="4C8F28B7FFF0FFF3D5ACFF30FD15FF3D" box="[495,661,151,176]" metricMagnitude="-2" metricUnit="m" metricValue="3.15" metricValueMax="3.57" metricValueMin="2.73" pageId="8" pageNumber="309" unit="mm" value="31.5" valueMax="35.7" valueMin="27.3">27.335.7 mm</quantity>
; TDH/ED = 0.4 50.66; a weak supratympanic fold running from the rear corner of the eye to curve slightly over and behind the tympanum toward the axilla; two or three small superciliary spines; distinct, raised dorsolateral glands, and bulbous tibial glands. The pectoral girdle is distinctly narrower than that seen in its congeners (see
<figureCitation id="134C99D7FFF0FFF3D6FDFEA3FC87FE8E" box="[702,775,260,285]" captionStart="FIGURE 7" captionStartId="10.[151,250,1056,1078]" captionTargetBox="[174,1403,201,1035]" captionTargetId="figure@10.[164,1422,193,1035]" captionTargetPageId="10" captionText="FIGURE 7. Osteology of Rhombophryne guentherpetersi (MNHN 1953.165). Full skeleton in (a) dorsal, (b) ventral, and (c) lateral view; skull in (d) dorsal, (e) ventral, and (f) lateral view. Abbreviations: angspl = angulosplenial, col = columella, exoc = exoccipital, fpar = frontoparietal, max = maxilla, mmk = mentomeckelian, npl = neopalatine, pmx = premaxilla, povom = postchoanal vomer, proot = prootic, prvom = prechoanal vomer, prsph = parasphenoid, pter = pterygoid, qj = quadratojugal, smax = septomaxilla, spheth = sphenethmoid, sq = squamosal." httpUri="https://zenodo.org/record/803063/files/figure.png" pageId="8" pageNumber="309">Fig. 7</figureCitation>
; 8.29.4% of SVL). Furthermore,
<taxonomicName id="4C77FED1FFF0FFF3D0EBFEA2FAFCFE8E" box="[1192,1404,260,284]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="8" pageNumber="309" phylum="Chordata" rank="species" species="guentherpetersi">
<emphasis id="B9035940FFF0FFF3D0EBFEA2FAFCFE8E" box="[1192,1404,260,284]" italics="true" pageId="8" pageNumber="309">R. guentherpetersi</emphasis>
</taxonomicName>
is separated from all other
<taxonomicName id="4C77FED1FFF0FFF3D5E5FE8FFDD1FED2" box="[422,593,296,320]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="8" pageNumber="309" phylum="Chordata" rank="genus">
<emphasis id="B9035940FFF0FFF3D5E5FE8FFDD1FED2" box="[422,593,296,320]" italics="true" pageId="8" pageNumber="309">Rhombophryne</emphasis>
</taxonomicName>
species except
<taxonomicName id="4C77FED1FFF0FFF3D6BDFE8EFCE8FED2" box="[766,872,297,320]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="8" pageNumber="309" phylum="Chordata" rank="species" species="ornata">
<emphasis id="B9035940FFF0FFF3D6BDFE8EFCE8FED2" box="[766,872,297,320]" italics="true" pageId="8" pageNumber="309">R. ornata</emphasis>
</taxonomicName>
and
<taxonomicName id="4C77FED1FFF0FFF3D7DEFE8EFB0AFED2" box="[925,1162,296,320]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="8" pageNumber="309" phylum="Chordata" rank="species" species="serratopalpebrosa">
<emphasis id="B9035940FFF0FFF3D7DEFE8EFB0AFED2" box="[925,1162,296,320]" italics="true" pageId="8" pageNumber="309">R. serratopalpebrosa</emphasis>
</taxonomicName>
by uncorrected pairwise distances of at least 4.4% in a segment of the
<emphasis id="B9035940FFF0FFF3D6FEFEEBFD68FEF6" box="[701,744,332,356]" italics="true" pageId="8" pageNumber="309">16S</emphasis>
rRNA mitochondrial gene (see
<tableCitation id="C6F5B0E9FFF0FFF3D021FEEBFB3AFEF6" box="[1122,1210,332,357]" captionStart="TABLE 1" captionStartId="6.[151,239,836,859]" captionText="TABLE 1. Uncorrected p-distances (in percent) in the ca. 550 bp 3 - terminal segment of the 16 S gene (lower diagonal) and in a ca. 600 bp segment of the cox 1 gene (upper diagonal). Grey cells are within-group distances in 16 S. Note that not all available sequences were included in the comparisons because some sequences of much shorter length were excluded. NA, not applicable (due to missing sequences for distance calculation)." pageId="8" pageNumber="309">Table 1</tableCitation>
; no molecular data available for
<taxonomicName id="4C77FED1FFF0FFF3D568FED6FD98FE1A" box="[299,536,368,392]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="8" pageNumber="309" phylum="Chordata" rank="species" species="serratopalpebrosa">
<emphasis id="B9035940FFF0FFF3D568FED6FD98FE1A" box="[299,536,368,392]" italics="true" pageId="8" pageNumber="309">R. serratopalpebrosa</emphasis>
</taxonomicName>
)—for distinction from
<taxonomicName id="4C77FED1FFF0FFF3D75CFED6FC09FE1A" box="[799,905,369,392]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="8" pageNumber="309" phylum="Chordata" rank="species" species="ornata">
<emphasis id="B9035940FFF0FFF3D75CFED6FC09FE1A" box="[799,905,369,392]" italics="true" pageId="8" pageNumber="309">R. ornata</emphasis>
</taxonomicName>
, see below.
</paragraph>
</subSubSection>
<subSubSection id="C36DD6D9FFF0FFF0D484FE33FD54FC87" lastPageId="11" lastPageNumber="312" pageId="8" pageNumber="309" type="description">
<paragraph id="8BC88552FFF0FFF3D484FE33FC87FCAA" blockId="8.[151,1437,151,896]" pageId="8" pageNumber="309">
<taxonomicName id="4C77FED1FFF0FFF3D484FE33FDBCFE3E" box="[199,572,404,428]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="8" pageNumber="309" phylum="Chordata" rank="species" species="guentherpetersi">
<emphasis id="B9035940FFF0FFF3D484FE33FDBCFE3E" box="[199,572,404,428]" italics="true" pageId="8" pageNumber="309">Rhombophryne guentherpetersi</emphasis>
</taxonomicName>
is distinguished from all
<taxonomicName id="4C77FED1FFF0FFF3D7F5FE33FBE1FE3E" box="[950,1121,404,428]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="8" pageNumber="309" phylum="Chordata" rank="genus">
<emphasis id="B9035940FFF0FFF3D7F5FE33FBE1FE3E" box="[950,1121,404,428]" italics="true" pageId="8" pageNumber="309">Rhombophryne</emphasis>
</taxonomicName>
species except the
<taxonomicName id="4C77FED1FFF0FFF3D1C7FE32FEE5FE42" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="8" pageNumber="309" phylum="Chordata" rank="species" species="serratopalpebrosa">
<emphasis id="B9035940FFF0FFF3D1C7FE32FEE5FE42" italics="true" pageId="8" pageNumber="309">R. serratopalpebrosa</emphasis>
</taxonomicName>
group by the possession of superciliary spines, and from all members of this group, and indeed all cophyline microhylids, by the possession of bulbous tibial glands. Genetically,
<taxonomicName id="4C77FED1FFF0FFF3D028FE7AFABFFE66" box="[1131,1343,476,500]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="8" pageNumber="309" phylum="Chordata" rank="species" species="guentherpetersi">
<emphasis id="B9035940FFF0FFF3D028FE7AFABFFE66" box="[1131,1343,476,500]" italics="true" pageId="8" pageNumber="309">R. guentherpetersi</emphasis>
</taxonomicName>
is very closely related to
<taxonomicName id="4C77FED1FFF0FFF3D521FDA6FE4EFD8A" box="[354,462,513,536]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="8" pageNumber="309" phylum="Chordata" rank="species" species="ornata">
<emphasis id="B9035940FFF0FFF3D521FDA6FE4EFD8A" box="[354,462,513,536]" italics="true" pageId="8" pageNumber="309">R. ornata</emphasis>
</taxonomicName>
—they are separated by just 2.43.4% in one fragment of the
<emphasis id="B9035940FFF0FFF3D0D2FDA7FB3BFD8A" box="[1169,1211,512,536]" italics="true" pageId="8" pageNumber="309">16S</emphasis>
gene (see
<tableCitation id="C6F5B0E9FFF0FFF3D174FE58FA0AFD8A" box="[1335,1418,511,536]" captionStart="TABLE 1" captionStartId="6.[151,239,836,859]" captionText="TABLE 1. Uncorrected p-distances (in percent) in the ca. 550 bp 3 - terminal segment of the 16 S gene (lower diagonal) and in a ca. 600 bp segment of the cox 1 gene (upper diagonal). Grey cells are within-group distances in 16 S. Note that not all available sequences were included in the comparisons because some sequences of much shorter length were excluded. NA, not applicable (due to missing sequences for distance calculation)." pageId="8" pageNumber="309">Table 1</tableCitation>
), which is lower than our standard threshold for candidate species recognition (
<bibRefCitation id="EFE6F8A3FFF0FFF3D7B7FD83FB45FDAE" author="Vieites" box="[1012,1221,548,573]" pageId="8" pageNumber="309" 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 United States of America, 106, 8267 - 8272." type="journal article" year="2009">
Vieites
<emphasis id="B9035940FFF0FFF3D009FD82FB02FDAE" box="[1098,1154,548,572]" italics="true" pageId="8" pageNumber="309">et al.</emphasis>
2009
</bibRefCitation>
). However, the two species are highly morphologically distinct, differing not just in the presence or absence of tibial glands, but also in the shorter relative hindlimb length of
<taxonomicName id="4C77FED1FFF0FFF3D63DFDCAFCD6FD16" box="[638,854,620,644]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="8" pageNumber="309" phylum="Chordata" rank="species" species="guentherpetersi">
<emphasis id="B9035940FFF0FFF3D63DFDCAFCD6FD16" box="[638,854,620,644]" italics="true" pageId="8" pageNumber="309">R. guentherpetersi</emphasis>
</taxonomicName>
(HIL/SVL 1.331.45 vs. 1.461.64), tibiotarsal articulation reaching the axilla or tympanum (vs. between the tympanum and the eye), absence (vs. presence) of red colouration in the inguinal region, partially ossified limb epiphyses and carpals (vs. unossified), and much narrower pectoral girdle (length of coracoid &lt;10% of SVL vs.&gt;12%; see
<figureCitation id="134C99D7FFF0FFF3D794FD70FB9BFD62" box="[983,1051,727,752]" captionStart="FIGURE 7" captionStartId="10.[151,250,1056,1078]" captionTargetBox="[174,1403,201,1035]" captionTargetId="figure@10.[164,1422,193,1035]" captionTargetPageId="10" captionText="FIGURE 7. Osteology of Rhombophryne guentherpetersi (MNHN 1953.165). Full skeleton in (a) dorsal, (b) ventral, and (c) lateral view; skull in (d) dorsal, (e) ventral, and (f) lateral view. Abbreviations: angspl = angulosplenial, col = columella, exoc = exoccipital, fpar = frontoparietal, max = maxilla, mmk = mentomeckelian, npl = neopalatine, pmx = premaxilla, povom = postchoanal vomer, proot = prootic, prvom = prechoanal vomer, prsph = parasphenoid, pter = pterygoid, qj = quadratojugal, smax = septomaxilla, spheth = sphenethmoid, sq = squamosal." httpUri="https://zenodo.org/record/803063/files/figure.png" pageId="8" pageNumber="309">Fig. 7</figureCitation>
). A differential diagnosis from all other cophyline microhylids is unnecessary because the tibial glands allow easy unambiguous distinction. These are noticeable, if not well developed, even in subadults.
</paragraph>
<paragraph id="8BC88552FFF0FFF3D484FCE3FCC8FCED" blockId="8.[151,1437,151,896]" pageId="8" pageNumber="309">
<emphasis id="B9035940FFF0FFF3D484FCE3FDD1FCCF" bold="true" box="[199,593,836,861]" pageId="8" pageNumber="309">Re-description of the holotype.</emphasis>
(
<figureCitation id="134C99D7FFF0FFF3D62BFCE3FD3AFCCE" box="[616,698,836,861]" captionStart="FIGURE 4" captionStartId="7.[151,250,1697,1719]" captionTargetBox="[151,1436,414,1675]" captionTargetId="figure@7.[151,1436,414,1675]" captionTargetPageId="7" captionText="FIGURE 4. Photographs of the holotypes of the species treated in this manuscript, in dorsal (top row) and ventral (bottom row) views." httpUri="https://zenodo.org/record/803057/files/figure.png" pageId="8" pageNumber="309">Figs 4</figureCitation>
,
<figureCitation id="134C99D7FFF0FFF3D68CFCE3FD5DFCCF" box="[719,733,836,861]" captionStart="FIGURE 5" captionStartId="8.[151,250,1879,1901]" captionTargetBox="[151,1436,949,1858]" captionTargetId="figure@8.[151,1436,949,1858]" captionTargetPageId="8" captionText="FIGURE 5. Comparative morphology of the heads, feet, and hands of members of the Rhombophryne serratopalpebrosa species group. All specimens are the holotypes of their respective species, except R. coronata, which is paratype ZSM 694 / 2001. Asterisks indicate mirrored images. Not to scale." httpUri="https://zenodo.org/record/803059/files/figure.png" pageId="8" pageNumber="309">5</figureCitation>
) An adult female specimen in moderately good state of preservation. Ventral incision and left lateral incision present.
</paragraph>
<caption id="DF08D5DAFFF0FFF3D4D4F8F0FD44F838" httpUri="https://zenodo.org/record/803059/files/figure.png" pageId="8" pageNumber="309" targetBox="[151,1436,949,1858]" targetPageId="8">
<paragraph id="8BC88552FFF0FFF3D4D4F8F0FD44F838" blockId="8.[151,1436,1879,1962]" pageId="8" pageNumber="309">
<emphasis id="B9035940FFF0FFF3D4D4F8F0FE96F8FE" bold="true" box="[151,278,1879,1901]" pageId="8" pageNumber="309">FIGURE 5.</emphasis>
Comparative morphology of the heads, feet, and hands of members of the
<taxonomicName id="4C77FED1FFF0FFF3D07EF8F0FA1CF8FF" box="[1085,1436,1879,1901]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="8" pageNumber="309" phylum="Chordata" rank="species" species="serratopalpebrosa">
<emphasis id="B9035940FFF0FFF3D07EF8F0FA1CF8FF" box="[1085,1436,1879,1901]" italics="true" pageId="8" pageNumber="309">Rhombophryne serratopalpebrosa</emphasis>
</taxonomicName>
species group. All specimens are the holotypes of their respective species, except
<taxonomicName id="4C77FED1FFF0FFF3D7AFF8D0FBE4F81E" box="[1004,1124,1911,1932]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="8" pageNumber="309" phylum="Chordata" rank="species" species="coronata">
<emphasis id="B9035940FFF0FFF3D7AFF8D0FBE4F81E" box="[1004,1124,1911,1932]" italics="true" pageId="8" pageNumber="309">R. coronata</emphasis>
</taxonomicName>
, which is paratype ZSM 694/ 2001. Asterisks indicate mirrored images. Not to scale.
</paragraph>
</caption>
<caption id="DF08D5DAFFF1FFF2D4D4F822FEE2F84A" httpUri="https://zenodo.org/record/803061/files/figure.png" pageId="9" pageNumber="310" targetBox="[170,1414,193,1885]" targetPageId="9">
<paragraph id="8BC88552FFF1FFF2D4D4F822FEE2F84A" blockId="9.[151,1436,1925,2008]" pageId="9" pageNumber="310">
<emphasis id="B9035940FFF1FFF2D4D4F822FE94F808" bold="true" box="[151,276,1925,1947]" pageId="9" pageNumber="310">FIGURE 6.</emphasis>
<taxonomicName id="4C77FED1FFF1FFF2D55FF822FDDEF809" box="[284,606,1925,1947]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="9" pageNumber="310" phylum="Chordata" rank="species" species="guentherpetersi">
<emphasis id="B9035940FFF1FFF2D55FF822FDDEF809" box="[284,606,1925,1947]" italics="true" pageId="9" pageNumber="310">Rhombophryne guentherpetersi</emphasis>
</taxonomicName>
in life. (ac) ZSM 607/2014 in (a) dorsolateral (with inset showing superciliary spines), (b) dorsal, and (c) ventral view; and (d, e) other specimens in dorsolateral view (assignment to field and collection numbers unknown).
</paragraph>
</caption>
<caption id="DF08D5DAFFF2FFF1D4D4FB87FC90FB23" httpUri="https://zenodo.org/record/803063/files/figure.png" pageId="10" pageNumber="311" targetBox="[174,1403,201,1035]" targetPageId="10">
<paragraph id="8BC88552FFF2FFF1D4D4FB87FC90FB23" blockId="10.[151,1436,1056,1201]" pageId="10" pageNumber="311">
<emphasis id="B9035940FFF2FFF1D4D4FB87FE94FBA7" bold="true" box="[151,276,1056,1078]" pageId="10" pageNumber="311">FIGURE 7.</emphasis>
Osteology of
<taxonomicName id="4C77FED1FFF2FFF1D5EBFB87FD69FBA4" box="[424,745,1056,1078]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="10" pageNumber="311" phylum="Chordata" rank="species" species="guentherpetersi">
<emphasis id="B9035940FFF2FFF1D5EBFB87FD69FBA4" box="[424,745,1056,1078]" italics="true" pageId="10" pageNumber="311">Rhombophryne guentherpetersi</emphasis>
</taxonomicName>
(MNHN 1953.165). Full skeleton in (a) dorsal, (b) ventral, and (c) lateral view; skull in (d) dorsal, (e) ventral, and (f) lateral view. Abbreviations: angspl = angulosplenial, col = columella, exoc = exoccipital, fpar = frontoparietal, max = maxilla, mmk = mentomeckelian, npl = neopalatine, pmx = premaxilla, povom = postchoanal vomer, proot = prootic, prvom = prechoanal vomer, prsph = parasphenoid, pter = pterygoid, qj = quadratojugal, smax = septomaxilla, spheth = sphenethmoid, sq = squamosal.
</paragraph>
</caption>
<paragraph id="8BC88552FFF2FFF1D484FB51FBB5FA0D" blockId="10.[151,1436,1270,2015]" pageId="10" pageNumber="311">Body robust. Head wider than long (HW/HL = 1.35). Pupils horizontally oval. Snout rounded in dorsal and lateral views. Canthus rostralis concave. Loreal region concave. Nostrils nearer to snout tip than to eye (END/NSD = 1.24), directed laterally, slightly protuberant. Tympanum distinct, TDH/ED = 0.47. Weak supratympanic fold, from middle rear of eye curving slightly over and behind tympanum toward the axilla. Two small superciliary spines above each eye. Vomerine teeth present, curved, not meeting at the midline.</paragraph>
<paragraph id="8BC88552FFF2FFF1D484FA0CFB9EF9E5" blockId="10.[151,1436,1270,2015]" pageId="10" pageNumber="311">Arms robust. Fingers without webbing; relative lengths 1&lt;2&lt;4&lt;3, fourth finger distinctly longer than second; finger tips not expanded; fingers not reduced; nuptial pads absent; inner metacarpal tubercle present, outer metacarpal tubercle divided, faint; subarticular tubercles distinct, undivided. Hindlimbs short and thick; tibiotarsal articulation reaches the axilla; TIBL/SVL = 0.35. Strongly developed tibial glands cover almost the entirety of the dorsal tibia. Inner metatarsal tubercle present, enlarged, outer metatarsal tubercle absent. Toes unwebbed; relative lengths 1&lt;2&lt;5&lt;3&lt;4, fifth toe distinctly shorter than third. Toe tips not expanded.</paragraph>
<paragraph id="8BC88552FFF2FFF1D484F924FC20F976" blockId="10.[151,1436,1270,2015]" pageId="10" pageNumber="311">Dorsal skin slightly granular except on the head, where it is slightly rougher, with a slightly raised ridge on the midline of the head. Dorsolateral folds absent. A porous glandular formation extends from the suprascapular region to the inguinal region on either side of the body. Ventral skin smooth.</paragraph>
<paragraph id="8BC88552FFF2FFF1D484F949FC25F8E6" blockId="10.[151,1436,1270,2015]" pageId="10" pageNumber="311">
Colouration of the
<typeStatus id="54CC3BF0FFF2FFF1D5D8F949FD81F895" box="[411,513,1774,1799]" pageId="10" pageNumber="311" type="holotype">holotype</typeStatus>
: (
<figureCitation id="134C99D7FFF2FFF1D650F949FDDCF895" box="[531,604,1774,1799]" captionStart="FIGURE 4" captionStartId="7.[151,250,1697,1719]" captionTargetBox="[151,1436,414,1675]" captionTargetId="figure@7.[151,1436,414,1675]" captionTargetPageId="7" captionText="FIGURE 4. Photographs of the holotypes of the species treated in this manuscript, in dorsal (top row) and ventral (bottom row) views." httpUri="https://zenodo.org/record/803057/files/figure.png" pageId="10" pageNumber="311">Figs 4</figureCitation>
,
<figureCitation id="134C99D7FFF2FFF1D62BF949FDF6F895" box="[616,630,1774,1799]" captionStart="FIGURE 5" captionStartId="8.[151,250,1879,1901]" captionTargetBox="[151,1436,949,1858]" captionTargetId="figure@8.[151,1436,949,1858]" captionTargetPageId="8" captionText="FIGURE 5. Comparative morphology of the heads, feet, and hands of members of the Rhombophryne serratopalpebrosa species group. All specimens are the holotypes of their respective species, except R. coronata, which is paratype ZSM 694 / 2001. Asterisks indicate mirrored images. Not to scale." httpUri="https://zenodo.org/record/803059/files/figure.png" pageId="10" pageNumber="311">5</figureCitation>
) Light reddish-brown above with irregular small dark speckling. Loreal and tympanic regions lighter than dorsal head, without speckling. Glandular formations on tibiae and dorsum darker brown, as are the anterior surfaces of head and above eyes. Flanks fading from dorsal colour to ventral colour. Venter uniformly tan to cream. Limbs coloured as the dorsum.
</paragraph>
<paragraph id="8BC88552FFF2FFF0D484F8D9FECAFE66" blockId="10.[151,1436,1270,2015]" lastBlockId="11.[151,1437,151,1982]" lastPageId="11" lastPageNumber="312" pageId="10" pageNumber="311">
<emphasis id="B9035940FFF2FFF1D484F8D9FEBFF805" bold="true" box="[199,319,1918,1943]" pageId="10" pageNumber="311">Variation.</emphasis>
The type series is composed of two juveniles, one subadult, and one poorly preserved adult, in addition to the holotype. Our newly added material consists of three well-preserved adults, including the first male specimen. All specimens, including the juveniles, possess superciliary spines, but they are small and variable in number, and in poorly preserved specimens cannot be well distinguished (but can be distinguished in life; see
<figureCitation id="134C99D7FFF3FFF0D133FF30FF27FF47" captionStart="FIGURE 6" captionStartId="9.[151,250,1925,1947]" captionTargetBox="[170,1414,193,1885]" captionTargetId="figure@9.[170,1414,193,1885]" captionTargetPageId="9" captionText="FIGURE 6. Rhombophryne guentherpetersi in life. (a c) ZSM 607 / 2014 in (a) dorsolateral (with inset showing superciliary spines), (b) dorsal, and (c) ventral view; and (d, e) other specimens in dorsolateral view (assignment to field and collection numbers unknown)." httpUri="https://zenodo.org/record/803061/files/figure.png" pageId="11" pageNumber="312">Fig. 6</figureCitation>
). ZSM 606/2014 has three superciliary spines, whereas all other specimens have two. The paratype MNHN 1953.165B is darker than the rest of the type series and has much rougher skin (but is also particularly poorly preserved). All specimens, including the juveniles, have tibial glands, but these are less bulbous in the juveniles and subadult, becoming large only in the adults (confirming that all members of the type series are indeed conspecifics). The one male specimen is smaller than the adult females, but is in all other aspects highly similar in morphology (see Appendix 1). The tibiotarsal articulation reaches either the axilla or the tympanum. A prepollex is well developed in the one available male specimen, ZSM 607/2014. The species may have some degree of sexual size dimorphism, as the only male specimen is slightly smaller than all adult female specimens available (27.3 vs.
<quantity id="4C8F28B7FFF3FFF0D4D4FE7BFEBEFE66" box="[151,318,476,501]" metricMagnitude="-2" metricUnit="m" metricValue="3.2299999999999995" metricValueMax="3.5700000000000003" metricValueMin="2.8899999999999997" pageId="11" pageNumber="312" unit="mm" value="32.3" valueMax="35.7" valueMin="28.9">28.935.7 mm</quantity>
).
</paragraph>
<paragraph id="8BC88552FFF3FFF0D484FE58FD54FC87" blockId="11.[151,1437,151,1982]" pageId="11" pageNumber="312">
Colouration is relatively homogeneous, but shade and pattern differ somewhat (
<figureCitation id="134C99D7FFF3FFF0D014FE58FB1EFD8A" box="[1111,1182,511,536]" captionStart="FIGURE 6" captionStartId="9.[151,250,1925,1947]" captionTargetBox="[170,1414,193,1885]" captionTargetId="figure@9.[170,1414,193,1885]" captionTargetPageId="9" captionText="FIGURE 6. Rhombophryne guentherpetersi in life. (a c) ZSM 607 / 2014 in (a) dorsolateral (with inset showing superciliary spines), (b) dorsal, and (c) ventral view; and (d, e) other specimens in dorsolateral view (assignment to field and collection numbers unknown)." httpUri="https://zenodo.org/record/803061/files/figure.png" pageId="11" pageNumber="312">Fig. 6</figureCitation>
). The newly collected specimens are much darker brown in colour than the type series, but generally have commonalities to their colour patterns: a pair of blackish oblong markings above the suprascapular region (
<figureCitation id="134C99D7FFF3FFF0D044FDE0FBCDFDF2" box="[1031,1101,583,608]" captionStart="FIGURE 6" captionStartId="9.[151,250,1925,1947]" captionTargetBox="[170,1414,193,1885]" captionTargetId="figure@9.[170,1414,193,1885]" captionTargetPageId="9" captionText="FIGURE 6. Rhombophryne guentherpetersi in life. (a c) ZSM 607 / 2014 in (a) dorsolateral (with inset showing superciliary spines), (b) dorsal, and (c) ventral view; and (d, e) other specimens in dorsolateral view (assignment to field and collection numbers unknown)." httpUri="https://zenodo.org/record/803061/files/figure.png" pageId="11" pageNumber="312">Fig. 6</figureCitation>
). These markings are present only in one member of the type series (MNHN 1973.592). The lateral body has several dark-brown flecks. These are present in all of the paratypes but not in the holotype. The venter is mottled dark and light brown in some specimens but is almost solid dark brown in others. A light brown interocular bar is sometimes present, anterior to which the snout is typically lighter brown than the dorsum. The dorsolateral glands can also be this light brown, or they can be continuous with the rest of the dorsum.
</paragraph>
</subSubSection>
<subSubSection id="C36DD6D9FFF3FFF0D484FCB8FDAEFB57" pageId="11" pageNumber="312" type="discussion">
<paragraph id="8BC88552FFF3FFF0D484FCB8FED8FBA7" blockId="11.[151,1437,151,1982]" pageId="11" pageNumber="312">
<emphasis id="B9035940FFF3FFF0D484FCB8FEBBFCAA" bold="true" box="[199,315,799,824]" pageId="11" pageNumber="312">Remarks.</emphasis>
The following inconsistencies exist between our re-description and the original description of this species by
<bibRefCitation id="EFE6F8A3FFF3FFF0D559FCE3FE38FCCE" author="Guibe" box="[282,440,836,861]" pageId="11" pageNumber="312" refString="Guibe, J. (1974) Batraciens nouveaux de Madagascar. Bulletin du Museum national d'Histoire Naturelle, 3, 1170 - 1192." type="journal article" year="1974">Guibé (1974)</bibRefCitation>
: Superciliary spines are present; these were apparently overlooked in his description. Tibiotarsal articulation was given as reaching the eye. This could not be verified in any member of the type series except the poorly preserved paratype MNHN 1973.592, the vertebral column of which is inverted (i.e. convex instead of concave), possibly misleading tibiotarsal articulation. The holotype is
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longer than described (
<quantity id="4C8F28B7FFF3FFF0D129FC08FF43FC7E" metricMagnitude="-2" metricUnit="m" metricValue="3.46" pageId="11" pageNumber="312" unit="mm" value="34.6">34.6 mm</quantity>
vs.
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).
<bibRefCitation id="EFE6F8A3FFF3FFF0D514FC73FE68FC7E" author="Guibe" box="[343,488,980,1005]" pageId="11" pageNumber="312" refString="Guibe, J. (1974) Batraciens nouveaux de Madagascar. Bulletin du Museum national d'Histoire Naturelle, 3, 1170 - 1192." type="journal article" year="1974">Guibé (1974</bibRefCitation>
,
<bibRefCitation id="EFE6F8A3FFF3FFF0D5B0FC73FDACFC7E" author="Guibe" box="[499,556,980,1004]" pageId="11" pageNumber="312" refString="Guibe, J. (1978) Les batraciens de Madagascar. Bonner zoologische Monographien, 11, 1 - 140." type="journal article" year="1978">1978</bibRefCitation>
) stated that the prepollex was developed in males, but the type series does not contain any males. Nevertheless, we have confirmed this observation with a newly collected male specimen (see Variation above).
</paragraph>
<paragraph id="8BC88552FFF3FFF0D484FB98FDAEFB57" blockId="11.[151,1437,151,1982]" pageId="11" pageNumber="312">
<emphasis id="B9035940FFF3FFF0D484FB98FE09FBCA" bold="true" box="[199,393,1087,1112]" pageId="11" pageNumber="312">Natural history.</emphasis>
At Andranomadio, specimens were found during the day in high-altitude rainforest with rather open canopy (
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a.s.l.), burrowed several centimetres deep in the ground. Another individual (ZSM 608/2014) was collected above the treeline (
<quantity id="4C8F28B7FFF3FFF0D6DEFB2FFD77FB32" box="[669,759,1160,1184]" metricMagnitude="3" metricUnit="m" metricValue="2.732" pageId="11" pageNumber="312" unit="m" value="2732.0">2732 m</quantity>
a.s.l.), in an area of grassland with some scattered heath, hidden under a stone during the day.
</paragraph>
</subSubSection>
<subSubSection id="C36DD6D9FFF3FFF0D484FB68FBF6F82C" pageId="11" pageNumber="312" type="distribution">
<paragraph id="8BC88552FFF3FFF0D484FB68FBF6F82C" blockId="11.[151,1437,151,1982]" pageId="11" pageNumber="312">
<emphasis id="B9035940FFF3FFF0D484FB68FD0AFB7A" bold="true" box="[199,650,1231,1256]" pageId="11" pageNumber="312">Distribution and conservation status.</emphasis>
<taxonomicName id="4C77FED1FFF3FFF0D6D7FB77FC78FB7A" box="[660,1016,1232,1256]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="11" pageNumber="312" phylum="Chordata" rank="species" species="guentherpetersi">
<emphasis id="B9035940FFF3FFF0D6D7FB77FC78FB7A" box="[660,1016,1232,1256]" italics="true" pageId="11" pageNumber="312">Rhombophryne guentherpetersi</emphasis>
</taxonomicName>
is currently listed in the IUCN Red List as Endangered under criterion B1ab(iii) (IUCN SSC Amphibian Specialist Group 2016a), meaning an extent of occurrence of &lt;
<quantity id="4C8F28B7FFF3FFF0D52FFABDFE50FAA1" box="[364,464,1306,1331]" metricMagnitude="6" metricUnit="m" metricValue="5.0" pageId="11" pageNumber="312" unit="km" value="5000.0">5000 km</quantity>
2 (B1), a severely fragmented habitat or known from Ẽ5 locations (a), and an observed, estimated, inferred, or projected on-going decline in the area, extent and/or quality of habitat (b(iii)). At present,
<taxonomicName id="4C77FED1FFF3FFF0D1C7FAE7FEC7FAE9" class="Amphibia" family="Microhylidae" genus="Rhombophryne" kingdom="Animalia" order="Anura" pageId="11" pageNumber="312" phylum="Chordata" rank="species" species="guentherpetersi">
<emphasis id="B9035940FFF3FFF0D1C7FAE7FEC7FAE9" italics="true" pageId="11" pageNumber="312">R. guentherpetersi</emphasis>
</taxonomicName>
is known only from the Tsaratanana Massif. The exact collecting locality of the
<typeStatus id="54CC3BF0FFF3FFF0D096FAC4FA85FAE9" box="[1237,1285,1379,1403]" pageId="11" pageNumber="312">type</typeStatus>
specimens is not clear (
<bibRefCitation id="EFE6F8A3FFF3FFF0D544FA21FE10FA0D" author="Guibe" box="[263,400,1414,1439]" pageId="11" pageNumber="312" refString="Guibe, J. (1974) Batraciens nouveaux de Madagascar. Bulletin du Museum national d'Histoire Naturelle, 3, 1170 - 1192." type="journal article" year="1974">Guibé 1974</bibRefCitation>
), but three specimens were collected on a recent expedition to the Tsaratanana Massif (ZSM 606607/2014 at
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,
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at
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a.s.l; ZSM 608/2014 at
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,
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,
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a.s.l.;
<figureCitation id="134C99D7FFF3FFF0D4D4FA69FF5BFA75" box="[151,219,1486,1511]" captionStart="FIGURE 8" captionStartId="12.[151,250,1199,1221]" captionTargetBox="[151,1436,195,1178]" captionTargetId="figure@12.[151,1436,195,1178]" captionTargetPageId="12" captionText="FIGURE 8. Map of the north of Madagascar showing the collection localities of specimens of the focal species of this paper, Rhombophryne guentherpetersi, R. regalis sp. nov., and R. diadema sp. nov. Basemap from www. vegmad. org. Hashing indicates protected areas; note that the full extent of Tsaratanana Strict Nature Reserve is not shown." httpUri="https://zenodo.org/record/803065/files/figure.png" pageId="11" pageNumber="312">Fig. 8</figureCitation>
). These two localities are well inside the protected area of Tsaratanana Strict Nature Reserve, to which only researchers and conservation workers are permitted access. Although it is possible that the species may occur in the forest corridor between Marojejy and Tsaratanana (COMATSA; see
<bibRefCitation id="EFE6F8A3FFF3FFF0D7EAF9B1FB42F9BD" author="Rabearivony" box="[937,1218,1558,1583]" pageId="11" pageNumber="312" refString="Rabearivony, J., Rasamoelina, M., Raveloson, J., Rakotomanana, H., Raselimanana, A. P., Raminosoa, N. R. &amp; Zaonarivelo, J. R. (2015) Roles of a forest corridor between Marojejy, Anjanaharibe-Sud and Tsaratanana protected areas, northern Madagascar, in maintaining endemic and threatened Malagasy taxa. Madagascar Conservation &amp; Development, 10, 85 - 92." type="journal article" year="2015">
Rabearivony
<emphasis id="B9035940FFF3FFF0D001F9BFFBFCF9BD" box="[1090,1148,1559,1583]" italics="true" pageId="11" pageNumber="312">et al.</emphasis>
2015
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), which is pending official protected status, we consider this unlikely: we suspect that
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<emphasis id="B9035940FFF3FFF0D7FFF99CFB11F9C1" box="[956,1169,1595,1619]" italics="true" pageId="11" pageNumber="312">R. guentherpetersi</emphasis>
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is found only at high elevations, possibly near to and above the tree line, which in Tsaratanana is around
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a.s.l. No part of the COMATSA corridor exceeds
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a.s.l. The likely extent of occurrence of this species is therefore limited to an area of
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2. Deforestation and habitat degradation pressure at such high altitude is minimal relative to the lower reaches of the forest, especially due to the strict conservation status of this forest. However, it is not free from threats: Climate change may be a significant factor for species living near the tops of mountains, as their niche space diminishes with rising temperatures. Fires in the high mountain grassland could swiftly eradicate the suitable open habitat and leave only forest habitat available. To balance (1) the small range of this species, (2) the highly protected status it enjoys, and (3) the threats that persist despite this protection, we propose to maintain a status of Endangered B1ab(iii) (IUCN 2012), but encourage further surveys within COMATSA and Tsaratanana Strict Nature Reserve in order to better understand the distribution and ecology of this species.
</paragraph>
</subSubSection>
</treatment>
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