treatments-xml/data/03/E2/87/03E2878D6C3BFFCAFF5BDD0EFA92CDFE.xml

<document id="202184E740E618B98B9B9887D349DDB5" ID-CLB-Dataset="39209" ID-DOI="10.11646/zootaxa.4059.3.8" ID-GBIF-Dataset="e44b5825-b650-4af5-99b2-f17cb415837d" ID-ISSN="1175-5326" ID-Zenodo-Dep="244218" ID-ZooBank="A9E81D14-85B7-41FE-8180-9C3770DB8FEB" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1461364716946" checkinUser="plazi" docAuthor="Vences, Miguel, Jovanovic, Olga, Safarek, Goran, Glaw, Frank &amp; Köhler, Jörn" docDate="2015" docId="03E2878D6C3BFFCAFF5BDD0EFA92CDFE" docLanguage="en" docName="zt04059p580.pdf" docOrigin="Zootaxa 4059 (3)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Guibemantis diphonus Vences, Jovanovic, Safarek, Glaw &amp; Köhler, 2015, sp. nov." docType="treatment" docUuid="6D0ED257-C1AB-4E37-9F51-171B7D5E87D2" docUuidSource="ZooBank" docVersion="9" lastPageNumber="577" masterDocId="FFDBFFF56C3AFFC2FFCCDB5AFFE0C95E" masterDocTitle="A new arboreal frog of the genus Guibemantis from the southeast of Madagascar (Anura: Mantellidae)" masterLastPageNumber="580" masterPageNumber="569" pageNumber="570" updateTime="1698661921841" updateUser="ExternalLinkService">
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<mods:title id="8C80745A6121590BF6C0D259CB6E6794">A new arboreal frog of the genus Guibemantis from the southeast of Madagascar (Anura: Mantellidae)</mods:title>
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<mods:namePart id="B975E1DDBA0016E22A98D7A618E545AE">Vences, Miguel</mods:namePart>
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<mods:namePart id="0C46BFBDE23EBD41FA09D72340A94A4E">Jovanovic, Olga</mods:namePart>
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<mods:namePart id="34DFEA324FB12EA042EB09FCBD492BCC">Safarek, Goran</mods:namePart>
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<mods:namePart id="35146BFA886E1AB181278817DE30D279">Glaw, Frank</mods:namePart>
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<treatment id="03E2878D6C3BFFCAFF5BDD0EFA92CDFE" ID-DOI="http://doi.org/10.5281/zenodo.5693742" ID-GBIF-Taxon="119649146" ID-Zenodo-Dep="5693742" LSID="urn:lsid:zoobank.org:act:6D0ED257-C1AB-4E37-9F51-171B7D5E87D2" httpUri="http://treatment.plazi.org/id/03E2878D6C3BFFCAFF5BDD0EFA92CDFE" lastPageId="8" lastPageNumber="577" pageId="1" pageNumber="570">
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<paragraph id="8BF4369B6C3BFFC3FF5BDD0EFDEECF30" blockId="1.[151,526,1620,1646]" box="[151,526,1620,1646]" pageId="1" pageNumber="570">
<heading id="D0BC81F76C3BFFC3FF5BDD0EFDEECF30" bold="true" box="[151,526,1620,1646]" fontSize="11" level="1" pageId="1" pageNumber="570" reason="1">
<emphasis id="B93FEA896C3BFFC3FF5BDD0EFDEECF30" bold="true" box="[151,526,1620,1646]" pageId="1" pageNumber="570">
<taxonomicName id="4C4B4D186C3BFFC3FF5BDD0EFE4ACF30" ID-CoL="6L5H7" box="[151,426,1620,1646]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="1" pageNumber="570" phylum="Chordata" rank="species" species="diphonus" status="sp. nov.">
<emphasis id="B93FEA896C3BFFC3FF5BDD0EFE4ACF30" bold="true" box="[151,426,1620,1646]" italics="true" pageId="1" pageNumber="570">Guibemantis diphonus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A20C57F26C3BFFC3FE7CDD0EFDEECF30" box="[432,526,1620,1646]" pageId="1" pageNumber="570" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
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</paragraph>
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<subSubSection id="C35165106C3BFFC3FF5BDDCDFAB6CE43" pageId="1" pageNumber="570" type="materials_examined">
<paragraph id="8BF4369B6C3BFFC3FF5BDDCDFBECCFA6" blockId="1.[151,1436,1687,2037]" pageId="1" pageNumber="570">
<emphasis id="B93FEA896C3BFFC3FF5BDDCDFEEBCFEE" bold="true" box="[151,267,1687,1712]" pageId="1" pageNumber="570">
<typeStatus id="54F088396C3BFFC3FF5BDDCDFEE6CFEE" box="[151,262,1687,1712]" pageId="1" pageNumber="570" type="holotype">Holotype</typeStatus>
.
</emphasis>
<collectionCode id="ED5AAE5E6C3BFFC3FEDFDDC2FEADCFEE" box="[275,333,1688,1712]" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="1" pageNumber="570">ZSM</collectionCode>
2665/2007 (field number 
<collectionCode id="ED5AAE5E6C3BFFC3FDB6DDC2FD2BCFEE" box="[634,715,1688,1712]" pageId="1" pageNumber="570">ZCMV</collectionCode>
5496), adult male (
<figureCitation id="13702A1E6C3BFFC3FC60DDCDFC0CCFEE" box="[940,1004,1687,1712]" captionStart="FIGURE 1" captionStartId="2.[151,250,1723,1745]" captionTargetBox="[150,1435,193,1699]" captionTargetId="figure@2.[150,1436,193,1702]" captionTargetPageId="2" captionText="FIGURE 1. Guibemantis diphonus sp. nov.: A − B, male holotype ZSM 2665 / 2007 from Manombo in life in dorsolateral and ventral view; C − D, male holotype ZSM 2665 / 2007 in preservative (photographed in May 2015); E − F, male paratype ZSM 2664 / 2007 shortly after preservation (photographed on 24 February 2007). Scale bars = 5 mm." httpUri="https://zenodo.org/record/244219/files/figure.png" pageId="1" pageNumber="570">Fig. 1</figureCitation>
A −D), collected in Manombo Special Reserve (coordinates not taken, but about -23.01 / 47.73), Fianarantsoa Province, southeastern 
<collectingCountry id="F35C760B6C3BFFC3FB0BDDE6FAB0CF8B" box="[1223,1360,1724,1749]" name="Madagascar" pageId="1" pageNumber="570">Madagascar</collectingCountry>
, on 
<date id="FFF5105B6C3BFFC3FA4CDDE6FEDECFA6" pageId="1" pageNumber="570" value="2007-02-23">23 February 2007</date>
, by M. Vences, G. Safarek, E. Rajeriarison and T. Rajoafiarison.
</paragraph>
<paragraph id="8BF4369B6C3BFFC3FF0BDC5EFAB6CE43" blockId="1.[151,1436,1687,2037]" box="[199,1366,1796,1821]" pageId="1" pageNumber="570">
<emphasis id="B93FEA896C3BFFC3FF0BDC5EFEDBCE43" bold="true" box="[199,315,1796,1821]" pageId="1" pageNumber="570">
<typeStatus id="54F088396C3BFFC3FF0BDC5EFED6CE43" box="[199,310,1796,1821]" pageId="1" pageNumber="570" type="paratype">Paratype</typeStatus>
.
</emphasis>
<collectionCode id="ED5AAE5E6C3BFFC3FE8EDC5EFE9CCE42" box="[322,380,1796,1820]" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="1" pageNumber="570">ZSM</collectionCode>
2664/2007 (field number 
<collectionCode id="ED5AAE5E6C3BFFC3FD6DDC5EFD12CE42" box="[673,754,1796,1820]" pageId="1" pageNumber="570">ZCMV</collectionCode>
5416), adult male (
<figureCitation id="13702A1E6C3BFFC3FC01DC5EFBEBCE42" box="[973,1035,1796,1821]" captionStart="FIGURE 1" captionStartId="2.[151,250,1723,1745]" captionTargetBox="[150,1435,193,1699]" captionTargetId="figure@2.[150,1436,193,1702]" captionTargetPageId="2" captionText="FIGURE 1. Guibemantis diphonus sp. nov.: A − B, male holotype ZSM 2665 / 2007 from Manombo in life in dorsolateral and ventral view; C − D, male holotype ZSM 2665 / 2007 in preservative (photographed in May 2015); E − F, male paratype ZSM 2664 / 2007 shortly after preservation (photographed on 24 February 2007). Scale bars = 5 mm." httpUri="https://zenodo.org/record/244219/files/figure.png" pageId="1" pageNumber="570">Fig. 1</figureCitation>
E −F), same data as 
<typeStatus id="54F088396C3BFFC3FB20DC5EFAB2CE43" box="[1260,1362,1796,1821]" pageId="1" pageNumber="570" type="holotype">holotype</typeStatus>
.
</paragraph>
</subSubSection>
<subSubSection id="C35165106C3BFFC3FF0BDC7DFE5BCEAB" pageId="1" pageNumber="570" type="discussion">
<paragraph id="8BF4369B6C3BFFC3FF0BDC7DFE5BCEAB" blockId="1.[151,1436,1687,2037]" pageId="1" pageNumber="570">
<emphasis id="B93FEA896C3BFFC3FF0BDC7DFEDBCE1E" bold="true" box="[199,315,1831,1856]" pageId="1" pageNumber="570">Remarks.</emphasis>
This species has been referred to as 
<taxonomicName id="4C4B4D186C3BFFC3FD11DC72FC8CCE1E" box="[733,876,1832,1856]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="1" pageNumber="570" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C3BFFC3FD11DC72FC8CCE1E" box="[733,876,1832,1856]" italics="true" pageId="1" pageNumber="570">Guibemantis</emphasis>
</taxonomicName>
sp. 20 by 
<bibRefCitation id="EFDA4B6A6C3BFFC3FC28DC7DFB26CE1E" box="[996,1222,1831,1856]" pageId="1" pageNumber="570" 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="B93FEA896C3BFFC3FBF6DC73FB94CE1E" box="[1082,1140,1832,1856]" italics="true" pageId="1" pageNumber="570">et al.</emphasis>
(2009)
</bibRefCitation>
and 
<bibRefCitation id="EFDA4B6A6C3BFFC3FB32DC7DFEE7CE3A" pageId="1" pageNumber="570" refString="Wollenberg, K. C., 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), 1 - 15. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 11 - 217" type="journal article">
Wollenberg 
<emphasis id="B93FEA896C3BFFC3FA44DC73FF54CE3A" italics="true" pageId="1" pageNumber="570">et al.</emphasis>
(2011)
</bibRefCitation>
, as 
<taxonomicName id="4C4B4D186C3BFFC3FEFEDC16FE21CE3A" box="[306,449,1868,1892]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="1" pageNumber="570" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C3BFFC3FEFEDC16FE21CE3A" box="[306,449,1868,1892]" italics="true" pageId="1" pageNumber="570">Guibemantis</emphasis>
</taxonomicName>
sp. 
<taxonomicName id="4C4B4D186C3BFFC3FE3DDC16FD90CE3A" box="[497,624,1868,1893]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="1" pageNumber="570" phylum="Chordata" rank="species" species="timidus">
aff. 
<emphasis id="B93FEA896C3BFFC3FDD2DC16FD90CE3A" box="[542,624,1868,1892]" italics="true" pageId="1" pageNumber="570">timidus</emphasis>
</taxonomicName>
by Glaw &amp; Vences (2007: 446) and as 
<taxonomicName id="4C4B4D186C3BFFC3FBE2DC16FB5DCE3A" box="[1070,1213,1868,1892]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="1" pageNumber="570" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C3BFFC3FBE2DC16FB5DCE3A" box="[1070,1213,1868,1892]" italics="true" pageId="1" pageNumber="570">Guibemantis</emphasis>
</taxonomicName>
sp. Ca20 by 
<bibRefCitation id="EFDA4B6A6C3BFFC3FA9FDC16FEE9CED6" pageId="1" pageNumber="570" 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="B93FEA896C3BFFC3FA44DC17FF54CED6" italics="true" pageId="1" pageNumber="570">et al.</emphasis>
(2014)
</bibRefCitation>
. DNA sequence data for the 
<typeStatus id="54F088396C3BFFC3FD80DC35FD4ECED6" box="[588,686,1903,1928]" pageId="1" pageNumber="570" type="holotype">holotype</typeStatus>
specimen ZSM 2665/2007 are available for the mitochondrial 16S rRNA gene from 
<bibRefCitation id="EFDA4B6A6C3BFFC3FEABDCCEFDB1CEF2" box="[359,593,1940,1965]" pageId="1" pageNumber="570" 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="B93FEA896C3BFFC3FE0CDCCFFE1CCEF2" box="[448,508,1940,1964]" italics="true" pageId="1" pageNumber="570">et al.</emphasis>
(2009)
</bibRefCitation>
under GenBank accession number 
<accessionNumber id="9418AB786C3BFFC3FC3EDCCEFB83CEF3" box="[1010,1123,1940,1965]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/FJ559212" pageId="1" pageNumber="570">FJ559212</accessionNumber>
, and for the mitochondrial cytochrome 
<emphasis id="B93FEA896C3BFFC3FEE9DCE2FED3CE8E" box="[293,307,1976,2000]" italics="true" pageId="1" pageNumber="570">b</emphasis>
and cytochrome oxidase subunit 1 genes from 
<bibRefCitation id="EFDA4B6A6C3BFFC3FC95DCE2FB93CE8E" box="[857,1139,1976,2001]" pageId="1" pageNumber="570" refString="Wollenberg, K. C., 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), 1 - 15. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 11 - 217" type="journal article">
Wollenberg 
<emphasis id="B93FEA896C3BFFC3FC29DCE3FBC0CE8E" box="[997,1056,1976,2000]" italics="true" pageId="1" pageNumber="570">et al.</emphasis>
(2011)
</bibRefCitation>
under accession numbers 
<accessionNumber id="9418AB786C3BFFC3FF5BDC86FEEACEAB" box="[151,266,2012,2037]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/JN133051" pageId="1" pageNumber="570">JN133051</accessionNumber>
and 
<accessionNumber id="9418AB786C3BFFC3FE8CDC86FE57CEAB" box="[320,439,2012,2037]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/JN133275" pageId="1" pageNumber="570">JN133275</accessionNumber>
.
</paragraph>
</subSubSection>
<caption id="DF3466136C38FFC0FF5BDDE1FBB5CE51" httpUri="https://zenodo.org/record/244219/files/figure.png" pageId="2" pageNumber="571" targetBox="[150,1435,193,1699]" targetPageId="2">
<paragraph id="8BF4369B6C38FFC0FF5BDDE1FBB5CE51" blockId="2.[151,1435,1723,1807]" pageId="2" pageNumber="571">
<emphasis id="B93FEA896C38FFC0FF5BDDE1FEF4CF8E" bold="true" box="[151,276,1723,1745]" pageId="2" pageNumber="571">FIGURE 1.</emphasis>
<taxonomicName id="4C4B4D186C38FFC0FED7DDE1FDE1CF8F" box="[283,513,1723,1745]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="2" pageNumber="571" phylum="Chordata" rank="species" species="diphonus" status="sp. nov.">
<emphasis id="B93FEA896C38FFC0FED7DDE1FDE1CF8F" box="[283,513,1723,1745]" italics="true" pageId="2" pageNumber="571">Guibemantis diphonus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A20C57F26C38FFC0FDC4DDE6FDBACF8F" box="[520,602,1724,1745]" pageId="2" pageNumber="571" rank="species">
<emphasis id="B93FEA896C38FFC0FDC4DDE6FDBACF8F" bold="true" box="[520,602,1724,1745]" pageId="2" pageNumber="571">sp. nov.</emphasis>
</taxonomicNameLabel>
: A −B, male holotype ZSM 2665/2007 from Manombo in life in dorsolateral and ventral view; C −D, male holotype ZSM 2665/2007 in preservative (photographed in May 2015); E −F, male paratype ZSM 2664/2007 shortly after preservation (photographed on 24 February 2007). Scale bars = 5 mm.
</paragraph>
</caption>
<subSubSection id="C35165106C38FFC6FF0BDC6CFC06CAFB" lastPageId="4" lastPageNumber="573" pageId="2" pageNumber="571" type="diagnosis">
<paragraph id="8BF4369B6C38FFC1FF0BDC6CFD95C843" blockId="2.[151,1436,1846,2015]" lastBlockId="3.[151,1436,151,285]" lastPageId="3" lastPageNumber="572" pageId="2" pageNumber="571">
<emphasis id="B93FEA896C38FFC0FF0BDC6CFEA1CE11" bold="true" box="[199,321,1846,1871]" pageId="2" pageNumber="571">Diagnosis.</emphasis>
The species is allocated to the subgenus 
<taxonomicName id="4C4B4D186C38FFC0FCF8DC6CFC23CE10" box="[820,963,1846,1870]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="2" pageNumber="571" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C38FFC0FCF8DC6CFC23CE10" box="[820,963,1846,1870]" italics="true" pageId="2" pageNumber="571">Guibemantis</emphasis>
</taxonomicName>
in the genus 
<taxonomicName id="4C4B4D186C38FFC0FBA7DC6CFB1ACE10" box="[1131,1274,1846,1870]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="2" pageNumber="571" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C38FFC0FBA7DC6CFB1ACE10" box="[1131,1274,1846,1870]" italics="true" pageId="2" pageNumber="571">Guibemantis</emphasis>
</taxonomicName>
based on the structure of femoral glands in males (of 
<typeStatus id="54F088396C38FFC0FDB5DC01FD49CE2D" box="[633,681,1883,1907]" pageId="2" pageNumber="571">type</typeStatus>
1 sensu 
<bibRefCitation id="EFDA4B6A6C38FFC0FCD4DC00FC04CE2D" author="Glaw" box="[792,996,1882,1907]" pageId="2" pageNumber="571" refString="Glaw, F., Vences, M. &amp; Gossmann, V. (2000) A new species of Mantidactylus from Madagascar, with a comparative survey of internal femoral gland structure in the genus (Amphibia: Ranidae: Mantellinae). Journal of Natural History, 34, 1135 - 1154. http: // dx. doi. org / 10.1080 / 00222930050020140" type="journal article" year="2000">
Glaw 
<emphasis id="B93FEA896C38FFC0FCACDC01FC7DCE2C" box="[864,925,1882,1906]" italics="true" pageId="2" pageNumber="571">et al.</emphasis>
2000
</bibRefCitation>
; Vences 
<emphasis id="B93FEA896C38FFC0FB83DC01FB6CCE2C" box="[1103,1164,1882,1906]" italics="true" pageId="2" pageNumber="571">et al.</emphasis>
2007), arboreal habits, molecular phylogeny (
<bibRefCitation id="EFDA4B6A6C38FFC0FE5EDC24FD82CEC9" author="Vieites" box="[402,610,1918,1943]" pageId="2" pageNumber="571" 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" year="2009">
Vieites 
<emphasis id="B93FEA896C38FFC0FE24DC25FDC0CEC8" box="[488,544,1918,1942]" italics="true" pageId="2" pageNumber="571">et al.</emphasis>
2009
</bibRefCitation>
; 
<bibRefCitation id="EFDA4B6A6C38FFC0FDA0DC24FC8DCEC9" author="Wollenberg" box="[620,877,1918,1943]" pageId="2" pageNumber="571" refString="Wollenberg, K. C., 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), 1 - 15. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 11 - 217" type="journal article" year="2011">
Wollenberg 
<emphasis id="B93FEA896C38FFC0FD38DC25FCCCCEC8" box="[756,812,1918,1942]" italics="true" pageId="2" pageNumber="571">et al.</emphasis>
2011
</bibRefCitation>
; 
<bibRefCitation id="EFDA4B6A6C38FFC0FCB4DC24FBC6CEC9" author="Perl" box="[888,1062,1918,1943]" pageId="2" pageNumber="571" 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" year="2014">
Perl 
<emphasis id="B93FEA896C38FFC0FC60DC25FC04CEC8" box="[940,996,1918,1942]" italics="true" pageId="2" pageNumber="571">et al.</emphasis>
2014
</bibRefCitation>
), white single subgular vocal sac, smooth dorsal skin without dorsolateral ridges, enlarged terminal disks of fingers and toes and complete separation of lateral metatarsalia. The new species is distinguished from all species of the subgenus 
<taxonomicName id="4C4B4D186C38FFC0FB64DC9CFAB0CE80" box="[1192,1360,1990,2014]" class="Amphibia" family="Ranidae" genus="Pandanusicola" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="2" pageNumber="571" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C38FFC0FB64DC9CFAB0CE80" box="[1192,1360,1990,2014]" italics="true" pageId="2" pageNumber="571">Pandanusicola</emphasis>
</taxonomicName>
in the genus 
<taxonomicName id="4C4B4D186C39FFC1FF2CDBC2FE8FC9EE" box="[224,367,152,176]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C39FFC1FF2CDBC2FE8FC9EE" box="[224,367,152,176]" italics="true" pageId="3" pageNumber="572">Guibemantis</emphasis>
</taxonomicName>
by larger body size (male SVL 
<quantity id="4CB39B7E6C39FFC1FD12DBCDFCB9C9F1" box="[734,857,151,176]" metricMagnitude="-2" metricUnit="m" metricValue="3.5" metricValueMax="3.6" metricValueMin="3.4" pageId="3" pageNumber="572" unit="mm" value="35.0" valueMax="36.0" valueMin="34.0">34−36 mm</quantity>
vs. &lt;
<quantity id="4CB39B7E6C39FFC1FC6DDBC2FC12C9F1" box="[929,1010,152,176]" metricMagnitude="-2" metricUnit="m" metricValue="3.4" pageId="3" pageNumber="572" unit="mm" value="34.0">34 mm</quantity>
in males of 
<taxonomicName id="4C4B4D186C39FFC1FB4CDBC2FAC8C9EE" box="[1152,1320,152,176]" class="Amphibia" family="Ranidae" genus="Pandanusicola" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C39FFC1FB4CDBC2FAC8C9EE" box="[1152,1320,152,176]" italics="true" pageId="3" pageNumber="572">Pandanusicola</emphasis>
</taxonomicName>
, and &lt;
<quantity id="4CB39B7E6C39FFC1FA4CDBC2FF23C98A" metricMagnitude="-2" metricUnit="m" metricValue="3.1" pageId="3" pageNumber="572" unit="mm" value="31.0">31 mm</quantity>
in females of 
<taxonomicName id="4C4B4D186C39FFC1FEBFDBE6FDFBC98A" box="[371,539,188,212]" class="Amphibia" family="Ranidae" genus="Pandanusicola" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C39FFC1FEBFDBE6FDFBC98A" box="[371,539,188,212]" italics="true" pageId="3" pageNumber="572">Pandanusicola</emphasis>
</taxonomicName>
except 
<taxonomicName id="4C4B4D186C39FFC1FDB6DBE6FCDCC98A" box="[634,828,188,212]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="flavobrunneus">
<emphasis id="B93FEA896C39FFC1FDB6DBE6FCDCC98A" box="[634,828,188,212]" italics="true" pageId="3" pageNumber="572">G. flavobrunneus</emphasis>
</taxonomicName>
; Glaw &amp; Vences 2007; 
<bibRefCitation id="EFDA4B6A6C39FFC1FB92DBE6FAACC98A" author="Lehtinen" box="[1118,1356,188,213]" pageId="3" pageNumber="572" refString="Lehtinen, R. M., Glaw, F. &amp; Vences, M. (2011) Two new plant-breeding frog species (Anura: Mantellidae, Guibemantis) from southeastern Madagascar. Herpetological Journal, 21, 95 - 112." type="journal article" year="2011">
Lehtinen 
<emphasis id="B93FEA896C39FFC1FB00DBE7FAE8C98A" box="[1228,1288,188,212]" italics="true" pageId="3" pageNumber="572">et al.</emphasis>
2011
</bibRefCitation>
, 
<bibRefCitation id="EFDA4B6A6C39FFC1FA90DBE6FA78C98A" author="Lehtinen" box="[1372,1432,188,212]" pageId="3" pageNumber="572" refString="Lehtinen, R. M., Glaw, F., Andreone, F., Pabijan, M. &amp; Vences, M. (2012) A new species of putatively pond breeding frog of the genus Guibemantis from Madagascar. Copeia, 2012, 648 - 662. http: // dx. doi. org / 10.1643 / CH- 11 - 128" type="journal article" year="2012">2012</bibRefCitation>
; 
<bibRefCitation id="EFDA4B6A6C39FFC1FF5BDBBAFE8EC9A6" author="Vences" box="[151,366,224,248]" pageId="3" pageNumber="572" refString="Vences, M., Rakotoarison, A., Rakotondrazafy, A. M. A., Ratsoavina, F. M., Randrianiaina, R. D., Glaw, F., Lehtinen, R. M. &amp; Raxworthy, C. J. (2013) Assessing the diversity of phytotelmic frogs along Madagascar's east coast: redefinition of Guibemantis bicalcaratus (Boettger, 1913) and revalidation of Guibemantis methueni (Angel, 1929). Vertebrate Zoology, 63, 193 - 205." type="journal article" year="2013">
Vences 
<emphasis id="B93FEA896C39FFC1FF23DBBBFEC9C9A6" box="[239,297,224,248]" italics="true" pageId="3" pageNumber="572">et al.</emphasis>
2013
</bibRefCitation>
), numerous details of colouration, natural history, bioacoustics (where advertisement calls are known), and mitochondrial differentiation.
</paragraph>
<caption id="DF3466136C39FFC1FF5BDD44FB6FCFF0" httpUri="https://zenodo.org/record/244220/files/figure.png" pageId="3" pageNumber="572" targetBox="[151,1422,348,1544]" targetPageId="3">
<paragraph id="8BF4369B6C39FFC1FF5BDD44FB6FCFF0" blockId="3.[151,1436,1566,1710]" pageId="3" pageNumber="572">
<emphasis id="B93FEA896C39FFC1FF5BDD44FEF2CF6D" bold="true" box="[151,274,1566,1588]" pageId="3" pageNumber="572">FIGURE 2.</emphasis>
Views of palmar surfaces of left hands of species in the subgenus 
<taxonomicName id="4C4B4D186C39FFC1FC60DD44FBCECF6A" box="[940,1070,1566,1588]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C39FFC1FC60DD44FBCECF6A" box="[940,1070,1566,1588]" italics="true" pageId="3" pageNumber="572">Guibemantis</emphasis>
</taxonomicName>
(not to scale). Note the very distinct metacarpal tubercle of 
<taxonomicName id="4C4B4D186C39FFC1FE4CDD67FE19CF0C" box="[384,505,1596,1618]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C39FFC1FE4CDD67FE19CF0C" box="[384,505,1596,1618]" italics="true" pageId="3" pageNumber="572">G. diphonus</emphasis>
</taxonomicName>
which is smaller or more flattened in all other figured specimens (especially in the types of earlier names, e.g. 
<taxonomicName id="4C4B4D186C39FFC1FE97DD01FDBECF2F" box="[347,606,1627,1649]" class="Diplopoda" family="Chelodesmidae" genus="Rhacophorus" kingdom="Animalia" order="Polydesmida" pageId="3" pageNumber="572" phylum="Arthropoda" rank="species" species="mocquardii">
<emphasis id="B93FEA896C39FFC1FE97DD01FDBECF2F" box="[347,606,1627,1649]" italics="true" pageId="3" pageNumber="572">Rhacophorus mocquardii</emphasis>
</taxonomicName>
from the east coast locality Sahambendrana). Moreover, there are differences in webbing between fingers: although rudimentary in all species, it is almost absent in 
<taxonomicName id="4C4B4D186C39FFC1FBD5DD20FB75CFCE" box="[1049,1173,1658,1680]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C39FFC1FBD5DD20FB75CFCE" box="[1049,1173,1658,1680]" italics="true" pageId="3" pageNumber="572">G. diphonus</emphasis>
</taxonomicName>
, but rather distinct in 
<taxonomicName id="4C4B4D186C39FFC1FA4ADD20FF04CFF0" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="tornieri">
<emphasis id="B93FEA896C39FFC1FA4ADD20FF04CFF0" italics="true" pageId="3" pageNumber="572">G. tornieri</emphasis>
</taxonomicName>
and sometimes visible in 
<taxonomicName id="4C4B4D186C39FFC1FE20DDC3FD67CFF0" box="[492,647,1688,1710]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="depressiceps">
<emphasis id="B93FEA896C39FFC1FE20DDC3FD67CFF0" box="[492,647,1688,1710]" italics="true" pageId="3" pageNumber="572">G. depressiceps</emphasis>
</taxonomicName>
, and also recognizable in the 
<taxonomicName id="4C4B4D186C39FFC1FC79DDC2FBC9CFF0" box="[949,1065,1688,1710]" class="Diplopoda" family="Chelodesmidae" genus="Rhacophorus" kingdom="Animalia" order="Polydesmida" pageId="3" pageNumber="572" phylum="Arthropoda" rank="species" species="mocquardii">
<emphasis id="B93FEA896C39FFC1FC79DDC2FBC9CFF0" box="[949,1065,1688,1710]" italics="true" pageId="3" pageNumber="572">mocquardii</emphasis>
</taxonomicName>
holotype.
</paragraph>
</caption>
<paragraph id="8BF4369B6C39FFC1FF0BDD8CFA7CCE98" blockId="3.[151,1436,1750,1991]" pageId="3" pageNumber="572">
Within the subgenus 
<taxonomicName id="4C4B4D186C39FFC1FE7FDD8CFDA3CFB0" box="[435,579,1750,1774]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C39FFC1FE7FDD8CFDA3CFB0" box="[435,579,1750,1774]" italics="true" pageId="3" pageNumber="572">Guibemantis</emphasis>
</taxonomicName>
(comparative values from 
<bibRefCitation id="EFDA4B6A6C39FFC1FCA3DD8DFBBECFB1" author="Vences" box="[879,1118,1750,1775]" pageId="3" pageNumber="572" refString="Vences, M. &amp; Glaw, F. (2005) A new species of Mantidactylus from the east coast of Madagascar and its molecular phylogenetic relationships within the subgenus Guibemantis. Herpetological Journal, 15, 37 - 44." type="journal article" year="2005">Vences &amp; Glaw 2005</bibRefCitation>
and 
<tableCitation id="C6C903206C39FFC1FB58DD8CFB07CFB1" box="[1172,1255,1750,1775]" captionStart="TABLE 1" captionStartId="7.[125,202,454,474]" captionTargetBox="[142,1784,561,966]" captionTargetPageId="7" captionText="TABLE 1. Morphometric measurements of Guibemantis diphonus and comparative specimens of G. timidus and G. tornieri, all in mm. Abbreviations as in Materials and Methods except for: HT, holotype; PT, paratype; M, male; RHL, relative hindlimb length given as the point reached by tibiotarsal articulation when hindlimb is adpressed along body (1, posterior eye corner; 2, eye center; 3, anterior eye corner; 4, between eye and nostril)." httpUri="http://table.plazi.org/id/DF3466136C3DFFC5FFB1DA9CFC6ECB51" pageId="3" pageNumber="572" tableUuid="DF3466136C3DFFC5FFB1DA9CFC6ECB51">Table 1</tableCitation>
herein) the new species is distinguished from 
<taxonomicName id="4C4B4D186C39FFC1FE2CDDA0FD89CE4C" box="[480,617,1786,1810]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="kathrinae">
<emphasis id="B93FEA896C39FFC1FE2CDDA0FD89CE4C" box="[480,617,1786,1810]" italics="true" pageId="3" pageNumber="572">G. kathrinae</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C4B4D186C39FFC1FD52DDA0FCF1CE4C" box="[670,785,1786,1810]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="tornieri">
<emphasis id="B93FEA896C39FFC1FD52DDA0FCF1CE4C" box="[670,785,1786,1810]" italics="true" pageId="3" pageNumber="572">G. tornieri</emphasis>
</taxonomicName>
by its distinctly smaller body size (adult male SVL 
<quantity id="4CB39B7E6C39FFC1FA98DDA0FF23CE68" metricMagnitude="-2" metricUnit="m" metricValue="3.5" metricValueMax="3.6" metricValueMin="3.4" pageId="3" pageNumber="572" unit="mm" value="35.0" valueMax="36.0" valueMin="34.0">34−36 mm</quantity>
vs. 
<quantity id="4CB39B7E6C39FFC1FF3EDC44FE8ECE68" box="[242,366,1822,1847]" metricMagnitude="-2" metricUnit="m" metricValue="5.8" metricValueMax="5.9" metricValueMin="5.7" pageId="3" pageNumber="572" unit="mm" value="58.0" valueMax="59.0" valueMin="57.0">57−59 mm</quantity>
in 
<taxonomicName id="4C4B4D186C39FFC1FE5FDC45FDFECE68" box="[403,542,1822,1846]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="kathrinae">
<emphasis id="B93FEA896C39FFC1FE5FDC45FDFECE68" box="[403,542,1822,1846]" italics="true" pageId="3" pageNumber="572">G. kathrinae</emphasis>
</taxonomicName>
, 
<quantity id="4CB39B7E6C39FFC1FDE1DC44FD49CE68" box="[557,681,1822,1847]" metricMagnitude="-2" metricUnit="m" metricValue="5.0" metricValueMax="5.8" metricValueMin="4.2" pageId="3" pageNumber="572" unit="mm" value="50.0" valueMax="58.0" valueMin="42.0">42−58 mm</quantity>
in 
<taxonomicName id="4C4B4D186C39FFC1FD02DC45FCA3CE68" box="[718,835,1823,1846]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="tornieri">
<emphasis id="B93FEA896C39FFC1FD02DC45FCA3CE68" box="[718,835,1823,1846]" italics="true" pageId="3" pageNumber="572">G. tornieri</emphasis>
</taxonomicName>
). Its body size also seems to be smaller than in most specimens of 
<taxonomicName id="4C4B4D186C39FFC1FEF9DC18FE04CE04" box="[309,484,1858,1882]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="depressiceps">
<emphasis id="B93FEA896C39FFC1FEF9DC18FE04CE04" box="[309,484,1858,1882]" italics="true" pageId="3" pageNumber="572">G. depressiceps</emphasis>
</taxonomicName>
, despite overlap of values: male SVL is 
<quantity id="4CB39B7E6C39FFC1FC72DC18FBDBCE04" box="[958,1083,1858,1883]" metricMagnitude="-2" metricUnit="m" metricValue="3.85" metricValueMax="4.5" metricValueMin="3.2" pageId="3" pageNumber="572" unit="mm" value="38.5" valueMax="45.0" valueMin="32.0">32−45 mm</quantity>
in 
<taxonomicName id="4C4B4D186C39FFC1FBAFDC18FAF2CE04" box="[1123,1298,1858,1882]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="depressiceps">
<emphasis id="B93FEA896C39FFC1FBAFDC18FAF2CE04" box="[1123,1298,1858,1882]" italics="true" pageId="3" pageNumber="572">G. depressiceps</emphasis>
</taxonomicName>
, with many specimens reaching sizes&gt; 
<quantity id="4CB39B7E6C39FFC1FE1EDC3DFDC5CE20" box="[466,549,1895,1919]" metricMagnitude="-2" metricUnit="m" metricValue="4.0" pageId="3" pageNumber="572" unit="mm" value="40.0">40 mm</quantity>
. Moreover, rudimentary webbing is almost absent in 
<taxonomicName id="4C4B4D186C39FFC1FB4CDC3DFAE6CE20" box="[1152,1286,1894,1918]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C39FFC1FB4CDC3DFAE6CE20" box="[1152,1286,1894,1918]" italics="true" pageId="3" pageNumber="572">G. diphonus</emphasis>
</taxonomicName>
whereas it is rather distinct in 
<taxonomicName id="4C4B4D186C39FFC1FE91DCD0FE35CEFC" box="[349,469,1930,1954]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="tornieri">
<emphasis id="B93FEA896C39FFC1FE91DCD0FE35CEFC" box="[349,469,1930,1954]" italics="true" pageId="3" pageNumber="572">G. tornieri</emphasis>
</taxonomicName>
and often well visible in 
<emphasis id="B93FEA896C39FFC1FCCDDCD0FC58CEFC" box="[769,952,1930,1954]" italics="true" pageId="3" pageNumber="572">
<taxonomicName id="4C4B4D186C39FFC1FCCDDCD0FC55CEFC" box="[769,949,1930,1954]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="depressiceps">G. depressiceps</taxonomicName>
.
</emphasis>
In addition, the metacarpal tubercle of 
<taxonomicName id="4C4B4D186C39FFC1FA48DCD0FF1ECE98" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C39FFC1FA48DCD0FF1ECE98" italics="true" pageId="3" pageNumber="572">G. diphonus</emphasis>
</taxonomicName>
is larger and more distinct compared to many though not all specimens of 
<taxonomicName id="4C4B4D186C39FFC1FBF3DCF5FB0BCE98" box="[1087,1259,1966,1990]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="depressiceps">
<emphasis id="B93FEA896C39FFC1FBF3DCF5FB0BCE98" box="[1087,1259,1966,1990]" italics="true" pageId="3" pageNumber="572">G. depressiceps</emphasis>
</taxonomicName>
and 
<emphasis id="B93FEA896C39FFC1FAEDDCF5FA7CCE98" box="[1313,1436,1967,1990]" italics="true" pageId="3" pageNumber="572">
<taxonomicName id="4C4B4D186C39FFC1FAEDDCF5FA78CE98" box="[1313,1432,1967,1990]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="3" pageNumber="572" phylum="Chordata" rank="species" species="tornieri">G. tornieri</taxonomicName>
.
</emphasis>
</paragraph>
<paragraph id="8BF4369B6C3EFFC6FF0BDBC2FD50CBF6" blockId="4.[151,1437,151,2016]" pageId="4" pageNumber="573">
In a direct comparison with its sister species, 
<taxonomicName id="4C4B4D186C3EFFC6FD18DBC2FCA7C9EE" box="[724,839,152,176]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C3EFFC6FD18DBC2FCA7C9EE" box="[724,839,152,176]" italics="true" pageId="4" pageNumber="573">G. timidus</emphasis>
</taxonomicName>
, the new species 
<taxonomicName id="4C4B4D186C3EFFC6FBD8DBC2FB7BC9EE" box="[1044,1179,152,176]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="diphonus" status="sp. nov.">
<emphasis id="B93FEA896C3EFFC6FBD8DBC2FB7BC9EE" box="[1044,1179,152,176]" italics="true" pageId="4" pageNumber="573">G. diphonus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A20C57F26C3EFFC6FB69DBCDFAE0C9EE" box="[1189,1280,151,176]" pageId="4" pageNumber="573" rank="species">
<emphasis id="B93FEA896C3EFFC6FB69DBCDFAE0C9EE" bold="true" box="[1189,1280,151,176]" pageId="4" pageNumber="573">sp. nov.</emphasis>
</taxonomicNameLabel>
differs in the following morphological characters: (1) 
<taxonomicName id="4C4B4D186C3EFFC6FDAADBE6FD0EC98A" box="[614,750,188,212]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C3EFFC6FDAADBE6FD0EC98A" box="[614,750,188,212]" italics="true" pageId="4" pageNumber="573">G. diphonus</emphasis>
</taxonomicName>
has a smaller average body size of males (average SVL 
<quantity id="4CB39B7E6C3EFFC6FA4CDBE6FF23C9A9" metricMagnitude="-2" metricUnit="m" metricValue="3.5" pageId="4" pageNumber="573" unit="mm" value="35.0">35 mm</quantity>
vs. 
<quantity id="4CB39B7E6C3EFFC6FF3CDBBAFEA0C9A6" box="[240,320,224,248]" metricMagnitude="-2" metricUnit="m" metricValue="4.1" pageId="4" pageNumber="573" unit="mm" value="41.0">41 mm</quantity>
) despite overlapping values (
<quantity id="4CB39B7E6C3EFFC6FD44DB85FCE4C9A9" box="[648,772,223,248]" metricMagnitude="-2" metricUnit="m" metricValue="3.5" metricValueMax="3.6" metricValueMin="3.4" pageId="4" pageNumber="573" unit="mm" value="35.0" valueMax="36.0" valueMin="34.0">34−36 mm</quantity>
vs. 
<quantity id="4CB39B7E6C3EFFC6FCFEDB85FC4EC9A6" box="[818,942,223,248]" metricMagnitude="-2" metricUnit="m" metricValue="3.85" metricValueMax="4.5" metricValueMin="3.2" pageId="4" pageNumber="573" unit="mm" value="38.5" valueMax="45.0" valueMin="32.0">32−45 mm</quantity>
) (n = 2 vs. n = 10; 
<taxonomicName id="4C4B4D186C3EFFC6FB48DBBAFB15C9A6" box="[1156,1269,224,248]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C3EFFC6FB48DBBAFB15C9A6" box="[1156,1269,224,248]" italics="true" pageId="4" pageNumber="573">G. timidus</emphasis>
</taxonomicName>
measurements from 
<bibRefCitation id="EFDA4B6A6C3EFFC6FF1ADA5FFE2BC843" author="Vences" box="[214,459,260,285]" pageId="4" pageNumber="573" refString="Vences, M. &amp; Glaw, F. (2005) A new species of Mantidactylus from the east coast of Madagascar and its molecular phylogenetic relationships within the subgenus Guibemantis. Herpetological Journal, 15, 37 - 44." type="journal article" year="2005">Vences &amp; Glaw 2005</bibRefCitation>
and 
<tableCitation id="C6C903206C3EFFC6FDCADA5EFDBBC842" box="[518,603,260,285]" captionStart="TABLE 1" captionStartId="7.[125,202,454,474]" captionTargetBox="[142,1784,561,966]" captionTargetPageId="7" captionText="TABLE 1. Morphometric measurements of Guibemantis diphonus and comparative specimens of G. timidus and G. tornieri, all in mm. Abbreviations as in Materials and Methods except for: HT, holotype; PT, paratype; M, male; RHL, relative hindlimb length given as the point reached by tibiotarsal articulation when hindlimb is adpressed along body (1, posterior eye corner; 2, eye center; 3, anterior eye corner; 4, between eye and nostril)." httpUri="http://table.plazi.org/id/DF3466136C3DFFC5FFB1DA9CFC6ECB51" pageId="4" pageNumber="573" tableUuid="DF3466136C3DFFC5FFB1DA9CFC6ECB51">Table 1</tableCitation>
herein). (2) 
<taxonomicName id="4C4B4D186C3EFFC6FD21DA5EFC94C842" box="[749,884,260,284]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C3EFFC6FD21DA5EFC94C842" box="[749,884,260,284]" italics="true" pageId="4" pageNumber="573">G. diphonus</emphasis>
</taxonomicName>
has a very distinct metacarpal tubercle. Despite variation in this character, this tubercle is typically more flat and less distinct in 
<taxonomicName id="4C4B4D186C3EFFC6FBDFDA72FB64C81E" box="[1043,1156,296,320]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C3EFFC6FBDFDA72FB64C81E" box="[1043,1156,296,320]" italics="true" pageId="4" pageNumber="573">G. timidus</emphasis>
</taxonomicName>
(
<figureCitation id="13702A1E6C3EFFC6FB5EDA7DFB37C81E" box="[1170,1239,295,320]" captionStart="FIGURE 2" captionStartId="3.[151,250,1566,1588]" captionTargetBox="[151,1422,348,1544]" captionTargetId="figure@3.[151,1435,337,1545]" captionTargetPageId="3" captionText="FIGURE 2. Views of palmar surfaces of left hands of species in the subgenus Guibemantis (not to scale). Note the very distinct metacarpal tubercle of G. diphonus which is smaller or more flattened in all other figured specimens (especially in the types of earlier names, e. g. Rhacophorus mocquardii from the east coast locality Sahambendrana). Moreover, there are differences in webbing between fingers: although rudimentary in all species, it is almost absent in G. diphonus, but rather distinct in G. tornieri and sometimes visible in G. depressiceps, and also recognizable in the mocquardii holotype." httpUri="https://zenodo.org/record/244220/files/figure.png" pageId="4" pageNumber="573">Fig. 2</figureCitation>
). (3) Hand length in 
<taxonomicName id="4C4B4D186C3EFFC6FF7FDA16FEC4C83A" box="[179,292,332,356]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C3EFFC6FF7FDA16FEC4C83A" box="[179,292,332,356]" italics="true" pageId="4" pageNumber="573">G. timidus</emphasis>
</taxonomicName>
is typically shorter than in other species of the subgenus 
<taxonomicName id="4C4B4D186C3EFFC6FC6EDA16FBD1C83A" box="[930,1073,332,356]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C3EFFC6FC6EDA16FBD1C83A" box="[930,1073,332,356]" italics="true" pageId="4" pageNumber="573">Guibemantis</emphasis>
</taxonomicName>
(HAL/SVL ratio &lt;0.32, 
<bibRefCitation id="EFDA4B6A6C3EFFC6FA80DA17FED2C8D6" author="Vences" pageId="4" pageNumber="573" refString="Vences, M. &amp; Glaw, F. (2005) A new species of Mantidactylus from the east coast of Madagascar and its molecular phylogenetic relationships within the subgenus Guibemantis. Herpetological Journal, 15, 37 - 44." type="journal article" year="2005">Vences &amp; Glaw 2005</bibRefCitation>
), and this is also true for the two south-eastern 
<taxonomicName id="4C4B4D186C3EFFC6FCF3DA2AFC50C8D6" box="[831,944,368,392]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C3EFFC6FCF3DA2AFC50C8D6" box="[831,944,368,392]" italics="true" pageId="4" pageNumber="573">G. timidus</emphasis>
</taxonomicName>
specimens from Beronono measured herein (HAL/SVL 0.28−0.29; 
<tableCitation id="C6C903206C3EFFC6FE51DACEFE0EC8F2" box="[413,494,404,429]" captionStart="TABLE 1" captionStartId="7.[125,202,454,474]" captionTargetBox="[142,1784,561,966]" captionTargetPageId="7" captionText="TABLE 1. Morphometric measurements of Guibemantis diphonus and comparative specimens of G. timidus and G. tornieri, all in mm. Abbreviations as in Materials and Methods except for: HT, holotype; PT, paratype; M, male; RHL, relative hindlimb length given as the point reached by tibiotarsal articulation when hindlimb is adpressed along body (1, posterior eye corner; 2, eye center; 3, anterior eye corner; 4, between eye and nostril)." httpUri="http://table.plazi.org/id/DF3466136C3DFFC5FFB1DA9CFC6ECB51" pageId="4" pageNumber="573" tableUuid="DF3466136C3DFFC5FFB1DA9CFC6ECB51">Table 1</tableCitation>
). In the 
<typeStatus id="54F088396C3EFFC6FD80DACEFD4EC8F3" box="[588,686,404,429]" pageId="4" pageNumber="573" type="holotype">holotype</typeStatus>
of 
<taxonomicName id="4C4B4D186C3EFFC6FD1EDACEFCB7C8F2" box="[722,855,404,428]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C3EFFC6FD1EDACEFCB7C8F2" box="[722,855,404,428]" italics="true" pageId="4" pageNumber="573">G. diphonus</emphasis>
</taxonomicName>
relative hand length is 0.32 and thus longer than the range known from 
<emphasis id="B93FEA896C3EFFC6FEA3DAE2FE07C88E" box="[367,487,440,464]" italics="true" pageId="4" pageNumber="573">
<taxonomicName id="4C4B4D186C3EFFC6FEA3DAE2FE03C88E" box="[367,483,440,464]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="timidus">G. timidus</taxonomicName>
,
</emphasis>
although in the 
<typeStatus id="54F088396C3EFFC6FD6DDAE2FCE1C88E" box="[673,769,440,464]" pageId="4" pageNumber="573" type="paratype">paratype</typeStatus>
the value is within the 
<taxonomicName id="4C4B4D186C3EFFC6FBC7DAE2FB9DC88E" box="[1035,1149,440,464]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C3EFFC6FBC7DAE2FB9DC88E" box="[1035,1149,440,464]" italics="true" pageId="4" pageNumber="573">G. timidus</emphasis>
</taxonomicName>
range (0.29). Despite the low sample size it might be concluded that relative hand length values at least in some specimens of 
<taxonomicName id="4C4B4D186C3EFFC6FAD9DA86FA7CC8AA" box="[1301,1436,476,500]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C3EFFC6FAD9DA86FA7CC8AA" box="[1301,1436,476,500]" italics="true" pageId="4" pageNumber="573">G. diphonus</emphasis>
</taxonomicName>
are higher than in 
<taxonomicName id="4C4B4D186C3EFFC6FEBFD95AFE08CB46" box="[371,488,512,536]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C3EFFC6FEBFD95AFE08CB46" box="[371,488,512,536]" italics="true" pageId="4" pageNumber="573">G. timidus</emphasis>
</taxonomicName>
. (4) 
<taxonomicName id="4C4B4D186C3EFFC6FDEBD95AFD51CB46" box="[551,689,512,536]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C3EFFC6FDEBD95AFD51CB46" box="[551,689,512,536]" italics="true" pageId="4" pageNumber="573">G. diphonus</emphasis>
</taxonomicName>
has in general a more contrasted and vivid colouration than 
<emphasis id="B93FEA896C3EFFC6FA48D95AFF11CB62" italics="true" pageId="4" pageNumber="573">
<taxonomicName id="4C4B4D186C3EFFC6FA48D95AFF0DCB62" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="timidus">G. timidus</taxonomicName>
.
</emphasis>
Patterns like the dark dorsal crossbands on shanks (seen in the two 
<taxonomicName id="4C4B4D186C3EFFC6FBCCD97EFB68CB62" box="[1024,1160,548,572]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C3EFFC6FBCCD97EFB68CB62" box="[1024,1160,548,572]" italics="true" pageId="4" pageNumber="573">G. diphonus</emphasis>
</taxonomicName>
specimens), the reddish colour on forelimbs, dark dorsal patches, broad light dorsal stripe or dark colour on flanks (seen each in only one of the two 
<taxonomicName id="4C4B4D186C3EFFC6FF3BD936FE9FCBDA" box="[247,383,620,644]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C3EFFC6FF3BD936FE9FCBDA" box="[247,383,620,644]" italics="true" pageId="4" pageNumber="573">G. diphonus</emphasis>
</taxonomicName>
specimens) were never present in any of the relatively large number of 
<taxonomicName id="4C4B4D186C3EFFC6FB05D936FADECBDA" box="[1225,1342,620,644]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C3EFFC6FB05D936FADECBDA" box="[1225,1342,620,644]" italics="true" pageId="4" pageNumber="573">G. timidus</emphasis>
</taxonomicName>
that we observed as preserved vouchers and in the wild.
</paragraph>
<paragraph id="8BF4369B6C3EFFC6FF0BD9EEFB43CA03" blockId="4.[151,1437,151,2016]" pageId="4" pageNumber="573">
Molecular phylogenetic data (
<bibRefCitation id="EFDA4B6A6C3EFFC6FDEED9EEFCE0CB92" author="Vieites" box="[546,768,692,717]" pageId="4" pageNumber="573" 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" year="2009">
Vieites 
<emphasis id="B93FEA896C3EFFC6FDB1D9EFFD59CB92" box="[637,697,692,716]" italics="true" pageId="4" pageNumber="573">et al.</emphasis>
2009
</bibRefCitation>
; 
<bibRefCitation id="EFDA4B6A6C3EFFC6FCC3D9EEFBFECB92" author="Wollenberg" box="[783,1054,692,717]" pageId="4" pageNumber="573" refString="Wollenberg, K. C., 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), 1 - 15. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 11 - 217" type="journal article" year="2011">
Wollenberg 
<emphasis id="B93FEA896C3EFFC6FC50D9EFFC38CB92" box="[924,984,692,716]" italics="true" pageId="4" pageNumber="573">et al.</emphasis>
2011
</bibRefCitation>
; 
<bibRefCitation id="EFDA4B6A6C3EFFC6FBE1D9EEFB07CB92" author="Perl" box="[1069,1255,692,717]" pageId="4" pageNumber="573" 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" year="2014">
Perl 
<emphasis id="B93FEA896C3EFFC6FBA9D9EFFB42CB92" box="[1125,1186,692,716]" italics="true" pageId="4" pageNumber="573">et al.</emphasis>
2014
</bibRefCitation>
) place the new species with moderate to high support as sister species of 
<emphasis id="B93FEA896C3EFFC6FCEAD982FC40CBAE" box="[806,928,728,752]" italics="true" pageId="4" pageNumber="573">
<taxonomicName id="4C4B4D186C3EFFC6FCEAD982FC7CCBAE" box="[806,924,728,752]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="timidus">G. timidus</taxonomicName>
.
</emphasis>
Its pairwise genetic divergence to 
<taxonomicName id="4C4B4D186C3EFFC6FAE6D982FA7CCBAE" box="[1322,1436,728,752]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C3EFFC6FAE6D982FA7CCBAE" box="[1322,1436,728,752]" italics="true" pageId="4" pageNumber="573">G. timidus</emphasis>
</taxonomicName>
in the analysed 16S rRNA gene fragment is 4.4% (
<bibRefCitation id="EFDA4B6A6C3EFFC6FD25D9A6FC24CA4A" author="Vieites" box="[745,964,764,789]" pageId="4" pageNumber="573" 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" year="2009">
Vieites 
<emphasis id="B93FEA896C3EFFC6FC8FD9A7FC9ECA4A" box="[835,894,764,788]" italics="true" pageId="4" pageNumber="573">et al.</emphasis>
2009
</bibRefCitation>
), and differentiation to all other nominal species of 
<taxonomicName id="4C4B4D186C3EFFC6FED8D87AFE44CA66" box="[276,420,800,824]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C3EFFC6FED8D87AFE44CA66" box="[276,420,800,824]" italics="true" pageId="4" pageNumber="573">Guibemantis</emphasis>
</taxonomicName>
is&gt; 5%. This molecular divergence was also confirmed in comparison to a 
<taxonomicName id="4C4B4D186C3EFFC6FAEBD87AFA7CCA66" box="[1319,1436,800,824]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="4" pageNumber="573" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C3EFFC6FAEBD87AFA7CCA66" box="[1319,1436,800,824]" italics="true" pageId="4" pageNumber="573">G. timidus</emphasis>
</taxonomicName>
sample from a syntopic specimen from Manombo (GenBank accession number 
<accessionNumber id="9418AB786C3EFFC6FBD9D81EFB77CA03" box="[1045,1175,836,861]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/HQ456658" pageId="4" pageNumber="573">HQ456658</accessionNumber>
).
</paragraph>
<paragraph id="8BF4369B6C3EFFC6FF0BD83DFC06CAFB" blockId="4.[151,1437,151,2016]" pageId="4" pageNumber="573">
Furthermore, the species differs from all species in the subgenus by its advertisement call consisting of two distinctly different note 
<typeStatus id="54F088396C3EFFC6FE68D8D7FE3FCAFB" box="[420,479,909,933]" pageId="4" pageNumber="573">types</typeStatus>
(see call description and comparisons below).
</paragraph>
</subSubSection>
<subSubSection id="C35165106C3EFFCAFF0BD8F5FC32C88E" lastPageId="8" lastPageNumber="577" pageId="4" pageNumber="573" type="description">
<paragraph id="8BF4369B6C3EFFC6FF0BD8F5FE6FCF55" blockId="4.[151,1437,151,2016]" pageId="4" pageNumber="573">
<emphasis id="B93FEA896C3EFFC6FF0BD8F5FDFACA96" bold="true" box="[199,538,943,968]" pageId="4" pageNumber="573">
Description of the 
<typeStatus id="54F088396C3EFFC6FE60D8F5FDF5CA96" box="[428,533,943,968]" pageId="4" pageNumber="573" type="holotype">holotype</typeStatus>
.
</emphasis>
Adult male in excellent state of preservation. A small piece of muscle tissue from right thigh removed for molecular analysis. For measurements see 
<tableCitation id="C6C903206C3EFFC6FC2ED88EFBD9CAB2" box="[994,1081,980,1005]" captionStart="TABLE 1" captionStartId="7.[125,202,454,474]" captionTargetBox="[142,1784,561,966]" captionTargetPageId="7" captionText="TABLE 1. Morphometric measurements of Guibemantis diphonus and comparative specimens of G. timidus and G. tornieri, all in mm. Abbreviations as in Materials and Methods except for: HT, holotype; PT, paratype; M, male; RHL, relative hindlimb length given as the point reached by tibiotarsal articulation when hindlimb is adpressed along body (1, posterior eye corner; 2, eye center; 3, anterior eye corner; 4, between eye and nostril)." httpUri="http://table.plazi.org/id/DF3466136C3DFFC5FFB1DA9CFC6ECB51" pageId="4" pageNumber="573" tableUuid="DF3466136C3DFFC5FFB1DA9CFC6ECB51">Table 1</tableCitation>
. Body relatively slender; head longer than wide, wider than body; snout slightly pointed in dorsal and relatively truncate in lateral view; nostrils directed laterally, slightly protuberant; canthus rostralis distinct, straight; loreal region concave; tympanum distinct, relatively small, its diameter 64% of eye diameter; distinct supratympanic fold; tongue ovoid, distinctly bifid posteriorly; vomerine teeth as one distinct oblong group posterolateral of each choana; choanae medium-sized, rounded. Forelimbs slender; subarticular tubercles distinct and single; central metacarpal tubercle large and oval, outer metacarpal tubercle smaller and elongated; a small prepollex (which could also be considered as an inner metacarpal tubercle) at the base of the first finger recognizable. Fingers without webbing; relative finger length 1&lt;2&lt;4&lt;3; finger disks distinctly enlarged; nuptial pads absent. Limbs slender; when adpressed along body, the tibiotarsal articulation reaches beyond the eye; lateral metatarsalia separated; inner metatarsal tubercle distinct, larger than outer metatarsal tubercle; webbing formula of the foot 1(1), 
<date id="FFF5105B6C3EFFC6FC05DE66FC00CC0B" box="[969,992,1340,1365]" pageId="4" pageNumber="573">2i</date>
(1.5), 2e(0.5), 
<date id="FFF5105B6C3EFFC6FB48DE66FB79CC0B" box="[1156,1177,1340,1365]" pageId="4" pageNumber="573">3i</date>
(2), 3e(1), 
<date id="FFF5105B6C3EFFC6FAD8DE66FACCCC0B" box="[1300,1324,1340,1365]" pageId="4" pageNumber="573">4i</date>
(2), 4e(2), 5(1) (because only one subarticular tubercle is recognizable on the second toe, the relative extension of web on this toe can only be estimated); webbing according to the formula of 
<bibRefCitation id="EFDA4B6A6C3EFFC6FCBDDEDDFB9CCCC1" author="Savage" box="[881,1148,1415,1440]" pageId="4" pageNumber="573" refString="Savage, J. M. &amp; Heyer, W. R. (1997) Digital webbing formulae for anurans: a refinement. Herpetological Review, 28, 131." type="journal article" year="1997">Savage &amp; Heyer (1997)</bibRefCitation>
<date id="FFF5105B6C3EFFC6FB48DED2FB2DCCC1" box="[1156,1229,1415,1439]" pageId="4" pageNumber="573">I 2 – 2</date>
+ 
<date id="FFF5105B6C3EFFC6FB11DED2FAE5CCC1" box="[1245,1285,1415,1439]" pageId="4" pageNumber="573">II 1</date>
+ – 3 
<date id="FFF5105B6C3EFFC6FA8CDED2FF16CC9D" pageId="4" pageNumber="573">III 2 – 3 IV 3 – 2</date>
V. Relative toe length 1&lt;2&lt;3=5&lt;4. Skin on the dorsum smooth; ventral skin smooth on throat and chest, slightly granular on belly. Femoral glands indistinct from external view, but of 
<typeStatus id="54F088396C3EFFC6FBC4DE8AFBD8CCB6" box="[1032,1080,1488,1512]" pageId="4" pageNumber="573">type</typeStatus>
1 from internal view as defined by 
<bibRefCitation id="EFDA4B6A6C3EFFC6FF76DEA8FE6ACF55" box="[186,394,1522,1547]" pageId="4" pageNumber="573" refString="Glaw, F., Vences, M. &amp; Gossmann, V. (2000) A new species of Mantidactylus from Madagascar, with a comparative survey of internal femoral gland structure in the genus (Amphibia: Ranidae: Mantellinae). Journal of Natural History, 34, 1135 - 1154. http: // dx. doi. org / 10.1080 / 00222930050020140" type="journal article">
Glaw 
<emphasis id="B93FEA896C3EFFC6FF31DEA9FED6CF55" box="[253,310,1523,1547]" italics="true" pageId="4" pageNumber="573">et al.</emphasis>
(2000)
</bibRefCitation>
.
</paragraph>
<paragraph id="8BF4369B6C3EFFC6FF0BDD42FCD4CE11" blockId="4.[151,1437,151,2016]" pageId="4" pageNumber="573">
In preservative, dorsal ground colour uniformly brown, with a broad cream middorsal stripe of varying width (
<quantity id="4CB39B7E6C3EFFC6FF6CDD61FEE1CF0D" box="[160,257,1595,1619]" metricMagnitude="-3" metricUnit="m" metricValue="2.0" metricValueMax="3.0" metricValueMin="1.0" pageId="4" pageNumber="573" unit="mm" value="2.0" valueMax="3.0" valueMin="1.0">1−3 mm</quantity>
) and with irregular edges, running from snout tip to vent. Another transversal cream stripe of ca. 
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width between the anterior edges of the eyes. Tympanum brown. Whitish colour below the tympanun fading into grey towards the eye. Flanks brown marked with few greyish spots in the inguinal region. Upper arm and proximal part of the lower arm largely cream with few dark brown spots. Hands and fingers brown with dark brown spots. Hindlimbs brown with 3−4 dark brown crossbands. Ventral surfaces fading from bright white on the throat and cream white on the chest to light brown with some whitish marbling on the limbs. In general, colouration in life (
<figureCitation id="13702A1E6C3EFFC6FF53DC48FF00CE75" box="[159,224,1810,1835]" captionStart="FIGURE 1" captionStartId="2.[151,250,1723,1745]" captionTargetBox="[150,1435,193,1699]" captionTargetId="figure@2.[150,1436,193,1702]" captionTargetPageId="2" captionText="FIGURE 1. Guibemantis diphonus sp. nov.: A − B, male holotype ZSM 2665 / 2007 from Manombo in life in dorsolateral and ventral view; C − D, male holotype ZSM 2665 / 2007 in preservative (photographed in May 2015); E − F, male paratype ZSM 2664 / 2007 shortly after preservation (photographed on 24 February 2007). Scale bars = 5 mm." httpUri="https://zenodo.org/record/244219/files/figure.png" pageId="4" pageNumber="573">Fig. 1</figureCitation>
) is similar to that in preservative, but the yellow patches in the inguinal region and the reddish brown areas on the forelimbs have faded to grey and cream respectively.
</paragraph>
<paragraph id="8BF4369B6C3EFFC7FF0BDC00FBD1C83B" blockId="4.[151,1437,151,2016]" lastBlockId="5.[151,1437,151,861]" lastPageId="5" lastPageNumber="574" pageId="4" pageNumber="573">
<emphasis id="B93FEA896C3EFFC6FF0BDC00FE16CE2D" bold="true" box="[199,502,1882,1907]" pageId="4" pageNumber="573">Morphological variation.</emphasis>
Morphometric measurements of the 
<typeStatus id="54F088396C3EFFC6FC61DC00FBEFCE2D" box="[941,1039,1882,1907]" pageId="4" pageNumber="573" type="holotype">holotype</typeStatus>
and the 
<typeStatus id="54F088396C3EFFC6FBB1DC01FB3ECE2D" box="[1149,1246,1883,1907]" pageId="4" pageNumber="573" type="paratype">paratype</typeStatus>
are provided in 
<tableCitation id="C6C903206C3EFFC6FF5BDC24FF0CCEC9" box="[151,236,1918,1943]" captionStart="TABLE 1" captionStartId="7.[125,202,454,474]" captionTargetBox="[142,1784,561,966]" captionTargetPageId="7" captionText="TABLE 1. Morphometric measurements of Guibemantis diphonus and comparative specimens of G. timidus and G. tornieri, all in mm. Abbreviations as in Materials and Methods except for: HT, holotype; PT, paratype; M, male; RHL, relative hindlimb length given as the point reached by tibiotarsal articulation when hindlimb is adpressed along body (1, posterior eye corner; 2, eye center; 3, anterior eye corner; 4, between eye and nostril)." httpUri="http://table.plazi.org/id/DF3466136C3DFFC5FFB1DA9CFC6ECB51" pageId="4" pageNumber="573" tableUuid="DF3466136C3DFFC5FFB1DA9CFC6ECB51">Table 1</tableCitation>
. The 
<typeStatus id="54F088396C3EFFC6FEE2DCDAFE6ECEC6" box="[302,398,1920,1944]" pageId="4" pageNumber="573" type="paratype">paratype</typeStatus>
resembles the 
<typeStatus id="54F088396C3EFFC6FDF6DC24FD7CCEC9" box="[570,668,1918,1943]" pageId="4" pageNumber="573" type="holotype">holotype</typeStatus>
in morphology, but is slightly larger, has relative shorter hindlimbs (tibiotarsal articulation reaches the eye versus beyond the eye), slightly more webbing between the toes with webbing formula 1(1), 
<date id="FFF5105B6C3EFFC6FE51DC9CFE54CE81" box="[413,436,1990,2015]" pageId="4" pageNumber="573">2i</date>
(1.5), 2e(0.5), 
<date id="FFF5105B6C3EFFC6FD9ADC9CFD8CCE81" box="[598,620,1990,2015]" pageId="4" pageNumber="573">3i</date>
(2), 3e(0.5), 
<date id="FFF5105B6C3EFFC6FD37DC9CFCF2CE81" box="[763,786,1990,2015]" pageId="4" pageNumber="573">4i</date>
(2), 4e(2), 5(1), a slightly smaller tympanum (56% of eye diameter), and toe 3 slightly shorter than toe 5 (relative toe length 1&lt;2&lt;3&lt;5&lt;4). It moreover differs in colouration: Head and dorsum are light brown with some faint greenish tint partly recognizable. Few poorly distinct brown patches in the vertebral region and between the eyes, and a small dark spot in the middle of the snout. A light middorsal stripe or line and a stripe between the eyes are lacking. Flanks are darker brown with numerous white spots. Dorsal surfaces of arms and hindlimbs similar to the 
<typeStatus id="54F088396C3FFFC7FCF9DA7DFC7AC81E" box="[821,922,295,320]" pageId="5" pageNumber="574" type="holotype">holotype</typeStatus>
, but slighty lighter and with more contrasting crossbands. Ventral colour similar to the 
<typeStatus id="54F088396C3FFFC7FDADDA16FD26C83B" box="[609,710,332,357]" pageId="5" pageNumber="574" type="holotype">holotype</typeStatus>
, but slightly lighter on the belly.
</paragraph>
<paragraph id="8BF4369B6C3FFFC7FF0BDA35FD54CA03" blockId="5.[151,1437,151,861]" pageId="5" pageNumber="574">
<emphasis id="B93FEA896C3FFFC7FF0BDA35FE4EC8D6" bold="true" box="[199,430,367,392]" pageId="5" pageNumber="574">Advertisement call.</emphasis>
Vocalizations of two individuals of 
<taxonomicName id="4C4B4D186C3FFFC7FC8FDA2AFBA0C8D6" box="[835,1088,368,392]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="5" pageNumber="574" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C3FFFC7FC8FDA2AFBA0C8D6" box="[835,1088,368,392]" italics="true" pageId="5" pageNumber="574">Guibemantis diphonus</emphasis>
</taxonomicName>
emitted at a regular basis were heard at Manombo, where also 
<taxonomicName id="4C4B4D186C3FFFC7FE32DACEFD90C8F2" box="[510,624,404,428]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="5" pageNumber="574" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C3FFFC7FE32DACEFD90C8F2" box="[510,624,404,428]" italics="true" pageId="5" pageNumber="574">G. timidus</emphasis>
</taxonomicName>
was calling at the same time. Calls were obtained from one of the 
<typeStatus id="54F088396C3FFFC7FAA0DACFFA7CC8F3" box="[1388,1436,405,429]" pageId="5" pageNumber="574">type</typeStatus>
specimens (recording temperature not taken but estimated ca. 25°C) and are composed of two distinctly different note 
<typeStatus id="54F088396C3FFFC7FF1DDA87FEECC8AB" box="[209,268,477,501]" pageId="5" pageNumber="574">types</typeStatus>
(
<figureCitation id="13702A1E6C3FFFC7FED3DA86FE86C8AA" box="[287,358,476,501]" captionStart="FIGURE 3" captionStartId="5.[151,250,1929,1951]" captionTargetBox="[151,1435,943,1883]" captionTargetId="figure@5.[151,1435,943,1883]" captionTargetPageId="5" captionText="FIGURE 3. Audiospectrogram and corresponding oscillogram of the advertisement call of Guibemantis diphonus sp. nov., recorded at Manombo Special Reserve on 23 February 2007 (temperature not taken but estimated ca. 25 ° C)." httpUri="https://zenodo.org/record/244221/files/figure.png" pageId="5" pageNumber="574">Fig. 3</figureCitation>
). The first note 
<typeStatus id="54F088396C3FFFC7FDEADA87FDB6C8AB" box="[550,598,477,501]" pageId="5" pageNumber="574">type</typeStatus>
is rather long (note duration 740−998 ms) composed of multiple pulses (29−44 pulses/note) emitted with relatively low energy, followed by a single short note 
<typeStatus id="54F088396C3FFFC7FBBAD95AFB46CB46" box="[1142,1190,512,536]" pageId="5" pageNumber="574">type</typeStatus>
(note duration 10−24 ms) with comparatively much higher energy containing a single pulse only. The two different note 
<typeStatus id="54F088396C3FFFC7FAFFD97FFA8ECB63" box="[1331,1390,549,573]" pageId="5" pageNumber="574">types</typeStatus>
are separated by an interval of 155−181 ms. The duration of the call (here defined from the beginning of the first note 
<typeStatus id="54F088396C3FFFC7FF5BD937FF27CBDB" box="[151,199,621,645]" pageId="5" pageNumber="574">type</typeStatus>
to the end of the second note 
<typeStatus id="54F088396C3FFFC7FDE1D936FDBFCBDA" box="[557,607,620,644]" pageId="5" pageNumber="574">type</typeStatus>
) is 1055 ± 98 ms (941−1177 ms). Within the first note 
<typeStatus id="54F088396C3FFFC7FB25D937FAFCCBDB" box="[1257,1308,621,645]" pageId="5" pageNumber="574">type</typeStatus>
, pulses are somewhat irregularly arranged in 12−17 groups containing either two or three pulses, with each pulse group separated by slightly larger intervals from the following pulse group, compared to inter-pulse intervals within a pulse group. Pulse repetition rate within the first note 
<typeStatus id="54F088396C3FFFC7FCCAD982FCD6CBAE" box="[774,822,728,752]" pageId="5" pageNumber="574">type</typeStatus>
ranges from 35.5−50.1 pulses per second. Frequency ranges between 1100−3500 Hz, with a dominant frequency peak at approximately 2320 Hz. For detailed numerical parameters see 
<tableCitation id="C6C903206C3FFFC7FE88D845FE7ACA66" box="[324,410,799,824]" captionStart="TABLE 2" captionStartId="7.[125,202,1000,1020]" captionText="TABLE 2. Comparative temporal and spectral parameters of advertisement calls of species of Guibemantis (subgenus Guibemantis). Values given as mean ± standard deviation followed by range in parentheses. Data of G. kathrinae taken from the original description of the species (Glaw et al. 2000). For all species except G. diphonus, we consider the call consisting of a single, pulsed note. In these cases call duration equals note duration. In G. kathrinae these calls are arranged in regular series, whereas G. depressiceps, G. timidus and. tornieri mostly emit isolated calls but highly motivated males can emit irregular call series. NA, not applicable (calls consisting of single notes only); n, number of calls analyzed temporal variables." pageId="5" pageNumber="574">Table 2</tableCitation>
. Calls of the second specimen were very similar according to field notes but recordings are of poor quality and cannot be analyzed in detail.
</paragraph>
<caption id="DF3466136C3FFFC7FF5BDCD3FB3FCEE3" httpUri="https://zenodo.org/record/244221/files/figure.png" pageId="5" pageNumber="574" targetBox="[151,1435,943,1883]" targetPageId="5">
<paragraph id="8BF4369B6C3FFFC7FF5BDCD3FB3FCEE3" blockId="5.[151,1435,1929,1981]" pageId="5" pageNumber="574">
<emphasis id="B93FEA896C3FFFC7FF5BDCD3FEF5CEC0" bold="true" box="[151,277,1929,1951]" pageId="5" pageNumber="574">FIGURE 3.</emphasis>
Audiospectrogram and corresponding oscillogram of the advertisement call of 
<taxonomicName id="4C4B4D186C3FFFC7FB9EDCD3FAD9CEC1" box="[1106,1337,1929,1951]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="5" pageNumber="574" phylum="Chordata" rank="species" species="diphonus" status="sp. nov.">
<emphasis id="B93FEA896C3FFFC7FB9EDCD3FAD9CEC1" box="[1106,1337,1929,1951]" italics="true" pageId="5" pageNumber="574">Guibemantis diphonus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A20C57F26C3FFFC7FA8EDCD3FA75CEC0" box="[1346,1429,1929,1950]" pageId="5" pageNumber="574" rank="species">
<emphasis id="B93FEA896C3FFFC7FA8EDCD3FA75CEC0" bold="true" box="[1346,1429,1929,1950]" pageId="5" pageNumber="574">sp. nov.</emphasis>
</taxonomicNameLabel>
, recorded at Manombo Special Reserve on 23 February 2007 (temperature not taken but estimated ca. 25°C).
</paragraph>
</caption>
<caption id="DF3466136C3CFFC4FF5BDCDCFD64CE84" httpUri="https://zenodo.org/record/244222/files/figure.png" pageId="6" pageNumber="575" targetBox="[230,1335,193,1895]" targetPageId="6">
<paragraph id="8BF4369B6C3CFFC4FF5BDCDCFD64CE84" blockId="6.[151,1436,1926,2010]" pageId="6" pageNumber="575">
<emphasis id="B93FEA896C3CFFC4FF5BDCDCFEF2CEC5" bold="true" box="[151,274,1926,1948]" pageId="6" pageNumber="575">FIGURE 4.</emphasis>
Comparative oscillograms (2000 ms sections each) of advertisement calls of species in the subgenus 
<taxonomicName id="4C4B4D186C3CFFC4FAD8DCDCFA75CEC2" box="[1300,1429,1926,1948]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="6" pageNumber="575" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C3CFFC4FAD8DCDCFA75CEC2" box="[1300,1429,1926,1948]" italics="true" pageId="6" pageNumber="575">Guibemantis</emphasis>
</taxonomicName>
. For detailed call parameters see Table 2. Oscillogram of 
<taxonomicName id="4C4B4D186C3CFFC4FD2CDCFCFCBECEE5" box="[736,862,1957,1979]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="6" pageNumber="575" phylum="Chordata" rank="species" species="kathrinae">
<emphasis id="B93FEA896C3CFFC4FD2CDCFCFCBECEE5" box="[736,862,1957,1979]" italics="true" pageId="6" pageNumber="575">G. kathrinae</emphasis>
</taxonomicName>
taken from Glaw 
<emphasis id="B93FEA896C3CFFC4FBD0DCFCFBB1CEE5" box="[1052,1105,1957,1979]" italics="true" pageId="6" pageNumber="575">et al.</emphasis>
(2000) showing a series of four calls according to the definition followed herein.
</paragraph>
</caption>
<caption id="DF3466136C3DFFC5FFB1DA9CFC6ECB51" ID-Table-UUID="DF3466136C3DFFC5FFB1DA9CFC6ECB51" httpUri="http://table.plazi.org/id/DF3466136C3DFFC5FFB1DA9CFC6ECB51" pageId="7" pageNumber="576" targetBox="[142,1784,561,966]" targetIsTable="true" targetPageId="7">
<paragraph id="8BF4369B6C3DFFC5FFB1DA9CFC6ECB51" blockId="7.[125,1770,453,527]" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FFB1DA9CFF00C884" bold="true" box="[125,224,454,474]" pageId="7" pageNumber="576">TABLE 1.</emphasis>
Morphometric measurements of 
<taxonomicName id="4C4B4D186C3DFFC5FDDEDA9FFD01C884" box="[530,737,453,474]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C3DFFC5FDDEDA9FFD01C884" box="[530,737,453,474]" italics="true" pageId="7" pageNumber="576">Guibemantis diphonus</emphasis>
</taxonomicName>
and comparative specimens of 
<taxonomicName id="4C4B4D186C3DFFC5FBCEDA9FFB82C884" box="[1026,1122,453,474]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C3DFFC5FBCEDA9FFB82C884" box="[1026,1122,453,474]" italics="true" pageId="7" pageNumber="576">G. timidus</emphasis>
</taxonomicName>
and 
<emphasis id="B93FEA896C3DFFC5FB42DA9FFB16C884" box="[1166,1270,453,474]" italics="true" pageId="7" pageNumber="576">
<taxonomicName id="4C4B4D186C3DFFC5FB42DA9FFB12C884" box="[1166,1266,453,474]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="tornieri">G. tornieri</taxonomicName>
,
</emphasis>
all in mm. Abbreviations as in Materials and Methods except for: HT, holotype; PT, paratype; M, male; RHL, relative hindlimb length given as the point reached by tibiotarsal articulation when hindlimb is adpressed along body (1, posterior eye corner; 2, eye center; 3, anterior eye corner; 4, between eye and nostril).
</paragraph>
</caption>
<paragraph id="8BF4369B6C3DFFC5FF42D96BF931CAF2" pageId="7" pageNumber="576">
<table id="F94BC43B6C3D003DFF42D96BF918CA98" box="[142,1784,561,966]" gridcols="18" gridrows="5" pageId="7" pageNumber="576">
<tr id="357B34D96C3D003DFF42D96BF918CB1B" box="[142,1784,561,581]" gridrow="0" pageId="7" pageNumber="576">
<th id="76AA5DA56C3D003DFF42D96BFE91CB1B" box="[142,369,561,581]" gridcol="0" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FF42D96BFF02CB1B" bold="true" box="[142,226,561,581]" pageId="7" pageNumber="576">Voucher</emphasis>
</th>
<th id="76AA5DA56C3D003DFE6BD96BFE04CB1B" box="[423,484,561,581]" gridcol="1" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FE6BD96BFE04CB1B" bold="true" box="[423,484,561,581]" pageId="7" pageNumber="576">Status</emphasis>
</th>
<th id="76AA5DA56C3D003DFDC5D96BFDCCCB1B" box="[521,556,561,581]" gridcol="2" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FDC5D96BFDCCCB1B" bold="true" box="[521,556,561,581]" pageId="7" pageNumber="576">Sex</emphasis>
</th>
<th id="76AA5DA56C3D003DFD81D96BFD9ACB1B" box="[589,634,561,581]" gridcol="3" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FD81D96BFD9ACB1B" bold="true" box="[589,634,561,581]" pageId="7" pageNumber="576">SVL</emphasis>
</th>
<th id="76AA5DA56C3D003DFD50D96BFD26CB1B" box="[668,710,561,581]" gridcol="4" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FD51D96BFD26CB1B" bold="true" box="[669,710,561,581]" pageId="7" pageNumber="576">HW</emphasis>
</th>
<th id="76AA5DA56C3D003DFD24D96BFCF0CB1B" box="[744,784,561,581]" gridcol="5" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FD24D96BFCE9CB1B" bold="true" box="[744,777,561,581]" pageId="7" pageNumber="576">HL</emphasis>
</th>
<th id="76AA5DA56C3D003DFCFED96BFCB3CB1B" box="[818,851,561,581]" gridcol="6" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FCFFD96BFCB3CB1B" bold="true" box="[819,851,561,581]" pageId="7" pageNumber="576">TD</emphasis>
</th>
<th id="76AA5DA56C3D003DFCBFD96BFC74CB1B" box="[883,916,561,581]" gridcol="7" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FCB8D96BFC74CB1B" bold="true" box="[884,916,561,581]" pageId="7" pageNumber="576">ED</emphasis>
</th>
<th id="76AA5DA56C3D003DFC79D96BFC05CB1B" box="[949,997,561,581]" gridcol="8" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FC79D96BFC05CB1B" bold="true" box="[949,997,561,581]" pageId="7" pageNumber="576">END</emphasis>
</th>
<th id="76AA5DA56C3D003DFBC5D96BFBD7CB1B" box="[1033,1079,561,581]" gridcol="9" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FBC5D96BFBD7CB1B" bold="true" box="[1033,1079,561,581]" pageId="7" pageNumber="576">NSD</emphasis>
</th>
<th id="76AA5DA56C3D003DFB95D96BFB6CCB1B" box="[1113,1164,561,581]" gridcol="10" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FB96D96BFB6CCB1B" bold="true" box="[1114,1164,561,581]" pageId="7" pageNumber="576">NND</emphasis>
</th>
<th id="76AA5DA56C3D003DFB63D96BFB0ECB1B" box="[1199,1262,561,581]" gridcol="11" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FB63D96BFB0ECB1B" bold="true" box="[1199,1262,561,581]" pageId="7" pageNumber="576">FORL</emphasis>
</th>
<th id="76AA5DA56C3D003DFADFD96BFAA6CB1B" box="[1299,1350,561,581]" gridcol="12" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FAD8D96BFAA6CB1B" bold="true" box="[1300,1350,561,581]" pageId="7" pageNumber="576">HAL</emphasis>
</th>
<th id="76AA5DA56C3D003DFAA4D96BFA73CB1B" box="[1384,1427,561,581]" gridcol="13" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FAA5D96BFA73CB1B" bold="true" box="[1385,1427,561,581]" pageId="7" pageNumber="576">HIL</emphasis>
</th>
<th id="76AA5DA56C3D003DFA79D96BFA05CB1B" box="[1461,1509,561,581]" gridcol="14" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FA7AD96BFA05CB1B" bold="true" box="[1462,1509,561,581]" pageId="7" pageNumber="576">FOL</emphasis>
</th>
<th id="76AA5DA56C3D003DF9C4D96BF9A6CB1B" box="[1544,1606,561,581]" gridcol="15" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5F9C4D96BF9A6CB1B" bold="true" box="[1544,1606,561,581]" pageId="7" pageNumber="576">FOTL</emphasis>
</th>
<th id="76AA5DA56C3D003DF9A7D96BF942CB1B" box="[1643,1698,561,581]" gridcol="16" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5F9A7D96BF942CB1B" bold="true" box="[1643,1698,561,581]" pageId="7" pageNumber="576">TIBL</emphasis>
</th>
<th id="76AA5DA56C3D003DF909D96BF918CB1B" box="[1733,1784,561,581]" gridcol="17" gridrow="0" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5F90AD96BF918CB1B" bold="true" box="[1734,1784,561,581]" pageId="7" pageNumber="576">RHL</emphasis>
</th>
</tr>
<tr id="357B34D96C3D003DFF42D90FF918CB34" box="[142,1784,597,618]" gridrow="1" pageId="7" pageNumber="576" rowspan-1="1" rowspan-10="1" rowspan-11="1" rowspan-12="1" rowspan-13="1" rowspan-14="1" rowspan-15="1" rowspan-16="1" rowspan-17="1" rowspan-2="1" rowspan-3="1" rowspan-4="1" rowspan-5="1" rowspan-6="1" rowspan-7="1" rowspan-8="1" rowspan-9="1">
<th id="76AA5DA56C3D003DFF42D90FFE91CB34" box="[142,369,597,618]" gridcol="0" gridrow="1" pageId="7" pageNumber="576">
<taxonomicName id="4C4B4D186C3DFFC5FF42D90FFE91CB34" authority="(Manombo)" authorityName="Manombo" baseAuthorityName="Manombo" box="[142,369,597,618]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C3DFFC5FF42D90FFF1ECB34" box="[142,254,597,618]" italics="true" pageId="7" pageNumber="576">G. diphonus</emphasis>
(Manombo)
</taxonomicName>
</th>
</tr>
<tr id="357B34D96C3D003DFF42D920F918CBA7" box="[142,1784,634,761]" gridrow="2" pageId="7" pageNumber="576">
<th id="76AA5DA56C3D003DFF42D920FE91CBA7" box="[142,369,634,761]" gridcol="0" gridrow="2" pageId="7" pageNumber="576">
ZSM 2665/2007 (ZCMV 5496) ZSM 2664/2007 (ZCMV 5416) 
<taxonomicName id="4C4B4D186C3DFFC5FF42D9BEFE83CBA7" authority="(Manombo)" authorityName="Manombo" baseAuthorityName="Manombo" box="[142,355,740,761]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="tornieri">
<emphasis id="B93FEA896C3DFFC5FF42D9BEFF10CBA7" box="[142,240,740,761]" italics="true" pageId="7" pageNumber="576">G. tornieri</emphasis>
(Manombo)
</taxonomicName>
</th>
<td id="76AA5DA56C3D003DFE6BD920FE04CBA7" box="[423,484,634,761]" gridcol="1" gridrow="2" pageId="7" pageNumber="576">HT PT</td>
<td id="76AA5DA56C3D003DFDC5D920FDCCCBA7" box="[521,556,634,761]" gridcol="2" gridrow="2" pageId="7" pageNumber="576">M M</td>
<td id="76AA5DA56C3D003DFD81D920FD9ACBA7" box="[589,634,634,761]" gridcol="3" gridrow="2" pageId="7" pageNumber="576">33.5 36.1</td>
<td id="76AA5DA56C3D003DFD50D920FD26CBA7" box="[668,710,634,761]" gridcol="4" gridrow="2" pageId="7" pageNumber="576">11.9 12.0</td>
<td id="76AA5DA56C3D003DFD24D920FCF0CBA7" box="[744,784,634,761]" gridcol="5" gridrow="2" pageId="7" pageNumber="576">14.2 14.1</td>
<td id="76AA5DA56C3D003DFCFED920FCB3CBA7" box="[818,851,634,761]" gridcol="6" gridrow="2" pageId="7" pageNumber="576">2.8 2.5</td>
<td id="76AA5DA56C3D003DFCBFD920FC74CBA7" box="[883,916,634,761]" gridcol="7" gridrow="2" pageId="7" pageNumber="576">4.4 4.4</td>
<td id="76AA5DA56C3D003DFC79D920FC05CBA7" box="[949,997,634,761]" gridcol="8" gridrow="2" pageId="7" pageNumber="576">4.0 3.9</td>
<td id="76AA5DA56C3D003DFBC5D920FBD7CBA7" box="[1033,1079,634,761]" gridcol="9" gridrow="2" pageId="7" pageNumber="576">1.9 1.9</td>
<td id="76AA5DA56C3D003DFB95D920FB6CCBA7" box="[1113,1164,634,761]" gridcol="10" gridrow="2" pageId="7" pageNumber="576">3.2 3.2</td>
<td id="76AA5DA56C3D003DFB63D920FB0ECBA7" box="[1199,1262,634,761]" gridcol="11" gridrow="2" pageId="7" pageNumber="576">22.1 22.3</td>
<td id="76AA5DA56C3D003DFADFD920FAA6CBA7" box="[1299,1350,634,761]" gridcol="12" gridrow="2" pageId="7" pageNumber="576">10.8 10.4</td>
<td id="76AA5DA56C3D003DFAA4D920FA73CBA7" box="[1384,1427,634,761]" gridcol="13" gridrow="2" pageId="7" pageNumber="576">55.8 55.3</td>
<td id="76AA5DA56C3D003DFA79D920FA05CBA7" box="[1461,1509,634,761]" gridcol="14" gridrow="2" pageId="7" pageNumber="576">16.7 16.4</td>
<td id="76AA5DA56C3D003DF9C4D920F9A6CBA7" box="[1544,1606,634,761]" gridcol="15" gridrow="2" pageId="7" pageNumber="576">25.1 26.0</td>
<td id="76AA5DA56C3D003DF9A7D920F942CBA7" box="[1643,1698,634,761]" gridcol="16" gridrow="2" pageId="7" pageNumber="576">16.5 17.4</td>
<td id="76AA5DA56C3D003DF909D920F918CBA7" box="[1733,1784,634,761]" gridcol="17" gridrow="2" pageId="7" pageNumber="576">4 3</td>
</tr>
<tr id="357B34D96C3D003DFF42D853F918CA0D" box="[142,1784,777,851]" gridrow="3" pageId="7" pageNumber="576">
<th id="76AA5DA56C3D003DFF42D853FE91CA0D" box="[142,369,777,851]" gridcol="0" gridrow="3" pageId="7" pageNumber="576">
ZSM 2668/2007 (ZCMV 5478) 
<taxonomicName id="4C4B4D186C3DFFC5FF42D864FEBDCA0D" authority="(Beronono)" authorityName="Beronono" baseAuthorityName="Beronono" box="[142,349,830,851]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C3DFFC5FF42D864FF0ECA0D" box="[142,238,830,851]" italics="true" pageId="7" pageNumber="576">G. timidus</emphasis>
(Beronono)
</taxonomicName>
</th>
<td id="76AA5DA56C3D003DFE6BD853FE04CA0D" box="[423,484,777,851]" gridcol="1" gridrow="3" pageId="7" pageNumber="576">--</td>
<td id="76AA5DA56C3D003DFDC5D853FDCCCA0D" box="[521,556,777,851]" gridcol="2" gridrow="3" pageId="7" pageNumber="576">M</td>
<td id="76AA5DA56C3D003DFD81D853FD9ACA0D" box="[589,634,777,851]" gridcol="3" gridrow="3" pageId="7" pageNumber="576">58.4</td>
<td id="76AA5DA56C3D003DFD50D853FD26CA0D" box="[668,710,777,851]" gridcol="4" gridrow="3" pageId="7" pageNumber="576">18.6</td>
<td id="76AA5DA56C3D003DFD24D853FCF0CA0D" box="[744,784,777,851]" gridcol="5" gridrow="3" pageId="7" pageNumber="576">20.9</td>
<td id="76AA5DA56C3D003DFCFED853FCB3CA0D" box="[818,851,777,851]" gridcol="6" gridrow="3" pageId="7" pageNumber="576">3.6</td>
<td id="76AA5DA56C3D003DFCBFD853FC74CA0D" box="[883,916,777,851]" gridcol="7" gridrow="3" pageId="7" pageNumber="576">6.9</td>
<td id="76AA5DA56C3D003DFC79D853FC05CA0D" box="[949,997,777,851]" gridcol="8" gridrow="3" pageId="7" pageNumber="576">5.9</td>
<td id="76AA5DA56C3D003DFBC5D853FBD7CA0D" box="[1033,1079,777,851]" gridcol="9" gridrow="3" pageId="7" pageNumber="576">2.8</td>
<td id="76AA5DA56C3D003DFB95D853FB6CCA0D" box="[1113,1164,777,851]" gridcol="10" gridrow="3" pageId="7" pageNumber="576">5.4</td>
<td id="76AA5DA56C3D003DFB63D853FB0ECA0D" box="[1199,1262,777,851]" gridcol="11" gridrow="3" pageId="7" pageNumber="576">33.4</td>
<td id="76AA5DA56C3D003DFADFD853FAA6CA0D" box="[1299,1350,777,851]" gridcol="12" gridrow="3" pageId="7" pageNumber="576">16.3</td>
<td id="76AA5DA56C3D003DFAA4D853FA73CA0D" box="[1384,1427,777,851]" gridcol="13" gridrow="3" pageId="7" pageNumber="576">85.6</td>
<td id="76AA5DA56C3D003DFA79D853FA05CA0D" box="[1461,1509,777,851]" gridcol="14" gridrow="3" pageId="7" pageNumber="576">26.5</td>
<td id="76AA5DA56C3D003DF9C4D853F9A6CA0D" box="[1544,1606,777,851]" gridcol="15" gridrow="3" pageId="7" pageNumber="576">39.4</td>
<td id="76AA5DA56C3D003DF9A7D853F942CA0D" box="[1643,1698,777,851]" gridcol="16" gridrow="3" pageId="7" pageNumber="576">25.0</td>
<td id="76AA5DA56C3D003DF909D853F918CA0D" box="[1733,1784,777,851]" gridcol="17" gridrow="3" pageId="7" pageNumber="576">1</td>
</tr>
<tr id="357B34D96C3D003DFF42D839F918CA98" box="[142,1784,867,966]" gridrow="4" pageId="7" pageNumber="576">
<th id="76AA5DA56C3D003DFF42D839FE91CA98" box="[142,369,867,966]" gridcol="0" gridrow="4" pageId="7" pageNumber="576">ZSM 2666/2007 (ZCMV 5419) ZSM 2667/2007 (ZCMV 5427)</th>
<td id="76AA5DA56C3D003DFE6BD839FE04CA98" box="[423,484,867,966]" gridcol="1" gridrow="4" pageId="7" pageNumber="576">-- --</td>
<td id="76AA5DA56C3D003DFDC5D839FDCCCA98" box="[521,556,867,966]" gridcol="2" gridrow="4" pageId="7" pageNumber="576">M M</td>
<td id="76AA5DA56C3D003DFD81D839FD9ACA98" box="[589,634,867,966]" gridcol="3" gridrow="4" pageId="7" pageNumber="576">43.4 37.0</td>
<td id="76AA5DA56C3D003DFD50D839FD26CA98" box="[668,710,867,966]" gridcol="4" gridrow="4" pageId="7" pageNumber="576">15.2 13.1</td>
<td id="76AA5DA56C3D003DFD24D839FCF0CA98" box="[744,784,867,966]" gridcol="5" gridrow="4" pageId="7" pageNumber="576">16.7 15.0</td>
<td id="76AA5DA56C3D003DFCFED839FCB3CA98" box="[818,851,867,966]" gridcol="6" gridrow="4" pageId="7" pageNumber="576">2.7 2.7</td>
<td id="76AA5DA56C3D003DFCBFD839FC74CA98" box="[883,916,867,966]" gridcol="7" gridrow="4" pageId="7" pageNumber="576">4.5 4.8</td>
<td id="76AA5DA56C3D003DFC79D839FC05CA98" box="[949,997,867,966]" gridcol="8" gridrow="4" pageId="7" pageNumber="576">4.8 4.3</td>
<td id="76AA5DA56C3D003DFBC5D839FBD7CA98" box="[1033,1079,867,966]" gridcol="9" gridrow="4" pageId="7" pageNumber="576">2.3 2.0</td>
<td id="76AA5DA56C3D003DFB95D839FB6CCA98" box="[1113,1164,867,966]" gridcol="10" gridrow="4" pageId="7" pageNumber="576">3.9 3.5</td>
<td id="76AA5DA56C3D003DFB63D839FB0ECA98" box="[1199,1262,867,966]" gridcol="11" gridrow="4" pageId="7" pageNumber="576">26.5 23.7</td>
<td id="76AA5DA56C3D003DFADFD839FAA6CA98" box="[1299,1350,867,966]" gridcol="12" gridrow="4" pageId="7" pageNumber="576">12.3 10.8</td>
<td id="76AA5DA56C3D003DFAA4D839FA73CA98" box="[1384,1427,867,966]" gridcol="13" gridrow="4" pageId="7" pageNumber="576">61.8 55.8</td>
<td id="76AA5DA56C3D003DFA79D839FA05CA98" box="[1461,1509,867,966]" gridcol="14" gridrow="4" pageId="7" pageNumber="576">19.5 17.3</td>
<td id="76AA5DA56C3D003DF9C4D839F9A6CA98" box="[1544,1606,867,966]" gridcol="15" gridrow="4" pageId="7" pageNumber="576">28.5 25.9</td>
<td id="76AA5DA56C3D003DF9A7D839F942CA98" box="[1643,1698,867,966]" gridcol="16" gridrow="4" pageId="7" pageNumber="576">18.6 17.4</td>
<td id="76AA5DA56C3D003DF909D839F918CA98" box="[1733,1784,867,966]" gridcol="17" gridrow="4" pageId="7" pageNumber="576">1 2</td>
</tr>
</table>
</paragraph>
<caption id="DF3466136C3DFFC5FFB1D8B2FEAECD2B" pageId="7" pageNumber="576">
<paragraph id="8BF4369B6C3DFFC5FFB1D8B2FEAECD2B" blockId="7.[125,1791,999,1141]" pageId="7" pageNumber="576">
<emphasis id="B93FEA896C3DFFC5FFB1D8B2FF01CAA2" bold="true" box="[125,225,1000,1020]" pageId="7" pageNumber="576">TABLE 2.</emphasis>
Comparative temporal and spectral parameters of advertisement calls of species of 
<taxonomicName id="4C4B4D186C3DFFC5FC13D8BDFBB5CAA2" box="[991,1109,999,1020]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C3DFFC5FC13D8BDFBB5CAA2" box="[991,1109,999,1020]" italics="true" pageId="7" pageNumber="576">Guibemantis</emphasis>
</taxonomicName>
(subgenus 
<taxonomicName id="4C4B4D186C3DFFC5FB71D8BDFAD2CAA2" box="[1213,1330,999,1020]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C3DFFC5FB71D8BDFAD2CAA2" box="[1213,1330,999,1020]" italics="true" pageId="7" pageNumber="576">Guibemantis</emphasis>
</taxonomicName>
). Values given as mean ± standard deviation followed by range in parentheses. Data of 
<taxonomicName id="4C4B4D186C3DFFC5FDCCDF5FFD94CD44" box="[512,628,1029,1050]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="kathrinae">
<emphasis id="B93FEA896C3DFFC5FDCCDF5FFD94CD44" box="[512,628,1029,1050]" italics="true" pageId="7" pageNumber="576">G. kathrinae</emphasis>
</taxonomicName>
taken from the original description of the species (Glaw 
<emphasis id="B93FEA896C3DFFC5FBB2DF5FFB4DCD44" box="[1150,1197,1029,1050]" italics="true" pageId="7" pageNumber="576">et al.</emphasis>
2000). For all species except 
<emphasis id="B93FEA896C3DFFC5FA72DF5FF9D5CD44" box="[1470,1589,1029,1050]" italics="true" pageId="7" pageNumber="576">
<taxonomicName id="4C4B4D186C3DFFC5FA72DF5FF9D0CD44" box="[1470,1584,1029,1050]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="diphonus">G. diphonus</taxonomicName>
,
</emphasis>
we consider the call consisting of a single, pulsed note. In these cases call duration equals note duration. In 
<taxonomicName id="4C4B4D186C3DFFC5FC55DF7EFBECCD67" box="[921,1036,1060,1081]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="kathrinae">
<emphasis id="B93FEA896C3DFFC5FC55DF7EFBECCD67" box="[921,1036,1060,1081]" italics="true" pageId="7" pageNumber="576">G. kathrinae</emphasis>
</taxonomicName>
these calls are arranged in regular series, whereas 
<emphasis id="B93FEA896C3DFFC5FA10DF7EF939CD67" box="[1500,1753,1060,1081]" italics="true" pageId="7" pageNumber="576">
<taxonomicName id="4C4B4D186C3DFFC5FA10DF7EF98FCD67" box="[1500,1647,1060,1081]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="depressiceps">G. depressiceps</taxonomicName>
, 
<taxonomicName id="4C4B4D186C3DFFC5F9B5DF7EF939CD67" box="[1657,1753,1060,1081]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="timidus">G. timidus</taxonomicName>
</emphasis>
and. 
<taxonomicName id="4C4B4D186C3DFFC5FF54DF18FF3ECD09" box="[152,222,1090,1111]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="tornieri">
<emphasis id="B93FEA896C3DFFC5FF54DF18FF3ECD09" box="[152,222,1090,1111]" italics="true" pageId="7" pageNumber="576">tornieri</emphasis>
</taxonomicName>
mostly emit isolated calls but highly motivated males can emit irregular call series. NA, not applicable (calls consisting of single notes only); n, number of calls analyzed temporal variables.
</paragraph>
</caption>
<paragraph id="8BF4369B6C3DFFC5FF42DFC6FA17CC5E" blockId="7.[142,1789,1180,1675]" pageId="7" pageNumber="576">
Species and Note duration [ms] Pulses/ Pulses/second Inter-note Frequency range Dominant frequency Recording Locality note within call interval [ms] [Hz] peak [Hz] temperature [°C] 
<taxonomicName id="4C4B4D186C3DFFC5FF42DF8BFF1ECDB8" box="[142,254,1233,1254]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="diphonus" status="sp. nov.">
<emphasis id="B93FEA896C3DFFC5FF42DF8BFF1ECDB8" box="[142,254,1233,1254]" italics="true" pageId="7" pageNumber="576">G. diphonus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A20C57F26C3DFFC5FEC8DF88FEAECDB8" box="[260,334,1234,1254]" pageId="7" pageNumber="576" rank="species">
<emphasis id="B93FEA896C3DFFC5FEC8DF88FEAECDB8" bold="true" box="[260,334,1234,1254]" pageId="7" pageNumber="576">sp. nov.</emphasis>
</taxonomicNameLabel>
Note 
<typeStatus id="54F088396C3DFFC5FE2EDF88FDEACDB8" box="[482,522,1234,1254]" pageId="7" pageNumber="576">type</typeStatus>
Note 
<typeStatus id="54F088396C3DFFC5FD0ADF88FD0ECDB8" box="[710,750,1234,1254]" pageId="7" pageNumber="576">type</typeStatus>
1: Note 
<typeStatus id="54F088396C3DFFC5FCB1DF88FC45CDB8" box="[893,933,1234,1254]" pageId="7" pageNumber="576">type</typeStatus>
1: 165 ± 12 1100−3500 2320 ± 102 ca. 25 Manombo (n = 6) 1: 843 ± 85 33.8 ± 5.3 41.6 ± 7.6 (155−181) (
<date id="FFF5105B6C3DFFC5FA4ADFB6FA11CC5E" box="[1414,1521,1260,1280]" pageId="7" pageNumber="576" value="2174" valueMax="2444">2174−2444</date>
)
</paragraph>
<paragraph id="8BF4369B6C3DFFC5FE7CDE5CFC57CC44" blockId="7.[142,1789,1180,1675]" box="[432,951,1286,1306]" pageId="7" pageNumber="576">(740−998) (29−44) (35.5−50.1)</paragraph>
<paragraph id="8BF4369B6C3DFFC5FE7CDE7BFDC1CC6B" blockId="7.[142,1789,1180,1675]" box="[432,545,1313,1333]" pageId="7" pageNumber="576">
Note 
<typeStatus id="54F088396C3DFFC5FE2EDE7BFDEACC6B" box="[482,522,1313,1333]" pageId="7" pageNumber="576">type</typeStatus>
2:
</paragraph>
<paragraph id="8BF4369B6C3DFFC5FE7CDE61FE0BCC11" blockId="7.[142,1789,1180,1675]" box="[432,491,1339,1359]" pageId="7" pageNumber="576">10−24</paragraph>
<paragraph id="8BF4369B6C3DFFC5FF42DE0FFC58CFD5" blockId="7.[142,1789,1180,1675]" pageId="7" pageNumber="576">
<taxonomicName id="4C4B4D186C3DFFC5FF42DE0FFF0ECC34" box="[142,238,1365,1386]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C3DFFC5FF42DE0FFF0ECC34" box="[142,238,1365,1386]" italics="true" pageId="7" pageNumber="576">G. timidus</emphasis>
</taxonomicName>
125 ± 17 3.3 ± 0.8 48.1 ± 10.9 NA 800−2800 1599 ± 181 ca. 25 Manombo (n = 7) (100−147) (2−4) (36.1−58.8) (
<date id="FFF5105B6C3DFFC5FA4ADE35FA11CCDD" box="[1414,1521,1391,1411]" pageId="7" pageNumber="576" value="1280" valueMax="1719">1280−1719</date>
) 
<taxonomicName id="4C4B4D186C3DFFC5FF42DED3FF0ECCC0" box="[142,238,1417,1438]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C3DFFC5FF42DED3FF0ECCC0" box="[142,238,1417,1438]" italics="true" pageId="7" pageNumber="576">G. timidus</emphasis>
</taxonomicName>
106 ± 23 2.1 ± 0.4 27.0 ± 6.0 NA 1000−2500 1768 ± 52 -- 
<collectingCountry id="F35C760B6C3DFFC5FF42DEFEFF5ECCE6" box="[142,190,1444,1464]" name="Iceland" pageId="7" pageNumber="576">Nosy</collectingCountry>
Boraha (n = 7) (65−120) (2−3) (22.7−35.7) (
<date id="FFF5105B6C3DFFC5FA4ADEFEFA11CCE6" box="[1414,1521,1444,1464]" pageId="7" pageNumber="576" value="1713" valueMax="1821">1713−1821</date>
) 
<taxonomicName id="4C4B4D186C3DFFC5FF42DEE4FF10CC8D" box="[142,240,1470,1491]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="tornieri">
<emphasis id="B93FEA896C3DFFC5FF42DEE4FF10CC8D" box="[142,240,1470,1491]" italics="true" pageId="7" pageNumber="576">G. tornieri</emphasis>
</taxonomicName>
346 ± 41 4.8 ± 0.4 12.3 ± 0.5 NA 800−2100 1127 ± 103 ca. 25 Manombo (n = 5) (277−384) (4−5) (12.3−13.4) (973−1195) 
<taxonomicName id="4C4B4D186C3DFFC5FF42DEA8FEFFCF59" box="[142,287,1522,1543]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="depressiceps">
<emphasis id="B93FEA896C3DFFC5FF42DEA8FEFFCF59" box="[142,287,1522,1543]" italics="true" pageId="7" pageNumber="576">G. depressiceps</emphasis>
</taxonomicName>
113 ± 25 3.7 ± 0.8 30.5 ± 1.9 NA 1000−3200 1808 ± 23 19.5 Ranomafana (n = 6) (90−160) (3−5) (28.6−33.0) (
<date id="FFF5105B6C3DFFC5FA4ADD54FA11CF7C" box="[1414,1521,1550,1570]" pageId="7" pageNumber="576" value="1775" valueMax="1841">1775−1841</date>
) 
<taxonomicName id="4C4B4D186C3DFFC5FF42DD7DFEFFCF62" box="[142,287,1575,1596]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="depressiceps">
<emphasis id="B93FEA896C3DFFC5FF42DD7DFEFFCF62" box="[142,287,1575,1596]" italics="true" pageId="7" pageNumber="576">G. depressiceps</emphasis>
</taxonomicName>
139 ± 26 4.2 ± 0.8 31.3 ± 1.8 NA 1200−3100 1977 ± 143 19−20 Andasibe (n = 6) (91−161) (3−5) (29.9−33.3) (1820−2100) 
<taxonomicName id="4C4B4D186C3DFFC5FF42DD06FEE2CF2F" box="[142,258,1628,1649]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="576" phylum="Chordata" rank="species" species="kathrinae">
<emphasis id="B93FEA896C3DFFC5FF42DD06FEE2CF2F" box="[142,258,1628,1649]" italics="true" pageId="7" pageNumber="576">G. kathrinae</emphasis>
</taxonomicName>
329 ± 44 6.2 ± 0.9 19.0 ± 1.6 NA 850−1700 (1250−1550) 21 An’Ala (n = 4) (268−372) (5−7) (17.3−21.1)
</paragraph>
<paragraph id="8BF4369B6C32FFCAFF0BDBC2FC32C88E" blockId="8.[151,1437,151,1185]" pageId="8" pageNumber="577">
Among the species in the subgenus 
<taxonomicName id="4C4B4D186C32FFCAFDA1DBC2FD1CC9EE" box="[621,764,152,176]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C32FFCAFDA1DBC2FD1CC9EE" box="[621,764,152,176]" italics="true" pageId="8" pageNumber="577">Guibemantis</emphasis>
</taxonomicName>
, the advertisement call of 
<taxonomicName id="4C4B4D186C32FFCAFBFADBC2FB5FC9EE" box="[1078,1215,152,176]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C32FFCAFBFADBC2FB5FC9EE" box="[1078,1215,152,176]" italics="true" pageId="8" pageNumber="577">G. diphonus</emphasis>
</taxonomicName>
is unique in being composed of two distinctly different note 
<typeStatus id="54F088396C32FFCAFDB8DBE7FD52C98B" box="[628,690,189,213]" pageId="8" pageNumber="577">types</typeStatus>
, whereas all other species as far as known exhibit a single pulsed note 
<typeStatus id="54F088396C32FFCAFF03DBBAFF1FC9A6" box="[207,255,224,248]" pageId="8" pageNumber="577">type</typeStatus>
only (see 
<figureCitation id="13702A1E6C32FFCAFEB9DB85FE5AC9A6" box="[373,442,223,248]" captionStart="FIGURE 4" captionStartId="6.[151,250,1926,1948]" captionTargetBox="[230,1335,193,1895]" captionTargetId="figure@6.[230,1336,193,1895]" captionTargetPageId="6" captionText="FIGURE 4. Comparative oscillograms (2000 ms sections each) of advertisement calls of species in the subgenus Guibemantis. For detailed call parameters see Table 2. Oscillogram of G. kathrinae taken from Glaw et al. (2000) showing a series of four calls according to the definition followed herein." httpUri="https://zenodo.org/record/244222/files/figure.png" pageId="8" pageNumber="577">Fig. 4</figureCitation>
). Comparing the first pulsed note 
<typeStatus id="54F088396C32FFCAFCF1DBBAFC8DC9A6" box="[829,877,224,248]" pageId="8" pageNumber="577">type</typeStatus>
of 
<taxonomicName id="4C4B4D186C32FFCAFC58DBBAFBFBC9A6" box="[916,1051,224,248]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C32FFCAFC58DBBAFBFBC9A6" box="[916,1051,224,248]" italics="true" pageId="8" pageNumber="577">G. diphonus</emphasis>
</taxonomicName>
with pulsed notes of other related 
<taxonomicName id="4C4B4D186C32FFCAFF5BDA5EFEC6C842" box="[151,294,260,284]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="genus">
<emphasis id="B93FEA896C32FFCAFF5BDA5EFEC6C842" box="[151,294,260,284]" italics="true" pageId="8" pageNumber="577">Guibemantis</emphasis>
</taxonomicName>
species, it differs from all of these by a much longer note duration (740−998 ms versus maximum 384 ms or far less in other species) and a far higher number of pulses per note (29−44 versus 7 or less pulses/note), at a somehow comparable pulse repetition rate. Furthermore, in 
<taxonomicName id="4C4B4D186C32FFCAFC89DA16FC2CC83A" box="[837,972,332,356]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C32FFCAFC89DA16FC2CC83A" box="[837,972,332,356]" italics="true" pageId="8" pageNumber="577">G. diphonus</emphasis>
</taxonomicName>
the two note 
<typeStatus id="54F088396C32FFCAFBA2DA17FB49C83B" box="[1134,1193,333,357]" pageId="8" pageNumber="577">types</typeStatus>
do not only differ in temporal characters, but also with respect to their relative amplitude, with the second note 
<typeStatus id="54F088396C32FFCAFB5EDA2AFB22C8D6" box="[1170,1218,368,392]" pageId="8" pageNumber="577">type</typeStatus>
being emitted with distinctly higher energy compared to the first note of the call. Given these very distinct advertisement call characters, 
<taxonomicName id="4C4B4D186C32FFCAFEDADAE2FE7BC88E" box="[278,411,440,464]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C32FFCAFEDADAE2FE7BC88E" box="[278,411,440,464]" italics="true" pageId="8" pageNumber="577">G. diphonus</emphasis>
</taxonomicName>
can be readily identified by its vocalization alone.
</paragraph>
</subSubSection>
<subSubSection id="C35165106C32FFCAFF0BDA86FA64CA4A" pageId="8" pageNumber="577" type="etymology">
<paragraph id="8BF4369B6C32FFCAFF0BDA86FEE4CB3E" blockId="8.[151,1437,151,1185]" pageId="8" pageNumber="577">
<emphasis id="B93FEA896C32FFCAFF0BDA86FEAEC8AB" bold="true" box="[199,334,476,501]" pageId="8" pageNumber="577">Etymology.</emphasis>
The specific epithet is derived from the Greek root word “phon” meaning “sound” and the prefix “di-” meaning “two”. In phonetics, a diphone is an adjacent pair of phones. The advertisement call of 
<taxonomicName id="4C4B4D186C32FFCAFAC1D95AFF1ECB62" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C32FFCAFAC1D95AFF1ECB62" italics="true" pageId="8" pageNumber="577">Guibemantis diphonus</emphasis>
</taxonomicName>
is characterized by the distinct combination of two note 
<typeStatus id="54F088396C32FFCAFC42D97FFC2CCB63" box="[910,972,549,573]" pageId="8" pageNumber="577">types</typeStatus>
. The specific epithet is used as latinized adjective.
</paragraph>
<paragraph id="8BF4369B6C32FFCAFF0BD936FA64CA4A" blockId="8.[151,1437,151,1185]" pageId="8" pageNumber="577">
<emphasis id="B93FEA896C32FFCAFF0BD936FE66CBDB" bold="true" box="[199,390,620,645]" pageId="8" pageNumber="577">Natural history.</emphasis>
The two males of the new species were collected at night on 
<date id="FFF5105B6C32FFCAFB8CD936FAE9CBDA" box="[1088,1289,620,645]" pageId="8" pageNumber="577" value="2007-02-23">23 February 2007</date>
along a very small moving shallow stream of ca. 
<quantity id="4CB39B7E6C32FFCAFDE3D9CAFD98CBF9" box="[559,632,656,680]" metricMagnitude="0" metricUnit="m" metricValue="2.5" metricValueMax="3.0" metricValueMin="2.0" pageId="8" pageNumber="577" unit="m" value="2.5" valueMax="3.0" valueMin="2.0">2−3 m</quantity>
in width and 
<quantity id="4CB39B7E6C32FFCAFCD8D9D5FCB9CBF9" box="[788,857,655,680]" metricMagnitude="-1" metricUnit="m" metricValue="5.0" pageId="8" pageNumber="577" unit="cm" value="50.0">50 cm</quantity>
in depth within a disturbed area of the low-altitude rainforest of Manombo Special Reserve. Specimens were sitting on perches about 
<quantity id="4CB39B7E6C32FFCAFB8CD9EEFB66CB92" box="[1088,1158,692,717]" metricMagnitude="-1" metricUnit="m" metricValue="5.0" pageId="8" pageNumber="577" unit="cm" value="50.0">50 cm</quantity>
above the ground in the low vegetation, directly next to the stream. Along the same stream, a dense chorus of 
<taxonomicName id="4C4B4D186C32FFCAFB42D982FAE3CBAE" box="[1166,1283,728,752]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="timidus">
<emphasis id="B93FEA896C32FFCAFB42D982FAE3CBAE" box="[1166,1283,728,752]" italics="true" pageId="8" pageNumber="577">G. timidus</emphasis>
</taxonomicName>
was present, together with a few individuals of 
<taxonomicName id="4C4B4D186C32FFCAFDD4D9A6FD6DCA4A" box="[536,653,764,788]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="tornieri">
<emphasis id="B93FEA896C32FFCAFDD4D9A6FD6DCA4A" box="[536,653,764,788]" italics="true" pageId="8" pageNumber="577">G. tornieri</emphasis>
</taxonomicName>
calling. The site was close to a swamp area with numerous 
<taxonomicName id="4C4B4D186C32FFCAFAE0D9A6FA9DCA4A" box="[1324,1405,764,788]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="liber">
<emphasis id="B93FEA896C32FFCAFAE0D9A6FA9DCA4A" box="[1324,1405,764,788]" italics="true" pageId="8" pageNumber="577">G. liber</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C35165106C32FFCAFF0BD845FA92CDFE" pageId="8" pageNumber="577" type="distribution">
<paragraph id="8BF4369B6C32FFCAFF0BD845FAA6CA66" blockId="8.[151,1437,151,1185]" box="[199,1350,799,824]" pageId="8" pageNumber="577">
<emphasis id="B93FEA896C32FFCAFF0BD845FE80CA66" bold="true" box="[199,352,799,824]" pageId="8" pageNumber="577">Distribution.</emphasis>
The species is only known from Manombo Special Reserve in southeastern 
<collectingCountry id="F35C760B6C32FFCAFB75D845FAA2CA66" box="[1209,1346,799,824]" name="Madagascar" pageId="8" pageNumber="577">Madagascar</collectingCountry>
.
</paragraph>
<paragraph id="8BF4369B6C32FFCAFF0BD81EFA92CDFE" blockId="8.[151,1437,151,1185]" pageId="8" pageNumber="577">
<emphasis id="B93FEA896C32FFCAFF0BD81EFE08CA03" bold="true" box="[199,488,836,861]" pageId="8" pageNumber="577">Available earlier names.</emphasis>
<taxonomicName id="4C4B4D186C32FFCAFE23D81EFC2FCA03" authority="Boulenger, 1896" authorityName="Boulenger" authorityYear="1896" box="[495,975,836,861]" class="Diplopoda" family="Chelodesmidae" genus="Rhacophorus" kingdom="Animalia" order="Polydesmida" pageId="8" pageNumber="577" phylum="Arthropoda" rank="species" species="mocquardii">
<emphasis id="B93FEA896C32FFCAFE23D81EFCECCA02" box="[495,780,836,860]" italics="true" pageId="8" pageNumber="577">Rhacophorus mocquardii</emphasis>
Boulenger, 1896
</taxonomicName>
, and 
<taxonomicName id="4C4B4D186C32FFCAFBC0D81EFA78CA02" authority="Ahl, 1929" authorityName="Ahl" authorityYear="1929" box="[1036,1432,836,861]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="acuticeps">
<emphasis id="B93FEA896C32FFCAFBC0D81EFAFDCA02" box="[1036,1309,836,860]" italics="true" pageId="8" pageNumber="577">Mantidactylus acuticeps</emphasis>
Ahl, 1929
</taxonomicName>
, both considered to be junior synonyms of 
<taxonomicName id="4C4B4D186C32FFCAFD5DD832FC5ACADE" box="[657,954,872,896]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="depressiceps">
<emphasis id="B93FEA896C32FFCAFD5DD832FC5ACADE" box="[657,954,872,896]" italics="true" pageId="8" pageNumber="577">Guibemantis depressiceps</emphasis>
</taxonomicName>
, could potentially be available as earlier names for 
<emphasis id="B93FEA896C32FFCAFED6D8D6FE4DCAFA" box="[282,429,908,932]" italics="true" pageId="8" pageNumber="577">
<taxonomicName id="4C4B4D186C32FFCAFED6D8D6FE49CAFA" box="[282,425,908,932]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="diphonus">G. diphonus</taxonomicName>
.
</emphasis>
However, the new species can be distinguished from the 
<typeStatus id="54F088396C32FFCAFBBED8D6FB34CAFB" box="[1138,1236,908,933]" pageId="8" pageNumber="577" type="holotype">holotype</typeStatus>
of 
<taxonomicName id="4C4B4D186C32FFCAFACAD8D6FEF7CA96" class="Diplopoda" family="Chelodesmidae" genus="Rhacophorus" kingdom="Animalia" order="Polydesmida" pageId="8" pageNumber="577" phylum="Arthropoda" rank="species" species="mocquardii">
<emphasis id="B93FEA896C32FFCAFACAD8D6FEF7CA96" italics="true" pageId="8" pageNumber="577">Rhacophorus mocquardii</emphasis>
</taxonomicName>
(BMNH 1947.2.8.62) by the almost complete lack of webbing between fingers 1 and 2 (see 
<figureCitation id="13702A1E6C32FFCAFAD0D8F5FABFCA96" box="[1308,1375,943,968]" captionStart="FIGURE 2" captionStartId="3.[151,250,1566,1588]" captionTargetBox="[151,1422,348,1544]" captionTargetId="figure@3.[151,1435,337,1545]" captionTargetPageId="3" captionText="FIGURE 2. Views of palmar surfaces of left hands of species in the subgenus Guibemantis (not to scale). Note the very distinct metacarpal tubercle of G. diphonus which is smaller or more flattened in all other figured specimens (especially in the types of earlier names, e. g. Rhacophorus mocquardii from the east coast locality Sahambendrana). Moreover, there are differences in webbing between fingers: although rudimentary in all species, it is almost absent in G. diphonus, but rather distinct in G. tornieri and sometimes visible in G. depressiceps, and also recognizable in the mocquardii holotype." httpUri="https://zenodo.org/record/244220/files/figure.png" pageId="8" pageNumber="577">Fig. 2</figureCitation>
), and from the 
<typeStatus id="54F088396C32FFCAFECDD88EFE83CAB3" box="[257,355,980,1005]" pageId="8" pageNumber="577" type="holotype">holotype</typeStatus>
of 
<taxonomicName id="4C4B4D186C32FFCAFE40D88EFD7FCAB2" box="[396,671,980,1004]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="acuticeps">
<emphasis id="B93FEA896C32FFCAFE40D88EFD7FCAB2" box="[396,671,980,1004]" italics="true" pageId="8" pageNumber="577">Mantidactylus acuticeps</emphasis>
</taxonomicName>
(ZMB 30496) by the complete lack of dark ventral spots and dark colour on limbs (colour patterns typical for specimens assigned to 
<taxonomicName id="4C4B4D186C32FFCAFCB0D8A2FBC8CD4E" box="[892,1064,1016,1040]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="depressiceps">
<emphasis id="B93FEA896C32FFCAFCB0D8A2FBC8CD4E" box="[892,1064,1016,1040]" italics="true" pageId="8" pageNumber="577">G. depressiceps</emphasis>
</taxonomicName>
). Furthermore, as documented in 
<figureCitation id="13702A1E6C32FFCAFF5BDF46FF18CD6A" box="[151,248,1052,1077]" captionStart="FIGURE 2" captionStartId="3.[151,250,1566,1588]" captionTargetBox="[151,1422,348,1544]" captionTargetId="figure@3.[151,1435,337,1545]" captionTargetPageId="3" captionText="FIGURE 2. Views of palmar surfaces of left hands of species in the subgenus Guibemantis (not to scale). Note the very distinct metacarpal tubercle of G. diphonus which is smaller or more flattened in all other figured specimens (especially in the types of earlier names, e. g. Rhacophorus mocquardii from the east coast locality Sahambendrana). Moreover, there are differences in webbing between fingers: although rudimentary in all species, it is almost absent in G. diphonus, but rather distinct in G. tornieri and sometimes visible in G. depressiceps, and also recognizable in the mocquardii holotype." httpUri="https://zenodo.org/record/244220/files/figure.png" pageId="8" pageNumber="577">Figure 2</figureCitation>
, the metacarpal tubercle in these two 
<typeStatus id="54F088396C32FFCAFD64DF47FD38CD6B" box="[680,728,1053,1077]" pageId="8" pageNumber="577">type</typeStatus>
specimens, and in the 
<typeStatus id="54F088396C32FFCAFC12DF46FBA5CD6B" box="[990,1093,1052,1077]" pageId="8" pageNumber="577" type="lectotype">lectotype</typeStatus>
of 
<taxonomicName id="4C4B4D186C32FFCAFBA1DF46FB1BCD6A" box="[1133,1275,1052,1076]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="depressiceps">
<emphasis id="B93FEA896C32FFCAFBA1DF46FB1BCD6A" box="[1133,1275,1052,1076]" italics="true" pageId="8" pageNumber="577">depressiceps</emphasis>
</taxonomicName>
, are more flat and less distinct compared to 
<taxonomicName id="4C4B4D186C32FFCAFE27DF1AFD92CD06" box="[491,626,1088,1112]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="diphonus">
<emphasis id="B93FEA896C32FFCAFE27DF1AFD92CD06" box="[491,626,1088,1112]" italics="true" pageId="8" pageNumber="577">G. diphonus</emphasis>
</taxonomicName>
. Although this is not a fully diagnostic character, we have observed in numerous more freshly preserved specimens of 
<taxonomicName id="4C4B4D186C32FFCAFD7FDF3EFC80CD22" box="[691,864,1124,1148]" class="Amphibia" family="Mantellidae" genus="Guibemantis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="577" phylum="Chordata" rank="species" species="depressiceps">
<emphasis id="B93FEA896C32FFCAFD7FDF3EFC80CD22" box="[691,864,1124,1148]" italics="true" pageId="8" pageNumber="577">G. depressiceps</emphasis>
</taxonomicName>
the more flat condition of this tubercle (e.g., ZSM 668/2001, 703/2003, 329/2010, 330/2010; see also 
<figureCitation id="13702A1E6C32FFCAFD1ADFDDFCF9CDFE" box="[726,793,1159,1184]" captionStart="FIGURE 2" captionStartId="3.[151,250,1566,1588]" captionTargetBox="[151,1422,348,1544]" captionTargetId="figure@3.[151,1435,337,1545]" captionTargetPageId="3" captionText="FIGURE 2. Views of palmar surfaces of left hands of species in the subgenus Guibemantis (not to scale). Note the very distinct metacarpal tubercle of G. diphonus which is smaller or more flattened in all other figured specimens (especially in the types of earlier names, e. g. Rhacophorus mocquardii from the east coast locality Sahambendrana). Moreover, there are differences in webbing between fingers: although rudimentary in all species, it is almost absent in G. diphonus, but rather distinct in G. tornieri and sometimes visible in G. depressiceps, and also recognizable in the mocquardii holotype." httpUri="https://zenodo.org/record/244220/files/figure.png" pageId="8" pageNumber="577">Fig. 2</figureCitation>
), similar to what is seen in the 
<typeStatus id="54F088396C32FFCAFBBADFD3FB46CDFF" box="[1142,1190,1161,1185]" pageId="8" pageNumber="577">type</typeStatus>
of 
<emphasis id="B93FEA896C32FFCAFB07DFD3FA92CDFE" box="[1227,1394,1160,1184]" italics="true" pageId="8" pageNumber="577">
<taxonomicName id="4C4B4D186C32FFCAFB07DFD3FA8ECDFE" box="[1227,1390,1160,1184]" class="Diplopoda" family="Chelodesmidae" genus="Rhacophorus" kingdom="Animalia" order="Polydesmida" pageId="8" pageNumber="577" phylum="Arthropoda" rank="species" species="mocquardii">R. mocquardii</taxonomicName>
.
</emphasis>
</paragraph>
</subSubSection>
</treatment>
</document>