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<document id="4FA72C36E225A8AEE3E0BFA07A3DC084" ID-DOI="10.11646/zootaxa.3717.2.8" ID-GBIF-Dataset="f327e088-45ea-4ad8-a005-cbc6528ea6db" ID-ISSN="1175-5326" ID-Zenodo-Dep="223536" ID-ZooBank="91AE5C97-8380-43B1-A97E-323CC64B9778" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1460683468232" checkinUser="plazi" docAuthor="Klages, Johannes, Glaw, Frank, Köhler, Jörn, Müller, Johannes, Hipsley, Christy A. &amp; Vences, Miguel" docDate="2013" docId="03D34717DC373250F682FF75FA0D161A" docLanguage="en" docName="zt03717p300.pdf" docOrigin="Zootaxa 3717 (2)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Stumpffia analamaina Klages, Glaw, Köhler, Müller, Hipsley &amp; Vences, 2013, sp. nov." docType="treatment" docUuid="CED11ABE-2814-4B2F-8506-5F22E5BF6710" docUuidSource="ZooBank" docVersion="8" lastPageNumber="294" masterDocId="FFEA3F6FDC31325EF615FFE2FF85110D" masterDocTitle="Molecular, morphological and osteological differentiation of a new species of microhylid frog of the genus Stumpffia from northwestern Madagascar" masterLastPageNumber="300" masterPageNumber="280" pageNumber="286" updateTime="1698350811545" updateUser="plazi">
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<mods:title id="50DD4C728C414196DAFEE8F2E91AE1E5">Molecular, morphological and osteological differentiation of a new species of microhylid frog of the genus Stumpffia from northwestern Madagascar</mods:title>
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<mods:namePart id="FDE3FCF4E9E0528CB7417D1D575CF5E7">Klages, Johannes</mods:namePart>
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<mods:namePart id="96CB9CCF03264DF99699B89973E47D8F">Glaw, Frank</mods:namePart>
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<mods:namePart id="3FAB137853CA9B3362D7EB923CF7C09F">Müller, Johannes</mods:namePart>
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<mods:namePart id="FADD3B31167252332B53029A746EF682">Vences, Miguel</mods:namePart>
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<treatment id="03D34717DC373250F682FF75FA0D161A" ID-DOI="http://doi.org/10.5281/zenodo.5695699" ID-GBIF-Taxon="119577131" ID-Zenodo-Dep="5695699" LSID="urn:lsid:zoobank.org:act:CED11ABE-2814-4B2F-8506-5F22E5BF6710" httpUri="http://treatment.plazi.org/id/03D34717DC373250F682FF75FA0D161A" lastPageId="14" lastPageNumber="294" pageId="6" pageNumber="286">
<subSubSection id="C360A58ADC373258F682FF75FE9411DE" pageId="6" pageNumber="286" type="nomenclature">
<paragraph id="8BC5F601DC373258F682FF75FD8F11BF" blockId="6.[151,522,151,211]" box="[151,522,151,178]" pageId="6" pageNumber="286">
<heading id="D08D416DDC373258F682FF75FD8F11BF" bold="true" box="[151,522,151,178]" fontSize="11" level="1" pageId="6" pageNumber="286" reason="1">
<emphasis id="B90E2A13DC373258F682FF75FD8F11BF" bold="true" box="[151,522,151,178]" pageId="6" pageNumber="286">
<taxonomicName id="4C7A8D82DC373258F682FF75FE2311BC" ID-CoL="537S6" box="[151,422,151,177]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="6" pageNumber="286" phylum="Chordata" rank="species" species="analamaina" status="sp. nov.">
<emphasis id="B90E2A13DC373258F682FF75FE2311BC" bold="true" box="[151,422,151,177]" italics="true" pageId="6" pageNumber="286">Stumpffia analamaina</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D9768DC373258F7B8FF7AFD8F11BF" box="[429,522,152,178]" pageId="6" pageNumber="286" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
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<paragraph id="8BC5F601DC373258F682FF58FE9411DE" blockId="6.[151,522,151,211]" box="[151,273,186,211]" pageId="6" pageNumber="286">
(
<figureCitation id="1341EA84DC373258F68AFF58FE8C11DE" box="[159,265,186,211]" captionStart-0="FIGURE 3" captionStart-1="FIGURE 4" captionStart-2="FIGURE 5" captionStartId-0="6.[151,250,1945,1967]" captionStartId-1="7.[151,250,1852,1874]" captionStartId-2="8.[151,250,1647,1669]" captionTargetBox-0="[242,1344,445,1924]" captionTargetBox-1="[176,1425,260,1828]" captionTargetBox-2="[151,1436,193,1624]" captionTargetId-0="figure@6.[242,1345,443,1924]" captionTargetId-1="figure@7.[166,1426,165,1829]" captionTargetId-2="figure@8.[151,1436,193,1626]" captionTargetPageId-0="6" captionTargetPageId-1="7" captionTargetPageId-2="8" captionText-0="FIGURE 3. Holotype (ZSM 542 / 2009) of Stumpffia analamaina sp. nov. in preservative in (A) dorsal view, (B) foot and (C) hand in ventral view, and (D) living holotype in dorsolateral view. Scale bar = 1 mm." captionText-1="FIGURE 4. Specimens and habitat of Stumpffia analamaina sp. nov. (A) Paratype specimen ZSM 2831 / 2010 (ZCMV 10037) in life (scale bar = 1 mm). (B) Frame from a video showing prey capture of S. analamaina, showing partially lateral tongue protrusion as also known from other microhylids. The whitish structure on the right side of the head (arrow) is the tongue which is protruded at an angle of 40 50 ° relative to the body axis. (C) Dry forest and (D) stream at type locality, specimens were found in the leaf litter at the border of the stream. (E) Foam nest produced by S. analamaina in captivity. (F) Additional specimen (catalogue number uncertain) showing color variation (beige ground color)." captionText-2="FIGURE 5. Skull in oblique right lateral view (above) and ventral view (below), and presacral vertebral column in dorsal view (right) of Stumpffia analamaina sp. nov., paratype specimen ZSM 2831 / 2010 (ZCMV 10037). Abbreviations: Den, dentary; FP, frontoparietals; Max, maxilla; Na, nasals; Npl—neopalatinum; ParSph, parasphenoid; PrMax, premaxilla; Pt—pterygoid; Qj—quadratojugale; Sac, sacrum; Sq, squamosal (Sq-AP and Sq-PP—anterior and posterior process of squamosal); Ur, urostyle (incompletely shown); V—Vomer; V 1 - V 8, presacral vertebrae (V 1 + V 2 includes atlas and axis)." httpUri-0="https://zenodo.org/record/223539/files/figure.png" httpUri-1="https://zenodo.org/record/223540/files/figure.png" httpUri-2="https://zenodo.org/record/223541/files/figure.png" pageId="6" pageNumber="286">Figs. 35</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="C360A58ADC373256F682FEE0FB4C1683" lastPageId="8" lastPageNumber="288" pageId="6" pageNumber="286" type="materials_examined">
<paragraph id="8BC5F601DC373258F682FEE0FF63108B" blockId="6.[151,1436,258,390]" pageId="6" pageNumber="286">
<emphasis id="B90E2A13DC373258F682FEE0FE8E1016" bold="true" box="[151,267,258,283]" pageId="6" pageNumber="286">
<typeStatus id="54C148A3DC373258F682FEE0FE831016" box="[151,262,258,283]" pageId="6" pageNumber="286" type="holotype">Holotype</typeStatus>
.
</emphasis>
Adult male (
<figureCitation id="1341EA84DC373258F788FEE0FE641016" box="[413,481,258,283]" captionStart="FIGURE 3" captionStartId="6.[151,250,1945,1967]" captionTargetBox="[242,1344,445,1924]" captionTargetId="figure@6.[242,1345,443,1924]" captionTargetPageId="6" captionText="FIGURE 3. Holotype (ZSM 542 / 2009) of Stumpffia analamaina sp. nov. in preservative in (A) dorsal view, (B) foot and (C) hand in ventral view, and (D) living holotype in dorsolateral view. Scale bar = 1 mm." httpUri="https://zenodo.org/record/223539/files/figure.png" pageId="6" pageNumber="286">Fig. 3</figureCitation>
), ZSM 542/2009 (field number ZCMV 11428), collected at a bridge located at km 27 on the national road from Antsohihy to Mandritsara,
<geoCoordinate id="EE4E90C6DC373258F4EBFEC4FC171032" box="[766,914,294,319]" direction="south" orientation="latitude" pageId="6" pageNumber="286" precision="1" value="-15.053223">15°03'11.6''S</geoCoordinate>
,
<geoCoordinate id="EE4E90C6DC373258F5B4FEC5FBB21032" box="[929,1079,295,319]" direction="east" orientation="longitude" pageId="6" pageNumber="286" precision="1" value="48.206417">48°12'23.1''E</geoCoordinate>
,
<quantity id="4C825BE4DC373258F253FEC5FB0A1033" box="[1094,1167,295,319]" metricMagnitude="2" metricUnit="m" metricValue="1.4" pageId="6" pageNumber="286" unit="m" value="140.0">140 m</quantity>
above sea level, on
<date id="FFC4D0C1DC373258F395FEC5FE8C106E" pageId="6" pageNumber="286" value="2009-06-18">18 June 2009</date>
by M. Vences, D. R. Vieites, F. Ratsoavina, R. D. Randrianiaina, E. Rajeriarison, T. Rajoafiarison, and J. Patton.
</paragraph>
<caption id="DF05A689DC373258F682F87BFC6216C3" httpUri="https://zenodo.org/record/223539/files/figure.png" pageId="6" pageNumber="286" targetBox="[242,1344,445,1924]" targetPageId="6">
<paragraph id="8BC5F601DC373258F682F87BFC6216C3" blockId="6.[151,1436,1945,1998]" pageId="6" pageNumber="286">
<emphasis id="B90E2A13DC373258F682F87BFE9116A3" bold="true" box="[151,276,1945,1967]" pageId="6" pageNumber="286">FIGURE 3.</emphasis>
Holotype (ZSM 542/2009) of
<taxonomicName id="4C7A8D82DC373258F45AF87BFCB516A2" box="[591,816,1945,1967]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="6" pageNumber="286" phylum="Chordata" rank="species" species="analamaina" status="sp. nov.">
<emphasis id="B90E2A13DC373258F45AF87BFCB516A2" box="[591,816,1945,1967]" italics="true" pageId="6" pageNumber="286">Stumpffia analamaina</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D9768DC373258F522F878FC0D16A2" box="[823,904,1946,1967]" pageId="6" pageNumber="286" rank="species">
<emphasis id="B90E2A13DC373258F522F878FC0D16A2" bold="true" box="[823,904,1946,1967]" pageId="6" pageNumber="286">sp. nov.</emphasis>
</taxonomicNameLabel>
in preservative in (A) dorsal view, (B) foot and (C) hand in ventral view, and (D) living holotype in dorsolateral view. Scale bar=1 mm.
</paragraph>
</caption>
<caption id="DF05A689DC363259F682F8DEFC7E16E6" httpUri="https://zenodo.org/record/223540/files/figure.png" pageId="7" pageNumber="287" targetBox="[176,1425,260,1828]" targetPageId="7">
<paragraph id="8BC5F601DC363259F682F8DEFC7E16E6" blockId="7.[151,1436,1852,2027]" pageId="7" pageNumber="287">
<emphasis id="B90E2A13DC363259F682F8DEFE96165C" bold="true" box="[151,275,1852,1874]" pageId="7" pageNumber="287">FIGURE 4.</emphasis>
Specimens and habitat of
<taxonomicName id="4C7A8D82DC363259F435F8DEFC85165F" box="[544,768,1852,1874]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="7" pageNumber="287" phylum="Chordata" rank="species" species="analamaina" status="sp. nov.">
<emphasis id="B90E2A13DC363259F435F8DEFC85165F" box="[544,768,1852,1874]" italics="true" pageId="7" pageNumber="287">Stumpffia analamaina</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D9768DC363259F512F8DFFCDD165F" box="[775,856,1853,1874]" pageId="7" pageNumber="287" rank="species">
<emphasis id="B90E2A13DC363259F512F8DFFCDD165F" bold="true" box="[775,856,1853,1874]" pageId="7" pageNumber="287">sp. nov.</emphasis>
</taxonomicNameLabel>
(A) Paratype specimen ZSM 2831/2010 (ZCMV 10037) in life (scale bar=1 mm). (B) Frame from a video showing prey capture of
<emphasis id="B90E2A13DC363259F5BAF8BEFBCF167C" box="[943,1098,1883,1905]" italics="true" pageId="7" pageNumber="287">
<taxonomicName id="4C7A8D82DC363259F5BAF8BEFBC2167C" box="[943,1095,1883,1905]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="7" pageNumber="287" phylum="Chordata" rank="species" species="analamaina">S. analamaina</taxonomicName>
,
</emphasis>
showing partially lateral tongue protrusion as also known from other microhylids. The whitish structure on the right side of the head (arrow) is the tongue which is protruded at an angle of 4050° relative to the body axis. (C) Dry forest and (D) stream at type locality, specimens were found in the leaf litter at the border of the stream. (E) Foam nest produced by
<taxonomicName id="4C7A8D82DC363259F5F6F85AFBFD16C0" box="[995,1144,1975,1997]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="7" pageNumber="287" phylum="Chordata" rank="species" species="analamaina">
<emphasis id="B90E2A13DC363259F5F6F85AFBFD16C0" box="[995,1144,1975,1997]" italics="true" pageId="7" pageNumber="287">S. analamaina</emphasis>
</taxonomicName>
in captivity. (F) Additional specimen (catalogue number uncertain) showing color variation (beige ground color).
</paragraph>
</caption>
<caption id="DF05A689DC393256F682F98DFB39160D" httpUri="https://zenodo.org/record/223541/files/figure.png" pageId="8" pageNumber="288" targetBox="[151,1436,193,1624]" targetPageId="8">
<paragraph id="8BC5F601DC393256F682F98DFB39160D" blockId="8.[151,1436,1647,1792]" pageId="8" pageNumber="288">
<emphasis id="B90E2A13DC393256F682F98DFE971788" bold="true" box="[151,274,1647,1669]" pageId="8" pageNumber="288">FIGURE 5.</emphasis>
Skull in oblique right lateral view (above) and ventral view (below), and presacral vertebral column in dorsal view (right) of
<taxonomicName id="4C7A8D82DC393256F6E8F96CFE5A17A9" box="[253,479,1678,1700]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="8" pageNumber="288" phylum="Chordata" rank="species" species="analamaina" status="sp. nov.">
<emphasis id="B90E2A13DC393256F6E8F96CFE5A17A9" box="[253,479,1678,1700]" italics="true" pageId="8" pageNumber="288">Stumpffia analamaina</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D9768DC393256F7F2F96DFDBC17A9" box="[487,569,1679,1700]" pageId="8" pageNumber="288" rank="species">
<emphasis id="B90E2A13DC393256F7F2F96DFDBC17A9" bold="true" box="[487,569,1679,1700]" pageId="8" pageNumber="288">sp. nov.</emphasis>
</taxonomicNameLabel>
, paratype specimen ZSM 2831/2010 (ZCMV 10037). Abbreviations: Den, dentary; FP, frontoparietals; Max, maxilla; Na, nasals; Npl—neopalatinum; ParSph, parasphenoid; PrMax, premaxilla; Pt—pterygoid; Qj—quadratojugale; Sac, sacrum; Sq, squamosal (Sq-AP and Sq-PP—anterior and posterior process of squamosal); Ur, urostyle (incompletely shown); V—Vomer; V1-V8, presacral vertebrae (V1+V2 includes atlas and axis).
</paragraph>
</caption>
<paragraph id="8BC5F601DC393256F6D2F8CEFB4C1683" blockId="8.[151,1436,1836,2006]" pageId="8" pageNumber="288">
<emphasis id="B90E2A13DC393256F6D2F8CEFEC31648" bold="true" box="[199,326,1836,1861]" pageId="8" pageNumber="288">
<typeStatus id="54C148A3DC393256F6D2F8CEFEC41648" box="[199,321,1836,1861]" pageId="8" pageNumber="288" type="paratype">Paratypes</typeStatus>
.
</emphasis>
Three specimens, ZSM 2829/2010 (ZCMV 10034), ZSM 2830/2010 (ZCMV 10036) and ZSM 2831/2010 (ZCMV 10037), collected at the same locality as the
<typeStatus id="54C148A3DC393256F59BF8B2FC761664" box="[910,1011,1872,1897]" pageId="8" pageNumber="288" type="holotype">holotype</typeStatus>
, on
<date id="FFC4D0C1DC393256F23CF8B3FB431664" box="[1065,1222,1873,1897]" pageId="8" pageNumber="288" value="2010-06-29">29 June 2010</date>
by M. Vences, E. Rajeriarison, T. Rajoafiarison, A. Rakotoarison, S. Rasamison, F. Ratsoavina, and D. R. Vieites.
</paragraph>
</subSubSection>
<subSubSection id="C360A58ADC393257F6D2F87AFD5414E8" lastPageId="9" lastPageNumber="289" pageId="8" pageNumber="288" type="diagnosis">
<paragraph id="8BC5F601DC393257F6D2F87AFE6010DD" blockId="8.[151,1436,1836,2006]" lastBlockId="9.[151,1436,151,2013]" lastPageId="9" lastPageNumber="289" pageId="8" pageNumber="288">
<emphasis id="B90E2A13DC393256F6D2F87AFEC416BC" bold="true" box="[199,321,1944,1969]" pageId="8" pageNumber="288">Diagnosis.</emphasis>
The new species is assigned to the genus
<taxonomicName id="4C7A8D82DC393256F531F87BFC0A16BC" box="[804,911,1945,1969]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="8" pageNumber="288" phylum="Chordata" rank="genus">
<emphasis id="B90E2A13DC393256F531F87BFC0A16BC" box="[804,911,1945,1969]" italics="true" pageId="8" pageNumber="288">Stumpffia</emphasis>
</taxonomicName>
on the basis of its very small size, absence of vomerine and maxillary teeth, ossified part of clavicles reduced to thin rod-like elements, sternum and omosternum not ossified, similarities in external body shape to other species in the genus, and molecular phylogenetic relationships. Within
<taxonomicName id="4C7A8D82DC383257F79FFF5EFE7111D9" box="[394,500,188,212]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="genus">
<emphasis id="B90E2A13DC383257F79FFF5EFE7111D9" box="[394,500,188,212]" italics="true" pageId="9" pageNumber="289">Stumpffia</emphasis>
</taxonomicName>
, it is uniquely characterized by the following combination of characters: (1) small body size, SVL of males and females
<quantity id="4C825BE4DC383257F457FF02FD3A11FA" box="[578,703,224,248]" metricMagnitude="-2" metricUnit="m" metricValue="1.1" metricValueMax="1.2" metricValueMin="1.0" pageId="9" pageNumber="289" unit="mm" value="11.0" valueMax="12.0" valueMin="10.0">1012 mm</quantity>
; (2) dorsum smooth or slightly tubercular; (3) absence of distinct terminal discs on fingers and toes; (4) advertisement calls consisting of a series of short melodious chirps; (5) greyish or brownish coloration with variable but typically indistinct pattern and without contrasted ventral coloration, red color elements on ventral side, or sharp color border between dorsum and flanks; (6) manus with four fingers (first finger slightly reduced in size) and pes with five toes (first toe slightly reduced in size); (7) third toe slightly to distinctly longer than fifth toe; (8) relative hand length, HAL/SVL 0.220.26, relative foot + tarsus length FOTL/SVL 0.680.78.
</paragraph>
<paragraph id="8BC5F601DC383257F6D2FE3FFA0F1555" blockId="9.[151,1436,151,2013]" pageId="9" pageNumber="289">
In comparison to the other nominal species of
<taxonomicName id="4C7A8D82DC383257F4D8FE3EFCB210F9" box="[717,823,476,500]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="genus">
<emphasis id="B90E2A13DC383257F4D8FE3EFCB210F9" box="[717,823,476,500]" italics="true" pageId="9" pageNumber="289">Stumpffia</emphasis>
</taxonomicName>
, the new species differs from
<taxonomicName id="4C7A8D82DC383257F29DFE3FFB7E10F8" box="[1160,1275,476,501]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="helenae">
<emphasis id="B90E2A13DC383257F29DFE3FFB7E10F8" box="[1160,1275,476,501]" italics="true" pageId="9" pageNumber="289">S. helenae</emphasis>
</taxonomicName>
by absence of terminal discs on toes and fingers (vs. presence) and slightly smaller body size (SVL of males and females 1012 vs.
<quantity id="4C825BE4DC383257F6ABFDC6FEB91331" box="[190,316,548,573]" metricMagnitude="-2" metricUnit="m" metricValue="1.45" metricValueMax="1.5" metricValueMin="1.4" pageId="9" pageNumber="289" unit="mm" value="14.5" valueMax="15.0" valueMin="14.0">1415 mm</quantity>
); from
<emphasis id="B90E2A13DC383257F79BFDC7FE411330" box="[398,452,548,573]" italics="true" pageId="9" pageNumber="289">S. be</emphasis>
by distinctly smaller body size (SVL
<quantity id="4C825BE4DC383257F564FDC6FC6E1331" box="[881,1003,548,572]" metricMagnitude="-2" metricUnit="m" metricValue="1.1" metricValueMax="1.2" metricValueMin="1.0" pageId="9" pageNumber="289" unit="mm" value="11.0" valueMax="12.0" valueMin="10.0">1012 mm</quantity>
vs.
<quantity id="4C825BE4DC383257F20CFDC6FBE81331" box="[1049,1133,548,573]" metricMagnitude="-2" metricUnit="m" metricValue="2.5" pageId="9" pageNumber="289" unit="mm" value="25.0">25 mm</quantity>
), absence of terminal discs on fingers and toes (vs. presence); from
<taxonomicName id="4C7A8D82DC383257F476FDAAFD30136D" box="[611,693,583,608]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="hara">
<emphasis id="B90E2A13DC383257F476FDAAFD30136D" box="[611,693,583,608]" italics="true" pageId="9" pageNumber="289">S. hara</emphasis>
</taxonomicName>
by distinctly smaller body size (SVL
<quantity id="4C825BE4DC383257F272FDAAFB671352" box="[1127,1250,584,608]" metricMagnitude="-2" metricUnit="m" metricValue="1.1" metricValueMax="1.2" metricValueMin="1.0" pageId="9" pageNumber="289" unit="mm" value="11.0" valueMax="12.0" valueMin="10.0">1012 mm</quantity>
vs.
<quantity id="4C825BE4DC383257F307FDA5FA15136D" box="[1298,1424,583,608]" metricMagnitude="-2" metricUnit="m" metricValue="2.35" metricValueMax="2.5" metricValueMin="2.2" pageId="9" pageNumber="289" unit="mm" value="23.5" valueMax="25.0" valueMin="22.0">2225 mm</quantity>
), absence of terminal discs on fingers and toes (vs. presence); from
<taxonomicName id="4C7A8D82DC383257F56BFD8FFC7D1389" box="[894,1016,621,644]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="megsoni">
<emphasis id="B90E2A13DC383257F56BFD8FFC7D1389" box="[894,1016,621,644]" italics="true" pageId="9" pageNumber="289">S. megsoni</emphasis>
</taxonomicName>
by distinctly smaller body size (SVL
<quantity id="4C825BE4DC383257F682FD72FE9113AA" box="[151,276,656,680]" metricMagnitude="-2" metricUnit="m" metricValue="1.1" metricValueMax="1.2" metricValueMin="1.0" pageId="9" pageNumber="289" unit="mm" value="11.0" valueMax="12.0" valueMin="10.0">1012 mm</quantity>
vs.
<quantity id="4C825BE4DC383257F75FFD72FE4913A5" box="[330,460,656,680]" metricMagnitude="-2" metricUnit="m" metricValue="2.15" metricValueMax="2.2" metricValueMin="2.1" pageId="9" pageNumber="289" unit="mm" value="21.5" valueMax="22.0" valueMin="21.0">2122 mm</quantity>
), absence of terminal discs on fingers and toes (vs. presence); from
<taxonomicName id="4C7A8D82DC383257F2E3FD72FAF013A5" box="[1270,1397,655,680]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="staffordi">
<emphasis id="B90E2A13DC383257F2E3FD72FAF013A5" box="[1270,1397,655,680]" italics="true" pageId="9" pageNumber="289">S. staffordi</emphasis>
</taxonomicName>
by distinctly smaller body size (SVL
<quantity id="4C825BE4DC383257F437FD56FD1813C1" box="[546,669,692,716]" metricMagnitude="-2" metricUnit="m" metricValue="1.1" metricValueMax="1.2" metricValueMin="1.0" pageId="9" pageNumber="289" unit="mm" value="11.0" valueMax="12.0" valueMin="10.0">1012 mm</quantity>
vs.
<quantity id="4C825BE4DC383257F4DAFD56FCCA13C1" box="[719,847,692,716]" metricMagnitude="-2" metricUnit="m" metricValue="2.75" metricValueMax="2.8" metricValueMin="2.7" pageId="9" pageNumber="289" unit="mm" value="27.5" valueMax="28.0" valueMin="27.0">2728 mm</quantity>
), absence of terminal discs on fingers and toes (vs. presence); from
<taxonomicName id="4C7A8D82DC383257F75AFD3AFE4413FD" box="[335,449,727,752]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="grandis">
<emphasis id="B90E2A13DC383257F75AFD3AFE4413FD" box="[335,449,727,752]" italics="true" pageId="9" pageNumber="289">S. grandis</emphasis>
</taxonomicName>
by smaller body size (SVL 1012 vs.
<quantity id="4C825BE4DC383257F560FD3AFC7713FD" box="[885,1010,728,752]" metricMagnitude="-2" metricUnit="m" metricValue="2.15" metricValueMax="2.4" metricValueMin="1.9" pageId="9" pageNumber="289" unit="mm" value="21.5" valueMax="24.0" valueMin="19.0">1924 mm</quantity>
), and a different coloration (indistinct brown-greyish pattern vs. strongly contrasted pattern of brown patches and bluish venter with black markings); from
<taxonomicName id="4C7A8D82DC383257F6C3FCC2FE101235" box="[214,405,799,824]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="roseifemoralis">
<emphasis id="B90E2A13DC383257F6C3FCC2FE101235" box="[214,405,799,824]" italics="true" pageId="9" pageNumber="289">S. roseifemoralis</emphasis>
</taxonomicName>
by smaller body size (
<quantity id="4C825BE4DC383257F48EFCC2FC92123A" box="[667,791,800,824]" metricMagnitude="-2" metricUnit="m" metricValue="1.1" metricValueMax="1.2" metricValueMin="1.0" pageId="9" pageNumber="289" unit="mm" value="11.0" valueMax="12.0" valueMin="10.0">1012 mm</quantity>
vs.
<quantity id="4C825BE4DC383257F55DFCC2FC411235" box="[840,964,800,824]" metricMagnitude="-2" metricUnit="m" metricValue="1.85" metricValueMax="1.9" metricValueMin="1.8" pageId="9" pageNumber="289" unit="mm" value="18.5" valueMax="19.0" valueMin="18.0">1819 mm</quantity>
) and different coloration (lack of reddish color ventrally on limbs vs. presence); from
<taxonomicName id="4C7A8D82DC383257F49CFCA7FCB11250" box="[649,820,836,861]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="psologlosssa">
<emphasis id="B90E2A13DC383257F49CFCA7FCB11250" box="[649,820,836,861]" italics="true" pageId="9" pageNumber="289">S. psologlosssa</emphasis>
</taxonomicName>
by slightly smaller body size (SVL
<quantity id="4C825BE4DC383257F2DEFCA6FAC11251" box="[1227,1348,836,860]" metricMagnitude="-2" metricUnit="m" metricValue="1.1" metricValueMax="1.2" metricValueMin="1.0" pageId="9" pageNumber="289" unit="mm" value="11.0" valueMax="12.0" valueMin="10.0">1012 mm</quantity>
vs.
<quantity id="4C825BE4DC383257F367FCA6FF6E128D" metricMagnitude="-2" metricUnit="m" metricValue="1.5" metricValueMax="1.6" metricValueMin="1.4" pageId="9" pageNumber="289" unit="mm" value="15.0" valueMax="16.0" valueMin="14.0">14 16 mm</quantity>
), a more strongly reduced first toe (vs. slightly reduced), and a different structure of advertisement call (a series of short chirps vs. short trills); from
<taxonomicName id="4C7A8D82DC383257F466FC6FFD7E12A8" box="[627,763,908,933]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="tridactyla">
<emphasis id="B90E2A13DC383257F466FC6FFD7E12A8" box="[627,763,908,933]" italics="true" pageId="9" pageNumber="289">S. tridactyla</emphasis>
</taxonomicName>
by presence of four externally recognizable fingers and five toes (vs. a single triangular finger and three toes); from
<taxonomicName id="4C7A8D82DC383257F51FFC52FC2812C5" box="[778,941,943,968]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="tetradactyla">
<emphasis id="B90E2A13DC383257F51FFC52FC2812C5" box="[778,941,943,968]" italics="true" pageId="9" pageNumber="289">S. tetradactyla</emphasis>
</taxonomicName>
by presence of five recognizable toes albeit first toe very small (vs. first toe almost completely reduced, only visible as a small tubercle); from
<taxonomicName id="4C7A8D82DC383257F315FC37FAF212E0" box="[1280,1399,980,1005]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="gimmeli">
<emphasis id="B90E2A13DC383257F315FC37FAF212E0" box="[1280,1399,980,1005]" italics="true" pageId="9" pageNumber="289">S. gimmeli</emphasis>
</taxonomicName>
by smaller body size (SVL
<quantity id="4C825BE4DC383257F7A2FC1AFDB11502" box="[439,564,1016,1040]" metricMagnitude="-2" metricUnit="m" metricValue="1.1" metricValueMax="1.2" metricValueMin="1.0" pageId="9" pageNumber="289" unit="mm" value="11.0" valueMax="12.0" valueMin="10.0">1012 mm</quantity>
vs.
<quantity id="4C825BE4DC383257F47CFC15FD6F151D" box="[617,746,1015,1040]" metricMagnitude="-2" metricUnit="m" metricValue="1.65" metricValueMax="1.8" metricValueMin="1.5" pageId="9" pageNumber="289" unit="mm" value="16.5" valueMax="18.0" valueMin="15.0">1518 mm</quantity>
), dorsal skin smooth or only slightly tubercular in life (vs. typically distinctly tubercular), first finger and first toe slightly reduced (vs. well developed), absence of terminal discs on fingers and toes (vs. typical presence of at least weakly developed discs, especially in larger specimens).
</paragraph>
<paragraph id="8BC5F601DC383257F6D2FB86FA9D1475" blockId="9.[151,1436,151,2013]" pageId="9" pageNumber="289">
The new species is morphologically most similar to
<taxonomicName id="4C7A8D82DC383257F536FB87FC231571" box="[803,934,1125,1148]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="pygmaea">
<emphasis id="B90E2A13DC383257F536FB87FC231571" box="[803,934,1125,1148]" italics="true" pageId="9" pageNumber="289">S. pygmaea</emphasis>
</taxonomicName>
and
<emphasis id="B90E2A13DC383257F5F7FB87FB571570" box="[994,1234,1124,1149]" italics="true" pageId="9" pageNumber="289">
<taxonomicName id="4C7A8D82DC383257F5F7FB87FB4B1570" box="[994,1230,1124,1149]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="madagascariensis">S. madagascariensis</taxonomicName>
.
</emphasis>
It differs from
<taxonomicName id="4C7A8D82DC383257F392FB87FEE415AD" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="madagascariensis">
<emphasis id="B90E2A13DC383257F392FB87FEE415AD" italics="true" pageId="9" pageNumber="289">S. madagascariensis</emphasis>
</taxonomicName>
by the lack of a sharp border between dark flanks and light dorsum (vs. presence; figure in Köhler
<emphasis id="B90E2A13DC383257F6F8FB4FFEAD15C8" box="[237,296,1196,1221]" italics="true" pageId="9" pageNumber="289">et al.</emphasis>
2010). Furthermore, based on the limited morphometric data available to date (see Vences &amp; Glaw 1991 and
<tableCitation id="C6F8C3BADC383257F702FB2DFEF715E5" box="[279,370,1231,1256]" captionStart="TABLE 1" captionStartId="10.[151,239,980,1003]" captionTargetBox="[159,1403,1249,1642]" captionTargetPageId="10" captionText="TABLE 1. Morphometric data (in mm) of selected specimens of S. gimmeli, S. madagascariensis, S. pygmaea, S. sp. 31, and S. analamaina sp. nov. SVL, snout-vent length; HAL, hand length; FOTL, foot and tarsus length; FOL, foot length; TIBL, tibia length; HT, holotype; PT, paratype." httpUri="http://table.plazi.org/id/DF05A689DC3B3254F682FC36FD0B1526" pageId="9" pageNumber="289" tableUuid="DF05A689DC3B3254F682FC36FD0B1526">Table 1</tableCitation>
), it appears that the new species has longer hands and feet than
<taxonomicName id="4C7A8D82DC383257F2A1FB32FABB15E5" box="[1204,1342,1232,1256]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="pygmaea">
<emphasis id="B90E2A13DC383257F2A1FB32FABB15E5" box="[1204,1342,1232,1256]" italics="true" pageId="9" pageNumber="289">S. pygmaea</emphasis>
</taxonomicName>
and
<taxonomicName id="4C7A8D82DC383257F392FB32FEE41400" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="madagascariensis">
<emphasis id="B90E2A13DC383257F392FB32FEE41400" italics="true" pageId="9" pageNumber="289">S. madagascariensis</emphasis>
</taxonomicName>
: values for relative hand length and relative foot + tarsus length of
<taxonomicName id="4C7A8D82DC383257F27CFB17FA881400" box="[1129,1293,1268,1293]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="analamaina">
<emphasis id="B90E2A13DC383257F27CFB17FA881400" box="[1129,1293,1268,1293]" italics="true" pageId="9" pageNumber="289">S. analamaina</emphasis>
</taxonomicName>
(HAL/SVL, 0.220.26; FOTL/SVL, 0.680.81) are in the range known for
<taxonomicName id="4C7A8D82DC383257F549FAFAFC78143D" box="[860,1021,1303,1328]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="psologlossa">
<emphasis id="B90E2A13DC383257F549FAFAFC78143D" box="[860,1021,1303,1328]" italics="true" pageId="9" pageNumber="289">S. psologlossa</emphasis>
</taxonomicName>
(HAL/SVL, 0.180.23; FOTL/SVL, 0.630.71) and
<taxonomicName id="4C7A8D82DC383257F75EFADFFE411458" box="[331,452,1340,1365]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="gimmeli">
<emphasis id="B90E2A13DC383257F75EFADFFE411458" box="[331,452,1340,1365]" italics="true" pageId="9" pageNumber="289">S. gimmeli</emphasis>
</taxonomicName>
(HAL/SVL, 0.210.23; FOTL/SVL, 0.680.75) but larger than in
<taxonomicName id="4C7A8D82DC383257F2D1FADFFACD1459" box="[1220,1352,1341,1364]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="pygmaea">
<emphasis id="B90E2A13DC383257F2D1FADFFACD1459" box="[1220,1352,1341,1364]" italics="true" pageId="9" pageNumber="289">S. pygmaea</emphasis>
</taxonomicName>
(HAL/ SVL, 0.160.18; FOTL/SVL 0.530.63) and
<taxonomicName id="4C7A8D82DC383257F487FA82FCFD1475" box="[658,888,1375,1400]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="species" species="madagascariensis">
<emphasis id="B90E2A13DC383257F487FA82FCFD1475" box="[658,888,1375,1400]" italics="true" pageId="9" pageNumber="289">S. madagascariensis</emphasis>
</taxonomicName>
(HAL/SVL, 0.16; FOTL/SVL 0.61).
</paragraph>
<paragraph id="8BC5F601DC383257F6D2FA66FD5414E8" blockId="9.[151,1436,151,2013]" pageId="9" pageNumber="289">
The new species differs from all nominal species of
<taxonomicName id="4C7A8D82DC383257F509FA66FC031491" box="[796,902,1412,1436]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="9" pageNumber="289" phylum="Chordata" rank="genus">
<emphasis id="B90E2A13DC383257F509FA66FC031491" box="[796,902,1412,1436]" italics="true" pageId="9" pageNumber="289">Stumpffia</emphasis>
</taxonomicName>
by strong mitochondrial differentiation (&gt;10% uncorrected pairwise distance in the 16S rRNA gene) and apparently exclusive haplotypes in the nuclear RAG1 gene (
<specimenCount id="9D7C3D88DC383257F6CEFA2EFEE014E8" box="[219,357,1484,1509]" pageId="9" pageNumber="289" type="generic">4 specimens</specimenCount>
sequenced for the new species).
</paragraph>
</subSubSection>
<subSubSection id="C360A58ADC383253F6D2FA0DFB85104D" lastPageId="13" lastPageNumber="293" pageId="9" pageNumber="289" type="description">
<paragraph id="8BC5F601DC383254F6D2FA0DFEA21068" blockId="9.[151,1436,151,2013]" lastBlockId="10.[151,1436,151,933]" lastPageId="10" lastPageNumber="290" pageId="9" pageNumber="289">
<emphasis id="B90E2A13DC383257F6D2FA0DFD941705" bold="true" box="[199,529,1519,1544]" pageId="9" pageNumber="289">
Description of the
<typeStatus id="54C148A3DC383257F7B6FA0DFD891705" box="[419,524,1519,1544]" pageId="9" pageNumber="289" type="holotype">holotype</typeStatus>
.
</emphasis>
Adult male specimen in good state of preservation (
<figureCitation id="1341EA84DC383257F24EFA0DFB1A1705" box="[1115,1183,1519,1544]" captionStart="FIGURE 3" captionStartId="6.[151,250,1945,1967]" captionTargetBox="[242,1344,445,1924]" captionTargetId="figure@6.[242,1345,443,1924]" captionTargetPageId="6" captionText="FIGURE 3. Holotype (ZSM 542 / 2009) of Stumpffia analamaina sp. nov. in preservative in (A) dorsal view, (B) foot and (C) hand in ventral view, and (D) living holotype in dorsolateral view. Scale bar = 1 mm." httpUri="https://zenodo.org/record/223539/files/figure.png" pageId="9" pageNumber="289">Fig. 3</figureCitation>
); tissue removed from ventral part of right thigh and venter cut open for gonad inspection. SVL
<quantity id="4C825BE4DC383257F5FAF9F6FBD21721" box="[1007,1111,1556,1580]" metricMagnitude="-2" metricUnit="m" metricValue="1.1099999999999999" pageId="9" pageNumber="289" unit="mm" value="11.1">11.1 mm</quantity>
; body slender; head slightly longer (
<quantity id="4C825BE4DC383257F6E5F9DAFECE175D" box="[240,331,1592,1616]" metricMagnitude="-3" metricUnit="m" metricValue="3.7" pageId="9" pageNumber="289" unit="mm" value="3.7">3.7 mm</quantity>
) than wide (
<quantity id="4C825BE4DC383257F7CBF9DAFDB9175D" box="[478,572,1592,1616]" metricMagnitude="-3" metricUnit="m" metricValue="3.3" pageId="9" pageNumber="289" unit="mm" value="3.3">3.3 mm</quantity>
), head less wide than the body; snout rounded to slightly pointed in dorsal view, protruding and slightly pointed in profile; nostrils directed laterally, slightly protuberant, about equidistant to the tip of the snout (
<quantity id="4C825BE4DC383257F79AF962FE691795" box="[399,492,1664,1688]" metricMagnitude="-4" metricUnit="m" metricValue="8.0" pageId="9" pageNumber="289" unit="mm" value="0.8">0.8 mm</quantity>
) and to the eye (
<quantity id="4C825BE4DC383257F4A8F962FC981795" box="[701,797,1664,1688]" metricMagnitude="-4" metricUnit="m" metricValue="8.0" pageId="9" pageNumber="289" unit="mm" value="0.8">0.8 mm</quantity>
); eye relatively large (
<quantity id="4C825BE4DC383257F23CF962FB011795" box="[1065,1156,1664,1688]" metricMagnitude="-3" metricUnit="m" metricValue="1.4" pageId="9" pageNumber="289" unit="mm" value="1.4">1.4 mm</quantity>
diameter), pupil round; internarial distance (
<quantity id="4C825BE4DC383257F79DF946FE6217B1" box="[392,487,1700,1724]" metricMagnitude="-3" metricUnit="m" metricValue="1.3" pageId="9" pageNumber="289" unit="mm" value="1.3">1.3 mm</quantity>
) distinctly smaller than interorbital distance (
<quantity id="4C825BE4DC383257F213F946FBED17B1" box="[1030,1128,1700,1724]" metricMagnitude="-3" metricUnit="m" metricValue="2.2" pageId="9" pageNumber="289" unit="mm" value="2.2">2.2 mm</quantity>
); canthus rostralis weakly marked and rounded; loreal region straight to slightly convex; tympanum not recognizable; supratympanic fold indistinct; tongue long and narrow, tip broadly rounded; maxillary teeth absent; vomerine teeth absent; choanae small and round. Arms slender; arm length from axilla to tip of longest finger
<quantity id="4C825BE4DC383257F21DF8EDFBE41625" box="[1032,1121,1807,1832]" metricMagnitude="-3" metricUnit="m" metricValue="5.6" pageId="9" pageNumber="289" unit="mm" value="5.6">5.6 mm</quantity>
; hand length to tip of finger III:
<quantity id="4C825BE4DC383257F6D0F8D6FEA61641" box="[197,291,1844,1869]" metricMagnitude="-3" metricUnit="m" metricValue="2.6" pageId="9" pageNumber="289" unit="mm" value="2.6">2.6 mm</quantity>
; finger II with one and finger III with two small, flat, round, subarticular tubercles; three weakly marked, flat palmar tubercles, of which the inner tubercle is the largest and round; fingers without webbing; relative length of fingers I&lt;IV&lt;II&lt;III, second toe slightly but distinctly longer than fourth toe; finger I reduced in length, all fingers relatively short; finger discs not enlarged, broadly rounded; prepollex absent; nuptial pads absent; tibiotarsal articulation reaches the nostril when hindlimb is adpressed along the body; hindlimbs slender; tibia length
<quantity id="4C825BE4DC3B3254F73EFF75FE0F11A2" box="[299,394,151,176]" metricMagnitude="-3" metricUnit="m" metricValue="5.8" pageId="10" pageNumber="290" unit="mm" value="5.8">5.8 mm</quantity>
, foot length
<quantity id="4C825BE4DC3B3254F439FF75FD0911A2" box="[556,652,151,176]" metricMagnitude="-3" metricUnit="m" metricValue="5.9" pageId="10" pageNumber="290" unit="mm" value="5.9">5.9 mm</quantity>
; lateral metatarsalia strongly connected; inner metatarsal tubercle indistinct, flat, ovoid; outer metatarsal tubercle absent; toe tips slightly broader than rest of toe without forming discs except on toe IV where a small disc is recognizable; toe tips with circummarginal grooves; distinct small, round subarticular tubercles; relative length of toes I&lt;II&lt;III=V&lt;IV, fifth toe same length as third toe, first toe strongly reduced in length but still clearly visible; no pedal webbing. Skin on dorsum and venter smooth in preservative.
</paragraph>
<paragraph id="8BC5F601DC3B3254F6D2FE92FDD313C0" blockId="10.[151,1436,151,933]" pageId="10" pageNumber="290">In life, the ground color of dorsum and dorsal sides of head and limbs was reddish brown, with an indistinct darker X-shaped marking of brownish color, of which the anterior branches reached above the eyes and the lower two branches faded at the level of the forelimb insertions. A dark patch was present in the inguinal region, and well-delimited although poorly contrasted brown stripes and patches were also present on the limbs: one crossband on each lower arm, a brown area around the knee, one broad crossband centrally on each thigh, shank, and tarsus. The flanks were densely covered with small silvery-whitish spots, without a sharp border between dorsal and lateral color. A number of brown patches were aligned along the flanks, and brown color was also present along the supratympanic fold and between eye and snout tip, including the nostril area. The iris was copper brown. After two years in preservative, the color remains largely similar, with a slightly stronger contrast between the brown dorsal pattern and the red-brown ground color.</paragraph>
<paragraph id="8BC5F601DC3B3254F6D2FD35FBB812A9" blockId="10.[151,1436,151,933]" pageId="10" pageNumber="290">
<emphasis id="B90E2A13DC3B3254F6D2FD35FEBA13FD" bold="true" box="[199,319,727,752]" pageId="10" pageNumber="290">Variation.</emphasis>
Numerous specimens of
<taxonomicName id="4C7A8D82DC3B3254F44DFD3AFD7F13FD" box="[600,762,727,752]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="10" pageNumber="290" phylum="Chordata" rank="species" species="analamaina">
<emphasis id="B90E2A13DC3B3254F44DFD3AFD7F13FD" box="[600,762,727,752]" italics="true" pageId="10" pageNumber="290">S. analamaina</emphasis>
</taxonomicName>
from the
<typeStatus id="54C148A3DC3B3254F570FD3AFC1013FD" box="[869,917,728,752]" pageId="10" pageNumber="290">type</typeStatus>
locality were observed alive but not preserved and therefore are not included in the
<typeStatus id="54C148A3DC3B3254F424FD1FFD141218" box="[561,657,765,789]" pageId="10" pageNumber="290" type="paratype">paratype</typeStatus>
series. In general, all examined
<typeStatus id="54C148A3DC3B3254F5ECFD1FFBE11218" box="[1017,1124,765,789]" pageId="10" pageNumber="290" type="paratype">paratypes</typeStatus>
agreed with the
<typeStatus id="54C148A3DC3B3254F308FD1EFAFA1218" box="[1309,1407,764,789]" pageId="10" pageNumber="290" type="holotype">holotype</typeStatus>
in morphology (
<figureCitation id="1341EA84DC3B3254F721FCFDFEFA1235" box="[308,383,799,824]" captionStart="FIGURE 4" captionStartId="7.[151,250,1852,1874]" captionTargetBox="[176,1425,260,1828]" captionTargetId="figure@7.[166,1426,165,1829]" captionTargetPageId="7" captionText="FIGURE 4. Specimens and habitat of Stumpffia analamaina sp. nov. (A) Paratype specimen ZSM 2831 / 2010 (ZCMV 10037) in life (scale bar = 1 mm). (B) Frame from a video showing prey capture of S. analamaina, showing partially lateral tongue protrusion as also known from other microhylids. The whitish structure on the right side of the head (arrow) is the tongue which is protruded at an angle of 40 50 ° relative to the body axis. (C) Dry forest and (D) stream at type locality, specimens were found in the leaf litter at the border of the stream. (E) Foam nest produced by S. analamaina in captivity. (F) Additional specimen (catalogue number uncertain) showing color variation (beige ground color)." httpUri="https://zenodo.org/record/223540/files/figure.png" pageId="10" pageNumber="290">Fig. 4</figureCitation>
; see
<tableCitation id="C6F8C3BADC3B3254F7D4FCFDFD9C1235" box="[449,537,799,824]" captionStart="TABLE 1" captionStartId="10.[151,239,980,1003]" captionTargetBox="[159,1403,1249,1642]" captionTargetPageId="10" captionText="TABLE 1. Morphometric data (in mm) of selected specimens of S. gimmeli, S. madagascariensis, S. pygmaea, S. sp. 31, and S. analamaina sp. nov. SVL, snout-vent length; HAL, hand length; FOTL, foot and tarsus length; FOL, foot length; TIBL, tibia length; HT, holotype; PT, paratype." httpUri="http://table.plazi.org/id/DF05A689DC3B3254F682FC36FD0B1526" pageId="10" pageNumber="290" tableUuid="DF05A689DC3B3254F682FC36FD0B1526">Table 1</tableCitation>
for selected measurements of
<typeStatus id="54C148A3DC3B3254F59CFCC2FC7F1235" box="[905,1018,800,824]" pageId="10" pageNumber="290" type="paratype">paratypes</typeStatus>
). While the red-brown dorsal color characterized several specimens (
<figureCitation id="1341EA84DC3B3254F403FCA6FDD91251" box="[534,604,836,861]" captionStart="FIGURE 4" captionStartId="7.[151,250,1852,1874]" captionTargetBox="[176,1425,260,1828]" captionTargetId="figure@7.[166,1426,165,1829]" captionTargetPageId="7" captionText="FIGURE 4. Specimens and habitat of Stumpffia analamaina sp. nov. (A) Paratype specimen ZSM 2831 / 2010 (ZCMV 10037) in life (scale bar = 1 mm). (B) Frame from a video showing prey capture of S. analamaina, showing partially lateral tongue protrusion as also known from other microhylids. The whitish structure on the right side of the head (arrow) is the tongue which is protruded at an angle of 40 50 ° relative to the body axis. (C) Dry forest and (D) stream at type locality, specimens were found in the leaf litter at the border of the stream. (E) Foam nest produced by S. analamaina in captivity. (F) Additional specimen (catalogue number uncertain) showing color variation (beige ground color)." httpUri="https://zenodo.org/record/223540/files/figure.png" pageId="10" pageNumber="290">Fig. 4</figureCitation>
A), others had a more beige ground color (
<figureCitation id="1341EA84DC3B3254F245FCA6FB131251" box="[1104,1174,836,861]" captionStart="FIGURE 4" captionStartId="7.[151,250,1852,1874]" captionTargetBox="[176,1425,260,1828]" captionTargetId="figure@7.[166,1426,165,1829]" captionTargetPageId="7" captionText="FIGURE 4. Specimens and habitat of Stumpffia analamaina sp. nov. (A) Paratype specimen ZSM 2831 / 2010 (ZCMV 10037) in life (scale bar = 1 mm). (B) Frame from a video showing prey capture of S. analamaina, showing partially lateral tongue protrusion as also known from other microhylids. The whitish structure on the right side of the head (arrow) is the tongue which is protruded at an angle of 40 50 ° relative to the body axis. (C) Dry forest and (D) stream at type locality, specimens were found in the leaf litter at the border of the stream. (E) Foam nest produced by S. analamaina in captivity. (F) Additional specimen (catalogue number uncertain) showing color variation (beige ground color)." httpUri="https://zenodo.org/record/223540/files/figure.png" pageId="10" pageNumber="290">Fig. 4</figureCitation>
F). The dark dorsal Xshaped pattern in several specimens was extended to form an indistinct &quot;gummi-bear&quot; marking as is typical for other
<taxonomicName id="4C7A8D82DC3B3254F6C2FC6EFEC412A9" box="[215,321,908,932]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="10" pageNumber="290" phylum="Chordata" rank="genus">
<emphasis id="B90E2A13DC3B3254F6C2FC6EFEC412A9" box="[215,321,908,932]" italics="true" pageId="10" pageNumber="290">Stumpffia</emphasis>
</taxonomicName>
species, especially
<taxonomicName id="4C7A8D82DC3B3254F40BFC6FFD3A12A8" box="[542,703,908,933]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="10" pageNumber="290" phylum="Chordata" rank="species" species="psologlossa">
<emphasis id="B90E2A13DC3B3254F40BFC6FFD3A12A8" box="[542,703,908,933]" italics="true" pageId="10" pageNumber="290">S. psologlossa</emphasis>
</taxonomicName>
(photos in Glaw &amp; Vences 2007).
</paragraph>
<caption id="DF05A689DC3B3254F682FC36FD0B1526" ID-Table-UUID="DF05A689DC3B3254F682FC36FD0B1526" httpUri="http://table.plazi.org/id/DF05A689DC3B3254F682FC36FD0B1526" pageId="10" pageNumber="290" targetBox="[159,1403,1249,1642]" targetIsTable="true" targetPageId="10">
<paragraph id="8BC5F601DC3B3254F682FC36FD0B1526" blockId="10.[151,1436,979,1067]" pageId="10" pageNumber="290">
<emphasis id="B90E2A13DC3B3254F682FC36FE8C12E6" bold="true" box="[151,265,980,1003]" pageId="10" pageNumber="290">TABLE 1.</emphasis>
Morphometric data (in mm) of selected specimens of
<taxonomicName id="4C7A8D82DC3B3254F558FC31FC3812E6" box="[845,957,979,1003]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="10" pageNumber="290" phylum="Chordata" rank="species" species="gimmeli">
<emphasis id="B90E2A13DC3B3254F558FC31FC3812E6" box="[845,957,979,1003]" italics="true" pageId="10" pageNumber="290">S. gimmeli</emphasis>
</taxonomicName>
,
<emphasis id="B90E2A13DC3B3254F5DEFC31FACA12E7" box="[971,1359,979,1003]" italics="true" pageId="10" pageNumber="290">
<taxonomicName id="4C7A8D82DC3B3254F5DEFC31FB2D12E6" box="[971,1192,979,1003]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="10" pageNumber="290" phylum="Chordata" rank="species" species="madagascariensis">S. madagascariensis</taxonomicName>
,
<taxonomicName id="4C7A8D82DC3B3254F2A6FC31FAB512E6" box="[1203,1328,979,1003]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="10" pageNumber="290" phylum="Chordata" rank="species" species="pygmaea">S. pygmaea</taxonomicName>
, S.
</emphasis>
sp. 31, and
<taxonomicName id="4C7A8D82DC3B3254F6D1FC11FEDA1506" box="[196,351,1011,1035]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="10" pageNumber="290" phylum="Chordata" rank="species" species="analamaina" status="sp. nov.">
<emphasis id="B90E2A13DC3B3254F6D1FC11FEDA1506" box="[196,351,1011,1035]" italics="true" pageId="10" pageNumber="290">S. analamaina</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D9768DC3B3254F773FC16FE3C1506" box="[358,441,1012,1035]" pageId="10" pageNumber="290" rank="species">sp. nov.</taxonomicNameLabel>
SVL, snout-vent length; HAL, hand length; FOTL, foot and tarsus length; FOL, foot length; TIBL, tibia length; HT, holotype; PT, paratype.
</paragraph>
</caption>
<paragraph id="8BC5F601DC3B3254F68AFBA4FAEE15C3" pageId="10" pageNumber="290">
<table id="F97A04A1DC3BCDA1F68AFBA4FAFE15C3" box="[159,1403,1094,1230]" gridcols="9" gridrows="3" pageId="10" pageNumber="290">
<tr id="354AF443DC3BCDA1F68AFBA4FAFE1576" box="[159,1403,1094,1147]" gridrow="0" pageId="10" pageNumber="290">
<th id="769B9D3FDC3BCDA1F68AFBA4FD9A1576" box="[159,543,1094,1147]" gridcol="0" gridrow="0" pageId="10" pageNumber="290">Species Number</th>
<th id="769B9D3FDC3BCDA1F460FBA4FD781576" box="[629,765,1094,1147]" gridcol="1" gridrow="0" pageId="10" pageNumber="290">Field number</th>
<th id="769B9D3FDC3BCDA1F50FFBA4FCCC1576" box="[794,841,1094,1147]" gridcol="2" gridrow="0" pageId="10" pageNumber="290">SVL</th>
<th id="769B9D3FDC3BCDA1F577FBA4FC131576" box="[866,918,1094,1147]" gridcol="3" gridrow="0" pageId="10" pageNumber="290">HAL</th>
<th id="769B9D3FDC3BCDA1F5A8FBA4FC791576" box="[957,1020,1094,1147]" gridcol="4" gridrow="0" pageId="10" pageNumber="290">FOTL</th>
<th id="769B9D3FDC3BCDA1F20DFBA4FBCD1576" box="[1048,1096,1094,1147]" gridcol="5" gridrow="0" pageId="10" pageNumber="290">FOL</th>
<th id="769B9D3FDC3BCDA1F266FBA4FB291576" box="[1139,1196,1094,1147]" gridcol="6" gridrow="0" pageId="10" pageNumber="290">TIBL</th>
<th id="769B9D3FDC3BCDA1F2DBFBA4FA911576" box="[1230,1300,1094,1147]" gridcol="7" gridrow="0" pageId="10" pageNumber="290">FOTL/ SVL</th>
<th id="769B9D3FDC3BCDA1F32AFBA4FAFE1576" box="[1343,1403,1094,1147]" gridcol="8" gridrow="0" pageId="10" pageNumber="290">HAL/ SVL</th>
</tr>
<tr id="354AF443DC3BCDA1F68AFB6CFAFE15A9" box="[159,1403,1166,1188]" gridrow="1" pageId="10" pageNumber="290">
<th id="769B9D3FDC3BCDA1F68AFB6CFD9A15A9" box="[159,543,1166,1188]" gridcol="0" gridrow="1" pageId="10" pageNumber="290">
<taxonomicName id="4C7A8D82DC3B3254F68AFB6DFE8F15A9" box="[159,266,1166,1188]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="10" pageNumber="290" phylum="Chordata" rank="species" species="gimmeli">
<emphasis id="B90E2A13DC3B3254F68AFB6DFE8F15A9" box="[159,266,1166,1188]" italics="true" pageId="10" pageNumber="290">S. gimmeli</emphasis>
</taxonomicName>
ZSM 412/2000
</th>
<td id="769B9D3FDC3BCDA1F460FB6CFD7815A9" box="[629,765,1166,1188]" gridcol="1" gridrow="1" pageId="10" pageNumber="290">-</td>
<td id="769B9D3FDC3BCDA1F50FFB6CFCCC15A9" box="[794,841,1166,1188]" gridcol="2" gridrow="1" pageId="10" pageNumber="290">18.2</td>
<td id="769B9D3FDC3BCDA1F577FB6CFC1315A9" box="[866,918,1166,1188]" gridcol="3" gridrow="1" pageId="10" pageNumber="290">4.7</td>
<td id="769B9D3FDC3BCDA1F5A8FB6CFC7915A9" box="[957,1020,1166,1188]" gridcol="4" gridrow="1" pageId="10" pageNumber="290">13.0</td>
<td id="769B9D3FDC3BCDA1F20DFB6CFBCD15A9" box="[1048,1096,1166,1188]" gridcol="5" gridrow="1" pageId="10" pageNumber="290">8.3</td>
<td id="769B9D3FDC3BCDA1F266FB6CFB2915A9" box="[1139,1196,1166,1188]" gridcol="6" gridrow="1" pageId="10" pageNumber="290">8.9</td>
<td id="769B9D3FDC3BCDA1F2DBFB6CFA9115A9" box="[1230,1300,1166,1188]" gridcol="7" gridrow="1" pageId="10" pageNumber="290">0.71</td>
<td id="769B9D3FDC3BCDA1F32AFB6CFAFE15A9" box="[1343,1403,1166,1188]" gridcol="8" gridrow="1" pageId="10" pageNumber="290">0.26</td>
</tr>
<tr id="354AF443DC3BCDA1F68AFB5AFAFE15C3" box="[159,1403,1208,1230]" gridrow="2" pageId="10" pageNumber="290">
<th id="769B9D3FDC3BCDA1F68AFB5AFD9A15C3" box="[159,543,1208,1230]" gridcol="0" gridrow="2" pageId="10" pageNumber="290">
<taxonomicName id="4C7A8D82DC3B3254F68AFB5BFEEA15C3" box="[159,367,1208,1230]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="10" pageNumber="290" phylum="Chordata" rank="species" species="madagascariensis">
<emphasis id="B90E2A13DC3B3254F68AFB5BFEEA15C3" box="[159,367,1208,1230]" italics="true" pageId="10" pageNumber="290">S. madagascariensis</emphasis>
</taxonomicName>
ZSM 201/2004
</th>
<td id="769B9D3FDC3BCDA1F460FB5AFD7815C3" box="[629,765,1208,1230]" gridcol="1" gridrow="2" pageId="10" pageNumber="290">FGZC 389</td>
<td id="769B9D3FDC3BCDA1F50FFB5AFCCC15C3" box="[794,841,1208,1230]" gridcol="2" gridrow="2" pageId="10" pageNumber="290">11.5</td>
<td id="769B9D3FDC3BCDA1F577FB5AFC1315C3" box="[866,918,1208,1230]" gridcol="3" gridrow="2" pageId="10" pageNumber="290">1.8</td>
<td id="769B9D3FDC3BCDA1F5A8FB5AFC7915C3" box="[957,1020,1208,1230]" gridcol="4" gridrow="2" pageId="10" pageNumber="290">7.0</td>
<td id="769B9D3FDC3BCDA1F20DFB5AFBCD15C3" box="[1048,1096,1208,1230]" gridcol="5" gridrow="2" pageId="10" pageNumber="290">4.4</td>
<td id="769B9D3FDC3BCDA1F266FB5AFB2915C3" box="[1139,1196,1208,1230]" gridcol="6" gridrow="2" pageId="10" pageNumber="290">4.7</td>
<td id="769B9D3FDC3BCDA1F2DBFB5AFA9115C3" box="[1230,1300,1208,1230]" gridcol="7" gridrow="2" pageId="10" pageNumber="290">0.61</td>
<td id="769B9D3FDC3BCDA1F32AFB5AFAFE15C3" box="[1343,1403,1208,1230]" gridcol="8" gridrow="2" pageId="10" pageNumber="290">0.16</td>
</tr>
</table>
</paragraph>
<paragraph id="8BC5F601DC3B3254F68AFB00FAEE1767" pageId="10" pageNumber="290">
<table id="F97A04A1DC3BCDA1F68AFB03FAFE1767" box="[159,1403,1249,1642]" gridcols="8" gridrows="7" pageId="10" pageNumber="290">
<tr id="354AF443DC3BCDA1F68AFB03FAFE15FA" box="[159,1403,1249,1271]" gridrow="0" pageId="10" pageNumber="290">
<th id="769B9D3FDC3BCDA1F68AFB03FEB415FA" box="[159,305,1249,1271]" gridcol="0" gridrow="0" pageId="10" pageNumber="290">
<taxonomicName id="4C7A8D82DC3B3254F68AFB00FE9615FA" box="[159,275,1250,1271]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="10" pageNumber="290" phylum="Chordata" rank="species" species="pygmaea">
<emphasis id="B90E2A13DC3B3254F68AFB00FE9615FA" box="[159,275,1250,1271]" italics="true" pageId="10" pageNumber="290">S. pygmaea</emphasis>
</taxonomicName>
</th>
<th id="769B9D3FDC3BCDA1F790FB03FDA915FA" box="[389,556,1249,1271]" gridcol="1" gridrow="0" pageId="10" pageNumber="290">ZSM 557/1999</th>
<th id="769B9D3FDC3BCDA1F460FB03FC0715FA" box="[629,898,1249,1271]" gridcol="2" gridrow="0" pageId="10" pageNumber="290">ZFMK 52543 11.2 1.9</th>
<th id="769B9D3FDC3BCDA1F5A8FB03FC5815FA" box="[957,989,1249,1271]" gridcol="3" gridrow="0" pageId="10" pageNumber="290">6.7</th>
<th id="769B9D3FDC3BCDA1F20DFB03FBBD15FA" box="[1048,1080,1249,1271]" gridcol="4" gridrow="0" pageId="10" pageNumber="290">4.3</th>
<th id="769B9D3FDC3BCDA1F266FB03FB1615FA" box="[1139,1171,1249,1271]" gridcol="5" gridrow="0" pageId="10" pageNumber="290">4.3</th>
<th id="769B9D3FDC3BCDA1F2DBFB03FB7F15FA" box="[1230,1274,1249,1271]" gridcol="6" gridrow="0" pageId="10" pageNumber="290">0.60</th>
<th id="769B9D3FDC3BCDA1F32AFB03FAFE15FA" box="[1343,1403,1249,1271]" gridcol="7" gridrow="0" pageId="10" pageNumber="290">0.17</th>
</tr>
<tr id="354AF443DC3BCDA1F68AFAE8FAFE1432" box="[159,1403,1290,1343]" gridrow="1" pageId="10" pageNumber="290">
<th id="769B9D3FDC3BCDA1F68AFAE8FEB41432" box="[159,305,1290,1343]" gridcol="0" gridrow="1" pageId="10" pageNumber="290">
<taxonomicName id="4C7A8D82DC3B3254F68AFAE9FEB4142D" box="[159,305,1290,1312]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="10" pageNumber="290" phylum="Chordata" rank="species" species="analamaina" status="sp. nov.">
<emphasis id="B90E2A13DC3B3254F68AFAE9FEB4142D" box="[159,305,1290,1312]" italics="true" pageId="10" pageNumber="290">S. analamaina</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D9768DC3B3254F68AFACBFF6A1433" box="[159,239,1321,1342]" pageId="10" pageNumber="290" rank="species">
<emphasis id="B90E2A13DC3B3254F68AFACBFF6A1433" bold="true" box="[159,239,1321,1342]" pageId="10" pageNumber="290">sp. nov.</emphasis>
</taxonomicNameLabel>
(PT)
</th>
<td id="769B9D3FDC3BCDA1F790FAE8FDA91432" box="[389,556,1290,1343]" gridcol="1" gridrow="1" pageId="10" pageNumber="290">ZSM 2829/2010</td>
<td id="769B9D3FDC3BCDA1F460FAE8FC071432" box="[629,898,1290,1343]" gridcol="2" gridrow="1" pageId="10" pageNumber="290">ZCMV 10034 9.6 2.3</td>
<td id="769B9D3FDC3BCDA1F5A8FAE8FC581432" box="[957,989,1290,1343]" gridcol="3" gridrow="1" pageId="10" pageNumber="290">7.5</td>
<td id="769B9D3FDC3BCDA1F20DFAE8FBBD1432" box="[1048,1080,1290,1343]" gridcol="4" gridrow="1" pageId="10" pageNumber="290">5.1</td>
<td id="769B9D3FDC3BCDA1F266FAE8FB161432" box="[1139,1171,1290,1343]" gridcol="5" gridrow="1" pageId="10" pageNumber="290">5.4</td>
<td id="769B9D3FDC3BCDA1F2DBFAE8FB7F1432" box="[1230,1274,1290,1343]" gridcol="6" gridrow="1" pageId="10" pageNumber="290">0.78</td>
<td id="769B9D3FDC3BCDA1F32AFAE8FAFE1432" box="[1343,1403,1290,1343]" gridcol="7" gridrow="1" pageId="10" pageNumber="290">0.24</td>
</tr>
<tr id="354AF443DC3BCDA1F68AFAB0FAFE148A" box="[159,1403,1362,1415]" gridrow="2" pageId="10" pageNumber="290">
<th id="769B9D3FDC3BCDA1F68AFAB0FEB4148A" box="[159,305,1362,1415]" gridcol="0" gridrow="2" pageId="10" pageNumber="290">
<taxonomicName id="4C7A8D82DC3B3254F68AFAB1FEB41465" box="[159,305,1362,1384]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="10" pageNumber="290" phylum="Chordata" rank="species" species="analamaina" status="sp. nov.">
<emphasis id="B90E2A13DC3B3254F68AFAB1FEB41465" box="[159,305,1362,1384]" italics="true" pageId="10" pageNumber="290">S. analamaina</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D9768DC3B3254F68AFA93FF6A148B" box="[159,239,1393,1414]" pageId="10" pageNumber="290" rank="species">
<emphasis id="B90E2A13DC3B3254F68AFA93FF6A148B" bold="true" box="[159,239,1393,1414]" pageId="10" pageNumber="290">sp. nov.</emphasis>
</taxonomicNameLabel>
(PT)
</th>
<td id="769B9D3FDC3BCDA1F790FAB0FDA9148A" box="[389,556,1362,1415]" gridcol="1" gridrow="2" pageId="10" pageNumber="290">ZSM 2830/2010</td>
<td id="769B9D3FDC3BCDA1F460FAB0FC07148A" box="[629,898,1362,1415]" gridcol="2" gridrow="2" pageId="10" pageNumber="290">ZCMV 10036 10.5 2.3</td>
<td id="769B9D3FDC3BCDA1F5A8FAB0FC58148A" box="[957,989,1362,1415]" gridcol="3" gridrow="2" pageId="10" pageNumber="290">7.1</td>
<td id="769B9D3FDC3BCDA1F20DFAB0FBBD148A" box="[1048,1080,1362,1415]" gridcol="4" gridrow="2" pageId="10" pageNumber="290">4.9</td>
<td id="769B9D3FDC3BCDA1F266FAB0FB16148A" box="[1139,1171,1362,1415]" gridcol="5" gridrow="2" pageId="10" pageNumber="290">4.6</td>
<td id="769B9D3FDC3BCDA1F2DBFAB0FB7F148A" box="[1230,1274,1362,1415]" gridcol="6" gridrow="2" pageId="10" pageNumber="290">0.68</td>
<td id="769B9D3FDC3BCDA1F32AFAB0FAFE148A" box="[1343,1403,1362,1415]" gridcol="7" gridrow="2" pageId="10" pageNumber="290">0.22</td>
</tr>
<tr id="354AF443DC3BCDA1F68AFA78FAFE14C2" box="[159,1403,1434,1487]" gridrow="3" pageId="10" pageNumber="290">
<th id="769B9D3FDC3BCDA1F68AFA78FEB414C2" box="[159,305,1434,1487]" gridcol="0" gridrow="3" pageId="10" pageNumber="290">
<taxonomicName id="4C7A8D82DC3B3254F68AFA79FEB414BD" box="[159,305,1434,1456]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="10" pageNumber="290" phylum="Chordata" rank="species" species="analamaina" status="sp. nov.">
<emphasis id="B90E2A13DC3B3254F68AFA79FEB414BD" box="[159,305,1434,1456]" italics="true" pageId="10" pageNumber="290">S. analamaina</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D9768DC3B3254F68AFA5BFF6A14C3" box="[159,239,1465,1486]" pageId="10" pageNumber="290" rank="species">
<emphasis id="B90E2A13DC3B3254F68AFA5BFF6A14C3" bold="true" box="[159,239,1465,1486]" pageId="10" pageNumber="290">sp. nov.</emphasis>
</taxonomicNameLabel>
(PT)
</th>
<td id="769B9D3FDC3BCDA1F790FA78FDA914C2" box="[389,556,1434,1487]" gridcol="1" gridrow="3" pageId="10" pageNumber="290">ZSM 2831/2010</td>
<td id="769B9D3FDC3BCDA1F460FA78FC0714C2" box="[629,898,1434,1487]" gridcol="2" gridrow="3" pageId="10" pageNumber="290">ZCMV 10037 10.8 2.8</td>
<td id="769B9D3FDC3BCDA1F5A8FA78FC5814C2" box="[957,989,1434,1487]" gridcol="3" gridrow="3" pageId="10" pageNumber="290">8.0</td>
<td id="769B9D3FDC3BCDA1F20DFA78FBBD14C2" box="[1048,1080,1434,1487]" gridcol="4" gridrow="3" pageId="10" pageNumber="290">5.3</td>
<td id="769B9D3FDC3BCDA1F266FA78FB1614C2" box="[1139,1171,1434,1487]" gridcol="5" gridrow="3" pageId="10" pageNumber="290">5.2</td>
<td id="769B9D3FDC3BCDA1F2DBFA78FB7F14C2" box="[1230,1274,1434,1487]" gridcol="6" gridrow="3" pageId="10" pageNumber="290">0.74</td>
<td id="769B9D3FDC3BCDA1F32AFA78FAFE14C2" box="[1343,1403,1434,1487]" gridcol="7" gridrow="3" pageId="10" pageNumber="290">0.26</td>
</tr>
<tr id="354AF443DC3BCDA1F68AFA00FAFE171A" box="[159,1403,1506,1559]" gridrow="4" pageId="10" pageNumber="290">
<th id="769B9D3FDC3BCDA1F68AFA00FEB4171A" box="[159,305,1506,1559]" gridcol="0" gridrow="4" pageId="10" pageNumber="290">
<taxonomicName id="4C7A8D82DC3B3254F68AFA01FEB414F5" box="[159,305,1506,1528]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="10" pageNumber="290" phylum="Chordata" rank="species" species="analamaina" status="sp. nov.">
<emphasis id="B90E2A13DC3B3254F68AFA01FEB414F5" box="[159,305,1506,1528]" italics="true" pageId="10" pageNumber="290">S. analamaina</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D9768DC3B3254F68AF9E3FF6A171B" box="[159,239,1537,1558]" pageId="10" pageNumber="290" rank="species">
<emphasis id="B90E2A13DC3B3254F68AF9E3FF6A171B" bold="true" box="[159,239,1537,1558]" pageId="10" pageNumber="290">sp. nov.</emphasis>
</taxonomicNameLabel>
(HT)
</th>
<td id="769B9D3FDC3BCDA1F790FA00FDA9171A" box="[389,556,1506,1559]" gridcol="1" gridrow="4" pageId="10" pageNumber="290">ZSM 542/2009</td>
<td id="769B9D3FDC3BCDA1F460FA00FC07171A" box="[629,898,1506,1559]" gridcol="2" gridrow="4" pageId="10" pageNumber="290">ZCMV 11428 11.1 2.6</td>
<td id="769B9D3FDC3BCDA1F5A8FA00FC58171A" box="[957,989,1506,1559]" gridcol="3" gridrow="4" pageId="10" pageNumber="290">9.0</td>
<td id="769B9D3FDC3BCDA1F20DFA00FBBD171A" box="[1048,1080,1506,1559]" gridcol="4" gridrow="4" pageId="10" pageNumber="290">5.9</td>
<td id="769B9D3FDC3BCDA1F266FA00FB16171A" box="[1139,1171,1506,1559]" gridcol="5" gridrow="4" pageId="10" pageNumber="290">5.8</td>
<td id="769B9D3FDC3BCDA1F2DBFA00FB7F171A" box="[1230,1274,1506,1559]" gridcol="6" gridrow="4" pageId="10" pageNumber="290">0.81</td>
<td id="769B9D3FDC3BCDA1F32AFA00FAFE171A" box="[1343,1403,1506,1559]" gridcol="7" gridrow="4" pageId="10" pageNumber="290">0.23</td>
</tr>
<tr id="354AF443DC3BCDA1F68AF9C8FAFE174D" box="[159,1403,1578,1600]" gridrow="5" pageId="10" pageNumber="290">
<th id="769B9D3FDC3BCDA1F68AF9C8FEB4174D" box="[159,305,1578,1600]" gridcol="0" gridrow="5" pageId="10" pageNumber="290">
<emphasis id="B90E2A13DC3B3254F68AF9C9FF37174D" box="[159,178,1579,1600]" italics="true" pageId="10" pageNumber="290">S.</emphasis>
sp. 31
</th>
<td id="769B9D3FDC3BCDA1F790F9C8FDA9174D" box="[389,556,1578,1600]" gridcol="1" gridrow="5" pageId="10" pageNumber="290">ZSM 1825/2010</td>
<td id="769B9D3FDC3BCDA1F460F9C8FC07174D" box="[629,898,1578,1600]" gridcol="2" gridrow="5" pageId="10" pageNumber="290">ZCMV 12600 10.0 2.1</td>
<td id="769B9D3FDC3BCDA1F5A8F9C8FC58174D" box="[957,989,1578,1600]" gridcol="3" gridrow="5" pageId="10" pageNumber="290">6.9</td>
<td id="769B9D3FDC3BCDA1F20DF9C8FBBD174D" box="[1048,1080,1578,1600]" gridcol="4" gridrow="5" pageId="10" pageNumber="290">4.3</td>
<td id="769B9D3FDC3BCDA1F266F9C8FB16174D" box="[1139,1171,1578,1600]" gridcol="5" gridrow="5" pageId="10" pageNumber="290">4.3</td>
<td id="769B9D3FDC3BCDA1F2DBF9C8FB7F174D" box="[1230,1274,1578,1600]" gridcol="6" gridrow="5" pageId="10" pageNumber="290">0.69</td>
<td id="769B9D3FDC3BCDA1F32AF9C8FAFE174D" box="[1343,1403,1578,1600]" gridcol="7" gridrow="5" pageId="10" pageNumber="290">0.21</td>
</tr>
<tr id="354AF443DC3BCDA1F68AF9B6FAFE1767" box="[159,1403,1620,1642]" gridrow="6" pageId="10" pageNumber="290">
<th id="769B9D3FDC3BCDA1F68AF9B6FEB41767" box="[159,305,1620,1642]" gridcol="0" gridrow="6" pageId="10" pageNumber="290">
<emphasis id="B90E2A13DC3B3254F68AF9B6FF371764" box="[159,178,1620,1641]" italics="true" pageId="10" pageNumber="290">S.</emphasis>
sp. 31
</th>
<td id="769B9D3FDC3BCDA1F790F9B6FDA91767" box="[389,556,1620,1642]" gridcol="1" gridrow="6" pageId="10" pageNumber="290">uncatalogued</td>
<td id="769B9D3FDC3BCDA1F460F9B6FC071767" box="[629,898,1620,1642]" gridcol="2" gridrow="6" pageId="10" pageNumber="290">DRV 6413 10.4 2.1</td>
<td id="769B9D3FDC3BCDA1F5A8F9B6FC581767" box="[957,989,1620,1642]" gridcol="3" gridrow="6" pageId="10" pageNumber="290">7.1</td>
<td id="769B9D3FDC3BCDA1F20DF9B6FBBD1767" box="[1048,1080,1620,1642]" gridcol="4" gridrow="6" pageId="10" pageNumber="290">4.6</td>
<td id="769B9D3FDC3BCDA1F266F9B6FB161767" box="[1139,1171,1620,1642]" gridcol="5" gridrow="6" pageId="10" pageNumber="290">4.7</td>
<td id="769B9D3FDC3BCDA1F2DBF9B6FB7F1767" box="[1230,1274,1620,1642]" gridcol="6" gridrow="6" pageId="10" pageNumber="290">0.68</td>
<td id="769B9D3FDC3BCDA1F32AF9B6FAFE1767" box="[1343,1403,1620,1642]" gridcol="7" gridrow="6" pageId="10" pageNumber="290">0.20</td>
</tr>
</table>
</paragraph>
<paragraph id="8BC5F601DC3B3254F6D2F94CFE3B16EB" blockId="10.[151,1436,1710,2023]" pageId="10" pageNumber="290">
<emphasis id="B90E2A13DC3B3254F6D2F94CFEC617CA" bold="true" box="[199,323,1710,1735]" pageId="10" pageNumber="290">Osteology.</emphasis>
Osteological features are described on the basis of CT scans of specimen ZCMV 10034 (
<figureCitation id="1341EA84DC3B3254F338F94CFA1517CA" box="[1325,1424,1710,1735]" captionStart-0="FIGURE 5" captionStart-1="FIGURE 6" captionStart-2="FIGURE 7" captionStartId-0="8.[151,250,1647,1669]" captionStartId-1="11.[151,250,1567,1589]" captionStartId-2="12.[151,250,1923,1945]" captionTargetBox-0="[151,1436,193,1624]" captionTargetBox-1="[151,1436,183,1546]" captionTargetBox-2="[153,1436,393,1900]" captionTargetId-0="figure@8.[151,1436,193,1626]" captionTargetId-1="figure@11.[151,1436,183,1546]" captionTargetId-2="figure@12.[151,1436,393,1901]" captionTargetPageId-0="8" captionTargetPageId-1="11" captionTargetPageId-2="12" captionText-0="FIGURE 5. Skull in oblique right lateral view (above) and ventral view (below), and presacral vertebral column in dorsal view (right) of Stumpffia analamaina sp. nov., paratype specimen ZSM 2831 / 2010 (ZCMV 10037). Abbreviations: Den, dentary; FP, frontoparietals; Max, maxilla; Na, nasals; Npl—neopalatinum; ParSph, parasphenoid; PrMax, premaxilla; Pt—pterygoid; Qj—quadratojugale; Sac, sacrum; Sq, squamosal (Sq-AP and Sq-PP—anterior and posterior process of squamosal); Ur, urostyle (incompletely shown); V—Vomer; V 1 - V 8, presacral vertebrae (V 1 + V 2 includes atlas and axis)." captionText-1="FIGURE 6. Ventral views of left hands of different species of Stumpffia of the north / northwestern clade and S. tridactyla and S. tetradactyla, showing various degrees of external reduction of digits on the right, and CT scans of the hands of the same individuals on the left. External reduction of fingers is only partly paralleled by reduction of the skeletal elements (especially in S. tridactyla; see text). Scale bar = 0.5 mm. Asterisks mark right hands in mirrored view. Formulae show the number of phalanges on fingers I IV." captionText-2="FIGURE 7. Ventral views of feet of different species of Stumpffia of the north / northwestern clade and S. tridactyla and S. tetradactyla, showing various degrees of external reduction of digits on the right, and CT scans of the feet of the same individuals on the left. External reduction of toes is only partly paralleled by reduction of the skeletal elements (especially in S. tridactyla; see text). Scale bar = 1 mm. Formulae show the number of phalanges on toes I V." httpUri-0="https://zenodo.org/record/223541/files/figure.png" httpUri-1="https://zenodo.org/record/223542/files/figure.png" httpUri-2="https://zenodo.org/record/223543/files/figure.png" pageId="10" pageNumber="290">Fig. 57</figureCitation>
). Vertebral column largely in agreement with description of Guibé (1978) for three other species of
<taxonomicName id="4C7A8D82DC3B3254F2F6F931FAC817E6" box="[1251,1357,1747,1771]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="10" pageNumber="290" phylum="Chordata" rank="genus">
<emphasis id="B90E2A13DC3B3254F2F6F931FAC817E6" box="[1251,1357,1747,1771]" italics="true" pageId="10" pageNumber="290">Stumpffia</emphasis>
</taxonomicName>
; seven presacral vertebrae (first and second incompletely fused), wider than long, with relatively long diapophyses on the three anterior vertebrae and shorter on the four posterior ones. Sacrum with relatively broad diapophyses. No traces of teeth on premaxilla or maxilla. Nasals not in contact. Squamosal with a small zygomatic (anterior) process (much shorter than otic process). Coracoid well ossified, but clavicula only ossified as a thin rod; sternum and omosternum not visible in the CT scan, hence not calcified. Terminal phalanges of fingers and toes with a weakly expressed T-shaped expansion. Three free distal tarsals recognizable. Phalangeal formula 123
<quantity id="4C825BE4DC3B3254F2E5F849FA9B16CE" box="[1264,1310,1962,1987]" metricMagnitude="-2" metricUnit="m" metricValue="5.08" pageId="10" pageNumber="290" unit="in" value="2.0">2 in</quantity>
the manus and 1234
<quantity id="4C825BE4DC3B3254F72DF82DFEE416EA" box="[312,353,1998,2023]" metricMagnitude="-2" metricUnit="m" metricValue="7.62" pageId="10" pageNumber="290" unit="in" value="3.0">3 in</quantity>
the pes.
</paragraph>
<caption id="DF05A689DC3A3255F682F9FDFE2617BD" httpUri="https://zenodo.org/record/223542/files/figure.png" pageId="11" pageNumber="291" targetBox="[151,1436,183,1546]" targetPageId="11">
<paragraph id="8BC5F601DC3A3255F682F9FDFE2617BD" blockId="11.[151,1436,1567,1712]" pageId="11" pageNumber="291">
<emphasis id="B90E2A13DC3A3255F682F9FDFE971739" bold="true" box="[151,274,1567,1589]" pageId="11" pageNumber="291">FIGURE 6.</emphasis>
Ventral views of left hands of different species of
<taxonomicName id="4C7A8D82DC3A3255F519F9FDFCE91738" box="[780,876,1567,1589]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="11" pageNumber="291" phylum="Chordata" rank="genus">
<emphasis id="B90E2A13DC3A3255F519F9FDFCE91738" box="[780,876,1567,1589]" italics="true" pageId="11" pageNumber="291">Stumpffia</emphasis>
</taxonomicName>
of the north/northwestern clade and
<taxonomicName id="4C7A8D82DC3A3255F2C8F9C2FADC1738" box="[1245,1369,1567,1589]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="11" pageNumber="291" phylum="Chordata" rank="species" species="tridactyla">
<emphasis id="B90E2A13DC3A3255F2C8F9C2FADC1738" box="[1245,1369,1567,1589]" italics="true" pageId="11" pageNumber="291">S. tridactyla</emphasis>
</taxonomicName>
and
<emphasis id="B90E2A13DC3A3255F39CF9C2FE921759" italics="true" pageId="11" pageNumber="291">
<taxonomicName id="4C7A8D82DC3A3255F39CF9C2FE911759" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="11" pageNumber="291" phylum="Chordata" rank="species" species="tetradactyla">S. tetradactyla</taxonomicName>
,
</emphasis>
showing various degrees of external reduction of digits on the right, and CT scans of the hands of the same individuals on the left. External reduction of fingers is only partly paralleled by reduction of the skeletal elements (especially in
<taxonomicName id="4C7A8D82DC3A3255F682F99EFE92179F" box="[151,279,1660,1682]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="11" pageNumber="291" phylum="Chordata" rank="species" species="tridactyla">
<emphasis id="B90E2A13DC3A3255F682F99EFE92179F" box="[151,279,1660,1682]" italics="true" pageId="11" pageNumber="291">S. tridactyla</emphasis>
</taxonomicName>
; see text). Scale bar=0.5 mm. Asterisks mark right hands in mirrored view. Formulae show the number of phalanges on fingers IIV.
</paragraph>
</caption>
<paragraph id="8BC5F601DC3A3252F6D2F93FFD5F1068" blockId="11.[151,1436,1757,2034]" lastBlockId="12.[151,1436,151,357]" lastPageId="12" lastPageNumber="292" pageId="11" pageNumber="291">
The degree of reduction of various digits in the manus and pes has served as an important diagnostic character to describe various new species such as
<taxonomicName id="4C7A8D82DC3A3255F479F8E0FC971617" box="[620,786,1793,1818]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="11" pageNumber="291" phylum="Chordata" rank="species" species="tetradactyla">
<emphasis id="B90E2A13DC3A3255F479F8E0FC971617" box="[620,786,1793,1818]" italics="true" pageId="11" pageNumber="291">S. tetradactyla</emphasis>
</taxonomicName>
which has its first toe almost completely reduced, or
<taxonomicName id="4C7A8D82DC3A3255F392F8E0FF791630" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="11" pageNumber="291" phylum="Chordata" rank="species" species="pygmaea">
<emphasis id="B90E2A13DC3A3255F392F8E0FF791630" italics="true" pageId="11" pageNumber="291">S. pygmaea</emphasis>
</taxonomicName>
, a highly miniaturized species with only slight digital reduction (Vences &amp; Glaw 1991). While in
<taxonomicName id="4C7A8D82DC3A3255F392F8C4FE81166F" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="11" pageNumber="291" phylum="Chordata" rank="species" species="tridactyla">
<emphasis id="B90E2A13DC3A3255F392F8C4FE81166F" italics="true" pageId="11" pageNumber="291">S. tridactyla</emphasis>
</taxonomicName>
the bony anatomy of manus and pes was also found to be affected by reductions of certain phalangeal elements (Guibé 1978), this has so far remained unstudied in most other species. We here provide a first survey of hand and foot skeletons of
<taxonomicName id="4C7A8D82DC3A3255F7D1F870FDE316A7" box="[452,614,1937,1962]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="11" pageNumber="291" phylum="Chordata" rank="species" species="analamaina">
<emphasis id="B90E2A13DC3A3255F7D1F870FDE316A7" box="[452,614,1937,1962]" italics="true" pageId="11" pageNumber="291">S. analamaina</emphasis>
</taxonomicName>
and a set of superficially similar and related species, mainly belonging to the north/northwestern clade of
<taxonomicName id="4C7A8D82DC3A3255F401F854FDFA16C3" box="[532,639,1974,1998]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="11" pageNumber="291" phylum="Chordata" rank="genus">
<emphasis id="B90E2A13DC3A3255F401F854FDFA16C3" box="[532,639,1974,1998]" italics="true" pageId="11" pageNumber="291">Stumpffia</emphasis>
</taxonomicName>
(
<figureCitation id="1341EA84DC3A3255F481F857FC8216C3" box="[660,775,1973,1998]" captionStart-0="FIGURE 6" captionStart-1="FIGURE 7" captionStartId-0="11.[151,250,1567,1589]" captionStartId-1="12.[151,250,1923,1945]" captionTargetBox-0="[151,1436,183,1546]" captionTargetBox-1="[153,1436,393,1900]" captionTargetId-0="figure@11.[151,1436,183,1546]" captionTargetId-1="figure@12.[151,1436,393,1901]" captionTargetPageId-0="11" captionTargetPageId-1="12" captionText-0="FIGURE 6. Ventral views of left hands of different species of Stumpffia of the north / northwestern clade and S. tridactyla and S. tetradactyla, showing various degrees of external reduction of digits on the right, and CT scans of the hands of the same individuals on the left. External reduction of fingers is only partly paralleled by reduction of the skeletal elements (especially in S. tridactyla; see text). Scale bar = 0.5 mm. Asterisks mark right hands in mirrored view. Formulae show the number of phalanges on fingers I IV." captionText-1="FIGURE 7. Ventral views of feet of different species of Stumpffia of the north / northwestern clade and S. tridactyla and S. tetradactyla, showing various degrees of external reduction of digits on the right, and CT scans of the feet of the same individuals on the left. External reduction of toes is only partly paralleled by reduction of the skeletal elements (especially in S. tridactyla; see text). Scale bar = 1 mm. Formulae show the number of phalanges on toes I V." httpUri-0="https://zenodo.org/record/223542/files/figure.png" httpUri-1="https://zenodo.org/record/223543/files/figure.png" pageId="11" pageNumber="291">Figs. 67</figureCitation>
). The phalangeal formula for the manus of the studied specimens of
<taxonomicName id="4C7A8D82DC3A3255F724F838FE5616FF" box="[305,467,2009,2034]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="11" pageNumber="291" phylum="Chordata" rank="species" species="analamaina">
<emphasis id="B90E2A13DC3A3255F724F838FE5616FF" box="[305,467,2009,2034]" italics="true" pageId="11" pageNumber="291">S. analamaina</emphasis>
</taxonomicName>
(ZCMV 10034),
<emphasis id="B90E2A13DC3A3255F48DF838FD2B16FC" box="[664,686,2010,2033]" italics="true" pageId="11" pageNumber="291">S.</emphasis>
sp. 31 (ZCMV 10031; terminal phalanx of finger IV possibly very small),
<taxonomicName id="4C7A8D82DC3D3252F6F8FF7AFE5011BD" box="[237,469,151,176]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="12" pageNumber="292" phylum="Chordata" rank="species" species="madagascariensis">
<emphasis id="B90E2A13DC3D3252F6F8FF7AFE5011BD" box="[237,469,151,176]" italics="true" pageId="12" pageNumber="292">S. madagascariensis</emphasis>
</taxonomicName>
(FGZC 389),
<emphasis id="B90E2A13DC3D3252F469FF7AFD1411A2" box="[636,657,152,175]" italics="true" pageId="12" pageNumber="292">S.</emphasis>
sp. 26 (FGMV 2002.594; phalangeal formula poorly recognizable), and
<taxonomicName id="4C7A8D82DC3D3252F6D2FF5FFEEE11D8" box="[199,363,188,213]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="12" pageNumber="292" phylum="Chordata" rank="species" species="tetradactyla">
<emphasis id="B90E2A13DC3D3252F6D2FF5FFEEE11D8" box="[199,363,188,213]" italics="true" pageId="12" pageNumber="292">S. tetradactyla</emphasis>
</taxonomicName>
(ZCMV 3395) was determined as 1232 (
<figureCitation id="1341EA84DC3D3252F57DFF5EFC2811D8" box="[872,941,188,213]" captionStart="FIGURE 6" captionStartId="11.[151,250,1567,1589]" captionTargetBox="[151,1436,183,1546]" captionTargetId="figure@11.[151,1436,183,1546]" captionTargetPageId="11" captionText="FIGURE 6. Ventral views of left hands of different species of Stumpffia of the north / northwestern clade and S. tridactyla and S. tetradactyla, showing various degrees of external reduction of digits on the right, and CT scans of the hands of the same individuals on the left. External reduction of fingers is only partly paralleled by reduction of the skeletal elements (especially in S. tridactyla; see text). Scale bar = 0.5 mm. Asterisks mark right hands in mirrored view. Formulae show the number of phalanges on fingers I IV." httpUri="https://zenodo.org/record/223542/files/figure.png" pageId="12" pageNumber="292">Fig. 6</figureCitation>
). A deviant phalangeal formula of 2233 was present in the two sister species,
<emphasis id="B90E2A13DC3D3252F42EFF02FDD511FA" box="[571,592,224,247]" italics="true" pageId="12" pageNumber="292">S.</emphasis>
sp. 27 (FGZC 433) and
<emphasis id="B90E2A13DC3D3252F571FF02FCFC11FA" box="[868,889,224,247]" italics="true" pageId="12" pageNumber="292">S.</emphasis>
sp. 28 (FGZC 1049), and a third combination of 223
<quantity id="4C825BE4DC3D3252F6FEFEE6FE921010" box="[235,279,260,285]" metricMagnitude="-2" metricUnit="m" metricValue="5.08" pageId="12" pageNumber="292" unit="in" value="2.0">2 in</quantity>
<taxonomicName id="4C7A8D82DC3D3252F70AFEE7FE131010" box="[287,406,260,285]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="12" pageNumber="292" phylum="Chordata" rank="species" species="gimmeli">
<emphasis id="B90E2A13DC3D3252F70AFEE7FE131010" box="[287,406,260,285]" italics="true" pageId="12" pageNumber="292">S. gimmeli</emphasis>
</taxonomicName>
(FGMV 2000.151) (
<figureCitation id="1341EA84DC3D3252F491FEE6FD4F1010" box="[644,714,260,285]" captionStart="FIGURE 6" captionStartId="11.[151,250,1567,1589]" captionTargetBox="[151,1436,183,1546]" captionTargetId="figure@11.[151,1436,183,1546]" captionTargetPageId="11" captionText="FIGURE 6. Ventral views of left hands of different species of Stumpffia of the north / northwestern clade and S. tridactyla and S. tetradactyla, showing various degrees of external reduction of digits on the right, and CT scans of the hands of the same individuals on the left. External reduction of fingers is only partly paralleled by reduction of the skeletal elements (especially in S. tridactyla; see text). Scale bar = 0.5 mm. Asterisks mark right hands in mirrored view. Formulae show the number of phalanges on fingers I IV." httpUri="https://zenodo.org/record/223542/files/figure.png" pageId="12" pageNumber="292">Fig. 6</figureCitation>
). A reduced number of phalanges was observed in
<taxonomicName id="4C7A8D82DC3D3252F307FEE7FA191010" box="[1298,1436,260,285]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="12" pageNumber="292" phylum="Chordata" rank="species" species="tridactyla">
<emphasis id="B90E2A13DC3D3252F307FEE7FA191010" box="[1298,1436,260,285]" italics="true" pageId="12" pageNumber="292">S. tridactyla</emphasis>
</taxonomicName>
(FGZC 2844) as already reported by Guibé (1978): 0131 (or 1131 if the tiny terminal knob of the first finger is counted as rudiment of a phalanx) (
<figureCitation id="1341EA84DC3D3252F492FEAEFD491068" box="[647,716,332,357]" captionStart="FIGURE 6" captionStartId="11.[151,250,1567,1589]" captionTargetBox="[151,1436,183,1546]" captionTargetId="figure@11.[151,1436,183,1546]" captionTargetPageId="11" captionText="FIGURE 6. Ventral views of left hands of different species of Stumpffia of the north / northwestern clade and S. tridactyla and S. tetradactyla, showing various degrees of external reduction of digits on the right, and CT scans of the hands of the same individuals on the left. External reduction of fingers is only partly paralleled by reduction of the skeletal elements (especially in S. tridactyla; see text). Scale bar = 0.5 mm. Asterisks mark right hands in mirrored view. Formulae show the number of phalanges on fingers I IV." httpUri="https://zenodo.org/record/223542/files/figure.png" pageId="12" pageNumber="292">Fig. 6</figureCitation>
).
</paragraph>
<caption id="DF05A689DC3D3252F682F861FBBD16F8" httpUri="https://zenodo.org/record/223543/files/figure.png" pageId="12" pageNumber="292" targetBox="[153,1436,393,1900]" targetPageId="12">
<paragraph id="8BC5F601DC3D3252F682F861FBBD16F8" blockId="12.[151,1436,1923,2037]" pageId="12" pageNumber="292">
<emphasis id="B90E2A13DC3D3252F682F861FE901695" bold="true" box="[151,277,1923,1945]" pageId="12" pageNumber="292">FIGURE 7.</emphasis>
Ventral views of feet of different species of
<taxonomicName id="4C7A8D82DC3D3252F4FAF861FCCA1694" box="[751,847,1923,1945]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="12" pageNumber="292" phylum="Chordata" rank="genus">
<emphasis id="B90E2A13DC3D3252F4FAF861FCCA1694" box="[751,847,1923,1945]" italics="true" pageId="12" pageNumber="292">Stumpffia</emphasis>
</taxonomicName>
of the north/northwestern clade and
<taxonomicName id="4C7A8D82DC3D3252F2C1F866FAD61694" box="[1236,1363,1923,1945]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="12" pageNumber="292" phylum="Chordata" rank="species" species="tridactyla">
<emphasis id="B90E2A13DC3D3252F2C1F866FAD61694" box="[1236,1363,1923,1945]" italics="true" pageId="12" pageNumber="292">S. tridactyla</emphasis>
</taxonomicName>
and
<emphasis id="B90E2A13DC3D3252F39CF866FE9216B5" italics="true" pageId="12" pageNumber="292">
<taxonomicName id="4C7A8D82DC3D3252F39CF866FE9116B5" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="12" pageNumber="292" phylum="Chordata" rank="species" species="tetradactyla">S. tetradactyla</taxonomicName>
,
</emphasis>
showing various degrees of external reduction of digits on the right, and CT scans of the feet of the same individuals on the left. External reduction of toes is only partly paralleled by reduction of the skeletal elements (especially in
<taxonomicName id="4C7A8D82DC3D3252F39CF823FF7F16F8" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="12" pageNumber="292" phylum="Chordata" rank="species" species="tridactyla">
<emphasis id="B90E2A13DC3D3252F39CF823FF7F16F8" italics="true" pageId="12" pageNumber="292">S. tridactyla</emphasis>
</taxonomicName>
; see text). Scale bar=1 mm. Formulae show the number of phalanges on toes IV.
</paragraph>
</caption>
<paragraph id="8BC5F601DC3C3253F6D2FF7AFB85104D" blockId="13.[151,1436,151,2013]" pageId="13" pageNumber="293">
In the pes (
<figureCitation id="1341EA84DC3C3253F753FF75FE0911BD" box="[326,396,151,176]" captionStart="FIGURE 7" captionStartId="12.[151,250,1923,1945]" captionTargetBox="[153,1436,393,1900]" captionTargetId="figure@12.[151,1436,393,1901]" captionTargetPageId="12" captionText="FIGURE 7. Ventral views of feet of different species of Stumpffia of the north / northwestern clade and S. tridactyla and S. tetradactyla, showing various degrees of external reduction of digits on the right, and CT scans of the feet of the same individuals on the left. External reduction of toes is only partly paralleled by reduction of the skeletal elements (especially in S. tridactyla; see text). Scale bar = 1 mm. Formulae show the number of phalanges on toes I V." httpUri="https://zenodo.org/record/223543/files/figure.png" pageId="13" pageNumber="293">Fig. 7</figureCitation>
), we observed a phalangeal formula of 1234
<quantity id="4C825BE4DC3C3253F5D5FF7AFC6E11BD" box="[960,1003,151,176]" metricMagnitude="-2" metricUnit="m" metricValue="7.62" pageId="13" pageNumber="293" unit="in" value="3.0">3 in</quantity>
<taxonomicName id="4C7A8D82DC3C3253F5E6FF7AFB1311BD" box="[1011,1174,151,176]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="analamaina">
<emphasis id="B90E2A13DC3C3253F5E6FF7AFB1311BD" box="[1011,1174,151,176]" italics="true" pageId="13" pageNumber="293">S. analamaina</emphasis>
</taxonomicName>
(ZCMV 10034),
<emphasis id="B90E2A13DC3C3253F34AFF7AFAF111A2" box="[1375,1396,152,175]" italics="true" pageId="13" pageNumber="293">S.</emphasis>
sp. 28 (FGZC 1049),
<taxonomicName id="4C7A8D82DC3C3253F76EFF5FFDA311D8" box="[379,550,188,213]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="tetradactyla">
<emphasis id="B90E2A13DC3C3253F76EFF5FFDA311D8" box="[379,550,188,213]" italics="true" pageId="13" pageNumber="293">S. tetradactyla</emphasis>
</taxonomicName>
(ZCMV 3395), and probably in
<taxonomicName id="4C7A8D82DC3C3253F5DDFF5FFB3311D8" box="[968,1206,188,213]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="madagascariensis">
<emphasis id="B90E2A13DC3C3253F5DDFF5FFB3311D8" box="[968,1206,188,213]" italics="true" pageId="13" pageNumber="293">S. madagascariensis</emphasis>
</taxonomicName>
(FGZC 389). The generalized, plesiomorphic anuran phalangeal formula of 22343 (Alberch &amp; Gale 1985) without losses was present in
<emphasis id="B90E2A13DC3C3253F719FEE7FEA41011" box="[268,289,261,284]" italics="true" pageId="13" pageNumber="293">S.</emphasis>
sp. 26 (FGMV 2002.594),
<emphasis id="B90E2A13DC3C3253F441FEE7FDEC1011" box="[596,617,261,284]" italics="true" pageId="13" pageNumber="293">S.</emphasis>
sp. 27 (FGZC 433), and
<taxonomicName id="4C7A8D82DC3C3253F596FEE7FC7C1010" box="[899,1017,260,285]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="gimmeli">
<emphasis id="B90E2A13DC3C3253F596FEE7FC7C1010" box="[899,1017,260,285]" italics="true" pageId="13" pageNumber="293">S. gimmeli</emphasis>
</taxonomicName>
(FGMV 2000.151) (
<figureCitation id="1341EA84DC3C3253F2F6FEE6FAA21011" box="[1251,1319,260,285]" captionStart="FIGURE 7" captionStartId="12.[151,250,1923,1945]" captionTargetBox="[153,1436,393,1900]" captionTargetId="figure@12.[151,1436,393,1901]" captionTargetPageId="12" captionText="FIGURE 7. Ventral views of feet of different species of Stumpffia of the north / northwestern clade and S. tridactyla and S. tetradactyla, showing various degrees of external reduction of digits on the right, and CT scans of the feet of the same individuals on the left. External reduction of toes is only partly paralleled by reduction of the skeletal elements (especially in S. tridactyla; see text). Scale bar = 1 mm. Formulae show the number of phalanges on toes I V." httpUri="https://zenodo.org/record/223543/files/figure.png" pageId="13" pageNumber="293">Fig. 7</figureCitation>
). Again, a stronger phalangeal reduction is apparent in
<taxonomicName id="4C7A8D82DC3C3253F49DFECAFC94104D" box="[648,785,295,320]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="tridactyla">
<emphasis id="B90E2A13DC3C3253F49DFECAFC94104D" box="[648,785,295,320]" italics="true" pageId="13" pageNumber="293">S. tridactyla</emphasis>
</taxonomicName>
: 01243 (
<figureCitation id="1341EA84DC3C3253F5B8FEC5FC77104D" box="[941,1010,295,320]" captionStart="FIGURE 7" captionStartId="12.[151,250,1923,1945]" captionTargetBox="[153,1436,393,1900]" captionTargetId="figure@12.[151,1436,393,1901]" captionTargetPageId="12" captionText="FIGURE 7. Ventral views of feet of different species of Stumpffia of the north / northwestern clade and S. tridactyla and S. tetradactyla, showing various degrees of external reduction of digits on the right, and CT scans of the feet of the same individuals on the left. External reduction of toes is only partly paralleled by reduction of the skeletal elements (especially in S. tridactyla; see text). Scale bar = 1 mm. Formulae show the number of phalanges on toes I V." httpUri="https://zenodo.org/record/223543/files/figure.png" pageId="13" pageNumber="293">Fig. 7</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C360A58ADC3C3253F6D2FEAEFE0A13A5" pageId="13" pageNumber="293" type="etymology">
<paragraph id="8BC5F601DC3C3253F6D2FEAEFC6910A0" blockId="13.[151,1436,151,2013]" pageId="13" pageNumber="293">
<emphasis id="B90E2A13DC3C3253F6D2FEAEFECB1068" bold="true" box="[199,334,332,357]" pageId="13" pageNumber="293">Etymology.</emphasis>
The species epithet is composed by the
<collectingCountry id="F36DB691DC3C3253F522FEAEFC211068" box="[823,932,332,357]" name="Madagascar" pageId="13" pageNumber="293">Malagasy</collectingCountry>
words
<emphasis id="B90E2A13DC3C3253F215FEAEFBA11068" box="[1024,1060,332,357]" italics="true" pageId="13" pageNumber="293">ala</emphasis>
(forest; an ala meaning &quot;of the forest&quot;) and
<emphasis id="B90E2A13DC3C3253F732FE93FEE81085" box="[295,365,369,392]" italics="true" pageId="13" pageNumber="293">maina</emphasis>
meaning dry, making reference to the occurrence of the new species in dry forest habitat in northwestern
<collectingCountry id="F36DB691DC3C3253F725FE76FE3F10A0" box="[304,442,404,429]" name="Madagascar" pageId="13" pageNumber="293">Madagascar</collectingCountry>
. The species epithet is used as noun in apposition.
</paragraph>
<paragraph id="8BC5F601DC3C3253F6D2FE55FE0A13A5" blockId="13.[151,1436,151,2013]" pageId="13" pageNumber="293">
<emphasis id="B90E2A13DC3C3253F6D2FE55FE7E10DD" bold="true" box="[199,507,439,464]" pageId="13" pageNumber="293">Molecular differentiation.</emphasis>
The mtDNA tree (
<figureCitation id="1341EA84DC3C3253F4D9FE55FC8910DD" box="[716,780,439,464]" captionStart="FIGURE 1" captionStartId="4.[151,250,1873,1895]" captionTargetBox="[247,1327,193,1822]" captionTargetId="figure@4.[247,1328,193,1823]" captionTargetPageId="4" captionText="FIGURE 1. Phylogenetic tree of 210 individuals of Stumpffia, showing in detail the north / northwestern clade, based on DNA sequences of a fragment of the mitochondrial 16 S rRNA gene (50 % majority rule consensus tree from a Bayesian inference analysis). Rhombophryne minuta was used as outgroup (not shown). All Stumpffia species and subclades not belonging to the north / northwestern clade are collapsed since they are not the subject of the present study. Bayesian support values are given above branches (only values of 0.95 and higher are shown). Scale bar represents changes per site." httpUri="https://zenodo.org/record/223537/files/figure.png" pageId="13" pageNumber="293">Fig. 1</figureCitation>
) places
<taxonomicName id="4C7A8D82DC3C3253F57FFE5AFB8910DD" box="[874,1036,439,464]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="analamaina">
<emphasis id="B90E2A13DC3C3253F57FFE5AFB8910DD" box="[874,1036,439,464]" italics="true" pageId="13" pageNumber="293">S. analamaina</emphasis>
</taxonomicName>
with maximum support into a clade with
<emphasis id="B90E2A13DC3C3253F6C5FE3FFF6310F9" box="[208,230,477,500]" italics="true" pageId="13" pageNumber="293">S.</emphasis>
sp. 29 and
<emphasis id="B90E2A13DC3C3253F77CFE3FFEFB10F9" box="[361,382,477,500]" italics="true" pageId="13" pageNumber="293">S.</emphasis>
sp. 31. Differentiation from these two candidate species in the 16S gene is 11.5% and 10.5% (uncorrected p-distance). Among the eight specimens of
<taxonomicName id="4C7A8D82DC3C3253F507FDE2FC361315" box="[786,947,511,536]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="analamaina" status="sp. nov.">
<emphasis id="B90E2A13DC3C3253F507FDE2FC361315" box="[786,947,511,536]" italics="true" pageId="13" pageNumber="293">S. analamaina</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D9768DC3C3253F5AFFE1DFB961315" box="[954,1043,511,536]" pageId="13" pageNumber="293" rank="species">
<emphasis id="B90E2A13DC3C3253F5AFFE1DFB961315" bold="true" box="[954,1043,511,536]" pageId="13" pageNumber="293">sp. nov.</emphasis>
</taxonomicNameLabel>
sequenced for 16S, two specimens differed by two substitutions (maximum sequence divergence 0.4%). In the RAG1 haplotype network (
<figureCitation id="1341EA84DC3C3253F33EFDC6FAF51331" box="[1323,1392,548,573]" captionStart="FIGURE 2" captionStartId="5.[151,250,1153,1175]" captionTargetBox="[152,1436,193,1130]" captionTargetId="figure@5.[151,1436,193,1131]" captionTargetPageId="5" captionText="FIGURE 2. Haplotype network based on DNA sequences of a fragment of the nuclear RAG 1 gene of species and candidate species of the north / northwestern clade of Stumpffia. Circles represent distinct haplotypes, with sizes proportional to the number of individuals within each haplotype. Colors represent species (or candidate species) assignment of individuals on the basis of the mtDNA tree (Fig. 1). Haplotypes connected by a simple line are separated by one nucleotide difference, small black dots represent additional substitutional steps. Note that the Stumpffia sp. 31 cluster contains a number of samples not sequenced for 16 S and therefore not reliably assigned to a mtDNA lineage." httpUri="https://zenodo.org/record/223538/files/figure.png" pageId="13" pageNumber="293">Fig. 2</figureCitation>
),
<taxonomicName id="4C7A8D82DC3C3253F392FDC7FE98136D" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="analamaina" status="sp. nov.">
<emphasis id="B90E2A13DC3C3253F392FDC7FE98136D" italics="true" pageId="13" pageNumber="293">S. analamaina</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D9768DC3C3253F731FDA5FEFB136D" box="[292,382,583,608]" pageId="13" pageNumber="293" rank="species">
<emphasis id="B90E2A13DC3C3253F731FDA5FEFB136D" bold="true" box="[292,382,583,608]" pageId="13" pageNumber="293">sp. nov.</emphasis>
</taxonomicNameLabel>
and the two candidate species form three distinct clusters, with a minimum of seven and nine mutational steps between them and with a maximum differentiation of three mutational steps between haplotypes within
<emphasis id="B90E2A13DC3C3253F6F0FD72FE0A13A5" box="[229,399,655,680]" italics="true" pageId="13" pageNumber="293">
<taxonomicName id="4C7A8D82DC3C3253F6F0FD72FE0E13A5" box="[229,395,655,680]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="analamaina">S. analamaina</taxonomicName>
.
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="C360A58ADC3C3250F6D2FD56FB211010" lastPageId="14" lastPageNumber="294" pageId="13" pageNumber="293" type="description">
<paragraph id="8BC5F601DC3C3253F6D2FD56FC0412A8" blockId="13.[151,1436,151,2013]" pageId="13" pageNumber="293">
<emphasis id="B90E2A13DC3C3253F6D2FD56FD7513C0" bold="true" box="[199,752,692,717]" pageId="13" pageNumber="293">Natural habitat and observations in captivity.</emphasis>
Specimens of
<taxonomicName id="4C7A8D82DC3C3253F58EFD56FB1113C0" box="[923,1172,692,717]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="analamaina">
<emphasis id="B90E2A13DC3C3253F58EFD56FB1113C0" box="[923,1172,692,717]" italics="true" pageId="13" pageNumber="293">Stumpffia analamaina</emphasis>
</taxonomicName>
were collected in June
<date id="FFC4D0C1DC3C3253F682FD3AFEB813FD" box="[151,317,727,752]" pageId="13" pageNumber="293" value="2009" valueMax="2010">
2009 and
<quantity id="4C825BE4DC3C3253F710FD3AFEB813FD" box="[261,317,728,752]" metricMagnitude="1" metricUnit="m" metricValue="5.1053999999999995" pageId="13" pageNumber="293" unit="in" value="2010.0">2010</quantity>
</date>
in the leaf litter directly adjacent to a small stream in a fragment of dry deciduous forest where they occurred in high densities. The forest of the area was heavily degraded and the forest fragment was approximately
<quantity id="4C825BE4DC3C3253F682FCC2FF7F123A" box="[151,250,800,824]" metricMagnitude="1" metricUnit="m" metricValue="1.5" metricValueMax="2.0" metricValueMin="1.0" pageId="13" pageNumber="293" unit="m" value="15.0" valueMax="20.0" valueMin="10.0">1020 m</quantity>
wide on each side along the stream (
<figureCitation id="1341EA84DC3C3253F48FFCFDFD5B1235" box="[666,734,799,824]" captionStart="FIGURE 4" captionStartId="7.[151,250,1852,1874]" captionTargetBox="[176,1425,260,1828]" captionTargetId="figure@7.[166,1426,165,1829]" captionTargetPageId="7" captionText="FIGURE 4. Specimens and habitat of Stumpffia analamaina sp. nov. (A) Paratype specimen ZSM 2831 / 2010 (ZCMV 10037) in life (scale bar = 1 mm). (B) Frame from a video showing prey capture of S. analamaina, showing partially lateral tongue protrusion as also known from other microhylids. The whitish structure on the right side of the head (arrow) is the tongue which is protruded at an angle of 40 50 ° relative to the body axis. (C) Dry forest and (D) stream at type locality, specimens were found in the leaf litter at the border of the stream. (E) Foam nest produced by S. analamaina in captivity. (F) Additional specimen (catalogue number uncertain) showing color variation (beige ground color)." httpUri="https://zenodo.org/record/223540/files/figure.png" pageId="13" pageNumber="293">Fig. 4</figureCitation>
CD). The time of collection was during the dry austral winter and therefore the whole area except the immediate stream surroundings was characterized by very low humidity, probably causing the specimens to gather along the stream. Additional frogs found at this site were
<emphasis id="B90E2A13DC3C3253F2E9FC85FD2912A8" italics="true" pageId="13" pageNumber="293">
<taxonomicName id="4C7A8D82DC3C3253F2E9FC85FE8112A8" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="ulcerosus">Mantidactylus ulcerosus</taxonomicName>
,
<taxonomicName id="4C7A8D82DC3C3253F71AFC6EFE5312A8" box="[271,470,908,933]" class="Amphibia" family="Mantellidae" genus="Mantella" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="ebenaui">Mantella ebenaui</taxonomicName>
,
<taxonomicName id="4C7A8D82DC3C3253F7F7FC6EFD2C12A8" box="[482,681,908,933]" class="Amphibia" family="Mantellidae" genus="Boophis" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="tampoka">Boophis tampoka</taxonomicName>
,
</emphasis>
and
<emphasis id="B90E2A13DC3C3253F4F7FC6FFC0412A8" box="[738,897,908,933]" italics="true" pageId="13" pageNumber="293">
<taxonomicName id="4C7A8D82DC3C3253F4F7FC6FFCF912A8" box="[738,892,908,933]" class="Amphibia" family="Mantellidae" genus="Boophis" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="brachychir">B. brachychir</taxonomicName>
.
</emphasis>
</paragraph>
<paragraph id="8BC5F601DC3C3253F6D2FC4DFAB415C8" blockId="13.[151,1436,151,2013]" pageId="13" pageNumber="293">
Specimens were kept over several months in captivity, 3
<specimenCount id="9D7C3D88DC3C3253F565FC4DFCFB12C5" box="[880,894,943,968]" pageId="13" pageNumber="293" type="adult">6</specimenCount>
adults in a terrarium of
<date id="FFC4D0C1DC3C3253F2BCFC52FA8312C5" box="[1193,1286,944,968]" pageId="13" pageNumber="293" value="1930-10-30">30 x 30</date>
x
<quantity id="4C825BE4DC3C3253F33EFC52FAFD12CA" box="[1323,1400,944,968]" metricMagnitude="-1" metricUnit="m" metricValue="3.0" pageId="13" pageNumber="293" unit="cm" value="30.0">30 cm</quantity>
, at temperatures between 20 and 25°C and air humidity of 6393%, on a substrate of turf with leaf litter (
<taxonomicName id="4C7A8D82DC3C3253F337FC36FF74151D" class="Magnoliopsida" family="Combretaceae" genus="Terminalia" kingdom="Plantae" order="Myrtales" pageId="13" pageNumber="293" phylum="Tracheophyta" rank="species" species="catappa">
<emphasis id="B90E2A13DC3C3253F337FC36FF74151D" italics="true" pageId="13" pageNumber="293">Terminalia catappa</emphasis>
</taxonomicName>
leaves) and with plastic plates as shallow water reservoirs. They were fed collembolans, mites, and fruit flies. Activity was mainly nocturnal, the individuals typical moved slowly and only jumped when disturbed (up to
<quantity id="4C825BE4DC3C3253F682FBDDFF59155A" box="[151,220,1087,1112]" metricMagnitude="-1" metricUnit="m" metricValue="6.0" pageId="13" pageNumber="293" unit="cm" value="60.0">60 cm</quantity>
jumping distance was recorded outside of the terrarium). Calls were emitted from concealed positions under dead leaves. In one case, during a period of high calling intensity in one terrarium, a foam nest was discovered within one of the plastic plates, with an estimated diameter of
<quantity id="4C825BE4DC3C3253F55BFB65FC181592" box="[846,925,1159,1184]" metricMagnitude="-2" metricUnit="m" metricValue="5.0" pageId="13" pageNumber="293" unit="mm" value="50.0">50 mm</quantity>
(
<figureCitation id="1341EA84DC3C3253F5B9FB65FC7515AD" box="[940,1008,1159,1184]" captionStart="FIGURE 4" captionStartId="7.[151,250,1852,1874]" captionTargetBox="[176,1425,260,1828]" captionTargetId="figure@7.[166,1426,165,1829]" captionTargetPageId="7" captionText="FIGURE 4. Specimens and habitat of Stumpffia analamaina sp. nov. (A) Paratype specimen ZSM 2831 / 2010 (ZCMV 10037) in life (scale bar = 1 mm). (B) Frame from a video showing prey capture of S. analamaina, showing partially lateral tongue protrusion as also known from other microhylids. The whitish structure on the right side of the head (arrow) is the tongue which is protruded at an angle of 40 50 ° relative to the body axis. (C) Dry forest and (D) stream at type locality, specimens were found in the leaf litter at the border of the stream. (E) Foam nest produced by S. analamaina in captivity. (F) Additional specimen (catalogue number uncertain) showing color variation (beige ground color)." httpUri="https://zenodo.org/record/223540/files/figure.png" pageId="13" pageNumber="293">Fig. 4</figureCitation>
E). It was guarded during several days by one individual. No eggs were laid in the foam nest, and no further reproductive activity was observed.
</paragraph>
<paragraph id="8BC5F601DC3C3253F6D2FB2DFAC61400" blockId="13.[151,1436,151,2013]" pageId="13" pageNumber="293">
During prey capture we observed specimens exhibiting lateral tongue protrusion (
<figureCitation id="1341EA84DC3C3253F24DFB2DFB1915E5" box="[1112,1180,1231,1256]" captionStart="FIGURE 4" captionStartId="7.[151,250,1852,1874]" captionTargetBox="[176,1425,260,1828]" captionTargetId="figure@7.[166,1426,165,1829]" captionTargetPageId="7" captionText="FIGURE 4. Specimens and habitat of Stumpffia analamaina sp. nov. (A) Paratype specimen ZSM 2831 / 2010 (ZCMV 10037) in life (scale bar = 1 mm). (B) Frame from a video showing prey capture of S. analamaina, showing partially lateral tongue protrusion as also known from other microhylids. The whitish structure on the right side of the head (arrow) is the tongue which is protruded at an angle of 40 50 ° relative to the body axis. (C) Dry forest and (D) stream at type locality, specimens were found in the leaf litter at the border of the stream. (E) Foam nest produced by S. analamaina in captivity. (F) Additional specimen (catalogue number uncertain) showing color variation (beige ground color)." httpUri="https://zenodo.org/record/223540/files/figure.png" pageId="13" pageNumber="293">Fig. 4</figureCitation>
B), which is typical for microhylids (Meyers
<emphasis id="B90E2A13DC3C3253F79DFB17FE451400" box="[392,448,1268,1293]" italics="true" pageId="13" pageNumber="293">et al.</emphasis>
2004) but so far has not been reported for any representative of the
<taxonomicName id="4C7A8D82DC3C3253F2ACFB16FABB1400" box="[1209,1342,1268,1293]" class="Amphibia" family="Microhylidae" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="subFamily" subFamily="Cophylinae">Cophylinae</taxonomicName>
.
</paragraph>
<paragraph id="8BC5F601DC3C3253F6D2FAF5FB9D1778" blockId="13.[151,1436,151,2013]" pageId="13" pageNumber="293">
<emphasis id="B90E2A13DC3C3253F6D2FAF5FE31143D" bold="true" box="[199,436,1303,1328]" pageId="13" pageNumber="293">Advertisement call.</emphasis>
The following description is based on several series of notes emitted by a male in captivity, from a glass terrarium at an air temperature of 23°C. Vocalizations were arranged in series of 38 notes, although we suspect that similar to all other
<taxonomicName id="4C7A8D82DC3C3253F491FA82FD6A1475" box="[644,751,1376,1400]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="genus">
<emphasis id="B90E2A13DC3C3253F491FA82FD6A1475" box="[644,751,1376,1400]" italics="true" pageId="13" pageNumber="293">Stumpffia</emphasis>
</taxonomicName>
for which calls have been described,
<taxonomicName id="4C7A8D82DC3C3253F284FA82FAB61475" box="[1169,1331,1375,1400]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="analamaina">
<emphasis id="B90E2A13DC3C3253F284FA82FAB61475" box="[1169,1331,1375,1400]" italics="true" pageId="13" pageNumber="293">S. analamaina</emphasis>
</taxonomicName>
might be able to emit notes in long series of an undetermined number of notes, potentially enduring for many minutes. In the five recorded note series, and several other series heard from the terrarium, the notes were emitted at regular intervals and sounded very similar to each other; we are therefore confident that the call described here corresponds to the true advertisement call. However, due to the recording conditions, we cannot exclude that the note duration measured may be longer than actually emitted due to echo effects of the terrarium. Nevertheless, as the measured notes altogether are shorter in duration rather than longer compared to calls of most other
<taxonomicName id="4C7A8D82DC3C3253F295F9DAFB6E175D" box="[1152,1259,1592,1616]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="genus">
<emphasis id="B90E2A13DC3C3253F295F9DAFB6E175D" box="[1152,1259,1592,1616]" italics="true" pageId="13" pageNumber="293">Stumpffia</emphasis>
</taxonomicName>
, this should not influence the value of the call as a taxonomic character in the context used here.
</paragraph>
<paragraph id="8BC5F601DC3C3253F6D2F99DFEAD1640" blockId="13.[151,1436,151,2013]" pageId="13" pageNumber="293">
The notes are small chirps of a rather noisy appearance (
<figureCitation id="1341EA84DC3C3253F556F99DFC021795" box="[835,903,1663,1688]" captionStart="FIGURE 8" captionStartId="14.[151,250,1296,1318]" captionTargetBox="[163,1433,337,1272]" captionTargetId="figure@14.[150,1436,325,1275]" captionTargetPageId="14" captionText="FIGURE 8. Spectrogram (top) and corresponding oscillogram (bottom) of the advertisement call of Stumpffia analamaina sp. nov., recorded from a glass terrarium (air temperature 23 ° C)." httpUri="https://zenodo.org/record/223544/files/figure.png" pageId="13" pageNumber="293">Fig. 8</figureCitation>
), less tonal than in most other
<taxonomicName id="4C7A8D82DC3C3253F2F7F962FAC91795" box="[1250,1356,1664,1688]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="genus">
<emphasis id="B90E2A13DC3C3253F2F7F962FAC91795" box="[1250,1356,1664,1688]" italics="true" pageId="13" pageNumber="293">Stumpffia</emphasis>
</taxonomicName>
. In ten notes from two series, all recorded from the same individual male, note duration was 168228 ms (201
<emphasis id="B90E2A13DC3C3253F323F947FAC017B1" box="[1334,1349,1701,1724]" italics="true" pageId="13" pageNumber="293">±</emphasis>
22 ms) and duration of inter-note intervals was
<date id="FFC4D0C1DC3C3253F449F925FD5F17ED" box="[604,730,1735,1760]" pageId="13" pageNumber="293" value="2045" valueMax="2570">20452570</date>
ms (2315
<emphasis id="B90E2A13DC3C3253F541F92BFCE617ED" box="[852,867,1737,1760]" italics="true" pageId="13" pageNumber="293">±</emphasis>
187 ms), which results in a note repetition rate of 2127 notes per minute. The calls show frequency bands between 48 kHz, but we cannot exclude that this wide frequency range is an artefact due to the suboptimal recording conditions. Maximum call energy is distributed at 6.06.4 kHz.
</paragraph>
<paragraph id="8BC5F601DC3C3250F6D2F8BAFB211010" blockId="13.[151,1436,151,2013]" lastBlockId="14.[151,1436,151,285]" lastPageId="14" lastPageNumber="294" pageId="13" pageNumber="293">
In general, the call variables of
<taxonomicName id="4C7A8D82DC3C3253F43FF8BAFCA4167D" box="[554,801,1879,1904]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="analamaina">
<emphasis id="B90E2A13DC3C3253F43FF8BAFCA4167D" box="[554,801,1879,1904]" italics="true" pageId="13" pageNumber="293">Stumpffia analamaina</emphasis>
</taxonomicName>
are very similar to those of
<taxonomicName id="4C7A8D82DC3C3253F248F8BAFB5B167D" box="[1117,1246,1880,1904]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="pygmaea">
<emphasis id="B90E2A13DC3C3253F248F8BAFB5B167D" box="[1117,1246,1880,1904]" italics="true" pageId="13" pageNumber="293">S. pygmaea</emphasis>
</taxonomicName>
. This species, as described by Vences &amp; Glaw (1991), emits series of tonal notes of ca. 200 ms duration at a rate of 2223 notes per minute and at a spectral frequency of 5.86.0 kHz. However, other species of the genus (e.g.,
<taxonomicName id="4C7A8D82DC3C3253F2AFF843FAD816B4" box="[1210,1373,1952,1977]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="13" pageNumber="293" phylum="Chordata" rank="species" species="tetradactyla">
<emphasis id="B90E2A13DC3C3253F2AFF843FAD816B4" box="[1210,1373,1952,1977]" italics="true" pageId="13" pageNumber="293">S. tetradactyla</emphasis>
</taxonomicName>
) also emit structurally similar calls (Vences &amp; Glaw 1991; Vences
<emphasis id="B90E2A13DC3C3253F541F827FC0A16D0" box="[852,911,1988,2013]" italics="true" pageId="13" pageNumber="293">et al.</emphasis>
2006), which can be differentiated from each other only by subtle differences (see discussion in Ndriantsoa
<emphasis id="B90E2A13DC3F3250F59AFF7BFC4A11BD" box="[911,975,151,176]" italics="true" pageId="14" pageNumber="294">et al.</emphasis>
2013). Therefore, and because of the suboptimal recording conditions, we prefer not to perform any more detailed comparison of the call of
<taxonomicName id="4C7A8D82DC3F3250F392FF5FFE9811F5" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="14" pageNumber="294" phylum="Chordata" rank="species" species="analamaina">
<emphasis id="B90E2A13DC3F3250F392FF5FFE9811F5" italics="true" pageId="14" pageNumber="294">S. analamaina</emphasis>
</taxonomicName>
at this time. Clearly, recordings of highly motivated males under natural conditions are necessary to fully describe the temporal and spectral characteristics of the vocal repertoire of this species.
</paragraph>
</subSubSection>
<caption id="DF05A689DC3F3250F682FAF2FC841448" httpUri="https://zenodo.org/record/223544/files/figure.png" pageId="14" pageNumber="294" targetBox="[163,1433,337,1272]" targetPageId="14">
<paragraph id="8BC5F601DC3F3250F682FAF2FC841448" blockId="14.[151,1436,1296,1349]" pageId="14" pageNumber="294">
<emphasis id="B90E2A13DC3F3250F682FAF2FE961428" bold="true" box="[151,275,1296,1318]" pageId="14" pageNumber="294">FIGURE 8.</emphasis>
Spectrogram (top) and corresponding oscillogram (bottom) of the advertisement call of
<taxonomicName id="4C7A8D82DC3F3250F282FAF2FAF2142B" box="[1175,1399,1296,1318]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="14" pageNumber="294" phylum="Chordata" rank="species" species="analamaina" status="sp. nov.">
<emphasis id="B90E2A13DC3F3250F282FAF2FAF2142B" box="[1175,1399,1296,1318]" italics="true" pageId="14" pageNumber="294">Stumpffia analamaina</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23D9768DC3F3250F368FAF3FF461449" pageId="14" pageNumber="294" rank="species">
<emphasis id="B90E2A13DC3F3250F368FAF3FF461449" bold="true" pageId="14" pageNumber="294">sp. nov.</emphasis>
</taxonomicNameLabel>
, recorded from a glass terrarium (air temperature 23°C).
</paragraph>
</caption>
<subSubSection id="C360A58ADC3F3250F6D2FA90FA0D161A" pageId="14" pageNumber="294" type="distribution">
<paragraph id="8BC5F601DC3F3250F6D2FA90FA0D161A" blockId="14.[151,1436,1394,1815]" pageId="14" pageNumber="294">
<emphasis id="B90E2A13DC3F3250F6D2FA90FEE51486" bold="true" box="[199,352,1394,1419]" pageId="14" pageNumber="294">Distribution.</emphasis>
The new species is so far only known from its
<typeStatus id="54C148A3DC3F3250F591FA91FC311486" box="[900,948,1395,1419]" pageId="14" pageNumber="294">type</typeStatus>
locality (
<figureCitation id="1341EA84DC3F3250F237FA90FBED1487" box="[1058,1128,1394,1419]" captionStart="FIGURE 9" captionStartId="15.[151,250,1289,1311]" captionTargetBox="[151,1436,185,1277]" captionTargetId="figure@15.[151,1436,185,1277]" captionTargetPageId="15" captionText="FIGURE 9. Map of northern Madagascar showing the known distribution records of species and candidate species in the north / northwestern clade of Stumpffia. Shades in the map show remaining primary vegetation in 2003 2006, modified from the Madagascar Vegetation Mapping Project (http: // www. vegmad. org)." httpUri="https://zenodo.org/record/223545/files/figure.png" pageId="14" pageNumber="294">Fig. 9</figureCitation>
). It is likely that it is more widespread within a small range, in remaining fragments of dry deciduous forests north and northeast of Antsohihy. Records of
<taxonomicName id="4C7A8D82DC3F3250F78AFA58FD8F14DF" box="[415,522,1466,1490]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="14" pageNumber="294" phylum="Chordata" rank="genus">
<emphasis id="B90E2A13DC3F3250F78AFA58FD8F14DF" box="[415,522,1466,1490]" italics="true" pageId="14" pageNumber="294">Stumpffia</emphasis>
</taxonomicName>
also exist from Ankarafantsika National Park, considerably south of Antsohihy (e.g., Blommers Schlösser &amp; Blanc 1991, and Ramanamanjato &amp; Rabibisoa 2002, as
<taxonomicName id="4C7A8D82DC3F3250F269FA3CFAA514FB" box="[1148,1312,1501,1526]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="14" pageNumber="294" phylum="Chordata" rank="species" species="psologlossa">
<emphasis id="B90E2A13DC3F3250F269FA3CFAA514FB" box="[1148,1312,1501,1526]" italics="true" pageId="14" pageNumber="294">S. psologlossa</emphasis>
</taxonomicName>
and
<emphasis id="B90E2A13DC3F3250F349FA3CFAEF14F8" box="[1372,1386,1502,1525]" italics="true" pageId="14" pageNumber="294">S</emphasis>
. sp. respectively), and these populations have so far not been studied from a molecular perspective. Two mitochondrial lineages, here named
<emphasis id="B90E2A13DC3F3250F79FF9C4FE1A1730" box="[394,415,1574,1597]" italics="true" pageId="14" pageNumber="294">S.</emphasis>
sp. 29 and
<emphasis id="B90E2A13DC3F3250F434F9C4FDB31730" box="[545,566,1574,1597]" italics="true" pageId="14" pageNumber="294">S.</emphasis>
sp. 31, were placed together with
<taxonomicName id="4C7A8D82DC3F3250F5ACF9C4FBDE1733" box="[953,1115,1573,1598]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="14" pageNumber="294" phylum="Chordata" rank="species" species="analamaina">
<emphasis id="B90E2A13DC3F3250F5ACF9C4FBDE1733" box="[953,1115,1573,1598]" italics="true" pageId="14" pageNumber="294">S. analamaina</emphasis>
</taxonomicName>
in a clade with high support by the mtDNA data (
<figureCitation id="1341EA84DC3F3250F797F9A8FE46176F" box="[386,451,1610,1635]" captionStart="FIGURE 1" captionStartId="4.[151,250,1873,1895]" captionTargetBox="[247,1327,193,1822]" captionTargetId="figure@4.[247,1328,193,1823]" captionTargetPageId="4" captionText="FIGURE 1. Phylogenetic tree of 210 individuals of Stumpffia, showing in detail the north / northwestern clade, based on DNA sequences of a fragment of the mitochondrial 16 S rRNA gene (50 % majority rule consensus tree from a Bayesian inference analysis). Rhombophryne minuta was used as outgroup (not shown). All Stumpffia species and subclades not belonging to the north / northwestern clade are collapsed since they are not the subject of the present study. Bayesian support values are given above branches (only values of 0.95 and higher are shown). Scale bar represents changes per site." httpUri="https://zenodo.org/record/223537/files/figure.png" pageId="14" pageNumber="294">Fig. 1</figureCitation>
). These two lineages occur at sites not too distant from the
<taxonomicName id="4C7A8D82DC3F3250F275F9A9FA87176E" box="[1120,1282,1610,1635]" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="14" pageNumber="294" phylum="Chordata" rank="species" species="analamaina">
<emphasis id="B90E2A13DC3F3250F275F9A9FA87176E" box="[1120,1282,1610,1635]" italics="true" pageId="14" pageNumber="294">S. analamaina</emphasis>
</taxonomicName>
<typeStatus id="54C148A3DC3F3250F31CF9A9FABC176E" box="[1289,1337,1611,1635]" pageId="14" pageNumber="294">type</typeStatus>
locality; especially
<emphasis id="B90E2A13DC3F3250F718F98CFEA71788" box="[269,290,1646,1669]" italics="true" pageId="14" pageNumber="294">S.</emphasis>
sp. 31 which occurs less than
<quantity id="4C825BE4DC3F3250F461F98CFD4C178B" box="[628,713,1645,1670]" metricMagnitude="5" metricUnit="m" metricValue="1.0" pageId="14" pageNumber="294" unit="km" value="100.0">100 km</quantity>
northeast of it (
<figureCitation id="1341EA84DC3F3250F56DF98FFC39178B" box="[888,956,1645,1670]" captionStart="FIGURE 9" captionStartId="15.[151,250,1289,1311]" captionTargetBox="[151,1436,185,1277]" captionTargetId="figure@15.[151,1436,185,1277]" captionTargetPageId="15" captionText="FIGURE 9. Map of northern Madagascar showing the known distribution records of species and candidate species in the north / northwestern clade of Stumpffia. Shades in the map show remaining primary vegetation in 2003 2006, modified from the Madagascar Vegetation Mapping Project (http: // www. vegmad. org)." httpUri="https://zenodo.org/record/223545/files/figure.png" pageId="14" pageNumber="294">Fig. 9</figureCitation>
). No bioacoustic data on those populations are available, and the limited morphological data of
<emphasis id="B90E2A13DC3F3250F539F971FCC417A7" box="[812,833,1683,1706]" italics="true" pageId="14" pageNumber="294">S.</emphasis>
sp. 31 (
<tableCitation id="C6F8C3BADC3F3250F5A3F970FB9717A7" box="[950,1042,1682,1707]" captionStart="TABLE 1" captionStartId="10.[151,239,980,1003]" captionTargetBox="[159,1403,1249,1642]" captionTargetPageId="10" captionText="TABLE 1. Morphometric data (in mm) of selected specimens of S. gimmeli, S. madagascariensis, S. pygmaea, S. sp. 31, and S. analamaina sp. nov. SVL, snout-vent length; HAL, hand length; FOTL, foot and tarsus length; FOL, foot length; TIBL, tibia length; HT, holotype; PT, paratype." httpUri="http://table.plazi.org/id/DF05A689DC3B3254F682FC36FD0B1526" pageId="14" pageNumber="294" tableUuid="DF05A689DC3B3254F682FC36FD0B1526">Table 1</tableCitation>
) do not indicate morphological differentiation; however, these mtDNA lineages did not share haplotypes in RAG1 (
<figureCitation id="1341EA84DC3F3250F254F954FB0017C2" box="[1089,1157,1718,1743]" captionStart="FIGURE 2" captionStartId="5.[151,250,1153,1175]" captionTargetBox="[152,1436,193,1130]" captionTargetId="figure@5.[151,1436,193,1131]" captionTargetPageId="5" captionText="FIGURE 2. Haplotype network based on DNA sequences of a fragment of the nuclear RAG 1 gene of species and candidate species of the north / northwestern clade of Stumpffia. Circles represent distinct haplotypes, with sizes proportional to the number of individuals within each haplotype. Colors represent species (or candidate species) assignment of individuals on the basis of the mtDNA tree (Fig. 1). Haplotypes connected by a simple line are separated by one nucleotide difference, small black dots represent additional substitutional steps. Note that the Stumpffia sp. 31 cluster contains a number of samples not sequenced for 16 S and therefore not reliably assigned to a mtDNA lineage." httpUri="https://zenodo.org/record/223538/files/figure.png" pageId="14" pageNumber="294">Fig. 2</figureCitation>
), suggesting a consistent genetic differentiation across markers. We therefore do not consider these populations as conspecific with
<taxonomicName id="4C7A8D82DC3F3250F392F939FE98161A" class="Amphibia" family="Microhylidae" genus="Stumpffia" kingdom="Animalia" order="Anura" pageId="14" pageNumber="294" phylum="Chordata" rank="species" species="analamaina">
<emphasis id="B90E2A13DC3F3250F392F939FE98161A" italics="true" pageId="14" pageNumber="294">S. analamaina</emphasis>
</taxonomicName>
although as with all candidate species—their status remains pending a thorough integrative revision.
</paragraph>
</subSubSection>
</treatment>
</document>