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, female), collected on 
<date id="E10CB4BEFFF400244CC9FA1AFB2259CE" box="[1012,1155,1426,1450]" pageId="21" pageNumber="107" value="2010-06-10">
<collectingDate id="F1484D56FFF400244CC9FA1AFB2259CE" box="[1012,1155,1426,1450]" pageId="21" pageNumber="107" value="2010-06-10">10 June 2010</collectingDate>
</date>
on the 
<location id="906DC4A5FFF400244FCCFA3DFED259A9" LSID="urn:lsid:plazi:treatment:1D1B2368FFF400284E70FB0AFEB059EB:906DC4A5FFF400244FCCFA3DFED259A9" box="[241,371,1461,1485]" country="Madagascar" county="Mahajanga province" latitude="-14.17413" longLatPrecision="1" longitude="48.94521" municipality="Sofia Region" name="Tsaratanana" pageId="21" pageNumber="107">Tsaratanana</location>
massif, 
<location id="906DC4A5FFF400244EF6FA3DFDAA59A9" LSID="urn:lsid:plazi:treatment:1D1B2368FFF400284E70FB0AFEB059EB:906DC4A5FFF400244EF6FA3DFDAA59A9" box="[459,523,1461,1485]" country="Madagascar" county="Mahajanga province" latitude="-14.17413" longLatPrecision="1" longitude="48.94521" municipality="Sofia Region" name="Camp" pageId="21" pageNumber="107">Camp</location>
1 (
<geoCoordinate id="F086F4B9FFF400244D12FA3DFD0959A9" box="[559,680,1461,1485]" degrees="14.17413" direction="south" orientation="latitude" pageId="21" pageNumber="107" precision="1" value="-14.17413">14.17413°S</geoCoordinate>
, 
<geoCoordinate id="F086F4B9FFF400244D8EFA3DFC8A59A9" box="[691,811,1461,1485]" degrees="48.94521" direction="east" orientation="longitude" pageId="21" pageNumber="107" precision="1" value="48.94521">48.94521°E</geoCoordinate>
, 
<elevation id="1E9F754DFFF400244C0BFA3DFB9C59A9" box="[822,1085,1461,1485]" metricMagnitude="3" metricUnit="m" metricValue="1.589" pageId="21" pageNumber="107" unit="m" value="1589.0">
<quantity id="524A3F9BFFF400244C0BFA3DFC2B59A9" box="[822,906,1461,1485]" metricMagnitude="3" metricUnit="m" metricValue="1.589" pageId="21" pageNumber="107" unit="m" value="1589.0">1589 m</quantity>
above sea level
</elevation>
), 
<collectingMunicipality id="75690804FFF400244B73FA3DFF4F598B" pageId="21" pageNumber="107">Sofia Region</collectingMunicipality>
, former 
<collectingCounty id="7C6CEAF2FFF400244E78FA5FFD83598B" box="[325,546,1495,1519]" pageId="21" pageNumber="107">Mahajanga province</collectingCounty>
, northwestern 
<collectingCountry id="EDA5D2EEFFF400244DFFFA5FFCE7598B" box="[706,838,1495,1519]" name="Madagascar" pageId="21" pageNumber="107">Madagascar</collectingCountry>
, by M
</materialsCitation>
. Vences, D. 
<materialsCitation id="25DA9823FFF400244B3FFA5FFBC5598B" box="[1026,1124,1495,1519]" collectionCode="R" collectorName="Vieites" country="Madagascar" pageId="21" pageNumber="107" specimenCount="1" typeStatus="paratype">
<collectorName id="3847F7A8FFF400244B3FFA5FFBEA598B" box="[1026,1099,1495,1519]" pageId="21" pageNumber="107">Vieites</collectorName>
, 
<collectionCode id="F3A30ABBFFF400244B69FA5FFBC5598B" box="[1108,1124,1495,1519]" country="Chile" name="Departamento de Geologia, Universidad de Chile" pageId="21" pageNumber="107">R</collectionCode>
</materialsCitation>
. D. Randrianiaina, F. Ratsoavina, S. Rasamison, A. Rakotoarison, F. Randrianasolo, E. 
<materialsCitation id="25DA9823FFF400244CCCFA72FF0E5A51" collectionCode="T" collectorName="Rajeriarison" country="Madagascar" pageId="21" pageNumber="107" specimenCount="1" typeStatus="paratype">
<collectorName id="3847F7A8FFF400244CCCFA72FBCC5A76" box="[1009,1133,1530,1554]" pageId="21" pageNumber="107">Rajeriarison</collectorName>
&amp; 
<collectionCode id="F3A30ABBFFF400244F9DF995FF0E5A51" box="[160,175,1565,1589]" name="Tavera, Department of Geology and Geophysics" pageId="21" pageNumber="107">T</collectionCode>
</materialsCitation>
. Rajoafiarison. 
<materialsCitation id="25DA9823FFF400244E76F995FDE45A1E" collectingDate="2010-06-29" collectionCode="ZSM, ZCMV" country="Madagascar" elevation="1109" latitude="-14.48251" location="Bemanevika river" longLatPrecision="1" longitude="48.62723" municipality="Sofia Region" pageId="21" pageNumber="107" specimenCode="ZSM 1801, ZCMV 12,567" specimenCount="1" stateProvince="Mahajanga" typeStatus="paratype">
<specimenCode id="C5143A05FFF400244E76F995FE135A51" box="[331,434,1565,1589]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="21" pageNumber="107" type="Museum">ZSM 1801</specimenCode>
/2010 (field number 
<specimenCode id="C5143A05FFF400244DB2F995FCBE5A51" box="[655,799,1565,1589]" collectionCode="ZCMV" pageId="21" pageNumber="107">ZCMV 12,567</specimenCode>
, male), collected on 
<date id="E10CB4BEFFF400244CCBF995FB225A51" box="[1014,1155,1565,1589]" pageId="21" pageNumber="107" value="2010-06-29">
<collectingDate id="F1484D56FFF400244CCBF995FB225A51" box="[1014,1155,1565,1589]" pageId="21" pageNumber="107" value="2010-06-29">29 June 2010</collectingDate>
</date>
at 
<location id="906DC4A5FFF400244F80F9B7FED95A33" LSID="urn:lsid:plazi:treatment:1D1B2368FFF400284E70FB0AFEB059EB:906DC4A5FFF400244F80F9B7FED95A33" box="[189,376,1599,1623]" country="Madagascar" latitude="-14.48251" longLatPrecision="1" longitude="48.62723" municipality="Sofia Region" name="Bemanevika river" pageId="21" pageNumber="107" stateProvince="Mahajanga">Bemanevika river</location>
(
<geoCoordinate id="F086F4B9FFF400244EB8F9B7FE5E5A33" box="[389,511,1599,1623]" degrees="14.48251" direction="south" orientation="latitude" pageId="21" pageNumber="107" precision="1" value="-14.48251">14.48251°S</geoCoordinate>
, 
<geoCoordinate id="F086F4B9FFF400244D37F9B7FD235A33" box="[522,642,1599,1623]" degrees="48.62723" direction="east" orientation="longitude" pageId="21" pageNumber="107" precision="1" value="48.62723">48.62723°E</geoCoordinate>
, 
<elevation id="1E9F754DFFF400244DB0F9B7FC355A33" box="[653,916,1599,1623]" metricMagnitude="3" metricUnit="m" metricValue="1.109" pageId="21" pageNumber="107" unit="m" value="1109.0">
<quantity id="524A3F9BFFF400244DB0F9B7FD415A33" box="[653,736,1599,1623]" metricMagnitude="3" metricUnit="m" metricValue="1.109" pageId="21" pageNumber="107" unit="m" value="1109.0">1109 m</quantity>
above sea level
</elevation>
), 
<collectingMunicipality id="75690804FFF400244C98F9B7FB915A33" box="[933,1072,1599,1623]" pageId="21" pageNumber="107">Sofia Region</collectingMunicipality>
, former 
<collectingRegion id="57765C9CFFF400244F9DF9EAFEDD5A1E" box="[160,380,1634,1658]" country="Madagascar" name="Mahajanga" pageId="21" pageNumber="107">Mahajanga province</collectingRegion>
, 
<collectingCountry id="EDA5D2EEFFF400244EB9F9EAFDA95A1E" box="[388,520,1634,1658]" name="Madagascar" pageId="21" pageNumber="107">Madagascar</collectingCountry>
, by M
</materialsCitation>
. Vences, D. 
<materialsCitation id="25DA9823FFF400244D83F9EAFCBF5A1E" box="[702,798,1634,1658]" collectionCode="R" collectorName="Vieites" country="Madagascar" pageId="21" pageNumber="107" specimenCount="1" stateProvince="Mahajanga" typeStatus="paratype">
<collectorName id="3847F7A8FFF400244D83F9EAFCA65A1E" box="[702,775,1634,1658]" pageId="21" pageNumber="107">Vieites</collectorName>
, 
<collectionCode id="F3A30ABBFFF400244C33F9EAFCBF5A1E" box="[782,798,1634,1658]" country="Chile" name="Departamento de Geologia, Universidad de Chile" pageId="21" pageNumber="107">R</collectionCode>
</materialsCitation>
. D. Randrianiaina, F. Ratsoavina, S. Rasamison, A. Rakotoarison, F. Randrianasolo, E. 
<materialsCitation id="25DA9823FFF400244DA1F90DFCE05AF9" box="[668,833,1669,1693]" collectionCode="T" collectorName="Rajeriarison" country="Madagascar" pageId="21" pageNumber="107" specimenCount="1" stateProvince="Mahajanga" typeStatus="paratype">
<collectorName id="3847F7A8FFF400244DA1F90DFCB95AF9" box="[668,792,1669,1693]" pageId="21" pageNumber="107">Rajeriarison</collectorName>
&amp; 
<collectionCode id="F3A30ABBFFF400244C0FF90DFCE05AF9" box="[818,833,1669,1693]" name="Tavera, Department of Geology and Geophysics" pageId="21" pageNumber="107">T</collectionCode>
</materialsCitation>
. Rajoafiarison. 
<materialsCitation id="25DA9823FFF400274CE1F90DFECC5CAD" collectingDate="2010-06-28" collectionCode="ZSM, ZCMV" country="Madagascar" county="Sofia Region" elevation="1538" lastPageId="22" lastPageNumber="108" latitude="-14.35991" location="Bemanevika" longLatPrecision="1" longitude="48.59022" municipality="Camp" pageId="21" pageNumber="107" specimenCode="ZSM 1802, ZCMV 12,596" specimenCount="1" stateProvince="Mahajanga" typeStatus="paratype">
<specimenCode id="C5143A05FFF400244CE1F90DFBE75AF9" box="[988,1094,1669,1693]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="21" pageNumber="107" type="Museum">ZSM 1802</specimenCode>
/2010 (field number 
<specimenCode id="C5143A05FFF400244E7CF92FFE775ADB" box="[321,470,1703,1727]" collectionCode="ZCMV" pageId="21" pageNumber="107">ZCMV 12,596</specimenCode>
, female), collected on 
<date id="E10CB4BEFFF400244DEBF92FFCCF5ADB" box="[726,878,1703,1727]" pageId="21" pageNumber="107" value="2010-06-28">
<collectingDate id="F1484D56FFF400244DEBF92FFCCF5ADB" box="[726,878,1703,1727]" pageId="21" pageNumber="107" value="2010-06-28">28 June 2010</collectingDate>
</date>
at 
<location id="906DC4A5FFF400244CA4F92FFBBF5ADB" LSID="urn:lsid:plazi:treatment:1D1B2368FFF400284E70FB0AFEB059EB:906DC4A5FFF400244CA4F92FFBBF5ADB" box="[921,1054,1703,1727]" country="Madagascar" county="Sofia Region" latitude="-14.35991" longLatPrecision="1" longitude="48.59022" municipality="Camp" name="Bemanevika" pageId="21" pageNumber="107" stateProvince="Mahajanga">Bemanevika</location>
, 
<collectingMunicipality id="75690804FFF400244B16F92FFBCA5ADB" box="[1067,1131,1703,1727]" pageId="21" pageNumber="107">Camp</collectingMunicipality>
2 (
<geoCoordinate id="F086F4B9FFF700274F9BFF06FE815CC2" box="[166,288,142,166]" degrees="14.35991" direction="south" orientation="latitude" pageId="22" pageNumber="108" precision="1" value="-14.35991">14.35991°S</geoCoordinate>
, 
<geoCoordinate id="F086F4B9FFF700274E14FF06FE035CC2" box="[297,418,142,166]" degrees="48.59022" direction="east" orientation="longitude" pageId="22" pageNumber="108" precision="1" value="48.59022">48.59022°E</geoCoordinate>
, 
<elevation id="1E9F754DFFF700274E96FF06FD0F5CC2" box="[427,686,142,166]" metricMagnitude="3" metricUnit="m" metricValue="1.538" pageId="22" pageNumber="108" unit="m" value="1538.0">
<quantity id="524A3F9BFFF700274E96FF06FE5C5CC2" box="[427,509,142,166]" metricMagnitude="3" metricUnit="m" metricValue="1.538" pageId="22" pageNumber="108" unit="m" value="1538.0">1538 m</quantity>
above sea level
</elevation>
), 
<collectingCounty id="7C6CEAF2FFF700274D83FF06FCE95CC2" box="[702,840,142,166]" pageId="22" pageNumber="108">Sofia Region</collectingCounty>
, former 
<collectingRegion id="57765C9CFFF700274C9FFF06FB215CC2" box="[930,1152,142,166]" country="Madagascar" name="Mahajanga" pageId="22" pageNumber="108">Mahajanga province</collectingRegion>
, 
<collectingCountry id="EDA5D2EEFFF700274F9DFF39FE855CAD" box="[160,292,177,201]" name="Madagascar" pageId="22" pageNumber="108">Madagascar</collectingCountry>
, by M
</materialsCitation>
. Vences, D. 
<materialsCitation id="25DA9823FFF700274EC5FF39FDFF5CAD" box="[504,606,177,201]" collectionCode="R" collectorName="Vieites" country="Madagascar" pageId="22" pageNumber="108" specimenCount="1" stateProvince="Mahajanga" typeStatus="paratype">
<collectorName id="3847F7A8FFF700274EC5FF39FDE05CAD" box="[504,577,177,201]" pageId="22" pageNumber="108">Vieites</collectorName>
, 
<collectionCode id="F3A30ABBFFF700274D73FF39FDFF5CAD" box="[590,606,177,201]" country="Chile" name="Departamento de Geologia, Universidad de Chile" pageId="22" pageNumber="108">R</collectionCode>
</materialsCitation>
. D. Randrianiaina, F. Ratsoavina, S. Rasamison, A. Rakotoarison, F. Randrianasolo, E. 
<materialsCitation id="25DA9823FFF700274D37FF5CFD105C88" box="[522,689,212,236]" collectionCode="T" collectorName="Rajeriarison" country="Madagascar" pageId="22" pageNumber="108" specimenCount="1" stateProvince="Mahajanga" typeStatus="paratype">
<collectorName id="3847F7A8FFF700274D37FF5CFD275C88" box="[522,646,212,236]" pageId="22" pageNumber="108">Rajeriarison</collectorName>
&amp; 
<collectionCode id="F3A30ABBFFF700274D9FFF5CFD105C88" box="[674,689,212,236]" name="Tavera, Department of Geology and Geophysics" pageId="22" pageNumber="108">T</collectionCode>
</materialsCitation>
. Rajoafiarison. 
<materialsCitation id="25DA9823FFF700274C72FF5CFCD95D12" collectingDate="2010-06-09" collectionCode="ZSM, DRV" country="Madagascar" county="Sofia Region" elevation="1162" latitude="-14.23989" location="Tsaratanana" longLatPrecision="1" longitude="48.97208" municipality="Ankijagna Lagnana" pageId="22" pageNumber="108" specimenCode="ZSM 581, DRV 6072, ZSM 582, DRV 6073, ZSM 583, DRV 6074, ZSM 584, DRV 6097" specimenCount="1" stateProvince="Mahajanga" typeStatus="paratype">
<specimenCode id="C5143A05FFF700274C72FF5CFC0A5C88" box="[847,939,212,236]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="22" pageNumber="108" type="Museum">ZSM 581</specimenCode>
/2014 (field number 
<specimenCode id="C5143A05FFF700274F9DFF7EFEAA5D6A" box="[160,267,246,270]" collectionCode="DRV" pageId="22" pageNumber="108">DRV 6072</specimenCode>
, female), 
<specimenCode id="C5143A05FFF700274E4CFF7EFE715D6A" box="[369,464,246,270]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="22" pageNumber="108" type="Museum">ZSM 582</specimenCode>
/2014 (field number 
<specimenCode id="C5143A05FFF700274D93FF7EFCB85D6A" box="[686,793,246,270]" collectionCode="DRV" pageId="22" pageNumber="108">DRV 6073</specimenCode>
, male), 
<specimenCode id="C5143A05FFF700274C56FF7EFC6B5D6A" box="[875,970,246,270]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="22" pageNumber="108" type="Museum">ZSM 583</specimenCode>
/2014 (
<specimenCode id="C5143A05FFF700274B2EFF7EFB215D6A" box="[1043,1152,246,270]" collectionCode="DRV" pageId="22" pageNumber="108">DRV 6074</specimenCode>
, male) and 
<specimenCode id="C5143A05FFF700274E23FE91FE235D55" box="[286,386,281,305]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="22" pageNumber="108" type="Museum">ZSM 584</specimenCode>
/2014 (field number 
<specimenCode id="C5143A05FFF700274D4DFE91FD405D55" box="[624,737,281,305]" collectionCode="DRV" pageId="22" pageNumber="108">DRV 6097</specimenCode>
, male), collected on 
<date id="E10CB4BEFFF700274CE9FE91FBC35D55" box="[980,1122,281,305]" pageId="22" pageNumber="108" value="2010-06-09">
<collectingDate id="F1484D56FFF700274CE9FE91FBC35D55" box="[980,1122,281,305]" pageId="22" pageNumber="108" value="2010-06-09">9 June 2010</collectingDate>
</date>
at 
<location id="906DC4A5FFF700274F9DFEB4FE835D30" LSID="urn:lsid:plazi:treatment:1D1B2368FFF400284E70FB0AFEB059EB:906DC4A5FFF700274F9DFEB4FE835D30" box="[160,290,316,340]" country="Madagascar" county="Sofia Region" latitude="-14.23989" longLatPrecision="1" longitude="48.97208" municipality="Ankijagna Lagnana" name="Tsaratanana" pageId="22" pageNumber="108" stateProvince="Mahajanga">Tsaratanana</location>
massif, camp 0 (
<collectingMunicipality id="75690804FFF700274EE5FEB4FD0D5D30" box="[472,684,316,340]" pageId="22" pageNumber="108">Ankijagna Lagnana</collectingMunicipality>
, 
<geoCoordinate id="F086F4B9FFF700274D8BFEB4FC8F5D30" box="[694,814,316,340]" degrees="14.23989" direction="south" orientation="latitude" pageId="22" pageNumber="108" precision="1" value="-14.23989">14.23989°S</geoCoordinate>
, 
<geoCoordinate id="F086F4B9FFF700274C05FEB4FC105D30" box="[824,945,316,340]" degrees="48.97208" direction="east" orientation="longitude" pageId="22" pageNumber="108" precision="1" value="48.97208">48.97208°E</geoCoordinate>
, 
<elevation id="1E9F754DFFF700274C86FEB4FF745D12" metricMagnitude="3" metricUnit="m" metricValue="1.162" pageId="22" pageNumber="108" unit="m" value="1162.0">
<quantity id="524A3F9BFFF700274C86FEB4FBAF5D30" box="[955,1038,316,340]" metricMagnitude="3" metricUnit="m" metricValue="1.162" pageId="22" pageNumber="108" unit="m" value="1162.0">1162 m</quantity>
above sea level
</elevation>
), 
<collectingCounty id="7C6CEAF2FFF700274FDBFED6FED15D12" box="[230,368,350,374]" pageId="22" pageNumber="108">Sofia Region</collectingCounty>
, former 
<collectingRegion id="57765C9CFFF700274EF6FED6FD0B5D12" box="[459,682,350,374]" country="Madagascar" name="Mahajanga" pageId="22" pageNumber="108">Mahajanga province</collectingRegion>
, 
<collectingCountry id="EDA5D2EEFFF700274D88FED6FC985D12" box="[693,825,350,374]" name="Madagascar" pageId="22" pageNumber="108">Madagascar</collectingCountry>
, by D
</materialsCitation>
. Vieites, M. 
<materialsCitation id="25DA9823FFF700274CC4FED6FBC05D12" box="[1017,1121,350,374]" collectionCode="R" collectorName="Vences" country="Madagascar" pageId="22" pageNumber="108" specimenCount="1" stateProvince="Mahajanga" typeStatus="paratype">
<collectorName id="3847F7A8FFF700274CC4FED6FBE95D12" box="[1017,1096,350,374]" pageId="22" pageNumber="108">Vences</collectorName>
, 
<collectionCode id="F3A30ABBFFF700274B6CFED6FBC05D12" box="[1105,1121,350,374]" country="Chile" name="Departamento de Geologia, Universidad de Chile" pageId="22" pageNumber="108">R</collectionCode>
</materialsCitation>
. D. Randrianiaina, S. Rasamison, A. Rakotoarison, F. Randrianasolo, E. 
<materialsCitation id="25DA9823FFF700274C82FE09FB215DFD" box="[959,1152,385,409]" collectionCode="T" collectorName="Rajeriarison" country="Madagascar" pageId="22" pageNumber="108" specimenCount="1" stateProvince="Mahajanga" typeStatus="paratype">
<collectorName id="3847F7A8FFF700274C82FE09FB9A5DFD" box="[959,1083,385,409]" pageId="22" pageNumber="108">Rajeriarison</collectorName>
&amp; 
<collectionCode id="F3A30ABBFFF700274B4CFE09FB215DFD" box="[1137,1152,385,409]" name="Tavera, Department of Geology and Geophysics" pageId="22" pageNumber="108">T</collectionCode>
</materialsCitation>
. Rajoafiarison. 
<materialsCitation id="25DA9823FFF700274E07FE2CFC415E65" collectingDate="2016-01-13" collectionCode="ZSM, MSZC" country="Madagascar" elevation="1038" latitude="-14.71788" location="Ampandrana forest" longLatPrecision="1" longitude="48.57548" municipality="Sofia Region" pageId="22" pageNumber="108" specimenCode="ZSM 95, MSZC 0189" specimenCount="1" stateProvince="Mahajanga" typeStatus="paratype">
<specimenCode id="C5143A05FFF700274E07FE2CFE2E5DD8" box="[314,399,420,444]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="22" pageNumber="108" type="Museum">ZSM 95</specimenCode>
/2016 (field number 
<specimenCode id="C5143A05FFF700274D41FE2CFCA15DD8" box="[636,768,420,444]" collectionCode="MSZC" pageId="22" pageNumber="108">MSZC 0189</specimenCode>
), collected on 
<date id="E10CB4BEFFF700274C94FE2CFBC35DD8" box="[937,1122,420,444]" pageId="22" pageNumber="108" value="2016-01-13">
<collectingDate id="F1484D56FFF700274C94FE2CFBC35DD8" box="[937,1122,420,444]" pageId="22" pageNumber="108" value="2016-01-13">13 January 2016</collectingDate>
</date>
at 
<location id="906DC4A5FFF700274F9DFE4EFED05DBA" LSID="urn:lsid:plazi:treatment:1D1B2368FFF400284E70FB0AFEB059EB:906DC4A5FFF700274F9DFE4EFED05DBA" box="[160,369,454,478]" country="Madagascar" latitude="-14.71788" longLatPrecision="1" longitude="48.57548" municipality="Sofia Region" name="Ampandrana forest" pageId="22" pageNumber="108" stateProvince="Mahajanga">Ampandrana forest</location>
(
<locationDeviation id="84BF8B5CFFF700274E43FE4EFD335DBA" box="[382,658,454,478]" location="Bealanana" pageId="22" pageNumber="108">
<quantity id="524A3F9BFFF700274E43FE4EFE745DBA" box="[382,469,454,478]" metricMagnitude="4" metricUnit="m" metricValue="2.66" pageId="22" pageNumber="108" unit="km" value="26.6">26.6 km</quantity>
SW of Bealanana
</locationDeviation>
on the RN31, 
<geoCoordinate id="F086F4B9FFF700274C10FE4EFC045DBA" box="[813,933,454,478]" degrees="14.71788" direction="south" orientation="latitude" pageId="22" pageNumber="108" precision="1" value="-14.71788">14.71788°S</geoCoordinate>
, 
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, 
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above sea level
</elevation>
), 
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, former 
<collectingRegion id="57765C9CFFF700274D0BFE61FCB25E65" box="[566,787,489,513]" country="Madagascar" name="Mahajanga" pageId="22" pageNumber="108">Mahajanga province</collectingRegion>
, 
<collectingCountry id="EDA5D2EEFFF700274C21FE61FC3E5E65" box="[796,927,489,513]" name="Madagascar" pageId="22" pageNumber="108">Madagascar</collectingCountry>
, by M
</materialsCitation>
. D. Scherz &amp; M. Rakotondratsima. 
<materialsCitation id="25DA9823FFF700274E5EFD84FC415E0D" collectingDate="2016-01-14" collectionCode="ZSM, MSZC" country="Madagascar" elevation="1204" latitude="-14.73654" location="Andranonafindra forest" longLatPrecision="1" longitude="48.54846" municipality="Sofia Region" pageId="22" pageNumber="108" specimenCode="ZSM 96, MSZC 0205" specimenCount="1" stateProvince="Mahajanga" typeStatus="paratype">
<specimenCode id="C5143A05FFF700274E5EFD84FE145E40" box="[355,437,524,548]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="22" pageNumber="108" type="Museum">ZSM 96</specimenCode>
/2016 (field number 
<specimenCode id="C5143A05FFF700274DAAFD84FCB75E40" box="[663,790,524,548]" collectionCode="MSZC" pageId="22" pageNumber="108">MSZC 0205</specimenCode>
), collected on 
<date id="E10CB4BEFFF700274C89FD84FBC75E40" box="[948,1126,524,548]" pageId="22" pageNumber="108" value="2016-01-14">
<collectingDate id="F1484D56FFF700274C89FD84FBC75E40" box="[948,1126,524,548]" pageId="22" pageNumber="108" value="2016-01-14">14 January 2016</collectingDate>
</date>
at 
<location id="906DC4A5FFF700274F9DFDA6FE385E22" LSID="urn:lsid:plazi:treatment:1D1B2368FFF400284E70FB0AFEB059EB:906DC4A5FFF700274F9DFDA6FE385E22" box="[160,409,558,582]" country="Madagascar" latitude="-14.73654" longLatPrecision="1" longitude="48.54846" municipality="Sofia Region" name="Andranonafindra forest" pageId="22" pageNumber="108" stateProvince="Mahajanga">Andranonafindra forest</location>
(
<locationDeviation id="84BF8B5CFFF700274E99FDA6FD3E5E22" box="[420,671,558,582]" location="Bealanana" pageId="22" pageNumber="108">
<quantity id="524A3F9BFFF700274E99FDA6FE495E22" box="[420,488,558,582]" metricMagnitude="4" metricUnit="m" metricValue="3.0" pageId="22" pageNumber="108" unit="km" value="30.0">30 km</quantity>
SW of Bealanana
</locationDeviation>
on the RN31, 
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, 
<geoCoordinate id="F086F4B9FFF700274C8EFDA6FB8A5E22" box="[947,1067,558,582]" degrees="48.54846" direction="east" orientation="longitude" pageId="22" pageNumber="108" precision="1" value="48.54846">48.54846°E</geoCoordinate>
, 
<elevation id="1E9F754DFFF700274B0EFDA6FEE65E0D" metricMagnitude="3" metricUnit="m" metricValue="1.204" pageId="22" pageNumber="108" unit="m" value="1204.0">
<quantity id="524A3F9BFFF700274B0EFDA6FB225E22" box="[1075,1155,558,582]" metricMagnitude="3" metricUnit="m" metricValue="1.204" pageId="22" pageNumber="108" unit="m" value="1204.0">1204 m</quantity>
above sea level
</elevation>
), 
<collectingMunicipality id="75690804FFF700274E6BFDD9FE7E5E0D" box="[342,479,593,617]" pageId="22" pageNumber="108">Sofia Region</collectingMunicipality>
, former 
<collectingRegion id="57765C9CFFF700274D0BFDD9FCB25E0D" box="[566,787,593,617]" country="Madagascar" name="Mahajanga" pageId="22" pageNumber="108">Mahajanga province</collectingRegion>
, 
<collectingCountry id="EDA5D2EEFFF700274C21FDD9FC3E5E0D" box="[796,927,593,617]" name="Madagascar" pageId="22" pageNumber="108">Madagascar</collectingCountry>
, by M
</materialsCitation>
. D. Scherz &amp; M. Rakotondratsima. 
<materialsCitation id="25DA9823FFF700274E5CFDFBFED75E97" collectingDate="2015-12-22" collectionCode="ZSM, MSZC" country="Madagascar" county="Sofia Region" elevation="1400" latitude="-14.41456" location="Ampotsidy mountains" longLatPrecision="1" longitude="48.71148" municipality="Beandrarezona" pageId="22" pageNumber="108" specimenCode="ZSM 97, MSZC 0080" specimenCount="1" stateProvince="Mahajanga" typeStatus="paratype">
<specimenCode id="C5143A05FFF700274E5CFDFBFE105EEF" box="[353,433,627,651]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="22" pageNumber="108" type="Museum">ZSM 97</specimenCode>
/2016 (field number 
<specimenCode id="C5143A05FFF700274DB6FDFBFCA65EEF" box="[651,775,627,651]" collectionCode="MSZC" pageId="22" pageNumber="108">MSZC 0080</specimenCode>
, female), collected on 
<date id="E10CB4BEFFF700274CCEFDFBFF745ECA" pageId="22" pageNumber="108" value="2015-12-22">
<collectingDate id="F1484D56FFF700274CCEFDFBFF745ECA" pageId="22" pageNumber="108" value="2015-12-22">22 December 2015</collectingDate>
</date>
on 
<location id="906DC4A5FFF700274E3DFD1EFE505ECA" LSID="urn:lsid:plazi:treatment:1D1B2368FFF400284E70FB0AFEB059EB:906DC4A5FFF700274E3DFD1EFE505ECA" box="[256,497,662,686]" country="Madagascar" county="Sofia Region" latitude="-14.41456" longLatPrecision="1" longitude="48.71148" municipality="Beandrarezona" name="Ampotsidy mountains" pageId="22" pageNumber="108" stateProvince="Mahajanga">Ampotsidy mountains</location>
, 
<locationDeviation id="84BF8B5CFFF700274EC6FD1EFC825ECA" box="[507,803,662,686]" location="Bealanana" pageId="22" pageNumber="108">
<quantity id="524A3F9BFFF700274EC6FD1EFDF05ECA" box="[507,593,662,686]" metricMagnitude="4" metricUnit="m" metricValue="1.5699999999999998" pageId="22" pageNumber="108" unit="km" value="15.7">15.7 km</quantity>
NNW of Bealanana
</locationDeviation>
(
<quantity id="524A3F9BFFF700274C12FD1EFCD85ECA" box="[815,889,662,686]" metricMagnitude="3" metricUnit="m" metricValue="8.7" pageId="22" pageNumber="108" unit="km" value="8.7">8.7 km</quantity>
NNW of 
<collectingMunicipality id="75690804FFF700274CE1FD1EFB215ECA" box="[988,1152,662,686]" pageId="22" pageNumber="108">Beandrarezona</collectingMunicipality>
, 
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, 
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, 
<elevation id="1E9F754DFFF700274E9BFD31FD0A5EB5" box="[422,683,697,721]" metricMagnitude="3" metricUnit="m" metricValue="1.4" pageId="22" pageNumber="108" unit="m" value="1400.0">
<quantity id="524A3F9BFFF700274E9BFD31FE595EB5" box="[422,504,697,721]" metricMagnitude="3" metricUnit="m" metricValue="1.4" pageId="22" pageNumber="108" unit="m" value="1400.0">1400 m</quantity>
above sea level
</elevation>
), 
<collectingCounty id="7C6CEAF2FFF700274D81FD31FCE75EB5" box="[700,838,697,721]" pageId="22" pageNumber="108">Sofia Region</collectingCounty>
, former 
<collectingRegion id="57765C9CFFF700274C9CFD31FB215EB5" box="[929,1152,697,721]" country="Madagascar" name="Mahajanga" pageId="22" pageNumber="108">Mahajanga province</collectingRegion>
, 
<collectingCountry id="EDA5D2EEFFF700274F9DFD53FE855E97" box="[160,292,731,755]" name="Madagascar" pageId="22" pageNumber="108">Madagascar</collectingCountry>
, by M
</materialsCitation>
. D. Scherz, J. Borrell, L. 
<materialsCitation id="25DA9823FFF700274D9DFD53FD485E97" box="[672,745,731,755]" collectionCode="T" collectorName="Ball" country="Madagascar" pageId="22" pageNumber="108" specimenCount="1" stateProvince="Mahajanga" typeStatus="paratype">
<collectorName id="3847F7A8FFF700274D9DFD53FD695E97" box="[672,712,731,755]" pageId="22" pageNumber="108">Ball</collectorName>
, 
<collectionCode id="F3A30ABBFFF700274DE7FD53FD485E97" box="[730,745,731,755]" name="Tavera, Department of Geology and Geophysics" pageId="22" pageNumber="108">T</collectionCode>
</materialsCitation>
. Starnes, E. Razafimandimby, D. H. Nomenjanahary &amp; J. Rabearivony.
</paragraph>
</subSubSection>
<subSubSection id="DDA8C1F5FFF700274F9DFCCCFCCA5A33" pageId="22" pageNumber="108" type="diagnosis">
<paragraph id="950D927EFFF700274F9DFCCCFEB55F38" blockId="22.[160,1156,836,1623]" box="[160,276,836,860]" pageId="22" pageNumber="108">
<emphasis id="A7C64E6CFFF700274F9DFCCCFEB55F38" bold="true" box="[160,276,836,860]" italics="true" pageId="22" pageNumber="108">Diagnosis</emphasis>
</paragraph>
<paragraph id="950D927EFFF700274F9DFCEFFD5958D3" blockId="22.[160,1156,836,1623]" pageId="22" pageNumber="108">
The new species is assigned to the genus 
<taxonomicName id="52B2E9FDFFF700274DACFCEEFC865F1B" authorityName="Boulenger" authorityYear="1895" box="[657,807,870,895]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="22" pageNumber="108" phylum="Chordata" rank="genus">
<emphasis id="A7C64E6CFFF700274DACFCEEFC865F1B" box="[657,807,870,895]" italics="true" pageId="22" pageNumber="108">Mantidactylus</emphasis>
</taxonomicName>
based on the presence of an intercalary element between terminal and subterminal phalanges of fingers and toes (verified by external observation only), of a central depression in femoral glands and of a (very) rudimentary femoral gland in the female. Within 
<taxonomicName id="52B2E9FDFFF700274C3BFC46FC3A5F83" authorityName="Boulenger" authorityYear="1895" box="[774,923,974,999]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="22" pageNumber="108" phylum="Chordata" rank="genus">
<emphasis id="A7C64E6CFFF700274C3BFC46FC3A5F83" box="[774,923,974,999]" italics="true" pageId="22" pageNumber="108">Mantidactylus</emphasis>
</taxonomicName>
, it is assigned to the nominal subgenus 
<taxonomicName id="52B2E9FDFFF700274E4EFC78FDA8586D" authorityName="Boulenger" authorityYear="1895" box="[371,521,1008,1033]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="22" pageNumber="108" phylum="Chordata" rank="genus">
<emphasis id="A7C64E6CFFF700274E4EFC78FDA8586D" box="[371,521,1008,1033]" italics="true" pageId="22" pageNumber="108">Mantidactylus</emphasis>
</taxonomicName>
by combination of (1) very large body size (male SVL 
<quantity id="524A3F9BFFF700274B66FC79FF4E5848" metricMagnitude="-2" metricUnit="m" metricValue="9.05" metricValueMax="9.3" metricValueMin="8.8" pageId="22" pageNumber="108" unit="mm" value="90.5" valueMax="93.0" valueMin="88.0">88– 93 mm</quantity>
, female SVL 
<quantity id="524A3F9BFFF700274E45FB9CFE5F5848" box="[376,510,1044,1068]" metricMagnitude="-2" metricUnit="m" metricValue="9.4" metricValueMax="10.3" metricValueMin="8.5" pageId="22" pageNumber="108" unit="mm" value="94.0" valueMax="103.0" valueMin="85.0">85–103 mm</quantity>
), (2) absence of dorsolateral colour border, (3) absence of a distinct frenal stripe, (4) absence of large yellowish patches or stripes in the inguinal region or between colouration of flanks and belly, (5) fully webbed feet and (6) riparian habits, living very close to or in streams. The assignment of the species to this group is also supported by its molecular phylogenetic relationships.
</paragraph>
<paragraph id="950D927EFFF700274FFDFB49FE155898" blockId="22.[160,1156,836,1623]" pageId="22" pageNumber="108">
From the two described species in the subgenus 
<taxonomicName id="52B2E9FDFFF700274C39FB48FC3C58BD" authorityName="Boulenger" authorityYear="1895" box="[772,925,1216,1241]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="22" pageNumber="108" phylum="Chordata" rank="genus">
<emphasis id="A7C64E6CFFF700274C39FB48FC3C58BD" box="[772,925,1216,1241]" italics="true" pageId="22" pageNumber="108">Mantidactylus</emphasis>
</taxonomicName>
, the new species is distinguished as follows:
</paragraph>
<paragraph id="950D927EFFF700274FFDFA8FFB2559A8" blockId="22.[160,1156,836,1623]" pageId="22" pageNumber="108">
From 
<taxonomicName id="52B2E9FDFFF700274E3DFA8EFE2E597B" baseAuthorityName="Boulenger" baseAuthorityYear="1881" box="[256,399,1286,1311]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="22" pageNumber="108" phylum="Chordata" rank="species" species="guttulatus">
<emphasis id="A7C64E6CFFF700274E3DFA8EFE2E597B" box="[256,399,1286,1311]" italics="true" pageId="22" pageNumber="108">M. guttulatus</emphasis>
</taxonomicName>
by a longer relative hindlimb length (HIL/SVL 1.46–1.60 versus 1.28– 1.43); larger femoral glands in males (FGL/SVL 0.13–0.20 versus 0.06–0.08), which are ovoid (versus round in 
<taxonomicName id="52B2E9FDFFF700274E5EFAC3FE545900" baseAuthorityName="Boulenger" baseAuthorityYear="1881" box="[355,501,1355,1380]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="22" pageNumber="108" phylum="Chordata" rank="species" species="guttulatus">
<emphasis id="A7C64E6CFFF700274E5EFAC3FE545900" box="[355,501,1355,1380]" italics="true" pageId="22" pageNumber="108">M. guttulatus</emphasis>
</taxonomicName>
), and indistinct femoral glands in females (versus always recognisable in 
<taxonomicName id="52B2E9FDFFF700274E6BFAE6FE4959E3" baseAuthorityName="Boulenger" baseAuthorityYear="1881" box="[342,488,1390,1415]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="22" pageNumber="108" phylum="Chordata" rank="species" species="guttulatus">
<emphasis id="A7C64E6CFFF700274E6BFAE6FE4959E3" box="[342,488,1390,1415]" italics="true" pageId="22" pageNumber="108">M. guttulatus</emphasis>
</taxonomicName>
); and possibly by an often-hidden tympanum and smaller relative tympanum size (TD/SVL 
<geoCoordinate id="F086F4B9FFF700274D2DFA19FDED59CD" box="[528,588,1425,1449]" degrees="0.032" direction="north" orientation="latitude" pageId="22" pageNumber="108" precision="55" value="0.032">0.032</geoCoordinate>
<geoCoordinate id="F086F4B9FFF700274D71FA19FD3759CD" box="[588,662,1425,1449]" degrees="0.047" direction="west" orientation="longitude" pageId="22" pageNumber="108" precision="55" value="-0.047">–0.047</geoCoordinate>
versus 
<geoCoordinate id="F086F4B9FFF700274DD1FA19FC8959CD" box="[748,808,1425,1449]" degrees="0.052" direction="north" orientation="latitude" pageId="22" pageNumber="108" precision="55" value="0.052">0.052</geoCoordinate>
<geoCoordinate id="F086F4B9FFF700274C14FA19FCD559CD" box="[809,884,1425,1449]" degrees="0.082" direction="west" orientation="longitude" pageId="22" pageNumber="108" precision="55" value="-0.082">–0.082</geoCoordinate>
) [however not applying to the 
<taxonomicName id="52B2E9FDFFF700274FD4FA3BFED959A8" baseAuthorityName="Boulenger" baseAuthorityYear="1881" box="[233,376,1459,1484]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="22" pageNumber="108" phylum="Chordata" rank="species" species="guttulatus">
<emphasis id="A7C64E6CFFF700274FD4FA3BFED959A8" box="[233,376,1459,1484]" italics="true" pageId="22" pageNumber="108">M. guttulatus</emphasis>
</taxonomicName>
<typeStatus id="4A092CDCFFF700274E42FA3CFE1059A8" box="[383,433,1460,1484]" pageId="22" pageNumber="108">type</typeStatus>
specimens (
<geoCoordinate id="F086F4B9FFF700274D04FA3CFDD659A8" box="[569,631,1460,1484]" degrees="0.035" direction="north" orientation="latitude" pageId="22" pageNumber="108" precision="55" value="0.035">0.035</geoCoordinate>
<geoCoordinate id="F086F4B9FFF700274D4AFA3CFD6459A8" box="[631,709,1460,1484]" degrees="0.049" direction="west" orientation="longitude" pageId="22" pageNumber="108" precision="55" value="-0.049">–0.049</geoCoordinate>
), possibly due to preservation artefacts].
</paragraph>
<paragraph id="950D927EFFF700274FFDFA5FFCCA5A33" blockId="22.[160,1156,836,1623]" pageId="22" pageNumber="108">
From 
<taxonomicName id="52B2E9FDFFF700274E3DFA5EFE37598B" authorityName="Mocquard" authorityYear="1895" box="[256,406,1494,1519]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="22" pageNumber="108" phylum="Chordata" rank="species" species="grandidieri">
<emphasis id="A7C64E6CFFF700274E3DFA5EFE37598B" box="[256,406,1494,1519]" italics="true" pageId="22" pageNumber="108">M. grandidieri</emphasis>
</taxonomicName>
by an often-hidden tympanum and smaller relative tympanum size (TD/SVL 
<geoCoordinate id="F086F4B9FFF700274E3CFA71FE9C5A75" box="[257,317,1529,1553]" degrees="0.032" direction="north" orientation="latitude" pageId="22" pageNumber="108" precision="55" value="0.032">0.032</geoCoordinate>
<geoCoordinate id="F086F4B9FFF700274E00FA71FE275A75" box="[317,390,1529,1553]" degrees="0.047" direction="west" orientation="longitude" pageId="22" pageNumber="108" precision="55" value="-0.047">–0.047</geoCoordinate>
versus 
<geoCoordinate id="F086F4B9FFF700274EE3FA71FDBB5A75" box="[478,538,1529,1553]" degrees="0.055" direction="north" orientation="latitude" pageId="22" pageNumber="108" precision="55" value="0.055">0.055</geoCoordinate>
<geoCoordinate id="F086F4B9FFF700274D27FA71FDC95A75" box="[538,616,1529,1553]" degrees="0.063" direction="west" orientation="longitude" pageId="22" pageNumber="108" precision="55" value="-0.063">–0.063</geoCoordinate>
); larger femoral glands in males (FGL/SVL 0.13– 0.20 versus 0.09); which are ovoid (versus round in 
<taxonomicName id="52B2E9FDFFF700274DDDF993FCD65A50" authorityName="Mocquard" authorityYear="1895" box="[736,887,1563,1588]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="22" pageNumber="108" phylum="Chordata" rank="species" species="grandidieri">
<emphasis id="A7C64E6CFFF700274DDDF993FCD65A50" box="[736,887,1563,1588]" italics="true" pageId="22" pageNumber="108">M. grandidieri</emphasis>
</taxonomicName>
); and indistinct femoral glands in females (versus always recognisable in 
<taxonomicName id="52B2E9FDFFF700274DFAF9B6FCFF5A33" authorityName="Mocquard" authorityYear="1895" box="[711,862,1598,1623]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="22" pageNumber="108" phylum="Chordata" rank="species" species="grandidieri">
<emphasis id="A7C64E6CFFF700274DFAF9B6FCFF5A33" box="[711,862,1598,1623]" italics="true" pageId="22" pageNumber="108">M. grandidieri</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="DDA8C1F5FFF700264F9DF90CFD175F72" lastPageId="23" lastPageNumber="109" pageId="22" pageNumber="108" type="description">
<paragraph id="950D927EFFF700274F9DF90CFB255ADB" blockId="22.[160,1156,1668,1727]" pageId="22" pageNumber="108">
<emphasis id="A7C64E6CFFF700274F9DF90CFE495AF8" bold="true" box="[160,488,1668,1692]" italics="true" pageId="22" pageNumber="108">
Description of the 
<typeStatus id="4A092CDCFFF700274E44F90CFE455AF8" box="[377,484,1668,1692]" pageId="22" pageNumber="108" type="holotype">holotype</typeStatus>
.
</emphasis>
Adult male in excellent state of preservation. For measurements, see 
<tableCitation id="D830A7C5FFF700274E1EF92FFED65ADB" box="[291,375,1703,1727]" captionStart="Table 1" captionStartId="8.[133,189,535,558]" captionText="Table 1. Morphometric measurements of specimens of the two nominal species Mantidactylus guttulatus and M. grandidieri, and the newly described species M. radaka sp. nov. (all in mm). Asterisks indicate name-bearing type specimens:Holotype of M. radaka sp. nov., syntypes of M. grandidieri and M. guttulatus, as well as the holotype of Rana pigra (here considered as junior synonym of M. guttulatus). Acronyms: M = male, F = female, SA = Sub-adult." pageId="22" pageNumber="108">Table 1</tableCitation>
. Body rather stout. Head wider than long, slightly wider than body.
</paragraph>
<paragraph id="950D927EFFF600264F9DFF06FC5E5E0D" blockId="23.[160,1156,142,790]" pageId="23" pageNumber="109">Snout blunt. Nostrils directed dorsolaterally, slightly protuberant, nearer to tip of snout than to eye. Canthus rostralis poorly distinct, loreal region concave. Upper part of the tympanum hidden under the tympanic fold. Tympanum small, horizonal diameter of tympanum 34% of horizontal eye diameter. Supratympanic fold distinct, straight above the tympanum, curving gently towards the axilla. Tongue ovoid, distinctly bifid posteriorly. Vomerine teeth form two rounded aggregations, positioned posterolateral to choanae. Choanae ovoid. Subarticular tubercles single. Outer metacarpal tubercle not recognisable, inner metacarpal tubercle present and distinct. Fingers without webbing. Relative length of fingers: I&lt;II&lt;IV&lt;III. Finger discs enlarged. Nuptial pads absent. Foot slightly shorter than tibia (97%). Lateral metatarsalia separated. Inner metatarsal tubercle present. Outer metatarsal tubercle not recognisable. Webbing formula: 1(0), 2i(0.5), 2e(0), 3i(1), 3e(0), 4i(1), 4e(0.5), 5(0). Relative length of toes: I&lt;II&lt;III=V&lt;IV. Skin on the upper surface strongly rugose with numerous small warts, these becoming larger on the flanks. Ventral side smooth. Femoral glands very large and distinct in external view.</paragraph>
<paragraph id="950D927EFFF600264FFDFDFBFD175F72" blockId="23.[160,1156,142,790]" pageId="23" pageNumber="109">
Colour in preservative (after 18 years in preservative) dorsally brown, with tiny yellowish dots on the warts. Venter beige, with indistinct mottling on the throat. The inner three fingers are lighter than the fourth one. Colouration in life (
<figureCitation id="0D898EFBFFF600264C6DFD31FC0E5EB5" box="[848,943,697,721]" captionStart="Figure 6" captionStartId="16.[160,225,1063,1086]" captionTargetBox="[166,1149,142,1031]" captionTargetId="figure-144@16.[166,1149,142,1031]" captionTargetPageId="16" captionText="Figure 6. Photographs of living specimens of Mantidactylus radaka sp. nov. (a, b) Male holotype ZSM 644/2001 (field number FGMV 2001.132) from Manarikoba forest, Tsaratanana Massif. (c–f) Female paratype ZSM 1800/2010 (ZCMV 12345) from Camp 1 (Antevialambazaha), Tsaratanana Massif. (g, h) Female paratype ZSM 97/2016 (MSZC 0080) from Ampotsidy. (i, j) Male paratype MSZC 0120 (uncatalogued in UADBA) from Ampotsidy. (k) Unidentified specimen from Camp 0 (Ankijagna Lagnana), Tsaratanana Massif. (l) Paratype ZSM 582/2014 (DRV 6073) from Camp 0 (Ankijagna Lagnana). (m, n) Unidentified female specimen from Manongarivo (Camp 0), probably preserved in UADBA collection." figureDoi="http://doi.org/10.5281/zenodo.5020665" httpUri="https://zenodo.org/record/5020665/files/figure.png" pageId="23" pageNumber="109">Figure 6</figureCitation>
) similar to that in preservative, with larger yellowish dots particularly on the flanks. Uniformly greenishbrown iris. Light tips on the inner three fingers.
</paragraph>
</subSubSection>
<subSubSection id="DDA8C1F5FFF600264F9DFCCBFC325FC5" pageId="23" pageNumber="109" type="etymology">
<paragraph id="950D927EFFF600264F9DFCCBFC325FC5" blockId="23.[160,1155,835,929]" pageId="23" pageNumber="109">
<emphasis id="A7C64E6CFFF600264F9DFCCBFE855F3F" bold="true" box="[160,292,835,859]" italics="true" pageId="23" pageNumber="109">Etymology.</emphasis>
The species name is a noun in apposition to the genus name, and is the Malagasy word for large frogs in general (as opposed to ‘sahona’, which usually refers to small frogs), and those of the subgenus 
<taxonomicName id="52B2E9FDFFF600264D58FC00FD5F5FC5" authorityName="Boulenger" authorityYear="1895" box="[613,766,904,929]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="23" pageNumber="109" phylum="Chordata" rank="genus">
<emphasis id="A7C64E6CFFF600264D58FC00FD5F5FC5" box="[613,766,904,929]" italics="true" pageId="23" pageNumber="109">Mantidactylus</emphasis>
</taxonomicName>
in particular.
</paragraph>
</subSubSection>
<subSubSection id="DDA8C1F5FFF600264F9DFC46FDFC5A33" pageId="23" pageNumber="109" type="description">
<paragraph id="950D927EFFF600264F9DFC46FC5B58D3" blockId="23.[160,1156,974,1207]" pageId="23" pageNumber="109">
<emphasis id="A7C64E6CFFF600264F9DFC46FEB45F82" bold="true" box="[160,277,974,998]" italics="true" pageId="23" pageNumber="109">Variation.</emphasis>
No obvious difference in tympanum diameter between males and females (TD/SVL female 
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<geoCoordinate id="F086F4B9FFF600264EA5FC79FE45586D" box="[408,484,1009,1033]" degrees="0.043" direction="west" orientation="longitude" pageId="23" pageNumber="109" precision="55" value="-0.043">–0.043</geoCoordinate>
, male 
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). The 
<typeStatus id="4A092CDCFFF600264C37FC79FCD9586D" box="[778,888,1009,1033]" pageId="23" pageNumber="109" type="paratype">paratypes</typeStatus>
and additional photographed specimens in general agree with the 
<typeStatus id="4A092CDCFFF600264D93FB9CFCB35848" box="[686,786,1044,1068]" pageId="23" pageNumber="109" type="holotype">holotype</typeStatus>
in morphology and colouration. In preserved females, femoral glands are not recognisable externally. In life, their femoral glands are visible as tiny black patches (cf. photos; 
<figureCitation id="0D898EFBFFF600264DF3FBD1FCF25815" box="[718,851,1113,1137]" captionStart="Figure 6" captionStartId="16.[160,225,1063,1086]" captionTargetBox="[166,1149,142,1031]" captionTargetId="figure-144@16.[166,1149,142,1031]" captionTargetPageId="16" captionText="Figure 6. Photographs of living specimens of Mantidactylus radaka sp. nov. (a, b) Male holotype ZSM 644/2001 (field number FGMV 2001.132) from Manarikoba forest, Tsaratanana Massif. (c–f) Female paratype ZSM 1800/2010 (ZCMV 12345) from Camp 1 (Antevialambazaha), Tsaratanana Massif. (g, h) Female paratype ZSM 97/2016 (MSZC 0080) from Ampotsidy. (i, j) Male paratype MSZC 0120 (uncatalogued in UADBA) from Ampotsidy. (k) Unidentified specimen from Camp 0 (Ankijagna Lagnana), Tsaratanana Massif. (l) Paratype ZSM 582/2014 (DRV 6073) from Camp 0 (Ankijagna Lagnana). (m, n) Unidentified female specimen from Manongarivo (Camp 0), probably preserved in UADBA collection." figureDoi="http://doi.org/10.5281/zenodo.5020665" httpUri="https://zenodo.org/record/5020665/files/figure.png" pageId="23" pageNumber="109">Figure 6F, H</figureCitation>
). Some females (ZSM 1800/ 
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particular) have very dark throat colouration. In a young male (ZSM 582/2014), femoral pores are already distinct (both externally and internally) and ovoid.
</paragraph>
<paragraph id="950D927EFFF600264F9DFB6CFDFC5A33" blockId="23.[160,1156,1252,1623]" pageId="23" pageNumber="109">
<emphasis id="A7C64E6CFFF600264F9DFB6CFEC05898" bold="true" box="[160,353,1252,1276]" italics="true" pageId="23" pageNumber="109">Natural History.</emphasis>
A large nocturnal frog typically found in relatively calm stretches of streams, including small and shallow streams with sandy bottom in rainforest (e.g., Tsaratanana), and more rarely also in shallow stretches of wider streams with rocky bottom in more open areas at the edge of rainforest (e.g., near Bemanevika). In Besariaka, individuals of this species also showed an apparently distinct natural history and behaviour, preferring running water (usually moving in the central parts of streams), while 
<taxonomicName id="52B2E9FDFFF600264CB9FA19FBB759CE" authorityName="Mocquard" authorityYear="1895" box="[900,1046,1425,1450]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="23" pageNumber="109" phylum="Chordata" rank="species" species="grandidieri">
<emphasis id="A7C64E6CFFF600264CB9FA19FBB759CE" box="[900,1046,1425,1450]" italics="true" pageId="23" pageNumber="109">M. grandidieri</emphasis>
</taxonomicName>
appeared associated more closely with stream banks. 
<taxonomicName id="52B2E9FDFFF600264D47FA3BFCC159A8" authority="Rancilhac &amp; Bruy &amp; Scherz &amp; Pereira &amp; Preick &amp; Straube &amp; Lyra &amp; Ohler &amp; Streicher &amp; Andreone &amp; Crottini &amp; Hutter &amp; Randrianantoandro &amp; Rakotoarison &amp; Glaw &amp; Hofreiter &amp; Vences, 2020" authorityName="Rancilhac &amp; Bruy &amp; Scherz &amp; Pereira &amp; Preick &amp; Straube &amp; Lyra &amp; Ohler &amp; Streicher &amp; Andreone &amp; Crottini &amp; Hutter &amp; Randrianantoandro &amp; Rakotoarison &amp; Glaw &amp; Hofreiter &amp; Vences" authorityYear="2020" box="[634,864,1459,1484]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="23" pageNumber="109" phylum="Chordata" rank="species" species="radaka" status="sp. nov.">
<emphasis id="A7C64E6CFFF600264D47FA3BFCC159A8" box="[634,864,1459,1484]" italics="true" pageId="23" pageNumber="109">Mantidactylus radaka</emphasis>
</taxonomicName>
<taxonomicNameLabel id="BCF5F317FFF600264C5BFA3CFC1B59A8" box="[870,954,1460,1484]" pageId="23" pageNumber="109" rank="species">sp. nov.</taxonomicNameLabel>
is the only species of the subgenus for which some natural history information is available: calls and tadpoles were described, from Tsaratanana and from a site between Bealanana and Antsohihy (close to the collection locality of ZSM 96/2016), respectively, by Vences et al. (2004) and 
<bibRefCitation id="F123EF8FFFF600264B29F994FEA45A33" author="Schulze A &amp; Randrianiaina RD &amp; Perl B &amp; Glaw F &amp; Vences M" pageId="23" pageNumber="109" pagination="119 - 125" refId="ref17527" refString="Schulze A, Randrianiaina RD, Perl B, Glaw F, Vences M. 2016. The unexpectedly dull tadpole of Madagascar' s largest frog Mantidactylus guttulatus. Acta Herpetologica. 11: 119 - 125." type="journal article" year="2016">Schulze et al. (2016)</bibRefCitation>
, under the name 
<taxonomicName id="52B2E9FDFFF600264EF4F9B6FDF75A33" baseAuthorityName="Boulenger" baseAuthorityYear="1881" box="[457,598,1598,1623]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="23" pageNumber="109" phylum="Chordata" rank="species" species="guttulatus">
<emphasis id="A7C64E6CFFF600264EF4F9B6FDF75A33" box="[457,598,1598,1623]" italics="true" pageId="23" pageNumber="109">M. guttulatus</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="DDA8C1F5FFF600284F9DF90CFEB059EB" lastPageId="25" lastPageNumber="111" pageId="23" pageNumber="109" type="distribution">
<paragraph id="950D927EFFF600294F9DF90CFD945D6A" blockId="23.[160,1155,1668,1727]" lastBlockId="24.[160,1156,142,270]" lastPageId="24" lastPageNumber="110" pageId="23" pageNumber="109">
<emphasis id="A7C64E6CFFF600264F9DF90CFE955AF8" bold="true" box="[160,308,1668,1692]" italics="true" pageId="23" pageNumber="109">Distribution.</emphasis>
Localities span an elevational range from 
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a.s.l. (Manongarivo Special Reserve, Camp 0) up to at least 
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a.s.l. (Tsaratanana Massif, Antevialambazaha/Camp 1). Based on genetic data herein, confirmed localities are (from north to south) the Tsaratanana Massif (localities Manarikoba forest and Antevialambazaha/Camp 1), Manongarivo, Ampotsidy, Bemanevika, several sites between Bealanana and Antsohihy (e.g. Ampandrana and Andranonafindra), and Besariaka.
</paragraph>
<paragraph id="950D927EFFF900284F9DFECBFD9D5C88" blockId="24.[160,1156,323,728]" lastBlockId="25.[160,1155,142,444]" lastPageId="25" lastPageNumber="111" pageId="24" pageNumber="110">
<emphasis id="A7C64E6CFFF900294F9DFECBFEE55D3F" bold="true" box="[160,324,323,347]" italics="true" pageId="24" pageNumber="110">Conservation.</emphasis>
During fieldwork in 2016, this species was found to be rather abundant, even where the forest was heavily degraded, as long as some riparian habitat remained. However, it is also actively hunted as a source of food by local people in the Bealanana District (
<figureCitation id="0D898EFBFFF900294FC0FE23FEC45DA7" box="[253,357,427,451]" captionStart="Figure 10" captionStartId="24.[160,225,1616,1639]" captionTargetBox="[166,1149,787,1585]" captionTargetId="figure-218@24.[166,1149,787,1585]" captionTargetPageId="24" captionText="Figure 10. Mantidactylus (Mantidactylus) radaka sp. nov. being prepared for human consumption. (a) Frogs and crabs are collected from broad streams. Then (b) the frogs are gutted and skinned, and the head, hands and feet removed. The frog is then rinsed in the stream, leaving (c) cleaned animals for cooking in a stew. Note the ovaries full with hundreds of eggs." figureDoi="http://doi.org/10.5281/zenodo.5020674" httpUri="https://zenodo.org/record/5020674/files/figure.png" pageId="24" pageNumber="110">Figure 10</figureCitation>
). When asked, two locals stated that they no longer hunted these frogs in their home village (Ampandrana, ca. 
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, 
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), because they are no longer found in sufficient abundance in its vicinity. We managed to collect specimens from the river that flows towards Ampandrana only by going upstream (
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, 
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), and these frogs were considerably more abundant 
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further SSW along the Route National 31 (at Andranonafindra), where there are no further villages for another 
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. Thus, it seems evident that anthropogenic pressures are having impacts on local populations. However, the extent of those impacts is currently unclear, and given the apparently very large clutches that these frogs have (see the egg-laden ovaries evident in 
<figureCitation id="0D898EFBFFF900294B39FD48FBDD5EBC" box="[1028,1148,704,728]" captionStart="Figure 10" captionStartId="24.[160,225,1616,1639]" captionTargetBox="[166,1149,787,1585]" captionTargetId="figure-218@24.[166,1149,787,1585]" captionTargetPageId="24" captionText="Figure 10. Mantidactylus (Mantidactylus) radaka sp. nov. being prepared for human consumption. (a) Frogs and crabs are collected from broad streams. Then (b) the frogs are gutted and skinned, and the head, hands and feet removed. The frog is then rinsed in the stream, leaving (c) cleaned animals for cooking in a stew. Note the ovaries full with hundreds of eggs." figureDoi="http://doi.org/10.5281/zenodo.5020674" httpUri="https://zenodo.org/record/5020674/files/figure.png" pageId="24" pageNumber="110">Figure 10C</figureCitation>
) and their stream-dwelling tadpoles, it is possible that up-stream populations may seed down-stream ones, and the species may thus be rather resilient to exploitation and able to recover quickly if pressures cease.
</paragraph>
<caption id="C1CDC2F6FFF900294F9DF9D8FCB25ADB" ID-DOI="http://doi.org/10.5281/zenodo.5020674" ID-Zenodo-Dep="5020674" httpUri="https://zenodo.org/record/5020674/files/figure.png" pageId="24" pageNumber="110" startId="24.[160,225,1616,1639]" targetBox="[166,1149,787,1585]" targetPageId="24">
<paragraph id="950D927EFFF900294F9DF9D8FCB25ADB" blockId="24.[160,1155,1615,1727]" pageId="24" pageNumber="110">
<emphasis id="A7C64E6CFFF900294F9DF9D8FEA65A03" bold="true" box="[160,263,1616,1639]" pageId="24" pageNumber="110">Figure 10.</emphasis>
<taxonomicName id="52B2E9FDFFF900294E30F9C7FDD65A02" authority="Rancilhac &amp; Bruy &amp; Scherz &amp; Pereira &amp; Preick &amp; Straube &amp; Lyra &amp; Ohler &amp; Streicher &amp; Andreone &amp; Crottini &amp; Hutter &amp; Randrianantoandro &amp; Rakotoarison &amp; Glaw &amp; Hofreiter &amp; Vences, 2020" authorityName="Rancilhac &amp; Bruy &amp; Scherz &amp; Pereira &amp; Preick &amp; Straube &amp; Lyra &amp; Ohler &amp; Streicher &amp; Andreone &amp; Crottini &amp; Hutter &amp; Randrianantoandro &amp; Rakotoarison &amp; Glaw &amp; Hofreiter &amp; Vences" authorityYear="2020" box="[269,631,1615,1638]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="24" pageNumber="110" phylum="Chordata" rank="species" species="radaka" status="sp. nov." subGenus="Mantidactylus">
<emphasis id="A7C64E6CFFF900294E30F9C7FDD65A02" box="[269,631,1615,1638]" italics="true" pageId="24" pageNumber="110">Mantidactylus (Mantidactylus) radaka</emphasis>
</taxonomicName>
<taxonomicNameLabel id="BCF5F317FFF900294D43F9D8FD6A5A03" box="[638,715,1616,1639]" pageId="24" pageNumber="110" rank="species">sp. nov.</taxonomicNameLabel>
being prepared for human consumption. (a) Frogs and crabs are collected from broad streams. Then (b) the frogs are gutted and skinned, and the head, hands and feet removed. The frog is then rinsed in the stream, leaving (c) cleaned animals for cooking in a stew. Note the ovaries full with hundreds of eggs.
</paragraph>
</caption>
<paragraph id="950D927EFFF800284FFDFF7EFC065DD8" blockId="25.[160,1155,142,444]" pageId="25" pageNumber="111">
At present, the distribution of this species appears to be rather large: a minimum convex polygon of known localities has an area of 
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<superScript id="62C73F36FFF800284C74FE9BFCF25D47" attach="left" box="[841,851,275,291]" fontSize="7" pageId="25" pageNumber="111">2</superScript>
. Over that area, there is no doubt that the habitat of the species is declining in extent and quality, and, although it is found in numerous locations, its habitat is also severely fragmented. Therefore, this species qualifies as Vulnerable under the International Union for Conservation of Nature (IUCN) Red List of Threatened Species, criterion B1ab(iii) (
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).
</paragraph>
<paragraph id="950D927EFFF800284F9DFE79FCDC5E6E" blockId="25.[160,1156,497,1423]" box="[160,893,497,522]" pageId="25" pageNumber="111">
<emphasis id="A7C64E6CFFF800284F9DFE79FCDC5E6E" bold="true" box="[160,893,497,522]" pageId="25" pageNumber="111">
Candidate species 
<taxonomicName id="52B2E9FDFFF800284EB8FE7AFDF65E6E" box="[389,599,497,522]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="25" pageNumber="111" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="A7C64E6CFFF800284EB8FE7AFD8C5E6E" bold="true" box="[389,557,498,522]" italics="true" pageId="25" pageNumber="111">Mantidactylus</emphasis>
sp.
</taxonomicName>
Ca55, Ca56, Ca66, Ca67
</emphasis>
</paragraph>
<paragraph id="950D927EFFF800284F9DFD9DFC5D5EF1" blockId="25.[160,1156,497,1423]" pageId="25" pageNumber="111">
Besides the three species now recognised, our analysis has identified four additional lineages in the subgenus 
<taxonomicName id="52B2E9FDFFF800284EF6FDBEFDC55E2B" authorityName="Boulenger" authorityYear="1895" box="[459,612,566,591]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="25" pageNumber="111" phylum="Chordata" rank="genus">
<emphasis id="A7C64E6CFFF800284EF6FDBEFDC55E2B" box="[459,612,566,591]" italics="true" pageId="25" pageNumber="111">Mantidactylus</emphasis>
</taxonomicName>
(
<figureCitation id="0D898EFBFFF800284D4BFDBFFD745E2B" box="[630,725,567,591]" captionStart="Figure 1" captionStartId="9.[160,225,1505,1528]" captionTargetBox="[166,1149,142,1473]" captionTargetId="figure-1@9.[166,1149,142,1474]" captionTargetPageId="9" captionText="Figure 1. Maximum likelihood phylogenetic tree obtained from 514 bp of the mitochondrial 16S rRNA gene. The values at the nodes are the bootstrap supports (not given for intra-lineage nodes for improved clarity). The type specimens of M. guttulatus and M. grandidieri from the London and Paris museum collections are highlighted in red and brown, respectively." figureDoi="http://doi.org/10.5281/zenodo.5020655" httpUri="https://zenodo.org/record/5020655/files/figure.png" pageId="25" pageNumber="111">Figure 1</figureCitation>
), for which we have herein used the candidate species names 
<taxonomicName id="52B2E9FDFFF800284EFEFDD1FD255E16" box="[451,644,601,626]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="25" pageNumber="111" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="A7C64E6CFFF800284EFEFDD1FDFD5E16" box="[451,604,601,626]" italics="true" pageId="25" pageNumber="111">Mantidactylus</emphasis>
sp.
</taxonomicName>
Ca55, Ca56, Ca66 and Ca67. At present, data are clearly insufficient to take taxonomic decisions regarding these lineages.
</paragraph>
<paragraph id="950D927EFFF800284FFDFD16FE6D581E" blockId="25.[160,1156,497,1423]" pageId="25" pageNumber="111">
<taxonomicName id="52B2E9FDFFF800284FFDFD16FE215ED3" box="[192,384,670,695]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="25" pageNumber="111" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="A7C64E6CFFF800284FFDFD16FEF85ED3" box="[192,345,670,695]" italics="true" pageId="25" pageNumber="111">Mantidactylus</emphasis>
sp.
</taxonomicName>
Ca55 and Ca56 occur sympatrically in Betampona, but no specimens of either candidate species were available for morphological examination in this study. Both of these candidate species are crepuscular-nocturnal. In Betampona, 
<taxonomicName id="52B2E9FDFFF800284CD7FD6CFF615F7B" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="25" pageNumber="111" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="A7C64E6CFFF800284CD7FD6CFB225E99" box="[1002,1155,740,765]" italics="true" pageId="25" pageNumber="111">Mantidactylus</emphasis>
sp.
</taxonomicName>
Ca56 seems to be both more abundant and more widespread, but precise data on its fine-scale distribution within this reserve are thus far lacking. 
<taxonomicName id="52B2E9FDFFF800284C41FCA1FBE05F26" box="[892,1089,809,834]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="25" pageNumber="111" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="A7C64E6CFFF800284C41FCA1FBB45F26" box="[892,1045,809,834]" italics="true" pageId="25" pageNumber="111">Mantidactylus</emphasis>
sp.
</taxonomicName>
Ca56 inhabits all large and fast-flowing streams across the reserve, while 
<taxonomicName id="52B2E9FDFFF800284CFDFCC4FB255F01" box="[960,1156,844,869]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="25" pageNumber="111" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="A7C64E6CFFF800284CFDFCC4FBF85F01" box="[960,1113,844,869]" italics="true" pageId="25" pageNumber="111">Mantidactylus</emphasis>
sp.
</taxonomicName>
Ca55 seems to prefer lateral branches of the fast-flowing streams, where water is shallower and runs much more slowly. Photos of genetically identified individuals of the two candidate species (
<figureCitation id="0D898EFBFFF800284E4FFC3DFE6C5FA9" box="[370,461,949,973]" captionStart="Figure 7" captionStartId="17.[160,225,1523,1546]" captionTargetBox="[166,1143,142,1491]" captionTargetId="figure-1@17.[166,1149,142,1491]" captionTargetPageId="17" captionText="Figure 7. Photographs of living specimens of Mantidactylus (Mantidactylus) guttulatus, M. (M.) grandidieri, and of three candidate species. (a, b) M. (M.) guttulatus, female ZSM 1013/2003 (FGMV 2002.438) from Ranomafana. (c) Unidentified specimen from Ranomafana, assigned tentatively to M. (M.) guttulatus (no genetic evidence). (d, e) M. (M.) guttulatus, specimen KU 340853 (CRH729) from Ranomafana. (f) M. (M.) grandidieri, specimen ZSM 5077/2005 (ZCMV 2159) from Nosy Mangabe. (g) M. (M.) grandidieri, specimen ZSM 276/2005 (FGZC 2682) from Vohidrazana. (h) M. (M.) grandidieri, unidentified specimen (probably subadult) from Andranofotsy. (i, j) M. (M.) grandidieri, specimen KU" figureDoi="http://doi.org/10.5281/zenodo.5020667" httpUri="https://zenodo.org/record/5020667/files/figure.png" pageId="25" pageNumber="111">Figure 7</figureCitation>
) do not immediately reveal obvious morphological differences between them. They do not appear to be sister species based on the mitochondrial data (
<figureCitation id="0D898EFBFFF800284F9AFC72FEA35876" box="[167,258,1018,1042]" captionStart="Figure 1" captionStartId="9.[160,225,1505,1528]" captionTargetBox="[166,1149,142,1473]" captionTargetId="figure-1@9.[166,1149,142,1474]" captionTargetPageId="9" captionText="Figure 1. Maximum likelihood phylogenetic tree obtained from 514 bp of the mitochondrial 16S rRNA gene. The values at the nodes are the bootstrap supports (not given for intra-lineage nodes for improved clarity). The type specimens of M. guttulatus and M. grandidieri from the London and Paris museum collections are highlighted in red and brown, respectively." figureDoi="http://doi.org/10.5281/zenodo.5020655" httpUri="https://zenodo.org/record/5020655/files/figure.png" pageId="25" pageNumber="111">Figure 1</figureCitation>
), and in the nuclear RAG-1 gene, the two examined samples do not share alleles. If confirmed with a larger number of samples, such a concordance between mitochondrial and nuclear gene signals in sympatry would clearly be indicative of a status as separate species (
<bibRefCitation id="F123EF8FFFF800284FC3FBEAFE61581E" author="Padial JM &amp; Miralles A &amp; De la Riva I &amp; Vences M" box="[254,448,1122,1146]" pageId="25" pageNumber="111" pagination="16" refId="ref16803" refString="Padial JM, Miralles A, De la Riva I, Vences M. 2010. The integrative future of taxonomy. Front Zool. 7 (1): 16. doi: 10.1186 / 1742 - 9994 - 7 - 16." type="journal article" year="2010">Padial et al. 2010</bibRefCitation>
).
</paragraph>
<paragraph id="950D927EFFF800284FFDFB0DFEB059EB" blockId="25.[160,1156,497,1423]" pageId="25" pageNumber="111">
In contrast, 
<taxonomicName id="52B2E9FDFFF800284E77FB0CFDAF58F9" box="[330,526,1156,1181]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="25" pageNumber="111" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="A7C64E6CFFF800284E77FB0CFE4258F9" box="[330,483,1156,1181]" italics="true" pageId="25" pageNumber="111">Mantidactylus</emphasis>
sp.
</taxonomicName>
Ca66 and Ca67 appear to be allopatric sister lineages distributed in the South East. Their mitochondrial divergence is low, and they may be conspecific. However, their distinctness from other species and candidate species of the subgenus 
<taxonomicName id="52B2E9FDFFF800284E29FB64FE0C5961" authorityName="Boulenger" authorityYear="1895" box="[276,429,1260,1285]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="25" pageNumber="111" phylum="Chordata" rank="genus">
<emphasis id="A7C64E6CFFF800284E29FB64FE0C5961" box="[276,429,1260,1285]" italics="true" pageId="25" pageNumber="111">Mantidactylus</emphasis>
</taxonomicName>
is supported by exclusive RAG-1 alleles of numerous individuals sequenced, and by their apparently very smooth skin. Thus, they are likely to represent at least one new and as yet undescribed species. In the forest at Manantantely, 
<taxonomicName id="52B2E9FDFFF800284CD7FAB9FF615909" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="25" pageNumber="111" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="A7C64E6CFFF800284CD7FAB9FB22592E" box="[1002,1155,1329,1354]" italics="true" pageId="25" pageNumber="111">Mantidactylus</emphasis>
sp.
</taxonomicName>
Ca66 was nocturnal and living in rocky streams in rather pristine low-elevation rainforest.
</paragraph>
</subSubSection>
</treatment>
</document>