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<document id="4069ECF8EAD2D122CAFE260C4EDE9CC0" ID-DOI="10.5281/zenodo.183039" ID-GBIF-Dataset="040a24cf-1d95-436f-81b1-eca726c2ba38" ID-ISSN="1175-5326" ID-Zenodo-Dep="183039" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1459887428451" checkinUser="plazi" docAuthor="Brown, Jason L., Twomey, Evan, Pepper, Mark &amp; Rodriguez, Manuel Sanchez" docDate="2008" docId="586D087BFFA3FFB7FF1FF944FE4EFDED" docLanguage="en" docName="zt01823p024.pdf" docOrigin="Zootaxa 1823" docStyle="DocumentStyle:6581673A57F01A3145754A1E615EDFF0.4:Zootaxa.2007-2008.journal_article" docStyleId="6581673A57F01A3145754A1E615EDFF0" docStyleName="Zootaxa.2007-2008.journal_article" docStyleVersion="4" docTitle="Ranitomeya benedicta Brown, Twomey, Pepper &amp; Rodriguez, 2008, sp. nov." docType="treatment" docVersion="8" lastPageNumber="15" masterDocId="A4547003FFA1FFB9FF88FFC6FFBBFFA9" masterDocTitle="Revision of the Ranitomeya fantastica species complex with description of two new species from Central Peru (Anura: Dendrobatidae)" masterLastPageNumber="24" masterPageNumber="1" pageNumber="3" updateTime="1698227885604" updateUser="plazi">
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<mods:title id="6F2FA50FF151F25AB0BD14621746D282">Revision of the Ranitomeya fantastica species complex with description of two new species from Central Peru (Anura: Dendrobatidae)</mods:title>
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<mods:namePart id="7BDCF56F7DD93B3FF32A571CB85CFE62">Brown, Jason L.</mods:namePart>
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<mods:namePart id="52A820729D939D184EDC5AA6BC14B911">Twomey, Evan</mods:namePart>
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<mods:role id="50AB085CAA0EBEC8C0F7B4433342DEB1">
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<mods:namePart id="08530DC7D93F902A2D71576E8D63CDAB">Pepper, Mark</mods:namePart>
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<mods:roleTerm id="47F49681C1837DED4BDD2AD53C5ED4F6">Author</mods:roleTerm>
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<mods:namePart id="C5F2ED3BC943EC6F66F1408F6E1DDD05">Rodriguez, Manuel Sanchez</mods:namePart>
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<mods:title id="69C68F9193198BB5D64397C420FEBFF1">Zootaxa</mods:title>
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<mods:date id="F8ECEF80491A2BDAA7B8F98AFEDF08E3">2008</mods:date>
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<mods:number id="9D15904E63B2A4BFB2AB225960B5327B">1823</mods:number>
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<treatment id="586D087BFFA3FFB7FF1FF944FE4EFDED" ID-DOI="http://doi.org/10.5281/zenodo.5611469" ID-GBIF-Taxon="119371710" ID-Zenodo-Dep="5611469" LSID="urn:lsid:plazi:treatment:586D087BFFA3FFB7FF1FF944FE4EFDED" httpUri="http://treatment.plazi.org/id/586D087BFFA3FFB7FF1FF944FE4EFDED" lastPageId="14" lastPageNumber="15" pageId="2" pageNumber="3">
<subSubSection id="98DEEAE6FFA3FFBBFF1FF944FED1F916" pageId="2" pageNumber="3" type="nomenclature">
<paragraph id="D07BB96DFFA3FFBBFF1FF944FDB5F935" blockId="2.[151,526,1666,1727]" box="[151,526,1666,1692]" pageId="2" pageNumber="3">
<heading id="8B330E01FFA3FFBBFF1FF944FDB5F935" bold="true" box="[151,526,1666,1692]" fontSize="11" level="1" pageId="2" pageNumber="3" reason="1">
<emphasis id="E2B0657FFFA3FFBBFF1FF944FDB5F935" bold="true" box="[151,526,1666,1692]" pageId="2" pageNumber="3">
<taxonomicName id="17C4C2EEFFA3FFBBFF1FF944FE19F935" ID-CoL="78T37" box="[151,418,1666,1692]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="benedicta" status="sp. nov.">
<emphasis id="E2B0657FFFA3FFBBFF1FF944FE19F935" bold="true" box="[151,418,1666,1692]" italics="true" pageId="2" pageNumber="3">Ranitomeya benedicta</emphasis>
</taxonomicName>
,
<taxonomicNameLabel id="F983D804FFA3FFBBFE38F944FDB5F935" box="[432,526,1666,1692]" pageId="2" pageNumber="3" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="D07BB96DFFA3FFBBFF1FF963FED1F916" blockId="2.[151,526,1666,1727]" box="[151,362,1701,1727]" pageId="2" pageNumber="3">
<figureCitation id="48FFA5E8FFA3FFBBFF1FF963FEBEF916" box="[151,261,1701,1727]" captionStart="FIGURE 1" captionStartId="3.[151,254,1345,1369]" captionTargetBox="[261,1328,264,1330]" captionTargetId="figure@3.[255,1333,258,1338]" captionTargetPageId="3" captionText="FIGURE 1. Drawing of species contained in the fantastica complex. Clockwise from left: Ranitomeya summersi sp. nov., R. fantastica sensu stricto, R. benedicta sp. nov. Illustration of R. fantastica is an artists rendering of the nominal population from Yurimaguas based on the original type series and recent field work nearby." httpUri="https://zenodo.org/record/183040/files/figure.png" pageId="2" pageNumber="3">Figures 1</figureCitation>
,
<figureCitation id="48FFA5E8FFA3FFBBFE9BF963FE9FF916" box="[275,292,1701,1727]" captionStart="FIGURE 4" captionStartId="6.[151,254,1989,2013]" captionTargetBox="[242,1336,206,1975]" captionTargetId="figure@6.[219,1379,182,2012]" captionTargetPageId="6" captionText="FIGURE 4. Type series of R. benedicta. (Top L-R: MUSM 26956 26958, Bottom L-R: MUSM 26959 26962)." httpUri="https://zenodo.org/record/183043/files/figure.png" pageId="2" pageNumber="3">4</figureCitation>
,
<figureCitation id="48FFA5E8FFA3FFBBFEB9F963FEFAF916" box="[305,321,1701,1727]" captionStart="FIGURE 5" captionStartId="9.[151,254,1864,1888]" captionTargetBox="[261,1339,325,1846]" captionTargetId="figure@9.[253,1346,314,1850]" captionTargetPageId="9" captionText="FIGURE 5. Map of known localities and estimated distributions of R. fantastica, R. summersi, and R. benedicta. Because some populations where close together, the 14 sampling localities were simplified into 10 geographic groups. Localities which were included in the phylogenetic analysis are indicated by numbers on the map: (1) Tarapoto; (2) Cainarachi Valley; (3) Convento; (4) Pongo de Cainarachi; (5) Varadero; (6) Lower Huallaga Canyon; (7) Shucushuyacu; (8) Pampa Hermosa; (9) Sauce; (10) Chipaota." httpUri="https://zenodo.org/record/183044/files/figure.png" pageId="2" pageNumber="3">5</figureCitation>
,
<figureCitation id="48FFA5E8FFA3FFBBFEC6F963FED1F916" box="[334,362,1701,1727]" captionStart="FIGURE 11" captionStartId="23.[151,254,1905,1929]" captionTargetBox="[157,1425,201,1871]" captionTargetId="figure@23.[151,1436,194,1881]" captionTargetPageId="23" captionText="FIGURE 11. Photos of R. benedicta. A E: Individuals from Pampa Hermosa, Loreto F M: Individuals from Shucushuyacu, Loreto." httpUri="https://zenodo.org/record/183050/files/figure.png" pageId="2" pageNumber="3">11</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="98DEEAE6FFA3FFBBFF1FF933FEE9F8E5" pageId="2" pageNumber="3" type="reference_group">
<paragraph id="D07BB96DFFA3FFBBFF1FF933FEE9F8E5" blockId="2.[151,1436,1781,1868]" pageId="2" pageNumber="3">
<taxonomicName id="17C4C2EEFFA3FFBBFF1FF933FE71F8A5" ID-CoL="34M9M" box="[151,458,1781,1804]" class="Amphibia" family="Dendrobatidae" genus="Dendrobates" kingdom="Animalia" order="Anura" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="quinquevittatus">
<emphasis id="E2B0657FFFA3FFBBFF1FF933FE71F8A5" box="[151,458,1781,1804]" italics="true" pageId="2" pageNumber="3">Dendrobates quinquevittatus</emphasis>
</taxonomicName>
(non Steindachner):
<bibRefCitation id="B455C49CFFA3FFBBFD22F933FCDEF8A5" author="Silverstone" box="[682,869,1781,1804]" pageId="2" pageNumber="18" refString="Silverstone, P. A. (1975) A revision of the poison-arrow frogs of the genus Dendrobates Wagler. Natural History Museum of Los Angeles County, 21, 1 - 55." type="journal article" year="1975">Silverstone, 1975</bibRefCitation>
(partim), p. 3336, Fig. 14 (drawing), patterns K, L.
<taxonomicName id="17C4C2EEFFA3FFBBFF1FF8D3FE21F885" ID-CoL="34M8B" box="[151,410,1813,1836]" class="Amphibia" family="Dendrobatidae" genus="Dendrobates" kingdom="Animalia" order="Anura" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="fantasticus">
<emphasis id="E2B0657FFFA3FFBBFF1FF8D3FE21F885" box="[151,410,1813,1836]" italics="true" pageId="2" pageNumber="3">Dendrobates fantasticus</emphasis>
</taxonomicName>
(non Boulenger):
<bibRefCitation id="B455C49CFFA3FFBBFDE8F8D3FD48F885" author="Schulte" box="[608,755,1813,1836]" pageId="2" pageNumber="18" refString="Schulte, R. (1999) Pfeilgiftfrosche &quot; Artenteil - Peru &quot;. INBICO, Wailblingen, Germany, 294 pp." type="book" year="1999">Schulte, 1999</bibRefCitation>
(partim), pp. 5769,
<figureCitation id="48FFA5E8FFA3FFBBFC50F8D3FBACF885" box="[984,1047,1813,1836]" captionStart="FIGURE 5" captionStartId="9.[151,254,1864,1888]" captionTargetBox="[261,1339,325,1846]" captionTargetId="figure@9.[253,1346,314,1850]" captionTargetPageId="9" captionText="FIGURE 5. Map of known localities and estimated distributions of R. fantastica, R. summersi, and R. benedicta. Because some populations where close together, the 14 sampling localities were simplified into 10 geographic groups. Localities which were included in the phylogenetic analysis are indicated by numbers on the map: (1) Tarapoto; (2) Cainarachi Valley; (3) Convento; (4) Pongo de Cainarachi; (5) Varadero; (6) Lower Huallaga Canyon; (7) Shucushuyacu; (8) Pampa Hermosa; (9) Sauce; (10) Chipaota." httpUri="https://zenodo.org/record/183044/files/figure.png" pageId="2" pageNumber="3">Fig. 5</figureCitation>
pattern K, L (reprinted from
<bibRefCitation id="B455C49CFFA3FFBBFADBF8D3FEFEF8E5" author="Silverstone" pageId="2" pageNumber="18" refString="Silverstone, P. A. (1975) A revision of the poison-arrow frogs of the genus Dendrobates Wagler. Natural History Museum of Los Angeles County, 21, 1 - 55." type="journal article" year="1975">Silverstone, 1975</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="98DEEAE6FFA3FFBAFF1FF8BBFDBAFF73" lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3" type="materials_examined">
<paragraph id="D07BB96DFFA3FFBBFF1FF8BBFB75F84E" blockId="2.[151,1436,1917,2023]" pageId="2" pageNumber="3">
<emphasis id="E2B0657FFFA3FFBBFF1FF8BBFEABF83E" bold="true" box="[151,272,1917,1943]" pageId="2" pageNumber="3">
<typeStatus id="0F7F07CFFFA3FFBBFF1FF8BBFEB0F83E" box="[151,267,1917,1943]" pageId="2" pageNumber="3" type="holotype">Holotype</typeStatus>
.
</emphasis>
<collectionCode id="B6D521A8FFA3FFBBFE90F8BBFEC9F83E" box="[280,370,1917,1943]" pageId="2" pageNumber="3">MUSM</collectionCode>
26957 (field number JLB07-594), an adult female collected by M. Pepper and E. Twomey near Shucushuyacu (alternative spellings: Shucushyacu and Shucush-yacu), a small town on the east bank of Rio Huallaga near Yurimaguas, Departamento Loreto,
<collectingCountry id="A8D3F9FDFFA3FFBBFC90F80BFCEAF84E" box="[792,849,1997,2023]" name="Peru" pageId="2" pageNumber="3">Peru</collectingCountry>
;
<quantity id="173C1488FFA3FFBBFCD5F80BFC1CF84E" box="[861,935,1997,2023]" metricMagnitude="2" metricUnit="m" metricValue="1.96" pageId="2" pageNumber="3" unit="m" value="196.0">196 m</quantity>
elevation;
<date id="A47A9FADFFA3FFBBFBA1F80BFB72F84E" box="[1065,1225,1997,2023]" pageId="2" pageNumber="3" value="2006-01">January 2006</date>
.
</paragraph>
<paragraph id="D07BB96DFFA2FFBAFF4EFF51FDBAFF73" blockId="3.[151,1436,151,218]" pageId="3" pageNumber="4">
<emphasis id="E2B0657FFFA2FFBAFF4EFF51FEC0FF18" bold="true" box="[198,379,151,177]" pageId="3" pageNumber="4">Paratopotypes</emphasis>
<collectionCode id="B6D521A8FFA2FFBAFE02FF5EFE5FFF1B" box="[394,484,152,178]" pageId="3" pageNumber="4">MUSM</collectionCode>
26956, 26958, 26959, 26962 (field numbers JLB07-592, 596, 598 and 600), collected along with the
<typeStatus id="0F7F07CFFFA2FFBAFE1BFF06FE46FF73" box="[403,509,192,218]" pageId="3" pageNumber="4" type="holotype">holotype</typeStatus>
.
</paragraph>
</subSubSection>
<caption id="84BBE9E5FFA2FFBAFF1FFA87FBDDFA37" httpUri="https://zenodo.org/record/183040/files/figure.png" pageId="3" pageNumber="4" targetBox="[261,1328,264,1330]" targetPageId="3">
<paragraph id="D07BB96DFFA2FFBAFF1FFA87FBDDFA37" blockId="3.[151,1436,1345,1438]" pageId="3" pageNumber="4">
<emphasis id="E2B0657FFFA2FFBAFF1FFA87FEA0FAF0" bold="true" box="[151,283,1345,1369]" pageId="3" pageNumber="4">FIGURE 1.</emphasis>
Drawing of species contained in the
<taxonomicName id="17C4C2EEFFA2FFBAFD32FA84FC98FAF0" box="[698,803,1346,1369]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA2FFBAFD32FA84FC98FAF0" box="[698,803,1346,1369]" italics="true" pageId="3" pageNumber="4">fantastica</emphasis>
</taxonomicName>
complex. Clockwise from left:
<taxonomicName id="17C4C2EEFFA2FFBAFB0CFA84FAC9FAF0" box="[1156,1394,1346,1369]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="summersi" status="sp. nov.">
<emphasis id="E2B0657FFFA2FFBAFB0CFA84FAC9FAF0" box="[1156,1394,1346,1369]" italics="true" pageId="3" pageNumber="4">Ranitomeya summersi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="F983D804FFA2FFBAFAF4FA87FF7DFAD5" pageId="3" pageNumber="4" rank="species">
<emphasis id="E2B0657FFFA2FFBAFAF4FA87FF7DFAD5" bold="true" pageId="3" pageNumber="4">sp. nov.</emphasis>
</taxonomicNameLabel>
,
<taxonomicName id="17C4C2EEFFA2FFBAFF5CFAA2FEE0FAD2" box="[212,347,1380,1403]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="species" sensu="stricto" species="fantastica">
<emphasis id="E2B0657FFFA2FFBAFF5CFAA2FEE0FAD2" box="[212,347,1380,1403]" italics="true" pageId="3" pageNumber="4">R. fantastica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="F983D804FFA2FFBAFEEAFAA2FE51FAD2" box="[354,490,1380,1403]" pageId="3" pageNumber="4" sensu="stricto">sensu stricto</taxonomicNameLabel>
,
<taxonomicName id="17C4C2EEFFA2FFBAFE7EFAA2FDC7FAD2" box="[502,636,1380,1403]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="benedicta" status="sp. nov.">
<emphasis id="E2B0657FFFA2FFBAFE7EFAA2FDC7FAD2" box="[502,636,1380,1403]" italics="true" pageId="3" pageNumber="4">R. benedicta</emphasis>
</taxonomicName>
<taxonomicNameLabel id="F983D804FFA2FFBAFD0BFAA2FD62FAD5" box="[643,729,1380,1404]" pageId="3" pageNumber="4" rank="species">
<emphasis id="E2B0657FFFA2FFBAFD0BFAA2FD62FAD5" bold="true" box="[643,729,1380,1404]" pageId="3" pageNumber="4">sp. nov.</emphasis>
</taxonomicNameLabel>
Illustration of
<taxonomicName id="17C4C2EEFFA2FFBAFCF2FAA2FBB9FAD2" box="[890,1026,1380,1403]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA2FFBAFCF2FAA2FBB9FAD2" box="[890,1026,1380,1403]" italics="true" pageId="3" pageNumber="4">R. fantastica</emphasis>
</taxonomicName>
is an artists rendering of the nominal population from Yurimaguas based on the original type series and recent field work nearby.
</paragraph>
</caption>
<subSubSection id="98DEEAE6FFA2FFBAFF4EFA17FE96F995" pageId="3" pageNumber="4" type="description">
<paragraph id="D07BB96DFFA2FFBAFF4EFA17FE96F995" blockId="3.[151,1437,1489,2036]" pageId="3" pageNumber="4">
<typeStatus id="0F7F07CFFFA2FFBAFF4EFA17FEF8FA42" box="[198,323,1489,1515]" pageId="3" pageNumber="4" type="paratype">
<emphasis id="E2B0657FFFA2FFBAFF4EFA17FEF8FA42" bold="true" box="[198,323,1489,1515]" pageId="3" pageNumber="4">Paratypes</emphasis>
</typeStatus>
MUSM 26960, 26961 (field numbers JLB07-602 and 604), collected by M. Sanchez and E. Twomey on the west bank of Rio Ucayali near Contamana, Departamento Loreto,
<collectingCountry id="A8D3F9FDFFA2FFBAFB00FA3CFB7AF9BD" box="[1160,1217,1530,1556]" name="Peru" pageId="3" pageNumber="4">Peru</collectingCountry>
;
<quantity id="173C1488FFA2FFBAFB58FA3CFAA5F9BD" box="[1232,1310,1530,1556]" metricMagnitude="2" metricUnit="m" metricValue="1.53" pageId="3" pageNumber="4" unit="m" value="153.0">153 m</quantity>
elevation;
<date id="A47A9FADFFA2FFBAFF1FF9E4FE92F995" box="[151,297,1570,1596]" pageId="3" pageNumber="4" value="2007-03">March 2007</date>
.
</paragraph>
</subSubSection>
<subSubSection id="98DEEAE6FFA2FFBDFF4EF98FFF5AFF73" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4" type="etymology">
<paragraph id="D07BB96DFFA2FFBAFF4EF98FFEECF8AD" blockId="3.[151,1437,1489,2036]" pageId="3" pageNumber="4">
<emphasis id="E2B0657FFFA2FFBAFF4EF98FFEE8F9CA" bold="true" box="[198,339,1609,1635]" pageId="3" pageNumber="4">Etymology.</emphasis>
The specific epithet is the feminine form of the Latin word
<emphasis id="E2B0657FFFA2FFBAFBE0F98CFB5EF9CA" box="[1128,1253,1610,1635]" italics="true" pageId="3" pageNumber="4">benedictus</emphasis>
, which means blessed. This name was chosen because the core range of
<taxonomicName id="17C4C2EEFFA2FFBAFCC6F9B4FC5AF922" box="[846,993,1650,1675]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA2FFBAFCC6F9B4FC5AF922" box="[846,993,1650,1675]" italics="true" pageId="3" pageNumber="4">R. benedicta</emphasis>
</taxonomicName>
occurs in the Pampas del Sacramento (The Holy Plains) in southern Loreto and eastern San Martin. The name also refers to quick and elusive nature of this species: if one is lucky enough to encounter it (as we were in 2005), one may consider themselves blessed.
</paragraph>
<paragraph id="D07BB96DFFA2FFBDFF4DF8D7FF5AFF73" blockId="3.[151,1437,1489,2036]" lastBlockId="4.[151,1436,152,778]" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4">
<emphasis id="E2B0657FFFA2FFBAFF4DF8D7FDB9F882" bold="true" box="[197,514,1809,1835]" pageId="3" pageNumber="4">Definition and diagnosis.</emphasis>
Assigned to genus
<taxonomicName id="17C4C2EEFFA2FFBAFD7FF8D4FC38F882" box="[759,899,1810,1835]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="E2B0657FFFA2FFBAFD7FF8D4FC38F882" box="[759,899,1810,1835]" italics="true" pageId="3" pageNumber="4">Ranitomeya</emphasis>
</taxonomicName>
by the following characteristics: first finger shorter than second, premaxillary and maxillary teeth absent, webbing between toes absent, pale limb reticulation present. A relatively large species of
<taxonomicName id="17C4C2EEFFA2FFBAFDF7F8A4FCB0F8D2" box="[639,779,1890,1915]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="E2B0657FFFA2FFBAFDF7F8A4FCB0F8D2" box="[639,779,1890,1915]" italics="true" pageId="3" pageNumber="4">Ranitomeya</emphasis>
</taxonomicName>
attaining a maximum adult SVL of
<quantity id="173C1488FFA2FFBAFB3BF8A4FAA4F8D5" box="[1203,1311,1890,1916]" metricMagnitude="-2" metricUnit="m" metricValue="2.02" pageId="3" pageNumber="4" unit="mm" value="20.2">20.2 mm</quantity>
. Body and limbs black with blue wash or reticulation, head bright red with black markings over eyes. In some populations, the blue reticulation on the dorsum and legs is diffuse, causing the legs and part of the dorsum to appear uniform blue (
<figureCitation id="48FFA5E8FFA2FFBAFEB7F81CFE2FF85D" box="[319,404,2010,2036]" captionStart="FIGURE 11" captionStartId="23.[151,254,1905,1929]" captionTargetBox="[157,1425,201,1871]" captionTargetId="figure@23.[151,1436,194,1881]" captionTargetPageId="23" captionText="FIGURE 11. Photos of R. benedicta. A E: Individuals from Pampa Hermosa, Loreto F M: Individuals from Shucushuyacu, Loreto." httpUri="https://zenodo.org/record/183050/files/figure.png" pageId="3" pageNumber="4">Fig. 11</figureCitation>
, AE). Ventral ground color black with blue reticulation (discrete or diffuse), underside of chin red. Teeth absent; first finger shorter than second; disc of third finger 22.3 times wider than finger width.
</paragraph>
</subSubSection>
<subSubSection id="98DEEAE6FFA5FFBEFF4DFF2EFB75FAC8" lastPageId="7" lastPageNumber="8" pageId="4" pageNumber="5" type="description">
<paragraph id="D07BB96DFFA5FFBDFF4DFF2EFED8FCA3" blockId="4.[151,1436,152,778]" pageId="4" pageNumber="5">
<taxonomicName id="17C4C2EEFFA5FFBDFF4DFF2EFE71FEA8" box="[197,458,232,257]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA5FFBDFF4DFF2EFE71FEA8" box="[197,458,232,257]" italics="true" pageId="4" pageNumber="5">Ranitomeya benedicta</emphasis>
</taxonomicName>
can be distinguished from other species of
<taxonomicName id="17C4C2EEFFA5FFBDFC4CFF2EFBEBFEA8" box="[964,1104,232,257]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis id="E2B0657FFFA5FFBDFC4CFF2EFBEBFEA8" box="[964,1104,232,257]" italics="true" pageId="4" pageNumber="5">Ranitomeya</emphasis>
</taxonomicName>
by the combination of a relatively large adult SVL and a characteristic red head.
<taxonomicName id="17C4C2EEFFA5FFBDFC9CFED6FBA1FE80" box="[788,1050,272,297]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="E2B0657FFFA5FFBDFC9CFED6FBA1FE80" box="[788,1050,272,297]" italics="true" pageId="4" pageNumber="5">Ranitomeya reticulata</emphasis>
</taxonomicName>
also possesses red coloration on the head, although this coloration extends well onto the dorsum whereas in
<taxonomicName id="17C4C2EEFFA5FFBDFB9CFEFEFB13FEF8" box="[1044,1192,312,337]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA5FFBDFB9CFEFEFB13FEF8" box="[1044,1192,312,337]" italics="true" pageId="4" pageNumber="5">R. benedicta</emphasis>
</taxonomicName>
the red coloration is confined to the head only. Furthermore,
<taxonomicName id="17C4C2EEFFA5FFBDFDF7FEA6FCAFFED0" box="[639,788,352,377]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="E2B0657FFFA5FFBDFDF7FEA6FCAFFED0" box="[639,788,352,377]" italics="true" pageId="4" pageNumber="5">R. reticulata</emphasis>
</taxonomicName>
is smaller than
<taxonomicName id="17C4C2EEFFA5FFBDFC51FEA6FBD5FED0" box="[985,1134,352,377]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA5FFBDFC51FEA6FBD5FED0" box="[985,1134,352,377]" italics="true" pageId="4" pageNumber="5">R. benedicta</emphasis>
</taxonomicName>
(SVL
<quantity id="173C1488FFA5FFBDFB4BFEA6FACBFED3" box="[1219,1392,352,378]" metricMagnitude="-2" metricUnit="m" metricValue="1.5150000000000001" metricValueMax="1.6600000000000001" metricValueMin="1.3699999999999999" pageId="4" pageNumber="5" unit="mm" value="15.15" valueMax="16.6" valueMin="13.7">13.716.6 mm</quantity>
vs. 15.0
<quantity id="173C1488FFA5FFBDFF50FE4EFEFDFE0B" box="[216,326,392,418]" metricMagnitude="-2" metricUnit="m" metricValue="2.02" pageId="4" pageNumber="5" unit="mm" value="20.2">20.2 mm</quantity>
).
<taxonomicName id="17C4C2EEFFA5FFBDFED1FE4EFDE4FE08" box="[345,607,392,417]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA5FFBDFED1FE4EFDE4FE08" box="[345,607,392,417]" italics="true" pageId="4" pageNumber="5">Ranitomeya benedicta</emphasis>
</taxonomicName>
is most similar in external morphology to
<taxonomicName id="17C4C2EEFFA5FFBDFBDAFE4EFB5DFE08" box="[1106,1254,392,417]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA5FFBDFBDAFE4EFB5DFE08" box="[1106,1254,392,417]" italics="true" pageId="4" pageNumber="5">R. fantastica</emphasis>
</taxonomicName>
and the second new species,
<taxonomicName id="17C4C2EEFFA5FFBDFEBAFE76FE79FE60" box="[306,450,432,457]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFA5FFBDFEBAFE76FE79FE60" box="[306,450,432,457]" italics="true" pageId="4" pageNumber="5">R. summersi</emphasis>
</taxonomicName>
, described below.
<taxonomicName id="17C4C2EEFFA5FFBDFD14FE76FC1AFE60" box="[668,929,432,457]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA5FFBDFD14FE76FC1AFE60" box="[668,929,432,457]" italics="true" pageId="4" pageNumber="5">Ranitomeya benedicta</emphasis>
</taxonomicName>
has comparatively shorter hind limbs, with a significantly shorter tibia than
<taxonomicName id="17C4C2EEFFA5FFBDFD99FE1EFD1EFE58" box="[529,677,472,497]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA5FFBDFD99FE1EFD1EFE58" box="[529,677,472,497]" italics="true" pageId="4" pageNumber="5">R. fantastica</emphasis>
</taxonomicName>
(mean tibia length/SVL =
<quantity id="173C1488FFA5FFBDFC56FE1EFB85FE5B" box="[990,1086,472,498]" metricMagnitude="-2" metricUnit="m" metricValue="1.12776" pageId="4" pageNumber="5" unit="in" value="0.444">0.444 in</quantity>
<emphasis id="E2B0657FFFA5FFBDFBCDFE1EFB65FE58" box="[1093,1246,472,497]" italics="true" pageId="4" pageNumber="5">
<taxonomicName id="17C4C2EEFFA5FFBDFBCDFE1EFB60FE58" box="[1093,1243,472,497]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="benedicta">R. benedicta</taxonomicName>
,
</emphasis>
<quantity id="173C1488FFA5FFBDFB6EFE1EFAFDFE5B" box="[1254,1350,472,498]" metricMagnitude="-2" metricUnit="m" metricValue="1.23444" pageId="4" pageNumber="5" unit="in" value="0.486">0.486 in</quantity>
<taxonomicName id="17C4C2EEFFA5FFBDFAC4FE1EFF5EFDB0" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA5FFBDFAC4FE1EFF5EFDB0" italics="true" pageId="4" pageNumber="5">R. fantastica</emphasis>
</taxonomicName>
,
<emphasis id="E2B0657FFFA5FFBDFF7EFDC6FEB3FDB0" box="[246,264,512,537]" italics="true" pageId="4" pageNumber="5">P</emphasis>
= 0.01,
<figureCitation id="48FFA5E8FFA5FFBDFEFAFDC6FE00FDB3" box="[370,443,512,538]" captionStart="FIGURE 2" captionStartId="4.[151,254,1295,1319]" captionTargetBox="[159,1425,822,1283]" captionTargetId="figure@4.[151,1436,814,1288]" captionTargetPageId="4" captionText="FIGURE 2. Left: Comparisons of tibia lengths for the two ± 2 standard errors. nificantly larger standardized tibia length (Tibia / SVL) than R. benedicta and R. summersi (One-way ANOVA, SS = 0.009, df = 2, F = 5.654. Between species, P = 0.008. Error: SS = 0.022, df = 29) Right: Drawing of tadpole mouthparts of R. summersi. Mouthparts of R. benedicta and R. fantastica sensu stricto are identical." httpUri="https://zenodo.org/record/183041/files/figure.png" pageId="4" pageNumber="5">Fig. 2</figureCitation>
). Certain color morphs of
<taxonomicName id="17C4C2EEFFA5FFBDFD77FDC6FC2DFDB0" box="[767,918,512,537]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA5FFBDFD77FDC6FC2DFDB0" box="[767,918,512,537]" italics="true" pageId="4" pageNumber="5">R. fantastica</emphasis>
</taxonomicName>
have head coloration approaching red (i.e. bright orange), but never a true red as in
<taxonomicName id="17C4C2EEFFA5FFBDFD0CFDEEFCA3FDE8" box="[644,792,552,577]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA5FFBDFD0CFDEEFCA3FDE8" box="[644,792,552,577]" italics="true" pageId="4" pageNumber="5">R. benedicta</emphasis>
</taxonomicName>
. Furthermore, the call of
<taxonomicName id="17C4C2EEFFA5FFBDFBC5FDEEFB5AFDE8" box="[1101,1249,552,577]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA5FFBDFBC5FDEEFB5AFDE8" box="[1101,1249,552,577]" italics="true" pageId="4" pageNumber="5">R. benedicta</emphasis>
</taxonomicName>
is much louder (audible from ~
<quantity id="173C1488FFA5FFBDFEC7FD96FEC0FDC3" box="[335,379,592,618]" metricMagnitude="0" metricUnit="m" metricValue="8.0" pageId="4" pageNumber="5" unit="m" value="8.0">8 m</quantity>
vs. ~
<quantity id="173C1488FFA5FFBDFE33FD96FE53FDC3" box="[443,488,592,618]" metricMagnitude="0" metricUnit="m" metricValue="3.0" pageId="4" pageNumber="5" unit="m" value="3.0">3 m</quantity>
in
<taxonomicName id="17C4C2EEFFA5FFBDFD85FD96FD1AFDC0" box="[525,673,592,617]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA5FFBDFD85FD96FD1AFDC0" box="[525,673,592,617]" italics="true" pageId="4" pageNumber="5">R. fantastica</emphasis>
</taxonomicName>
), decreases in tone in consecutive notes (vs. constant tone across notes in
<taxonomicName id="17C4C2EEFFA5FFBDFF73FDBEFE34FD38" box="[251,399,632,657]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA5FFBDFF73FDBEFE34FD38" box="[251,399,632,657]" italics="true" pageId="4" pageNumber="5">R. fantastica</emphasis>
</taxonomicName>
) and has shorter pulses than the call of
<taxonomicName id="17C4C2EEFFA5FFBDFCEAFDBEFC4CFD38" box="[866,1015,632,657]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA5FFBDFCEAFDBEFC4CFD38" box="[866,1015,632,657]" italics="true" pageId="4" pageNumber="5">R. fantastica</emphasis>
</taxonomicName>
(150 ± 20 ms vs. 290 ± 30 ms in
<taxonomicName id="17C4C2EEFFA5FFBDFA0BFDBEFEB0FD10" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA5FFBDFA0BFDBEFEB0FD10" italics="true" pageId="4" pageNumber="5">R. fantastica</emphasis>
</taxonomicName>
,
<figureCitation id="48FFA5E8FFA5FFBDFE91FD66FEE5FD13" box="[281,350,672,698]" captionStart="FIGURE 3" captionStartId="5.[152,255,1239,1263]" captionTargetBox="[165,1429,552,1216]" captionTargetId="figure@5.[155,1440,546,1225]" captionTargetPageId="5" captionText="FIGURE 3. Advertisement calls of (A) R. fantastica individual from Cainarachi Valley, recording at 24 C, (B) R. summersi individual from Sauce, recorded at 24.5 C in captivity, (C) R. benedicta individual from Shucushuyacu, recorded at 24 C in captivity." httpUri="https://zenodo.org/record/183042/files/figure.png" pageId="4" pageNumber="5">Fig. 3</figureCitation>
). For differences between
<taxonomicName id="17C4C2EEFFA5FFBDFD1CFD66FC9CFD10" box="[660,807,672,697]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA5FFBDFD1CFD66FC9CFD10" box="[660,807,672,697]" italics="true" pageId="4" pageNumber="5">R. benedicta</emphasis>
</taxonomicName>
and
<taxonomicName id="17C4C2EEFFA5FFBDFCD6FD66FC54FD10" box="[862,1007,672,697]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFA5FFBDFCD6FD66FC54FD10" box="[862,1007,672,697]" italics="true" pageId="4" pageNumber="5">R. summersi</emphasis>
</taxonomicName>
, see below.
<taxonomicName id="17C4C2EEFFA5FFBDFBF5FD66FA39FD10" box="[1149,1410,672,697]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA5FFBDFBF5FD66FA39FD10" box="[1149,1410,672,697]" italics="true" pageId="4" pageNumber="5">Ranitomeya benedicta</emphasis>
</taxonomicName>
is distinguished from its closest relatives on the basis of 8 unique unambiguous mitochondrial gene nucleotide site substitutions.
</paragraph>
<caption id="84BBE9E5FFA5FFBDFF1FFAC9FF41FA18" httpUri="https://zenodo.org/record/183041/files/figure.png" pageId="4" pageNumber="5" targetBox="[159,1425,822,1283]" targetPageId="4">
<paragraph id="D07BB96DFFA5FFBDFF1FFAC9FF41FA18" blockId="4.[151,1436,1295,1457]" pageId="4" pageNumber="5">
<emphasis id="E2B0657FFFA5FFBDFF1FFAC9FEA2FA8E" bold="true" box="[151,281,1295,1319]" pageId="4" pageNumber="5">FIGURE 2.</emphasis>
Left: Comparisons of tibia lengths for the two ± 2 standard errors. nificantly larger standardized tibia length (Tibia/ SVL) than
<taxonomicName id="17C4C2EEFFA5FFBDFB4DFAF4FAF6FAE0" box="[1221,1357,1330,1353]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA5FFBDFB4DFAF4FAF6FAE0" box="[1221,1357,1330,1353]" italics="true" pageId="4" pageNumber="5">R. benedicta</emphasis>
</taxonomicName>
and
<taxonomicName id="17C4C2EEFFA5FFBDFA0DFAF4FF46FAC5" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFA5FFBDFA0DFAF4FF46FAC5" italics="true" pageId="4" pageNumber="5">R. summersi</emphasis>
</taxonomicName>
(One-way ANOVA, SS = 0.009, df = 2, F = 5.654. Between species, P = 0.008. Error: SS = 0.022, df = 29) Right: Drawing of tadpole mouthparts of
<emphasis id="E2B0657FFFA5FFBDFDE8FAB1FD56FA27" box="[608,749,1399,1422]" italics="true" pageId="4" pageNumber="5">
<taxonomicName id="17C4C2EEFFA5FFBDFDE8FAB1FD51FA27" box="[608,746,1399,1422]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="summersi">R. summersi</taxonomicName>
.
</emphasis>
Mouthparts of
<taxonomicName id="17C4C2EEFFA5FFBDFC11FAB1FB9AFA27" box="[921,1057,1399,1422]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA5FFBDFC11FAB1FB9AFA27" box="[921,1057,1399,1422]" italics="true" pageId="4" pageNumber="5">R. benedicta</emphasis>
</taxonomicName>
and
<taxonomicName id="17C4C2EEFFA5FFBDFBD1FAB1FB59FA27" box="[1113,1250,1399,1422]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" sensu="stricto" species="fantastica">
<emphasis id="E2B0657FFFA5FFBDFBD1FAB1FB59FA27" box="[1113,1250,1399,1422]" italics="true" pageId="4" pageNumber="5">R. fantastica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="F983D804FFA5FFBDFB64FAB1FAC8FA27" box="[1260,1395,1399,1422]" pageId="4" pageNumber="5" sensu="stricto">sensu stricto</taxonomicNameLabel>
are identical.
</paragraph>
</caption>
<paragraph id="D07BB96DFFA5FFBDFF4DFA27FB75F9DD" blockId="4.[151,1437,1505,2012]" pageId="4" pageNumber="5">
<emphasis id="E2B0657FFFA5FFBDFF4DFA27FDC4FA52" bold="true" box="[197,639,1505,1531]" pageId="4" pageNumber="5">
Measurements (in mm) of
<typeStatus id="0F7F07CFFFA5FFBDFD85FA27FDC1FA52" box="[525,634,1505,1531]" pageId="4" pageNumber="5" type="holotype">holotype</typeStatus>
.
</emphasis>
The undissected
<typeStatus id="0F7F07CFFFA5FFBDFCC2FA24FC0BFA55" box="[842,944,1506,1532]" pageId="4" pageNumber="5" type="holotype">holotype</typeStatus>
(
<figureCitation id="48FFA5E8FFA5FFBDFC48FA24FBBDFA55" box="[960,1030,1506,1532]" captionStart="FIGURE 4" captionStartId="6.[151,254,1989,2013]" captionTargetBox="[242,1336,206,1975]" captionTargetId="figure@6.[219,1379,182,2012]" captionTargetPageId="6" captionText="FIGURE 4. Type series of R. benedicta. (Top L-R: MUSM 26956 26958, Bottom L-R: MUSM 26959 26962)." httpUri="https://zenodo.org/record/183043/files/figure.png" pageId="4" pageNumber="5">Fig. 4</figureCitation>
) is an adult female as shown by its relatively large size and lack of vocal slits. SVL 17.2; FL 8.1; TL 8.3; KK 15.1; FoL 7.3; HaL 5.0; HL 5.0; HW 5.5; BW 5.5; UEW 2.57; IOD 2.50;
<collectingCountry id="A8D3F9FDFFA5FFBDFDF5F9F4FD0AF9E5" box="[637,689,1586,1612]" name="India" pageId="4" pageNumber="5">IND</collectingCountry>
2.21; TD 0.75; ED 2.14; DET 0.50; L1F 1.86; L2F 2.86; W3D 0.96; W3F 0.43. Measurements of additional specimens are given in the appendix (
<tableCitation id="9D468CD6FFA5FFBDFBECF99CFB07F9DD" box="[1124,1212,1626,1652]" captionStart="TABLE 1" captionStartId="17.[151,239,818,842]" captionTargetBox="[159,1368,874,1785]" captionTargetPageId="17" captionText="TABLE 1. Measurements of the type series of R. benedicta." httpUri="http://table.plazi.org/id/84BBE9E5FFB0FFA8FF1FFCF4FCA3FCE0" pageId="4" pageNumber="5" tableUuid="84BBE9E5FFB0FFA8FF1FFCF4FCA3FCE0">Table 1</tableCitation>
).
</paragraph>
<paragraph id="D07BB96DFFA5FFBDFF4DF947FB29F895" blockId="4.[151,1437,1505,2012]" pageId="4" pageNumber="5">
<emphasis id="E2B0657FFFA5FFBDFF4DF947FE4BF932" bold="true" box="[197,496,1665,1691]" pageId="4" pageNumber="5">
Description of
<typeStatus id="0F7F07CFFFA5FFBDFEF5F947FE51F932" box="[381,490,1665,1691]" pageId="4" pageNumber="5" type="holotype">holotype</typeStatus>
.
</emphasis>
Widest part of head is at jaw articulations. Head width as wide as body. Tongue ovoid; teeth absent. In life, head bright red with comma-shaped black spots over the eyes and tympanum. Red coloration extends posteriorly to level of the axillae; black ground color with blue reticulation on dorsal body. Limbs and digits black, limbs reticulated with blue. Underside of head red, lacking paired gular spots. Venter black with discrete blue reticulation. Nares black surrounded by black spots. Iris black.
</paragraph>
<paragraph id="D07BB96DFFA5FFBCFF4DF88CFA28FF73" blockId="4.[151,1437,1505,2012]" lastBlockId="5.[151,1437,152,498]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">In life, skin texture nearly smooth on the dorsal surfaces of the body and head; limbs and rump weakly granular. Venter weakly granular on limbs and body, ventral surface of head nearly smooth. Snout sloping and rounded in lateral profile, round or slightly blunted in dorsal profile. Nares situated at tip of snout and directed laterally; both nares visible from ventral and anterior view but not from dorsal view. Canthus rostralis rounded, loreal region flat and nearly vertical. Upper eyelid approximately equal in width to interorbital distance; internarial distance roughly equal to eye width. Tympanum round, partially concealed posterodorsally.</paragraph>
<paragraph id="D07BB96DFFA4FFBCFF4DFF2EFED7FE5B" blockId="5.[151,1437,152,498]" pageId="5" pageNumber="6">
Hands relatively large, length 29 % of SVL. Relative length of appressed fingers III&gt; IV
<emphasis id="E2B0657FFFA4FFBCFB54FF2DFB52FEA9" bold="true" box="[1244,1257,235,256]" pageId="5" pageNumber="6"></emphasis>
II&gt; I; first finger 65 % length of second; finger discs moderately expanded, width of disc on finger III 2.2 times width of adjacent phalanx. An unpigmented median metacarpal tubercle is present on base of palm; inner metacarpal tubercle present near base of finger I but angled posteriorly; unpigmented proximal subarticular tubercles present on base of each digit, except on finger I, where tubercle is part-way up the digit; distal subarticular tubercle visible only on fingers III and IV. All tubercles raised above level of hands; scutes present on dorsal surface of fingers.
</paragraph>
<caption id="84BBE9E5FFA4FFBCFF10FB11FEF4FA9A" httpUri="https://zenodo.org/record/183042/files/figure.png" pageId="5" pageNumber="6" targetBox="[165,1429,552,1216]" targetPageId="5">
<paragraph id="D07BB96DFFA4FFBCFF10FB11FEF4FA9A" blockId="5.[151,1436,1239,1331]" pageId="5" pageNumber="6">
<emphasis id="E2B0657FFFA4FFBCFF10FB11FEA1FB46" bold="true" box="[152,282,1239,1263]" pageId="5" pageNumber="6">FIGURE 3.</emphasis>
Advertisement calls of (A)
<taxonomicName id="17C4C2EEFFA4FFBCFDC1FB11FD6AFB47" box="[585,721,1239,1262]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA4FFBCFDC1FB11FD6AFB47" box="[585,721,1239,1262]" italics="true" pageId="5" pageNumber="6">R. fantastica</emphasis>
</taxonomicName>
individual from Cainarachi Valley, recording at 24 C, (B)
<taxonomicName id="17C4C2EEFFA4FFBCFAC2FB11FF69FAB8" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFA4FFBCFAC2FB11FF69FAB8" italics="true" pageId="5" pageNumber="6">R. summersi</emphasis>
</taxonomicName>
individual from Sauce, recorded at 24.5 C in captivity, (C)
<taxonomicName id="17C4C2EEFFA4FFBCFCCFFB3CFC76FAB8" box="[839,973,1274,1297]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA4FFBCFCCFFB3CFC76FAB8" box="[839,973,1274,1297]" italics="true" pageId="5" pageNumber="6">R. benedicta</emphasis>
</taxonomicName>
individual from Shucushuyacu, recorded at 24 C in captivity.
</paragraph>
</caption>
<paragraph id="D07BB96DFFA4FFBCFF4DFAA1FCEFF968" blockId="5.[151,1437,1383,2009]" pageId="5" pageNumber="6">
<collectingCountry id="A8D3F9FDFFA4FFBCFF4DFAA1FEBBFA28" box="[197,256,1383,1409]" name="India" pageId="5" pageNumber="6">Hind</collectingCountry>
limbs moderate length, with heel of appressed hind limbs reaching level of eye. Femur and tibia roughly equal in length, tibia 102 % length of femur; knee-knee distance 88 % of SVL. Relative lengths of appressed toes IV&gt; V
<emphasis id="E2B0657FFFA4FFBCFE21FA7CFE0DFA66" bold="true" box="[425,438,1466,1487]" pageId="5" pageNumber="6"></emphasis>
III&gt; II&gt; I; first toe short with unexpanded disc; second toe with slightly expanded disc, discs on toes IIIV moderately expanded. Two unpigmented metatarsal tubercles present on base of foot, one situated medially near base of toe I, the other situated laterally at the base of the fifth metatarsal. Proximal subarticular tubercles present at base of each toe but most notable on toes I and II due to their lack of pigmentation. Toes III and V with two subarticular tubercles, toe IV with three subarticular tubercles. A tarsal keel is present starting below the knee and turning into the medial metatarsal tubercle at the foot. Tarsal tubercle absent; feet and hands lacking webbing and lateral fringing.
</paragraph>
<paragraph id="D07BB96DFFA4FFBCFF4DF909FD92F870" blockId="5.[151,1437,1383,2009]" pageId="5" pageNumber="6">
<emphasis id="E2B0657FFFA4FFBCFF4DF909FE84F940" bold="true" box="[197,319,1743,1769]" pageId="5" pageNumber="6">Variation.</emphasis>
Adults 15.0
<quantity id="173C1488FFA4FFBCFE6FF909FDE8F940" box="[487,595,1743,1769]" metricMagnitude="-2" metricUnit="m" metricValue="2.02" pageId="5" pageNumber="6" unit="mm" value="20.2">20.2 mm</quantity>
SVL, females slightly larger than males: three adult males 15.0
<quantity id="173C1488FFA4FFBCFAE2F909FF7EF8B8" metricMagnitude="-2" metricUnit="m" metricValue="1.75" pageId="5" pageNumber="6" unit="mm" value="17.5">17.5 mm</quantity>
(mean
<quantity id="173C1488FFA4FFBCFE94F931FE32F8B8" box="[284,393,1783,1809]" metricMagnitude="-2" metricUnit="m" metricValue="1.65" pageId="5" pageNumber="6" unit="mm" value="16.5">16.5 mm</quantity>
); four adult females
<quantity id="173C1488FFA4FFBCFDF1F931FC9FF8B8" box="[633,804,1783,1809]" metricMagnitude="-2" metricUnit="m" metricValue="1.85" metricValueMax="2.02" metricValueMin="1.68" pageId="5" pageNumber="6" unit="mm" value="18.5" valueMax="20.2" valueMin="16.8">16.820.2 mm</quantity>
(mean
<quantity id="173C1488FFA4FFBCFCF3F931FC53F8B8" box="[891,1000,1783,1809]" metricMagnitude="-2" metricUnit="m" metricValue="1.8399999999999999" pageId="5" pageNumber="6" unit="mm" value="18.4">18.4 mm</quantity>
). Widest part of head is at jaw articulations. Head about as wide as body except in two apparently gravid females whose bodies are wider than the head. Head width 92 % of body width in three males (range 80106 %), 94 % percent of body width in four females (range 74121 %). Head width 2734 % of SVL in adults. No apparent sexual dimorphism in external morphology except that males are smaller, possess faint vocal slits on the floor of the mouth, and have a slightly expanded subgular pouch.
</paragraph>
<caption id="84BBE9E5FFA7FFBFFF1FF803FAF9F874" box="[151,1346,1989,2013]" httpUri="https://zenodo.org/record/183043/files/figure.png" pageId="6" pageNumber="7" targetBox="[242,1336,206,1975]" targetPageId="6">
<paragraph id="D07BB96DFFA7FFBFFF1FF803FAF9F874" blockId="6.[151,1346,1989,2013]" box="[151,1346,1989,2013]" pageId="6" pageNumber="7">
<emphasis id="E2B0657FFFA7FFBFFF1FF803FEA3F874" bold="true" box="[151,280,1989,2013]" pageId="6" pageNumber="7">FIGURE 4.</emphasis>
Type series of
<emphasis id="E2B0657FFFA7FFBFFE33F800FDF3F874" box="[443,584,1990,2013]" italics="true" pageId="6" pageNumber="7">
<taxonomicName id="17C4C2EEFFA7FFBFFE33F800FDFFF874" box="[443,580,1990,2013]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="benedicta">R. benedicta</taxonomicName>
.
</emphasis>
(Top L-R: MUSM 2695626958, Bottom L-R: MUSM 2695926962).
</paragraph>
</caption>
<paragraph id="D07BB96DFFA6FFBEFF4DFF5EFB51FDB3" blockId="7.[151,1437,152,2018]" pageId="7" pageNumber="8">Live animals bear a diagnostic scarlet-red head with black spots over the eyes. These spots are medially fused in some individuals and form a W-shaped head crest which descends over the eyes creating a face-mask; in most individuals this mask also covers the tympanum. The red coloration on the head extends posterior to level of the axillae; from here the dorsal body coloration changes to a black ground color with blue reticulation. This reticulation is diffuse in some individuals, sometimes to the point of the body appearing uniform blue on the rump. Limbs and digits are colored as the dorsum: individuals with defined dorsal reticulation have limbs reticulated with blue; individuals with diffuse reticulation on the body have either diffuse limb reticulation or limbs colored uniformly blue. Underside of head red, most individuals possessing some amount of black marbling on chin. Venter black with diffuse or discrete blue reticulation. Nares black and surrounded by black spots, these spots are sometimes fused to form a bar across the loreal region.</paragraph>
<paragraph id="D07BB96DFFA6FFBEFF4DFDEEFB94FD13" blockId="7.[151,1437,152,2018]" pageId="7" pageNumber="8">
Hands relatively large, length 2429 % of SVL. First finger 5681 % length of second; finger discs moderately expanded in both males and females, width of disc on finger
<date id="A47A9FADFFA6FFBEFC3BFD96FC5EFDC3" box="[947,997,592,618]" pageId="7" pageNumber="8">III 2</date>
2.3 times width of adjacent phalanx. Tibia 85115 % length of femur (mean 99 %); knee-knee distance 8193 % of SVL (mean 87 %). In preservative, red coloration turning pale pink and blue reticulation turning purple-grey.
</paragraph>
<paragraph id="D07BB96DFFA6FFBEFF4DFD01FDBCFBE3" blockId="7.[151,1437,152,2018]" pageId="7" pageNumber="8">
<emphasis id="E2B0657FFFA6FFBEFF4DFD01FE8AFD48" bold="true" box="[197,305,711,737]" pageId="7" pageNumber="8">Tadpole.</emphasis>
One tadpole (captive bred from topotypic parents) was used for the mouthpart description. The body of the tadpole was destroyed after preservation making a full description impossible, but the mouthparts remained intact. Oral disc emarginate, anterior and posterior labia forming flaps free from body wall. Marginal papillae absent on anterior labium except for lateral-most portion (34 papillae present), present in one complete row on posterior labium. Papillae white, rounded; submarginal papillae absent. Jaw sheaths deep in longitudinal width, serrate, lacking indentations. Lateral processes short, extending barely past lower jaw. Labial tooth row formula is 2(2)/3. A-1 complete, A-2 with medial gap, same width as A-1. P-1, P-2, and P-3 complete; P-1 and P-2 equal width, P-3 slightly shorter. We examined two
<taxonomicName id="17C4C2EEFFA6FFBEFBC4FC26FB5DFC50" box="[1100,1254,992,1017]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA6FFBEFBC4FC26FB5DFC50" box="[1100,1254,992,1017]" italics="true" pageId="7" pageNumber="8">R. fantastica</emphasis>
</taxonomicName>
tadpoles from Tarapoto and the upper Cainarachi Valley (stages 27 and 29) and the mouthparts were indistinguishable from the single
<taxonomicName id="17C4C2EEFFA6FFBEFE87FBF6FE1AFBE0" box="[271,417,1072,1097]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA6FFBEFE87FBF6FE1AFBE0" box="[271,417,1072,1097]" italics="true" pageId="7" pageNumber="8">R. benedicta</emphasis>
</taxonomicName>
tadpole.
</paragraph>
<paragraph id="D07BB96DFFA6FFBEFF4DFB91FB75FAC8" blockId="7.[151,1437,152,2018]" pageId="7" pageNumber="8">
<emphasis id="E2B0657FFFA6FFBEFF4DFB91FEC9FBD8" bold="true" box="[197,370,1111,1137]" pageId="7" pageNumber="8">Vocalizations.</emphasis>
The advertisement call is a series of buzz-like notes (
<figureCitation id="48FFA5E8FFA6FFBEFC7EFB9EFB85FBDB" box="[1014,1086,1112,1138]" captionStart="FIGURE 3" captionStartId="5.[152,255,1239,1263]" captionTargetBox="[165,1429,552,1216]" captionTargetId="figure@5.[155,1440,546,1225]" captionTargetPageId="5" captionText="FIGURE 3. Advertisement calls of (A) R. fantastica individual from Cainarachi Valley, recording at 24 C, (B) R. summersi individual from Sauce, recorded at 24.5 C in captivity, (C) R. benedicta individual from Shucushuyacu, recorded at 24 C in captivity." httpUri="https://zenodo.org/record/183042/files/figure.png" pageId="7" pageNumber="8">Fig. 3</figureCitation>
). The dominant frequency of the call starts at 4240 Hz and gradually decreases in subsequent notes to 3190 Hz (air temperature 24° C). The notes in the calls were very short with a duration of 150 ± 20 ms (vs. 290 ± 30 ms in
<taxonomicName id="17C4C2EEFFA6FFBEFB01FB6EFAA6FB68" box="[1161,1309,1192,1217]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA6FFBEFB01FB6EFAA6FB68" box="[1161,1309,1192,1217]" italics="true" pageId="7" pageNumber="8">R. fantastica</emphasis>
</taxonomicName>
and 440 ± 30 ms in
<taxonomicName id="17C4C2EEFFA6FFBEFE8DFB16FE2DFB40" box="[261,406,1232,1257]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFA6FFBEFE8DFB16FE2DFB40" box="[261,406,1232,1257]" italics="true" pageId="7" pageNumber="8">R. summersi</emphasis>
</taxonomicName>
); brief pauses between each note lasting 140 ± 20 ms (vs. 200 ± 20 ms in
<taxonomicName id="17C4C2EEFFA6FFBEFA8FFB16FA20FB40" box="[1287,1435,1232,1257]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA6FFBEFA8FFB16FA20FB40" box="[1287,1435,1232,1257]" italics="true" pageId="7" pageNumber="8">R. fantastica</emphasis>
</taxonomicName>
and 150 ± 20 ms in
<taxonomicName id="17C4C2EEFFA6FFBEFE08FB3EFDABFAB8" box="[384,528,1272,1297]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFA6FFBEFE08FB3EFDABFAB8" box="[384,528,1272,1297]" italics="true" pageId="7" pageNumber="8">R. summersi</emphasis>
</taxonomicName>
). The ratio of note length to pause length is approximately 1:1 (vs 1.4:
<quantity id="173C1488FFA6FFBEFADAFB3EFAC0FABB" box="[1362,1403,1272,1298]" metricMagnitude="-2" metricUnit="m" metricValue="2.54" pageId="7" pageNumber="8" unit="in" value="1.0">1 in</quantity>
<taxonomicName id="17C4C2EEFFA6FFBEFA0BFB3EFEB0FA90" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA6FFBEFA0BFB3EFEB0FA90" italics="true" pageId="7" pageNumber="8">R. fantastica</emphasis>
</taxonomicName>
and 2.8:
<quantity id="173C1488FFA6FFBEFEFEFAE6FE1AFA93" box="[374,417,1312,1338]" metricMagnitude="-2" metricUnit="m" metricValue="2.54" pageId="7" pageNumber="8" unit="in" value="1.0">1 in</quantity>
<taxonomicName id="17C4C2EEFFA6FFBEFE21FAE6FD80FA90" box="[425,571,1312,1337]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFA6FFBEFE21FAE6FD80FA90" box="[425,571,1312,1337]" italics="true" pageId="7" pageNumber="8">R. summersi</emphasis>
</taxonomicName>
). The call can be heard from as far as ~8 meters which is comparatively loud for a member of the
<taxonomicName id="17C4C2EEFFA6FFBEFE4AFA8EFD8DFAC8" box="[450,566,1352,1377]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA6FFBEFE4AFA8EFD8DFAC8" box="[450,566,1352,1377]" italics="true" pageId="7" pageNumber="8">fantastica</emphasis>
</taxonomicName>
group (vs.
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in
<taxonomicName id="17C4C2EEFFA6FFBEFC85FA8EFC1AFAC8" box="[781,929,1352,1377]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA6FFBEFC85FA8EFC1AFAC8" box="[781,929,1352,1377]" italics="true" pageId="7" pageNumber="8">R. fantastica</emphasis>
</taxonomicName>
and
<quantity id="173C1488FFA6FFBEFC52FA8EFBB3FACB" box="[986,1032,1352,1378]" metricMagnitude="0" metricUnit="m" metricValue="1.0" pageId="7" pageNumber="8" unit="m" value="1.0">1 m</quantity>
in
<taxonomicName id="17C4C2EEFFA6FFBEFBA5FA8EFB06FAC8" box="[1069,1213,1352,1377]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFA6FFBEFBA5FA8EFB06FAC8" box="[1069,1213,1352,1377]" italics="true" pageId="7" pageNumber="8">R. summersi</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="98DEEAE6FFA6FFB1FF4DFAA9FED0F96B" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8" type="distribution">
<paragraph id="D07BB96DFFA6FFBEFF4DFAA9FDDFF8B3" blockId="7.[151,1437,152,2018]" pageId="7" pageNumber="8">
<emphasis id="E2B0657FFFA6FFBEFF4DFAA9FDE5FA20" bold="true" box="[197,606,1391,1417]" pageId="7" pageNumber="8">Distribution and natural history.</emphasis>
This frog appears to be widely distributed throughout the lowland forest of the Pampas del Sacramento in southern Loreto and eastern San Martin (
<figureCitation id="48FFA5E8FFA6FFBEFBA6FA5EFBCDFA1B" box="[1070,1142,1432,1458]" captionStart="FIGURE 5" captionStartId="9.[151,254,1864,1888]" captionTargetBox="[261,1339,325,1846]" captionTargetId="figure@9.[253,1346,314,1850]" captionTargetPageId="9" captionText="FIGURE 5. Map of known localities and estimated distributions of R. fantastica, R. summersi, and R. benedicta. Because some populations where close together, the 14 sampling localities were simplified into 10 geographic groups. Localities which were included in the phylogenetic analysis are indicated by numbers on the map: (1) Tarapoto; (2) Cainarachi Valley; (3) Convento; (4) Pongo de Cainarachi; (5) Varadero; (6) Lower Huallaga Canyon; (7) Shucushuyacu; (8) Pampa Hermosa; (9) Sauce; (10) Chipaota." httpUri="https://zenodo.org/record/183044/files/figure.png" pageId="7" pageNumber="8">Fig. 5</figureCitation>
). The Pampas del Sacramento are enclosed by Rio Ucayali to the east, Cordillera Azul and Rio Huallaga to the west and northwest, and the flooded forests of Pacaya-Samiria to the north.
<taxonomicName id="17C4C2EEFFA6FFBEFC91FA2EFBA5F9A8" box="[793,1054,1512,1537]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA6FFBEFC91FA2EFBA5F9A8" box="[793,1054,1512,1537]" italics="true" pageId="7" pageNumber="8">Ranitomeya benedicta</emphasis>
</taxonomicName>
appears to occur throughout this region in forests over ca.
<quantity id="173C1488FFA6FFBEFE58F9D6FDA5F983" box="[464,542,1552,1578]" metricMagnitude="2" metricUnit="m" metricValue="1.5" pageId="7" pageNumber="8" unit="m" value="150.0">150 m</quantity>
elevation. The southern margin of its range likely extends at least to Rio Cushabatay, but not reaching Pucallpa, as frequent surveys along the Aguaytia-Pucallpa road since 2006 have failed to encounter any members of the
<taxonomicName id="17C4C2EEFFA6FFBEFDEFF9A6FD60F9D0" box="[615,731,1632,1657]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA6FFBEFDEFF9A6FD60F9D0" box="[615,731,1632,1657]" italics="true" pageId="7" pageNumber="8">fantastica</emphasis>
</taxonomicName>
group.
<taxonomicName id="17C4C2EEFFA6FFBEFCBCF9A6FB82F9D0" box="[820,1081,1632,1657]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA6FFBEFCBCF9A6FB82F9D0" box="[820,1081,1632,1657]" italics="true" pageId="7" pageNumber="8">Ranitomeya benedicta</emphasis>
</taxonomicName>
has never been observed west of Rio Huallaga or east of Rio Ucayali, and both rivers appear to be boundaries restricting its westward and eastward distribution, respectively. The Ucayali and the lower Huallaga are generally surrounded by 1050 km-wide flood plains and wetlands; areas which are inhospitable to most poison frogs and may be more effective barriers than the rivers themselves.
</paragraph>
<paragraph id="D07BB96DFFA6FFB1FF4DF8EEFB57FE83" blockId="7.[151,1437,152,2018]" lastBlockId="8.[151,1437,152,1580]" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8">
These frogs are most frequently encountered amidst the tangled branches in or near tree-falls. They are extremely alert and agile, and when encountered, immediately flee to the ground and disappear amidst the leaf litter, jumping in erratic spurts. These frogs are rarely encountered and often after days of searching we were only able to find one or two individuals. Their distribution appears to be patchy but in some areas locally abundant. This species appears to favor slightly elevated areas within the forest and have not been found areas susceptible seasonal flooding. Although
<taxonomicName id="17C4C2EEFFA9FFB1FDF6FF5EFCA8FF18" box="[638,787,152,177]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA9FFB1FDF6FF5EFCA8FF18" box="[638,787,152,177]" italics="true" pageId="8" pageNumber="9">R. benedicta</emphasis>
</taxonomicName>
is primarily a lowland species, individuals have been found in higher elevation areas in the Huallaga Canyon at
<quantity id="173C1488FFA9FFB1FCC7FF06FC20FF73" box="[847,923,192,218]" metricMagnitude="2" metricUnit="m" metricValue="3.15" pageId="8" pageNumber="9" unit="m" value="315.0">315 m</quantity>
and in the upper Cushabatay at
<quantity id="173C1488FFA9FFB1FA95FF06FAD3FF73" box="[1309,1384,192,218]" metricMagnitude="2" metricUnit="m" metricValue="4.05" pageId="8" pageNumber="9" unit="m" value="405.0">405 m</quantity>
(M. Ramírez Zárate pers. comm.). These records may represent the upper altitudinal limits of this species, although further exploration of the Cordillera Azul may uncover additional montane localities.
</paragraph>
<paragraph id="D07BB96DFFA9FFB1FF4DFEFEFDA5FDC3" blockId="8.[151,1437,152,1580]" pageId="8" pageNumber="9">
In the northwestern extent of its distribution,
<taxonomicName id="17C4C2EEFFA9FFB1FD51FEFEFCD0FEF8" box="[729,875,312,337]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA9FFB1FD51FEFEFCD0FEF8" box="[729,875,312,337]" italics="true" pageId="8" pageNumber="9">R. benedicta</emphasis>
</taxonomicName>
is sympatric with
<taxonomicName id="17C4C2EEFFA9FFB1FBCCFEFEFB78FEF8" box="[1092,1219,312,337]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="imitator">
<emphasis id="E2B0657FFFA9FFB1FBCCFEFEFB78FEF8" box="[1092,1219,312,337]" italics="true" pageId="8" pageNumber="9">R. imitator</emphasis>
</taxonomicName>
and
<taxonomicName id="17C4C2EEFFA9FFB1FB74FEFEFF68FED0" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="ventrimaculata">
<emphasis id="E2B0657FFFA9FFB1FB74FEFEFF68FED0" italics="true" pageId="8" pageNumber="9">R. ventrimaculata</emphasis>
</taxonomicName>
. Reproduction occurs terrestrially, with clutches of 46 grey eggs being laid amongst humid leaves accumulated on the forest floor. Egg development lasts 1216 days (
<collectingCountry id="A8D3F9FDFFA9FFB1FCF6FE4EFC7BFE0B" box="[894,960,392,418]" name="Egypt" pageId="8" pageNumber="9">MSR</collectingCountry>
, pers. obsv.). Tadpoles are transported to larger phytotelmata, such as
<emphasis id="E2B0657FFFA9FFB1FE6CFE76FDE7FE60" box="[484,604,432,457]" italics="true" pageId="8" pageNumber="9">Aechemea</emphasis>
bromeliads, and (in captivity) metamorphose in about 100 days (
<collectingCountry id="A8D3F9FDFFA9FFB1FADDFE76FA23FE63" box="[1365,1432,432,458]" name="Egypt" pageId="8" pageNumber="9">MSR</collectingCountry>
, pers. obsv.). Due to the low phytotelmata abundance in the forests of the Pampas del Sacramento and the observation that
<taxonomicName id="17C4C2EEFFA9FFB1FED1FDC6FD91FDB0" box="[345,554,512,537]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="ventrimaculata">
<emphasis id="E2B0657FFFA9FFB1FED1FDC6FD91FDB0" box="[345,554,512,537]" italics="true" pageId="8" pageNumber="9">R. ventrimaculata</emphasis>
</taxonomicName>
are the tadpoles most commonly encountered, there is likely strong competition for phytotelmata access, and phytotelmata abundance may be a major limiting factor on population sizes in
<taxonomicName id="17C4C2EEFFA9FFB1FF3DFD96FDA1FDC3" authority="Donnelly, 1989" authorityName="Donnelly" authorityYear="1989" box="[181,538,592,618]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA9FFB1FF3DFD96FEF3FDC0" box="[181,328,592,617]" italics="true" pageId="8" pageNumber="9">R. benedicta</emphasis>
(
<bibRefCitation id="B455C49CFFA9FFB1FEDFFD96FDAAFDC3" author="Donnelly" box="[343,529,592,618]" pageId="8" pageNumber="17" refString="Donnelly, M. A. (1989) Demographic effects of reproductive resource supplementation in a territorial frog, Dendrobates pumilio. Ecological Monographs, 59, 207 - 221." type="journal article" year="1989">Donnelly, 1989</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="D07BB96DFFA9FFB1FF4DFDBEFCC7FCA3" blockId="8.[151,1437,152,1580]" pageId="8" pageNumber="9">
Aside from terrestrial reproduction and foraging, these frogs appear to be arboreal, which could partially explain why they are so difficult to find. Near the village of Shucushuyacu, farmers felling large trees frequently encounter this frog fleeing from the large canopy bromeliads (i.e.
<emphasis id="E2B0657FFFA9FFB1FBA6FD0EFB1DFD48" box="[1070,1190,712,737]" italics="true" pageId="8" pageNumber="9">Aechemea</emphasis>
). Timber harvesters along the Rio Ucayali have also reported the same observation.
</paragraph>
<paragraph id="D07BB96DFFA9FFB1FF4DFCDEFC46FCF3" blockId="8.[151,1437,152,1580]" pageId="8" pageNumber="9">These frogs also are extremely tolerant of heat. We observed one frog carrying tadpoles across a large, deforested area in full, midday sun (36° C) and was seemingly unaffected.</paragraph>
<paragraph id="D07BB96DFFA9FFB1FF4DFCA1FEB7FB35" blockId="8.[151,1437,152,1580]" pageId="8" pageNumber="9">
<emphasis id="E2B0657FFFA9FFB1FF4DFCA1FE7EFC28" bold="true" box="[197,453,871,897]" pageId="8" pageNumber="9">Conservation status.</emphasis>
Under the IUCN Red List criteria (IUCN, 2001), we suggest
<taxonomicName id="17C4C2EEFFA9FFB1FB16FCAEFA8AFC28" box="[1182,1329,872,897]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA9FFB1FB16FCAEFA8AFC28" box="[1182,1329,872,897]" italics="true" pageId="8" pageNumber="9">R. benedicta</emphasis>
</taxonomicName>
be listed as Vulnerable (VU) under the following criteria (1) we estimate its extent of occurrence at
<quantity id="173C1488FFA9FFB1FA99FC54FA2FFC05" box="[1297,1428,914,940]" metricMagnitude="7" metricUnit="m" metricValue="1.9" pageId="8" pageNumber="9" unit="km" value="19000.0">19,000 km</quantity>
2 (although the area of occupancy may be much lower), and ongoing deforestation along the lower Huallaga will reduce the amount of suitable habitat substantially over the coming years, (2) this species is found primarily in undisturbed rainforest and does not occur in areas disturbed by human activity, (3) populations appear to be patchily distributed, and (4) there will be a high demand for this species in the pet trade. This species does occur in a protected area (Cordillera Azul National Park), although the extent to which it occurs there is unknown.
</paragraph>
<paragraph id="D07BB96DFFA9FFB1FF4EFB6CFCE9F985" blockId="8.[151,1437,152,1580]" pageId="8" pageNumber="9">
We fear that by describing this species (and others), we are also putting them at risk of illegal smuggling, and expect that there will be a large influx of illegally acquired
<taxonomicName id="17C4C2EEFFA9FFB1FCF6FB15FBAAFB45" box="[894,1041,1235,1260]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA9FFB1FCF6FB15FBAAFB45" box="[894,1041,1235,1260]" italics="true" pageId="8" pageNumber="9">R. benedicta</emphasis>
</taxonomicName>
to the North American and European markets over the next few years. This is particularly frustrating for the authors who try to minimize these negative impacts through sustainable harvest and legal exportation (i.e. Understory Enterprises). This organization is concerned with preserving species in their native habitats, while sustainably harvesting individuals for the pet trade. We encourage anyone interested in owning any poison frog to exercise restraint and purchase only sustainably-harvested or captive-bred frogs of legal origin. Further, consumers must understand that when these projects are done correctly; they take a considerable amount of time and money. When a species is illegally smuggled, it reduces the potential market for the legal business, and as a result, reduces the amount of money these organizations can generate for land protection.
</paragraph>
<paragraph id="D07BB96DFFA9FFB1FF1FF946FED0F96B" blockId="8.[151,529,1664,1730]" pageId="8" pageNumber="9">
<taxonomicName id="17C4C2EEFFA9FFB1FF1FF946FE1FF933" box="[151,420,1664,1690]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="summersi" status="sp. nov.">
<emphasis id="E2B0657FFFA9FFB1FF1FF946FE1FF933" bold="true" box="[151,420,1664,1690]" italics="true" pageId="8" pageNumber="9">Ranitomeya summersi</emphasis>
</taxonomicName>
,
<taxonomicNameLabel id="F983D804FFA9FFB1FE3BF946FDAAF933" box="[435,529,1664,1690]" pageId="8" pageNumber="9" rank="species">
<emphasis id="E2B0657FFFA9FFB1FE3BF946FDAAF933" bold="true" box="[435,529,1664,1690]" pageId="8" pageNumber="9">sp. nov.</emphasis>
</taxonomicNameLabel>
<figureCitation id="48FFA5E8FFA9FFB1FF1FF96EFEBEF96B" box="[151,261,1704,1730]" captionStart="FIGURE 1" captionStartId="3.[151,254,1345,1369]" captionTargetBox="[261,1328,264,1330]" captionTargetId="figure@3.[255,1333,258,1338]" captionTargetPageId="3" captionText="FIGURE 1. Drawing of species contained in the fantastica complex. Clockwise from left: Ranitomeya summersi sp. nov., R. fantastica sensu stricto, R. benedicta sp. nov. Illustration of R. fantastica is an artists rendering of the nominal population from Yurimaguas based on the original type series and recent field work nearby." httpUri="https://zenodo.org/record/183040/files/figure.png" pageId="8" pageNumber="9">Figures 1</figureCitation>
,
<figureCitation id="48FFA5E8FFA9FFB1FE9BF96EFE98F96B" box="[275,291,1704,1730]" captionStart="FIGURE 5" captionStartId="9.[151,254,1864,1888]" captionTargetBox="[261,1339,325,1846]" captionTargetId="figure@9.[253,1346,314,1850]" captionTargetPageId="9" captionText="FIGURE 5. Map of known localities and estimated distributions of R. fantastica, R. summersi, and R. benedicta. Because some populations where close together, the 14 sampling localities were simplified into 10 geographic groups. Localities which were included in the phylogenetic analysis are indicated by numbers on the map: (1) Tarapoto; (2) Cainarachi Valley; (3) Convento; (4) Pongo de Cainarachi; (5) Varadero; (6) Lower Huallaga Canyon; (7) Shucushuyacu; (8) Pampa Hermosa; (9) Sauce; (10) Chipaota." httpUri="https://zenodo.org/record/183044/files/figure.png" pageId="8" pageNumber="9">5</figureCitation>
,
<figureCitation id="48FFA5E8FFA9FFB1FEB9F96EFEF9F96B" box="[305,322,1704,1730]" captionStart="FIGURE 6" captionStartId="10.[151,254,1958,1982]" captionTargetBox="[353,1250,213,1929]" captionTargetId="figure@10.[334,1268,194,1951]" captionTargetPageId="10" captionText="FIGURE 6. Type series of R. summersi. (Top L-R: MUSM 26949, MUSM 26967, MUSM 26991, Bottom L-R: MUSM 26992 26994)" httpUri="https://zenodo.org/record/183045/files/figure.png" pageId="8" pageNumber="9">6</figureCitation>
,
<figureCitation id="48FFA5E8FFA9FFB1FEC6F96EFED0F96B" box="[334,363,1704,1730]" captionStart="FIGURE 10" captionStartId="22.[151,254,1884,1908]" captionTargetBox="[159,1428,201,1852]" captionTargetId="figure@22.[151,1436,194,1861]" captionTargetPageId="22" captionText="FIGURE 10. Photos of R. summersi (A H). I L: Type series of R. fantastica (1947.2.15.1 4, order not known, photos from Natural History Museum London). Individual L (top): Inset photo in lower-right corner shows close-up of head spotting." httpUri="https://zenodo.org/record/183049/files/figure.png" pageId="8" pageNumber="9">10</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="98DEEAE6FFA9FFB1FF1FF93EFB29F806" pageId="8" pageNumber="9" type="reference_group">
<paragraph id="D07BB96DFFA9FFB1FF1FF93EFC09F826" blockId="8.[151,1436,1784,1967]" pageId="8" pageNumber="9">
<taxonomicName id="17C4C2EEFFA9FFB1FF1FF93EFE71F8A6" ID-CoL="34M9M" box="[151,458,1784,1807]" class="Amphibia" family="Dendrobatidae" genus="Dendrobates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="quinquevittatus">
<emphasis id="E2B0657FFFA9FFB1FF1FF93EFE71F8A6" box="[151,458,1784,1807]" italics="true" pageId="8" pageNumber="9">Dendrobates quinquevittatus</emphasis>
</taxonomicName>
(non Steindachner):
<bibRefCitation id="B455C49CFFA9FFB1FD22F93EFCDEF8A6" author="Silverstone" box="[682,869,1784,1807]" pageId="8" pageNumber="18" refString="Silverstone, P. A. (1975) A revision of the poison-arrow frogs of the genus Dendrobates Wagler. Natural History Museum of Los Angeles County, 21, 1 - 55." type="journal article" year="1975">Silverstone, 1975</bibRefCitation>
(partim), p. 3336, Fig. 14 (drawing), patterns G, H.
<taxonomicName id="17C4C2EEFFA9FFB1FF1FF8DEFE20F886" ID-CoL="34M8B" box="[151,411,1816,1839]" class="Amphibia" family="Dendrobatidae" genus="Dendrobates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="fantasticus">
<emphasis id="E2B0657FFFA9FFB1FF1FF8DEFE20F886" box="[151,411,1816,1839]" italics="true" pageId="8" pageNumber="9">Dendrobates fantasticus</emphasis>
</taxonomicName>
(non Boulenger):
<bibRefCitation id="B455C49CFFA9FFB1FDECF8DEFD43F886" author="Schulte" box="[612,760,1816,1839]" pageId="8" pageNumber="18" refString="Schulte, R. (1999) Pfeilgiftfrosche &quot; Artenteil - Peru &quot;. INBICO, Wailblingen, Germany, 294 pp." type="book" year="1999">Schulte, 1999</bibRefCitation>
(partim), pp. 5769,
<figureCitation id="48FFA5E8FFA9FFB1FC6AF8DEFB99F886" box="[994,1058,1816,1839]" captionStart="FIGURE 5" captionStartId="9.[151,254,1864,1888]" captionTargetBox="[261,1339,325,1846]" captionTargetId="figure@9.[253,1346,314,1850]" captionTargetPageId="9" captionText="FIGURE 5. Map of known localities and estimated distributions of R. fantastica, R. summersi, and R. benedicta. Because some populations where close together, the 14 sampling localities were simplified into 10 geographic groups. Localities which were included in the phylogenetic analysis are indicated by numbers on the map: (1) Tarapoto; (2) Cainarachi Valley; (3) Convento; (4) Pongo de Cainarachi; (5) Varadero; (6) Lower Huallaga Canyon; (7) Shucushuyacu; (8) Pampa Hermosa; (9) Sauce; (10) Chipaota." httpUri="https://zenodo.org/record/183044/files/figure.png" pageId="8" pageNumber="9">Fig. 5</figureCitation>
patterns H, L, Cordillera Oriental, “West flank” and “North spur” morphs, (H and L reprinted from
<bibRefCitation id="B455C49CFFA9FFB1FC05F8FEFBF6F8E6" author="Silverstone" box="[909,1101,1848,1871]" pageId="8" pageNumber="18" refString="Silverstone, P. A. (1975) A revision of the poison-arrow frogs of the genus Dendrobates Wagler. Natural History Museum of Los Angeles County, 21, 1 - 55." type="journal article" year="1975">Silverstone, 1975</bibRefCitation>
).
<bibRefCitation id="B455C49CFFA9FFB1FBE9F8FEFA8EF8E6" author="Symula" box="[1121,1333,1848,1871]" pageId="8" pageNumber="18" refString="Symula, R., Schulte, R. &amp; Summers, K. (2001) Molecular phylogenetic evidence for a mimetic radiation in Peruvian poison frogs supports a Mullerian mimicry hypothesis. Proceedings of the Royal Society of London Series B, 268, 2415 - 2421." type="journal article" year="2001">
Symula
<emphasis id="E2B0657FFFA9FFB1FB32F8FEFB50F8E6" box="[1210,1259,1848,1871]" italics="true" pageId="8" pageNumber="9">et al</emphasis>
., 2001
</bibRefCitation>
(partim), pp.
<date id="A47A9FADFFA9FFB1FF66F89EFED2F8C6" box="[238,361,1880,1903]" pageId="8" pageNumber="9" value="2415" valueMax="2420">24152420</date>
,
<figureCitation id="48FFA5E8FFA9FFB1FEFBF89EFE0AF8C6" box="[371,433,1880,1903]" captionStart="FIGURE 1" captionStartId="3.[151,254,1345,1369]" captionTargetBox="[261,1328,264,1330]" captionTargetId="figure@3.[255,1333,258,1338]" captionTargetPageId="3" captionText="FIGURE 1. Drawing of species contained in the fantastica complex. Clockwise from left: Ranitomeya summersi sp. nov., R. fantastica sensu stricto, R. benedicta sp. nov. Illustration of R. fantastica is an artists rendering of the nominal population from Yurimaguas based on the original type series and recent field work nearby." httpUri="https://zenodo.org/record/183040/files/figure.png" pageId="8" pageNumber="9">Fig. 1</figureCitation>
photo E,
<figureCitation id="48FFA5E8FFA9FFB1FD90F89EFDEEF8C6" box="[536,597,1880,1903]" captionStart="FIGURE 3" captionStartId="5.[152,255,1239,1263]" captionTargetBox="[165,1429,552,1216]" captionTargetId="figure@5.[155,1440,546,1225]" captionTargetPageId="5" captionText="FIGURE 3. Advertisement calls of (A) R. fantastica individual from Cainarachi Valley, recording at 24 C, (B) R. summersi individual from Sauce, recorded at 24.5 C in captivity, (C) R. benedicta individual from Shucushuyacu, recorded at 24 C in captivity." httpUri="https://zenodo.org/record/183042/files/figure.png" pageId="8" pageNumber="9">Fig. 3</figureCitation>
(phylogenetic tree/drawing).
<bibRefCitation id="B455C49CFFA9FFB1FC1AF89EFBDAF8C6" author="Symula" box="[914,1121,1880,1903]" pageId="8" pageNumber="18" refString="Symula, R., Schulte, R. &amp; Summers, K. (2003) Molecular systematics and phylogeography of Amazonian poison frogs of the genus Dendrobates. Molecular Phylogenetics and Evolution, 26, 452 - 75." type="journal article" year="2003">
Symula
<emphasis id="E2B0657FFFA9FFB1FC62F89EFBA3F8C6" box="[1002,1048,1880,1903]" italics="true" pageId="8" pageNumber="9">et al</emphasis>
., 2003
</bibRefCitation>
(partim), table 1 (
<taxonomicName id="17C4C2EEFFA9FFB1FAABF89EFE80F826" ID-CoL="34M8B" authority="Sauce" authorityName="Sauce" class="Amphibia" family="Dendrobatidae" genus="Dendrobates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="fantasticus">
<emphasis id="E2B0657FFFA9FFB1FAABF89EFF50F826" italics="true" pageId="8" pageNumber="9">D. fantasticus</emphasis>
, Sauce
</taxonomicName>
),
<figureCitation id="48FFA5E8FFA9FFB1FEC6F8BEFE30F826" box="[334,395,1912,1935]" captionStart="FIGURE 3" captionStartId="5.[152,255,1239,1263]" captionTargetBox="[165,1429,552,1216]" captionTargetId="figure@5.[155,1440,546,1225]" captionTargetPageId="5" captionText="FIGURE 3. Advertisement calls of (A) R. fantastica individual from Cainarachi Valley, recording at 24 C, (B) R. summersi individual from Sauce, recorded at 24.5 C in captivity, (C) R. benedicta individual from Shucushuyacu, recorded at 24 C in captivity." httpUri="https://zenodo.org/record/183042/files/figure.png" pageId="8" pageNumber="9">Fig. 3</figureCitation>
(phylogenetic tree/drawing;
<taxonomicName id="17C4C2EEFFA9FFB1FD48F8BEFC1DF826" ID-CoL="34M8B" authority="Sauce" authorityName="Sauce" box="[704,934,1912,1935]" class="Amphibia" family="Dendrobatidae" genus="Dendrobates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="fantasticus">
<emphasis id="E2B0657FFFA9FFB1FD48F8BEFCEEF826" box="[704,853,1912,1935]" italics="true" pageId="8" pageNumber="9">D. fantasticus</emphasis>
, Sauce
</taxonomicName>
).
</paragraph>
<paragraph id="D07BB96DFFA9FFB1FF1FF85EFB29F806" blockId="8.[151,1436,1784,1967]" box="[151,1170,1944,1967]" pageId="8" pageNumber="9">
<taxonomicName id="17C4C2EEFFA9FFB1FF1FF85EFE3DF806" ID-CoL="79536" box="[151,390,1944,1967]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA9FFB1FF1FF85EFE3DF806" box="[151,390,1944,1967]" italics="true" pageId="8" pageNumber="9">Ranitomeya fantastica</emphasis>
</taxonomicName>
(non Boulenger):
<bibRefCitation id="B455C49CFFA9FFB1FDC2F85EFCAFF806" author="Lotters" box="[586,788,1944,1967]" pageId="8" pageNumber="17" refString="Lotters, S., Jungfer, K. H., Schmidt, W. &amp; Henkel, F. W. (2007) Poison Frogs: Biology, Species and Captive Husbandry. Edition Chimaira, Frankfurt am Main, 667 pg." type="book chapter" year="2007">
Lötters
<emphasis id="E2B0657FFFA9FFB1FD14F85EFD62F806" box="[668,729,1944,1967]" italics="true" pageId="8" pageNumber="9">et al.,</emphasis>
2007
</bibRefCitation>
(partim), Fig. 592 (photo), pp. 473.
</paragraph>
</subSubSection>
<subSubSection id="98DEEAE6FFA9FFB2FF1FF81CFA22FF73" lastPageId="11" lastPageNumber="12" pageId="8" pageNumber="9" type="materials_examined">
<paragraph id="D07BB96DFFA9FFB0FF1FF81CFEBEFEAB" blockId="8.[151,1436,2010,2036]" lastBlockId="9.[151,1436,152,258]" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9">
<typeStatus id="0F7F07CFFFA9FFB1FF1FF81CFEB2F85D" box="[151,265,2010,2036]" pageId="8" pageNumber="9" type="holotype">
<emphasis id="E2B0657FFFA9FFB1FF1FF81CFEB2F85D" bold="true" box="[151,265,2010,2036]" pageId="8" pageNumber="9">Holotype</emphasis>
</typeStatus>
<collectionCode id="B6D521A8FFA9FFB1FE93F81CFECEF85D" box="[283,373,2010,2036]" pageId="8" pageNumber="9">MUSM</collectionCode>
26994 (field number JLB07-794), an adult male collected by J. Brown and E. Twomey near the town of Sauce, San Martin,
<collectingCountry id="A8D3F9FDFFA8FFB0FDCBFF5EFDC6FF1B" box="[579,637,152,178]" name="Peru" pageId="9" pageNumber="10">Peru</collectingCountry>
;
<geoCoordinate id="B5F0DFAAFFA8FFB0FD00FF5EFD58FF1B" box="[648,739,152,178]" direction="south" orientation="latitude" pageId="9" pageNumber="10" precision="925" value="-6.7166667">6°43 S</geoCoordinate>
,
<geoCoordinate id="B5F0DFAAFFA8FFB0FD78FF5EFCDDFF1B" box="[752,870,152,178]" direction="west" orientation="longitude" pageId="9" pageNumber="10" precision="925" value="-76.25">76°15 W</geoCoordinate>
;
<quantity id="173C1488FFA8FFB0FCFAFF5EFC07FF1B" box="[882,956,152,178]" metricMagnitude="2" metricUnit="m" metricValue="6.84" pageId="9" pageNumber="10" unit="m" value="684.0">684 m</quantity>
elevation;
<date id="A47A9FADFFA8FFB0FBB6FF5EFB70FF1B" box="[1086,1227,152,178]" pageId="9" pageNumber="10" value="2007-07-08">8 July 2007</date>
.
<emphasis id="E2B0657FFFA8FFB0FF4EFF79FEC0FF70" bold="true" box="[198,379,191,217]" pageId="9" pageNumber="10">Paratopotypes</emphasis>
<collectionCode id="B6D521A8FFA8FFB0FE05FF06FE5CFF73" box="[397,487,192,218]" pageId="9" pageNumber="10">MUSM</collectionCode>
26991-26993 (field numbers JLB07-788, 790, and 792), collected along with
<typeStatus id="0F7F07CFFFA8FFB0FF1FFF2EFEB9FEAB" box="[151,258,232,258]" pageId="9" pageNumber="10" type="holotype">holotype</typeStatus>
.
</paragraph>
<caption id="84BBE9E5FFA8FFB0FF1FF88EFDD8F843" httpUri="https://zenodo.org/record/183044/files/figure.png" pageId="9" pageNumber="10" targetBox="[261,1339,325,1846]" targetPageId="9">
<paragraph id="D07BB96DFFA8FFB0FF1FF88EFDD8F843" blockId="9.[151,1437,1864,2026]" pageId="9" pageNumber="10">
<emphasis id="E2B0657FFFA8FFB0FF1FF88EFEA5F8C9" bold="true" box="[151,286,1864,1888]" pageId="9" pageNumber="10">FIGURE 5.</emphasis>
Map of known localities and estimated distributions of
<taxonomicName id="17C4C2EEFFA8FFB0FC17F88EFB90F8F6" box="[927,1067,1864,1887]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFA8FFB0FC17F88EFB90F8F6" box="[927,1067,1864,1887]" italics="true" pageId="9" pageNumber="10">R. fantastica</emphasis>
</taxonomicName>
,
<taxonomicName id="17C4C2EEFFA8FFB0FBB5F88EFB7DF8F6" box="[1085,1222,1864,1887]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFA8FFB0FBB5F88EFB7DF8F6" box="[1085,1222,1864,1887]" italics="true" pageId="9" pageNumber="10">R. summersi</emphasis>
</taxonomicName>
, and
<taxonomicName id="17C4C2EEFFA8FFB0FA83F88EFA2EF8F6" box="[1291,1429,1864,1887]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFA8FFB0FA83F88EFA2EF8F6" box="[1291,1429,1864,1887]" italics="true" pageId="9" pageNumber="10">R. benedicta</emphasis>
</taxonomicName>
. Because some populations where close together, the 14 sampling localities were simplified into 10 geographic groups. Localities which were included in the phylogenetic analysis are indicated by numbers on the map: (1) Tarapoto; (2) Cainarachi Valley; (3) Convento; (4) Pongo de Cainarachi; (5) Varadero; (6) Lower Huallaga Canyon; (7) Shucushuyacu; (8) Pampa Hermosa; (9) Sauce; (10) Chipaota.
</paragraph>
</caption>
<caption id="84BBE9E5FFABFFB3FF1FF860FE89F84D" httpUri="https://zenodo.org/record/183045/files/figure.png" pageId="10" pageNumber="11" targetBox="[353,1250,213,1929]" targetPageId="10">
<paragraph id="D07BB96DFFABFFB3FF1FF860FE89F84D" blockId="10.[151,1435,1958,2020]" pageId="10" pageNumber="11">
<emphasis id="E2B0657FFFABFFB3FF1FF860FEA2F817" bold="true" box="[151,281,1958,1982]" pageId="10" pageNumber="11">FIGURE 6.</emphasis>
Type series of
<emphasis id="E2B0657FFFABFFB3FE34F861FDF3F817" box="[444,584,1959,1982]" italics="true" pageId="10" pageNumber="11">
<taxonomicName id="17C4C2EEFFABFFB3FE34F861FDFFF817" box="[444,580,1959,1982]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="summersi">R. summersi</taxonomicName>
.
</emphasis>
(Top L-R: MUSM 26949, MUSM 26967, MUSM 26991, Bottom L-R: MUSM 2699226994)
</paragraph>
</caption>
<paragraph id="D07BB96DFFAAFFB2FF4EFF51FA22FF73" blockId="11.[151,1437,151,2018]" pageId="11" pageNumber="12">
<typeStatus id="0F7F07CFFFAAFFB2FF4EFF51FEF8FF18" box="[198,323,151,177]" pageId="11" pageNumber="12" type="paratype">
<emphasis id="E2B0657FFFAAFFB2FF4EFF51FEF8FF18" bold="true" box="[198,323,151,177]" pageId="11" pageNumber="12">Paratypes</emphasis>
</typeStatus>
<collectionCode id="B6D521A8FFAAFFB2FED0FF5EFE09FF1B" box="[344,434,152,178]" pageId="11" pageNumber="12">MUSM</collectionCode>
26949, 26967(field numbers JLB07-540 and 628), collected by J. Brown and E. Twomey from near Chazuta, Rio Tunumtunumba drainage, San Martin,
<collectingCountry id="A8D3F9FDFFAAFFB2FC6DFF06FBA4FF73" box="[997,1055,192,218]" name="Peru" pageId="11" pageNumber="12">Peru</collectingCountry>
;
<quantity id="173C1488FFAAFFB2FBA2FF06FBCEFF73" box="[1066,1141,192,218]" metricMagnitude="2" metricUnit="m" metricValue="2.98" pageId="11" pageNumber="12" unit="m" value="298.0">298 m</quantity>
elevation;
<date id="A47A9FADFFAAFFB2FB7EFF06FA2EFF73" box="[1270,1429,192,218]" pageId="11" pageNumber="12" value="2007-06-11">11 June 2007</date>
.
</paragraph>
</subSubSection>
<subSubSection id="98DEEAE6FFAAFFB2FF4EFF21FE09FD13" pageId="11" pageNumber="12" type="etymology">
<paragraph id="D07BB96DFFAAFFB2FF4EFF21FE13FED3" blockId="11.[151,1437,151,2018]" pageId="11" pageNumber="12">
<emphasis id="E2B0657FFFAAFFB2FF4EFF21FEE8FEA8" bold="true" box="[198,339,231,257]" pageId="11" pageNumber="12">Etymology.</emphasis>
The specific epithet is a patronym for Dr. Kyle Summers, an evolutionary biologist who has studied the evolution, behavior, and ecology of poison frogs for over 25 years, contributing greatly to the knowledge of poison frogs. His work on this species demonstrated conclusively the first case of Müllerian mimicry in vertebrates.
</paragraph>
<paragraph id="D07BB96DFFAAFFB2FF4DFE41FE09FD13" blockId="11.[151,1437,151,2018]" pageId="11" pageNumber="12">
<emphasis id="E2B0657FFFAAFFB2FF4DFE41FDB9FE08" bold="true" box="[197,514,391,417]" pageId="11" pageNumber="12">Definition and diagnosis.</emphasis>
Assigned to genus
<taxonomicName id="17C4C2EEFFAAFFB2FD7FFE4EFC38FE08" box="[759,899,392,417]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">
<emphasis id="E2B0657FFFAAFFB2FD7FFE4EFC38FE08" box="[759,899,392,417]" italics="true" pageId="11" pageNumber="12">Ranitomeya</emphasis>
</taxonomicName>
by the following characteristics: first finger shorter than second, premaxillary and maxillary teeth absent, webbing between toes absent, pale limb reticulation present. A relatively large species of
<taxonomicName id="17C4C2EEFFAAFFB2FD1DFE1EFC9AFE58" box="[661,801,472,497]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">
<emphasis id="E2B0657FFFAAFFB2FD1DFE1EFC9AFE58" box="[661,801,472,497]" italics="true" pageId="11" pageNumber="12">Ranitomeya</emphasis>
</taxonomicName>
attaining a maximum adult SVL of
<quantity id="173C1488FFAAFFB2FB57FE1EFAF4FE5B" box="[1247,1359,472,498]" metricMagnitude="-2" metricUnit="m" metricValue="2.04" pageId="11" pageNumber="12" unit="mm" value="20.4">20.4 mm</quantity>
. Body black with one orange cross-band around the sacral region; head black with an orange band outlining the perimeter of the head; face orange with black mask over eyes; limbs primarily black with a longitudinal orange stripe on the dorsal surface. Ventral ground color black with coarse orange marbling, underside of head orange with large black spots. Teeth absent; first finger shorter than second; disc of third finger 22.7 times wider than finger width.
</paragraph>
</subSubSection>
<subSubSection id="98DEEAE6FFAAFFB4FF4DFD0EFEBDFACB" lastPageId="13" lastPageNumber="14" pageId="11" pageNumber="12" type="description">
<paragraph id="D07BB96DFFAAFFB2FF4DFD0EFD2FFB33" blockId="11.[151,1437,151,2018]" pageId="11" pageNumber="12">
<taxonomicName id="17C4C2EEFFAAFFB2FF4DFD0EFE71FD48" box="[197,458,712,737]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFAAFFB2FF4DFD0EFE71FD48" box="[197,458,712,737]" italics="true" pageId="11" pageNumber="12">Ranitomeya summersi</emphasis>
</taxonomicName>
can be distinguished from other species of
<taxonomicName id="17C4C2EEFFAAFFB2FC41FD0EFBEEFD48" box="[969,1109,712,737]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">
<emphasis id="E2B0657FFFAAFFB2FC41FD0EFBEEFD48" box="[969,1109,712,737]" italics="true" pageId="11" pageNumber="12">Ranitomeya</emphasis>
</taxonomicName>
by the combination of relatively large adult SVL, distinctly bicolored pattern consisting of continuous bands (as opposed to spots or dashes), and a buzz-call. This species is most similar in external morphology to
<taxonomicName id="17C4C2EEFFAAFFB2FBCCFCDEFB63FC98" box="[1092,1240,792,817]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFAAFFB2FBCCFCDEFB63FC98" box="[1092,1240,792,817]" italics="true" pageId="11" pageNumber="12">R. fantastica</emphasis>
</taxonomicName>
and some populations of
<taxonomicName id="17C4C2EEFFAAFFB2FE84FC86FE31FCF0" box="[268,394,832,857]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="imitator">
<emphasis id="E2B0657FFFAAFFB2FE84FC86FE31FCF0" box="[268,394,832,857]" italics="true" pageId="11" pageNumber="12">R. imitator</emphasis>
</taxonomicName>
.
<taxonomicName id="17C4C2EEFFAAFFB2FE10FC86FD27FCF0" box="[408,668,832,857]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFAAFFB2FE10FC86FD27FCF0" box="[408,668,832,857]" italics="true" pageId="11" pageNumber="12">Ranitomeya summersi</emphasis>
</taxonomicName>
can be distinguished from
<taxonomicName id="17C4C2EEFFAAFFB2FC51FC86FBD6FCF0" box="[985,1133,832,857]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFAAFFB2FC51FC86FBD6FCF0" box="[985,1133,832,857]" italics="true" pageId="11" pageNumber="12">R. fantastica</emphasis>
</taxonomicName>
using the following characteristics: shorter hind limbs, with a significantly smaller tibia/SVL ratio (mean tibia length/SVL =
<quantity id="173C1488FFAAFFB2FAB3FCAEFA27FC2B" box="[1339,1436,872,898]" metricMagnitude="-2" metricUnit="m" metricValue="1.16332" pageId="11" pageNumber="12" unit="in" value="0.458">0.458 in</quantity>
<taxonomicName id="17C4C2EEFFAAFFB2FF1FFC56FE93FC00" box="[151,296,912,937]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFAAFFB2FF1FFC56FE93FC00" box="[151,296,912,937]" italics="true" pageId="11" pageNumber="12">R. summersi</emphasis>
</taxonomicName>
,
<quantity id="173C1488FFAAFFB2FEBFFC56FE2CFC03" box="[311,407,912,938]" metricMagnitude="-2" metricUnit="m" metricValue="1.23444" pageId="11" pageNumber="12" unit="in" value="0.486">0.486 in</quantity>
<taxonomicName id="17C4C2EEFFAAFFB2FE17FC56FD88FC00" box="[415,563,912,937]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFAAFFB2FE17FC56FD88FC00" box="[415,563,912,937]" italics="true" pageId="11" pageNumber="12">R. fantastica</emphasis>
</taxonomicName>
,
<emphasis id="E2B0657FFFAAFFB2FDCAFC56FDEFFC00" box="[578,596,912,937]" italics="true" pageId="11" pageNumber="12">P</emphasis>
= 0.044;
<figureCitation id="48FFA5E8FFAAFFB2FD4EFC56FCB6FC03" box="[710,781,912,938]" captionStart="FIGURE 2" captionStartId="4.[151,254,1295,1319]" captionTargetBox="[159,1425,822,1283]" captionTargetId="figure@4.[151,1436,814,1288]" captionTargetPageId="4" captionText="FIGURE 2. Left: Comparisons of tibia lengths for the two ± 2 standard errors. nificantly larger standardized tibia length (Tibia / SVL) than R. benedicta and R. summersi (One-way ANOVA, SS = 0.009, df = 2, F = 5.654. Between species, P = 0.008. Error: SS = 0.022, df = 29) Right: Drawing of tadpole mouthparts of R. summersi. Mouthparts of R. benedicta and R. fantastica sensu stricto are identical." httpUri="https://zenodo.org/record/183041/files/figure.png" pageId="11" pageNumber="12">Fig. 2</figureCitation>
); the call of
<taxonomicName id="17C4C2EEFFAAFFB2FC29FC56FB89FC00" box="[929,1074,912,937]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFAAFFB2FC29FC56FB89FC00" box="[929,1074,912,937]" italics="true" pageId="11" pageNumber="12">R. summersi</emphasis>
</taxonomicName>
has longer pulses than the call of
<taxonomicName id="17C4C2EEFFAAFFB2FF30FC7EFEF5FC78" box="[184,334,952,977]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFAAFFB2FF30FC7EFEF5FC78" box="[184,334,952,977]" italics="true" pageId="11" pageNumber="12">R. fantastica</emphasis>
</taxonomicName>
(440 ± 30 ms vs. 290 ± 30 ms in
<taxonomicName id="17C4C2EEFFAAFFB2FD6FFC7EFCC7FC78" box="[743,892,952,977]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFAAFFB2FD6FFC7EFCC7FC78" box="[743,892,952,977]" italics="true" pageId="11" pageNumber="12">R. fantastica</emphasis>
</taxonomicName>
,
<figureCitation id="48FFA5E8FFAAFFB2FC05FC7EFC6FFC7B" box="[909,980,952,978]" captionStart="FIGURE 3" captionStartId="5.[152,255,1239,1263]" captionTargetBox="[165,1429,552,1216]" captionTargetId="figure@5.[155,1440,546,1225]" captionTargetPageId="5" captionText="FIGURE 3. Advertisement calls of (A) R. fantastica individual from Cainarachi Valley, recording at 24 C, (B) R. summersi individual from Sauce, recorded at 24.5 C in captivity, (C) R. benedicta individual from Shucushuyacu, recorded at 24 C in captivity." httpUri="https://zenodo.org/record/183042/files/figure.png" pageId="11" pageNumber="12">Fig. 3</figureCitation>
); and the ratio of note length to pause length is 2.8:1 (vs 1.4:
<quantity id="173C1488FFAAFFB2FE38FC26FE66FC53" box="[432,477,992,1018]" metricMagnitude="-2" metricUnit="m" metricValue="2.54" pageId="11" pageNumber="12" unit="in" value="1.0">1 in</quantity>
<taxonomicName id="17C4C2EEFFAAFFB2FE60FC26FD3BFC50" box="[488,640,992,1017]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFAAFFB2FE60FC26FD3BFC50" box="[488,640,992,1017]" italics="true" pageId="11" pageNumber="12">R. fantastica</emphasis>
</taxonomicName>
). All known
<taxonomicName id="17C4C2EEFFAAFFB2FCA0FC26FC07FC50" box="[808,956,992,1017]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFAAFFB2FCA0FC26FC07FC50" box="[808,956,992,1017]" italics="true" pageId="11" pageNumber="12">R. summersi</emphasis>
</taxonomicName>
individuals possess a bicolored pattern (orange and black), whereas
<taxonomicName id="17C4C2EEFFAAFFB2FE63FBCEFDC4FB88" box="[491,639,1032,1057]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFAAFFB2FE63FBCEFDC4FB88" box="[491,639,1032,1057]" italics="true" pageId="11" pageNumber="12">R. fantastica</emphasis>
</taxonomicName>
typically are tricolored (orange, black, and blue or white) and have reticulated leg netting (see
<figureCitation id="48FFA5E8FFAAFFB2FE5DFBF6FDF1FBE3" box="[469,586,1072,1098]" captionStart-0="FIGURE 8" captionStart-1="FIGURE 9" captionStart-2="FIGURE 10" captionStartId-0="20.[151,254,1886,1910]" captionStartId-1="21.[151,254,1909,1933]" captionStartId-2="22.[151,254,1884,1908]" captionTargetBox-0="[172,1422,204,1852]" captionTargetBox-1="[157,1436,201,1877]" captionTargetBox-2="[159,1428,201,1852]" captionTargetId-0="figure@20.[151,1436,197,1860]" captionTargetId-1="figure@21.[151,1436,194,1885]" captionTargetId-2="figure@22.[151,1436,194,1861]" captionTargetPageId-0="20" captionTargetPageId-1="21" captionTargetPageId-2="22" captionText-0="FIGURE 8. Photos of R. fantastica. A H: Individuals from Varadero, Loreto. I Q: Individuals from Cainarachi Valley, San Martin." captionText-1="FIGURE 9. Photos of R. fantastica. A J: Tarapoto, San Martin. K M: Lower Huallaga, San Martin. P S: Pongo de Cainarachi, San Martin." captionText-2="FIGURE 10. Photos of R. summersi (A H). I L: Type series of R. fantastica (1947.2.15.1 4, order not known, photos from Natural History Museum London). Individual L (top): Inset photo in lower-right corner shows close-up of head spotting." httpUri-0="https://zenodo.org/record/183047/files/figure.png" httpUri-1="https://zenodo.org/record/183048/files/figure.png" httpUri-2="https://zenodo.org/record/183049/files/figure.png" pageId="11" pageNumber="12">Fig. 810</figureCitation>
).
<taxonomicName id="17C4C2EEFFAAFFB2FDD6FBF6FCD9FBE0" box="[606,866,1072,1097]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFAAFFB2FDD6FBF6FCD9FBE0" box="[606,866,1072,1097]" italics="true" pageId="11" pageNumber="12">Ranitomeya summersi</emphasis>
</taxonomicName>
can be distinguished from
<taxonomicName id="17C4C2EEFFAAFFB2FB2AFBF6FA8EFBE0" box="[1186,1333,1072,1097]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFAAFFB2FB2AFBF6FA8EFBE0" box="[1186,1333,1072,1097]" italics="true" pageId="11" pageNumber="12">R. benedicta</emphasis>
</taxonomicName>
by adult coloration;
<taxonomicName id="17C4C2EEFFAAFFB2FE97FB9EFE0FFBD8" box="[287,436,1112,1137]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFAAFFB2FE97FB9EFE0FFBD8" box="[287,436,1112,1137]" italics="true" pageId="11" pageNumber="12">R. benedicta</emphasis>
</taxonomicName>
has a red head with blue dorsal and ventral reticulation (compared to the bicolored pattern in
<taxonomicName id="17C4C2EEFFAAFFB2FE85FB46FE25FB30" box="[269,414,1152,1177]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFAAFFB2FE85FB46FE25FB30" box="[269,414,1152,1177]" italics="true" pageId="11" pageNumber="12">R. summersi</emphasis>
</taxonomicName>
as described above).
</paragraph>
<paragraph id="D07BB96DFFAAFFB2FF4DFB6EFB82FA23" blockId="11.[151,1437,151,2018]" pageId="11" pageNumber="12">
<taxonomicName id="17C4C2EEFFAAFFB2FF4DFB6EFE76FB68" box="[197,461,1192,1217]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFAAFFB2FF4DFB6EFE76FB68" box="[197,461,1192,1217]" italics="true" pageId="11" pageNumber="12">Ranitomeya summersi</emphasis>
</taxonomicName>
is considered a Müllerian mimic of some sympatric populations of
<taxonomicName id="17C4C2EEFFAAFFB2FA90FB6EFA20FB68" box="[1304,1435,1192,1217]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="imitator">
<emphasis id="E2B0657FFFAAFFB2FA90FB6EFA20FB68" box="[1304,1435,1192,1217]" italics="true" pageId="11" pageNumber="12">R. imitator</emphasis>
</taxonomicName>
(
<bibRefCitation id="B455C49CFFAAFFB2FF17FB16FE3EFB43" author="Symula" box="[159,389,1232,1258]" pageId="11" pageNumber="18" refString="Symula, R., Schulte, R. &amp; Summers, K. (2001) Molecular phylogenetic evidence for a mimetic radiation in Peruvian poison frogs supports a Mullerian mimicry hypothesis. Proceedings of the Royal Society of London Series B, 268, 2415 - 2421." type="journal article" year="2001">
Symula
<emphasis id="E2B0657FFFAAFFB2FE89FB16FE80FB40" box="[257,315,1232,1257]" italics="true" pageId="11" pageNumber="12">et al.</emphasis>
, 2001
</bibRefCitation>
applying the name
<taxonomicName id="17C4C2EEFFAAFFB2FDE3FB16FC3CFB40" box="[619,903,1232,1257]" class="Amphibia" family="Dendrobatidae" genus="Dendrobates" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="fantasticus">
<emphasis id="E2B0657FFFAAFFB2FDE3FB16FC3CFB40" box="[619,903,1232,1257]" italics="true" pageId="11" pageNumber="12">Dendrobates fantasticus</emphasis>
</taxonomicName>
) and can be distinguished from these populations by its soft buzz call, audible from less than
<quantity id="173C1488FFAAFFB2FD6DFB3EFCAAFABB" box="[741,785,1272,1298]" metricMagnitude="0" metricUnit="m" metricValue="1.0" pageId="11" pageNumber="12" unit="m" value="1.0">1 m</quantity>
(vs. loud trill call, audible from greater than
<quantity id="173C1488FFAAFFB2FABAFB3EFAE5FABB" box="[1330,1374,1272,1298]" metricMagnitude="0" metricUnit="m" metricValue="5.0" pageId="11" pageNumber="12" unit="m" value="5.0">5 m</quantity>
in
<taxonomicName id="17C4C2EEFFAAFFB2FA0BFB3EFF4EFA90" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="imitator">
<emphasis id="E2B0657FFFAAFFB2FA0BFB3EFF4EFA90" italics="true" pageId="11" pageNumber="12">R. imitator</emphasis>
</taxonomicName>
). In our experience, the black eye-mask of
<taxonomicName id="17C4C2EEFFAAFFB2FD7EFAE6FCCDFA90" box="[758,886,1312,1337]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="imitator">
<emphasis id="E2B0657FFFAAFFB2FD7EFAE6FCCDFA90" box="[758,886,1312,1337]" italics="true" pageId="11" pageNumber="12">R. imitator</emphasis>
</taxonomicName>
usually does not cover the tympanum, and the black head spot of
<taxonomicName id="17C4C2EEFFAAFFB2FEFAFA8EFE4BFAC8" box="[370,496,1352,1377]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="imitator">
<emphasis id="E2B0657FFFAAFFB2FEFAFA8EFE4BFAC8" box="[370,496,1352,1377]" italics="true" pageId="11" pageNumber="12">R. imitator</emphasis>
</taxonomicName>
is ovoid, whereas the head spot in
<taxonomicName id="17C4C2EEFFAAFFB2FC0FFA8EFBACFAC8" box="[903,1047,1352,1377]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFAAFFB2FC0FFA8EFBACFAC8" box="[903,1047,1352,1377]" italics="true" pageId="11" pageNumber="12">R. summersi</emphasis>
</taxonomicName>
is pentagonal in shape.
<taxonomicName id="17C4C2EEFFAAFFB2FAA7FA8EFE83FA20" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFAAFFB2FAA7FA8EFE83FA20" italics="true" pageId="11" pageNumber="12">Ranitomeya summersi</emphasis>
</taxonomicName>
usually has paired black gular spots, rarely present in
<taxonomicName id="17C4C2EEFFAAFFB2FC3CFAB6FB89FA20" box="[948,1074,1392,1417]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="imitator">
<emphasis id="E2B0657FFFAAFFB2FC3CFAB6FB89FA20" box="[948,1074,1392,1417]" italics="true" pageId="11" pageNumber="12">R. imitator</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="D07BB96DFFAAFFB2FF4DFA5EFD92F983" blockId="11.[151,1437,151,2018]" pageId="11" pageNumber="12">
This species can be distinguished from its sister taxon,
<emphasis id="E2B0657FFFAAFFB2FCC4FA5EFC53FA18" box="[844,1000,1432,1457]" italics="true" pageId="11" pageNumber="12">
<taxonomicName id="17C4C2EEFFAAFFB2FCC4FA5EFC5FFA18" box="[844,996,1432,1457]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="fantastica">R. fantastica</taxonomicName>
,
</emphasis>
on the basis of 16 unique unambiguous mitochondrial gene nucleotide site substitutions, and can be distinguished from
<taxonomicName id="17C4C2EEFFAAFFB2FBFDFA06FAB3FA70" box="[1141,1288,1472,1497]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFAAFFB2FBFDFA06FAB3FA70" box="[1141,1288,1472,1497]" italics="true" pageId="11" pageNumber="12">R. benedicta</emphasis>
</taxonomicName>
on the basis of 20 unique unambiguous mitochondrial gene site substitutions, and is remarkably distant, genetically, to
<taxonomicName id="17C4C2EEFFAAFFB2FA0BFA2EFF4DF980" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="imitator">
<emphasis id="E2B0657FFFAAFFB2FA0BFA2EFF4DF980" italics="true" pageId="11" pageNumber="12">R. imitator</emphasis>
</taxonomicName>
(see
<bibRefCitation id="B455C49CFFAAFFB2FEBBF9D6FDA7F983" author="Symula" box="[307,540,1552,1578]" pageId="11" pageNumber="18" refString="Symula, R., Schulte, R. &amp; Summers, K. (2001) Molecular phylogenetic evidence for a mimetic radiation in Peruvian poison frogs supports a Mullerian mimicry hypothesis. Proceedings of the Royal Society of London Series B, 268, 2415 - 2421." type="journal article" year="2001">
Symula
<emphasis id="E2B0657FFFAAFFB2FE1CF9D6FE6DF980" box="[404,470,1552,1577]" italics="true" pageId="11" pageNumber="12">et al.,</emphasis>
2001
</bibRefCitation>
).
</paragraph>
<paragraph id="D07BB96DFFAAFFB2FF4DF9F1FBBCF963" blockId="11.[151,1437,151,2018]" pageId="11" pageNumber="12">
<emphasis id="E2B0657FFFAAFFB2FF4DF9F1FD3BF9F8" bold="true" box="[197,640,1591,1617]" pageId="11" pageNumber="12">
Measurements (in mm) of
<typeStatus id="0F7F07CFFFAAFFB2FD85F9F1FDC0F9F8" box="[525,635,1591,1617]" pageId="11" pageNumber="12" type="holotype">holotype</typeStatus>
.
</emphasis>
The undissected
<typeStatus id="0F7F07CFFFAAFFB2FCC3F9FEFC09F9FB" box="[843,946,1592,1618]" pageId="11" pageNumber="12" type="holotype">holotype</typeStatus>
(
<figureCitation id="48FFA5E8FFAAFFB2FC49F9FEFBBDF9FB" box="[961,1030,1592,1618]" captionStart="FIGURE 6" captionStartId="10.[151,254,1958,1982]" captionTargetBox="[353,1250,213,1929]" captionTargetId="figure@10.[334,1268,194,1951]" captionTargetPageId="10" captionText="FIGURE 6. Type series of R. summersi. (Top L-R: MUSM 26949, MUSM 26967, MUSM 26991, Bottom L-R: MUSM 26992 26994)" httpUri="https://zenodo.org/record/183045/files/figure.png" pageId="11" pageNumber="12">Fig. 6</figureCitation>
) is an adult male as determined by the presence of vocal slits. SVL 19.5; FL 8.1; TL 8.5; KK 16.7; FoL 8.0; HaL 5.5; HL 5.3; HW 6.3; BW 7.3; UEW 2.86; IOD 1.93;
<collectingCountry id="A8D3F9FDFFAAFFB2FE2CF94EFE63F90B" box="[420,472,1672,1698]" name="India" pageId="11" pageNumber="12">IND</collectingCountry>
2.29; TD 1.00; ED 2.07; DET 0.50; L1F I 2.07; L2F 3.14; W3D 0.96; W3F 0.36. Measurements of additional specimens are given in the appendix (
<tableCitation id="9D468CD6FFAAFFB2FC16F976FC4DF963" box="[926,1014,1712,1738]" captionStart="TABLE 1" captionStartId="17.[151,239,818,842]" captionTargetBox="[159,1368,874,1785]" captionTargetPageId="17" captionText="TABLE 1. Measurements of the type series of R. benedicta." httpUri="http://table.plazi.org/id/84BBE9E5FFB0FFA8FF1FFCF4FCA3FCE0" pageId="11" pageNumber="12" tableUuid="84BBE9E5FFB0FFA8FF1FFCF4FCA3FCE0">Table 1</tableCitation>
).
</paragraph>
<paragraph id="D07BB96DFFAAFFB5FF4DF911FE49FE63" blockId="11.[151,1437,151,2018]" lastBlockId="12.[151,1437,152,2018]" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12">
<emphasis id="E2B0657FFFAAFFB2FF4DF911FE4CF958" bold="true" box="[197,503,1751,1777]" pageId="11" pageNumber="12">
Description of
<typeStatus id="0F7F07CFFFAAFFB2FE0DF911FE49F958" box="[389,498,1751,1777]" pageId="11" pageNumber="12" type="holotype">holotype</typeStatus>
.
</emphasis>
Widest part of head is at jaw articulations. Head narrower than body. Head width 86 % of body width. Subgular pouch not visibly expanded. Tongue grey, ovoid, attaching anteriorly; teeth absent. Pattern bicolored; black on the head, body, and limbs, highlighted by several golden orange bands. The head is black and orange, the black confined to the top of the head and around the eyes and tympanum. The black eye mask covers the tympanum and does not connect with the black on top of the head. Dorsum jet black with one incomplete orange cross-band near the sacral region; broken medially left of vertebrae. The hind limbs are black and lack reticulation, but possess a solid orange band extending from the rump over the femur, down across the anterior surface of the knee, back across the posterior surface of the tibia, and across the posterior surface of the tarsus to the foot. The dorsal surface of the foot is half orange and half black, toes orange. The front limbs are orange dorsally; this orange is fused with the orange on the face. Dorsal surfaces of the hands are both orange and black, fingers orange, finger tips black. Ventrally, the belly is black with irregular and coarse orange marbling. From a ventral aspect, the hind limbs have irregular orange longitudinal bands. Ventral surfaces of the forelimbs are uniformly black. The underside of the head is orange; two well-defined black gular spots laterally. Black bar present on the anterior surface of the snout between the nares. Nares black; iris black.
</paragraph>
<paragraph id="D07BB96DFFADFFB5FF4DFE1EFB7DFD13" blockId="12.[151,1437,152,2018]" pageId="12" pageNumber="13">In life, skin texture granular on the dorsal surfaces of the body and head; dorsal surfaces of limbs weakly granular. Venter weakly granular on belly; limbs and ventral surface of head nearly smooth. Snout sloping and rounded in lateral profile, rounded in dorsal profile. Nares situated at tip of snout and directed laterally; both nares visible from ventral and anterior view but hidden in dorsal view. Canthus rostralis rounded, loreal region slightly concave and vertical. Upper eyelid approximately equal in width to interorbital distance; internarial distance roughly equal to eye width. Tympanum round, partially concealed posterodorsally.</paragraph>
<paragraph id="D07BB96DFFADFFB5FF4DFD0EFED7FC7B" blockId="12.[151,1437,152,2018]" pageId="12" pageNumber="13">Hands relatively large, length 28 % of SVL. Relative length of appressed fingers III&gt; IV&gt; II&gt; I; first finger 67 % length of second; finger discs moderately expanded, width of disc on finger III 2.6 times width of adjacent phalanx. An unpigmented median metacarpal tubercle is present on base of palm; inner metacarpal tubercle present near base of finger I but angled posteriorly; unpigmented proximal subarticular tubercles present on base of each digit, except on finger I, where tubercle is part-way up the digit; distal subarticular tubercle visible only on fingers III and IV. All tubercles raised above level of hands; scutes present on dorsal surface of fingers.</paragraph>
<paragraph id="D07BB96DFFADFFB5FF4DFC26FCC6FA93" blockId="12.[151,1437,152,2018]" pageId="12" pageNumber="13">
<collectingCountry id="A8D3F9FDFFADFFB5FF4DFC26FEBBFC53" box="[197,256,992,1018]" name="India" pageId="12" pageNumber="13">Hind</collectingCountry>
limbs moderate length, with heel of appressed hind limbs reaching reaching to eye. Femur and tibia roughly equal in length, tibia 105 % length of femur; knee-knee distance 86 % of SVL. Relative lengths of appressed toes IV&gt; III
<emphasis id="E2B0657FFFADFFB5FE39FBF5FE05FBE1" bold="true" box="[433,446,1075,1096]" pageId="12" pageNumber="13"></emphasis>
V&gt; II&gt; I; first toe short with unexpanded disc; second toe with slightly expanded disc, discs on toes IIIV moderately expanded. Two metatarsal tubercles present on base of foot, one situated medially near base of toe I, the other situated laterally at the base of the fifth metatarsal. Proximal subarticular tubercles present at base of each toe but most notable on toes I and II due to their lack of pigmentation. Toes III and V with two pigmented subarticular tubercles, toe IV with three subarticular tubercles. A faint tarsal keel is present starting below the knee and turning into the medial metatarsal tubercle at the foot. Tarsal tubercle absent; feet and hands lacking webbing and lateral fringing.
</paragraph>
<paragraph id="D07BB96DFFADFFB5FF4DFA81FC1AFA73" blockId="12.[151,1437,152,2018]" pageId="12" pageNumber="13">
<emphasis id="E2B0657FFFADFFB5FF4DFA81FE84FAC8" bold="true" box="[197,319,1351,1377]" pageId="12" pageNumber="13">Variation.</emphasis>
Adults range from
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SVL, females and males roughly equal in SVL. Head width 86 % of body width in
<typeStatus id="0F7F07CFFFADFFB5FE2BFAB6FE6EFA23" box="[419,469,1392,1418]" pageId="12" pageNumber="13">type</typeStatus>
series (range 66127 %). Head width 3033 % of SVL in adults. No apparent sexual dimorphism in external morphology except that males tend to be less rotund and possess vocal slits on the floor of the mouth. Subgular pouch not visibly expanded in males.
</paragraph>
<paragraph id="D07BB96DFFADFFB5FF4DFA2EFD2BF8C3" blockId="12.[151,1437,152,2018]" pageId="12" pageNumber="13">
The black eye mask invariably covers the tympanum and does not connect with the black on top of the head, but the head-black and eye-black can fuse with the black on the dorsum. Dorsum jet black with one orange cross-band near the sacral region. The cross-band is usually complete but in some individuals is broken medially. Some individuals have longitudinal extensions of the cross-band in line with the vertebrae, which can extend up the back, or posteriorly connecting with the orange bands of the hind limbs (
<figureCitation id="48FFA5E8FFADFFB5FA9CF94EFAD0F90B" box="[1300,1387,1672,1698]" captionStart="FIGURE 10" captionStartId="22.[151,254,1884,1908]" captionTargetBox="[159,1428,201,1852]" captionTargetId="figure@22.[151,1436,194,1861]" captionTargetPageId="22" captionText="FIGURE 10. Photos of R. summersi (A H). I L: Type series of R. fantastica (1947.2.15.1 4, order not known, photos from Natural History Museum London). Individual L (top): Inset photo in lower-right corner shows close-up of head spotting." httpUri="https://zenodo.org/record/183049/files/figure.png" pageId="12" pageNumber="13">Fig. 10</figureCitation>
, A G). The dorsal surface of the foot is usually half orange and half black; toes can be either orange or black. Dorsal surfaces of the hands are both orange and black, fingers are usually either all orange or all black. The underside of the head is orange; most individuals have two well-defined black gular spots laterally, in some individuals these spots are medially fused. Most individuals in the
<typeStatus id="0F7F07CFFFADFFB5FC2EF8EEFC62F8EB" box="[934,985,1832,1858]" pageId="12" pageNumber="13">type</typeStatus>
series possess a black bar on the anterior surface of the snout between the nares.
</paragraph>
<paragraph id="D07BB96DFFADFFB4FF4DF8BEFE68FF73" blockId="12.[151,1437,152,2018]" lastBlockId="13.[151,1437,152,2018]" lastPageId="13" lastPageNumber="14" pageId="12" pageNumber="13">
Hands relatively large, length 2631 % of SVL. First finger 6676 % length of second; finger discs moderately expanded in both males and females, width of disc on finger
<date id="A47A9FADFFADFFB5FC3BF866FC5EF813" box="[947,997,1952,1978]" pageId="12" pageNumber="13">III 2</date>
2.7 times width of adjacent phalanx. Femur and tibia roughly equal in length, tibia 100107 % length of femur (mean 105 %); knee-knee distance 8693 % of SVL (mean 90 %). In preservative little color change occurs, except that the orange coloration turns to pale gold or white.
</paragraph>
<paragraph id="D07BB96DFFACFFB4FF4EFF21FBB6FE5B" blockId="13.[151,1437,152,2018]" pageId="13" pageNumber="14">
<emphasis id="E2B0657FFFACFFB4FF4EFF21FE91FEA8" bold="true" box="[198,298,231,257]" pageId="13" pageNumber="14">Tadpole</emphasis>
. A stage 29 tadpole was used for the description (
<figureCitation id="48FFA5E8FFACFFB4FCF0FF2EFC7AFEAB" box="[888,961,232,258]" captionStart="FIGURE 2" captionStartId="4.[151,254,1295,1319]" captionTargetBox="[159,1425,822,1283]" captionTargetId="figure@4.[151,1436,814,1288]" captionTargetPageId="4" captionText="FIGURE 2. Left: Comparisons of tibia lengths for the two ± 2 standard errors. nificantly larger standardized tibia length (Tibia / SVL) than R. benedicta and R. summersi (One-way ANOVA, SS = 0.009, df = 2, F = 5.654. Between species, P = 0.008. Error: SS = 0.022, df = 29) Right: Drawing of tadpole mouthparts of R. summersi. Mouthparts of R. benedicta and R. fantastica sensu stricto are identical." httpUri="https://zenodo.org/record/183041/files/figure.png" pageId="13" pageNumber="14">Fig. 2</figureCitation>
). Total length 19.0 mm, body length
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, body width
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. Snout rounded when viewed from above; body ovoid in dorsal view. Eyes black, dorsal, angled laterally, pupils white in preservative. Nares not forming tube, situated half-way between eye and tip of snout, directed dorsolaterally. Spiracle sinistral; vent dextral. Tail
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in depth measured halfway along length; ventral tail fin begins at tail base, dorsal tail fin begins just posterior to plane of vent opening, ventral and dorsal fins relatively uniform in thickness throughout tail, tapering towards tip. Musculature depth uniform throughout, tapering towards tip, musculature depth 2.0 mm.
</paragraph>
<paragraph id="D07BB96DFFACFFB4FF4EFDC6FD30FC7B" blockId="13.[151,1437,152,2018]" pageId="13" pageNumber="14">
The mouth is directed anteroventrally. Oral disc emarginate, anterior and posterior labia forming flaps free from body wall,
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in width. Marginal papillae absent on anterior labium except for lateral-most portion (34 papillae present), present in one complete row on posterior labium. Papillae white, rounded; submarginal papillae absent. Jaw sheaths deep in longitudinal width, serrate, lacking indentations. Lateral processes very short, extending barely past lower jaw. Labial tooth row formula is 2(2)/3. A-1 complete, A-2 with medial gap, same width as A-1. P-1, P-2, and P-3 complete; P-1 and P-2 equal width, P-3 slightly shorter. In preservative, the head appears light brown. Pigmentation on dorsum is mottled brown with a white ground color. Ventral coloration is white with irregular brown flecking which is most dense near the mouth. Tail musculature brown, fins transparent with brown mottling. Life colors grey. One additional stage 27 tadpole was examined and agrees with this description. Additionally, the above description agrees with two
<taxonomicName id="17C4C2EEFFACFFB4FB12FCAEFA8AFC28" box="[1178,1329,872,897]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFACFFB4FB12FCAEFA8AFC28" box="[1178,1329,872,897]" italics="true" pageId="13" pageNumber="14">R. fantastica</emphasis>
</taxonomicName>
tadpoles (see above). Mouthparts are identical between these two species. It appears that tadpole morphology is consistent within the
<taxonomicName id="17C4C2EEFFACFFB4FECFFC7EFE60FC78" box="[327,475,952,977]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFACFFB4FECFFC7EFE60FC78" box="[327,475,952,977]" italics="true" pageId="13" pageNumber="14">R. fantastica</emphasis>
</taxonomicName>
species group.
</paragraph>
<paragraph id="D07BB96DFFACFFB4FF4DFC19FEBDFACB" blockId="13.[151,1437,152,2018]" pageId="13" pageNumber="14">
<emphasis id="E2B0657FFFACFFB4FF4DFC19FEC9FC50" bold="true" box="[197,370,991,1017]" pageId="13" pageNumber="14">Vocalizations.</emphasis>
The advertisement call is a series of soft buzz-like notes, similar to
<taxonomicName id="17C4C2EEFFACFFB4FB16FC26FA88FC50" box="[1182,1331,992,1017]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFACFFB4FB16FC26FA88FC50" box="[1182,1331,992,1017]" italics="true" pageId="13" pageNumber="14">R. fantastica</emphasis>
</taxonomicName>
(
<figureCitation id="48FFA5E8FFACFFB4FACCFC26FA37FC53" box="[1348,1420,992,1018]" captionStart="FIGURE 3" captionStartId="5.[152,255,1239,1263]" captionTargetBox="[165,1429,552,1216]" captionTargetId="figure@5.[155,1440,546,1225]" captionTargetPageId="5" captionText="FIGURE 3. Advertisement calls of (A) R. fantastica individual from Cainarachi Valley, recording at 24 C, (B) R. summersi individual from Sauce, recorded at 24.5 C in captivity, (C) R. benedicta individual from Shucushuyacu, recorded at 24 C in captivity." httpUri="https://zenodo.org/record/183042/files/figure.png" pageId="13" pageNumber="14">Fig. 3</figureCitation>
). The notes in the calls are 440 ± 30 ms (vs. 290 ± 30 ms in
<taxonomicName id="17C4C2EEFFACFFB4FCC8FBCEFC68FB88" box="[832,979,1032,1057]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFACFFB4FCC8FBCEFC68FB88" box="[832,979,1032,1057]" italics="true" pageId="13" pageNumber="14">R. fantastica</emphasis>
</taxonomicName>
) in duration with brief pauses of 150 ± 20 ms between each note (vs. 200 ± 20 ms in
<taxonomicName id="17C4C2EEFFACFFB4FD31FBF6FCF5FBE0" box="[697,846,1072,1097]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFACFFB4FD31FBF6FCF5FBE0" box="[697,846,1072,1097]" italics="true" pageId="13" pageNumber="14">R. fantastica</emphasis>
</taxonomicName>
). The ratio of note length to pause length is 2.8:1 (vs 1.4:
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<taxonomicName id="17C4C2EEFFACFFB4FEAAFB9EFE0DFBD8" box="[290,438,1112,1137]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFACFFB4FEAAFB9EFE0DFBD8" box="[290,438,1112,1137]" italics="true" pageId="13" pageNumber="14">R. fantastica</emphasis>
</taxonomicName>
). The call is audible from
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(vs.
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in
<taxonomicName id="17C4C2EEFFACFFB4FC58FB9EFBD8FBD8" box="[976,1123,1112,1137]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFACFFB4FC58FB9EFBD8FBD8" box="[976,1123,1112,1137]" italics="true" pageId="13" pageNumber="14">R. fantastica</emphasis>
</taxonomicName>
). The dominant frequency of the call is 2890 Hz at an air temperature of 24° C (vs. 3290 Hz in
<taxonomicName id="17C4C2EEFFACFFB4FC26FB46FBFAFB30" box="[942,1089,1152,1177]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="fantastica">
<emphasis id="E2B0657FFFACFFB4FC26FB46FBFAFB30" box="[942,1089,1152,1177]" italics="true" pageId="13" pageNumber="14">R. fantastica</emphasis>
</taxonomicName>
at 24° C). The call of
<taxonomicName id="17C4C2EEFFACFFB4FACBFB46FF63FB68" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFACFFB4FACBFB46FF63FB68" italics="true" pageId="13" pageNumber="14">R. summersi</emphasis>
</taxonomicName>
can be distinguished from
<taxonomicName id="17C4C2EEFFACFFB4FDA8FB6EFD0EFB68" box="[544,693,1192,1217]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFACFFB4FDA8FB6EFD0EFB68" box="[544,693,1192,1217]" italics="true" pageId="13" pageNumber="14">R. benedicta</emphasis>
</taxonomicName>
on the basis of the following parameters: audible from &lt;
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(vs. up to
<quantity id="173C1488FFACFFB4FE9CFB16FEF8FB43" box="[276,323,1232,1258]" metricMagnitude="0" metricUnit="m" metricValue="8.0" pageId="13" pageNumber="14" unit="m" value="8.0">8 m</quantity>
in
<taxonomicName id="17C4C2EEFFACFFB4FEE6FB16FDB8FB40" box="[366,515,1232,1257]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFACFFB4FEE6FB16FDB8FB40" box="[366,515,1232,1257]" italics="true" pageId="13" pageNumber="14">R. benedicta</emphasis>
</taxonomicName>
); note length&gt; 400 ms (vs. 150 ± 20 ms in
<taxonomicName id="17C4C2EEFFACFFB4FBABFB16FB02FB40" box="[1059,1209,1232,1257]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFACFFB4FBABFB16FB02FB40" box="[1059,1209,1232,1257]" italics="true" pageId="13" pageNumber="14">R. benedicta</emphasis>
</taxonomicName>
), and ratio of note length to pause length is&gt; 2:1 (vs. 1:
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<taxonomicName id="17C4C2EEFFACFFB4FDF5FB3EFCABFAB8" box="[637,784,1272,1297]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="benedicta">
<emphasis id="E2B0657FFFACFFB4FDF5FB3EFCABFAB8" box="[637,784,1272,1297]" italics="true" pageId="13" pageNumber="14">R. benedicta</emphasis>
</taxonomicName>
). This call can be distinguished from
<taxonomicName id="17C4C2EEFFACFFB4FB44FB3EFAF1FAB8" box="[1228,1354,1272,1297]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="imitator">
<emphasis id="E2B0657FFFACFFB4FB44FB3EFAF1FAB8" box="[1228,1354,1272,1297]" italics="true" pageId="13" pageNumber="14">R. imitator</emphasis>
</taxonomicName>
in that
<taxonomicName id="17C4C2EEFFACFFB4FF1FFAE6FE9CFA90" box="[151,295,1312,1337]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFACFFB4FF1FFAE6FE9CFA90" box="[151,295,1312,1337]" italics="true" pageId="13" pageNumber="14">R. summersi</emphasis>
</taxonomicName>
has soft buzz-like notes audible from &lt;
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(vs. loud, trill-like call easily audible from&gt;
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in
<taxonomicName id="17C4C2EEFFACFFB4FA0BFAE6FF4EFAC8" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="imitator">
<emphasis id="E2B0657FFFACFFB4FA0BFAE6FF4EFAC8" italics="true" pageId="13" pageNumber="14">R. imitator</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="98DEEAE6FFACFFB7FF4DFAA9FE4EFDED" lastPageId="14" lastPageNumber="15" pageId="13" pageNumber="14" type="distribution">
<paragraph id="D07BB96DFFACFFB4FF4DFAA9FA31F9AB" blockId="13.[151,1437,152,2018]" pageId="13" pageNumber="14">
<emphasis id="E2B0657FFFACFFB4FF4DFAA9FDE5FA20" bold="true" box="[197,606,1391,1417]" pageId="13" pageNumber="14">Distribution and natural history.</emphasis>
<taxonomicName id="17C4C2EEFFACFFB4FDEEFAB6FCD1FA20" box="[614,874,1392,1417]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFACFFB4FDEEFAB6FCD1FA20" box="[614,874,1392,1417]" italics="true" pageId="13" pageNumber="14">Ranitomeya summersi</emphasis>
</taxonomicName>
occurs throughout the central Huallaga Canyon (
<figureCitation id="48FFA5E8FFACFFB4FF17FA5EFF5CFA1B" box="[159,231,1432,1458]" captionStart="FIGURE 5" captionStartId="9.[151,254,1864,1888]" captionTargetBox="[261,1339,325,1846]" captionTargetId="figure@9.[253,1346,314,1850]" captionTargetPageId="9" captionText="FIGURE 5. Map of known localities and estimated distributions of R. fantastica, R. summersi, and R. benedicta. Because some populations where close together, the 14 sampling localities were simplified into 10 geographic groups. Localities which were included in the phylogenetic analysis are indicated by numbers on the map: (1) Tarapoto; (2) Cainarachi Valley; (3) Convento; (4) Pongo de Cainarachi; (5) Varadero; (6) Lower Huallaga Canyon; (7) Shucushuyacu; (8) Pampa Hermosa; (9) Sauce; (10) Chipaota." httpUri="https://zenodo.org/record/183044/files/figure.png" pageId="13" pageNumber="14">Fig. 5</figureCitation>
), extending into the southernmost tip of the Cordillera Escalera near Chazuta and to the northwestern edge of the Cordillera Azul. This species occurs on both sides of the Rio Huallaga, extending from Curiyacu westward to Sauce, where they persist in humid recesses of the rocky stream valleys of this semiarid region.
</paragraph>
<paragraph id="D07BB96DFFACFFB4FF4DF9D6FA91F84B" blockId="13.[151,1437,152,2018]" pageId="13" pageNumber="14">
This species is primarily terrestrial and is a denizen of relatively dry secondary and primary premontane forests. They are extremely alert and difficult to detect amidst the leaf litter and rocky terrain they inhabit. The forests they inhabit are characterized by a relatively open canopy and poor, rocky soils, which allows for abundant growth of
<taxonomicName id="17C4C2EEFFACFFB4FE0EF94EFD97F908" box="[390,556,1672,1697]" class="Liliopsida" family="Araceae" genus="Dieffenbachia" kingdom="Plantae" order="Alismatales" pageId="13" pageNumber="14" phylum="Tracheophyta" rank="genus">
<emphasis id="E2B0657FFFACFFB4FE0EF94EFD97F908" box="[390,556,1672,1697]" italics="true" pageId="13" pageNumber="14">Dieffenbachia</emphasis>
</taxonomicName>
in the understory. The relative dryness of the region is conducive to a terrestrial lifestyle, keeping them in contact with the more humid forest floor.
<taxonomicName id="17C4C2EEFFACFFB4FB81F976FAB7F960" box="[1033,1292,1712,1737]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFACFFB4FB81F976FAB7F960" box="[1033,1292,1712,1737]" italics="true" pageId="13" pageNumber="14">Ranitomeya summersi</emphasis>
</taxonomicName>
is primarily active during the early morning and late afternoon. In midday they are infrequently encountered on the ground, but can be found taking refuge in the humid axils of
<emphasis id="E2B0657FFFACFFB4FCFDF8C6FB98F8B0" box="[885,1059,1792,1817]" italics="true" pageId="13" pageNumber="14">
<taxonomicName id="17C4C2EEFFACFFB4FCFDF8C6FBA4F8B0" box="[885,1055,1792,1817]" class="Liliopsida" family="Araceae" genus="Dieffenbachia" kingdom="Plantae" order="Alismatales" pageId="13" pageNumber="14" phylum="Tracheophyta" rank="genus">Dieffenbachia</taxonomicName>
.
</emphasis>
Breeding occurs terrestrially in humid leaf litter; clutch sizes range from 4
<specimenCount id="C6C272E4FFACFFB4FD38F8EEFD05F8EB" box="[688,702,1832,1858]" pageId="13" pageNumber="14" type="egg">9</specimenCount>
eggs. Healthy females in captivity are capable of producing clutches every 3 or 4 days when cycled properly. Given the regions aridity, phytotelmata only retain water for a short period each season (MP, pers. obsv.). This truncated breeding season has also been observed in sympatric populations of
<taxonomicName id="17C4C2EEFFACFFB4FEF3F866FE47F810" box="[379,508,1952,1977]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="imitator">
<emphasis id="E2B0657FFFACFFB4FEF3F866FE47F810" box="[379,508,1952,1977]" italics="true" pageId="13" pageNumber="14">R. imitator</emphasis>
</taxonomicName>
(JLB, unpub. data). Tree holes,
<emphasis id="E2B0657FFFACFFB4FC0CF866FB8AF810" box="[900,1073,1952,1977]" italics="true" pageId="13" pageNumber="14">
<taxonomicName id="17C4C2EEFFACFFB4FC0CF866FB96F810" box="[900,1069,1952,1977]" class="Liliopsida" family="Araceae" genus="Dieffenbachia" kingdom="Plantae" order="Alismatales" pageId="13" pageNumber="14" phylum="Tracheophyta" rank="genus">Dieffenbachia</taxonomicName>
,
</emphasis>
and
<taxonomicName id="17C4C2EEFFACFFB4FBE6F866FABAF810" box="[1134,1281,1952,1977]" class="Liliopsida" family="Araceae" genus="Xanthosoma" kingdom="Animalia" order="Alismatales" pageId="13" pageNumber="14" phylum="Tracheophyta" rank="genus">
<emphasis id="E2B0657FFFACFFB4FBE6F866FABAF810" box="[1134,1281,1952,1977]" italics="true" pageId="13" pageNumber="14">Xanthosoma</emphasis>
</taxonomicName>
axils are the preferred pools for tadpole deposition. In captivity metamorphosis occurs between 90 and 110 days.
</paragraph>
<paragraph id="D07BB96DFFAFFFB7FF4DFF51FE4EFDED" blockId="14.[151,1437,151,580]" pageId="14" pageNumber="15">
<emphasis id="E2B0657FFFAFFFB7FF4DFF51FE06FF18" bold="true" box="[197,445,151,177]" pageId="14" pageNumber="15">Conservation status</emphasis>
. Under the IUCN Red List Criteria (IUCN, 2001), we suggest
<taxonomicName id="17C4C2EEFFAFFFB7FB29FF5EFA89FF18" box="[1185,1330,152,177]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFAFFFB7FB29FF5EFA89FF18" box="[1185,1330,152,177]" italics="true" pageId="14" pageNumber="15">R. summersi</emphasis>
</taxonomicName>
be listed as Endangered (EN) under the following criteria: (1) we estimate the extent of occurrence area of
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2 or less, most of which is unsuitable habitat, meaning the area of occupancy may be much smaller, (2) deforestation in this area has degraded its habitat, particularly in the Chipaotillo and Tunumtunumba drainages near Chazuta, and will further reduce the amount of suitable habitat for this species in the future, (3) illegal smuggling for the pet trade appears to have caused localized population declines in certain areas. Over the past 30 years, Chazuta has been a staging point for many smuggling operations and populations in close proximity to Chazuta have dramatically declined. However, in the few areas overlooked by illegal collectors, populations appear to flourishing. The constant encroachment by farmers looking to expand their farms continues to put this frog in increased peril. Unlike
<emphasis id="E2B0657FFFAFFFB7FDB3FDC5FD7BFDB5" box="[571,704,515,540]" italics="true" pageId="14" pageNumber="15">
<taxonomicName id="17C4C2EEFFAFFFB7FDB3FDC5FD07FDB5" box="[571,700,515,540]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="imitator">R. imitator</taxonomicName>
,
</emphasis>
which will adapt to use agricultural landscapes,
<taxonomicName id="17C4C2EEFFAFFFB7FA81FDC5FA27FDB5" box="[1289,1436,515,540]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="summersi">
<emphasis id="E2B0657FFFAFFFB7FA81FDC5FA27FDB5" box="[1289,1436,515,540]" italics="true" pageId="14" pageNumber="15">R. summersi</emphasis>
</taxonomicName>
does not appear to adapt well.
</paragraph>
</subSubSection>
</treatment>
</document>