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<mods:title id="E07D5A666EA62AE742CBA899889172B0">A review of Northern Peruvian glassfrogs (Centrolenidae), with the description of four new remarkable species</mods:title>
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<taxonomicName id="29E14D24FFF6FF90FF7CFF71D2ECFF5A" ID-CoL="6X9DP" box="[151,410,152,178]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="mcdiarmidi">
<emphasis id="DC95EAB5FFF6FF90FF7CFF71D2ECFF5A" bold="true" box="[151,410,152,178]" italics="true" pageId="47" pageNumber="48">Rulyrana mcdiarmidi</emphasis>
</taxonomicName>
(
<bibRefCitation id="8A704B56FFF6FF90FE41FF71D76CFF5A" author="Cisneros-Heredia" box="[426,1050,152,178]" pageId="47" pageNumber="48" refString="Cisneros-Heredia, D. F., Venegas, P. J., Rada, M. &amp; Schulte, R. (2008) A new species of glassfrog (Anura: Centrolenidae) from the foothill Andean forests of Ecuador and Peru. Herpetologica, 64, 341 - 353. http: // dx. doi. org / 10.1655 / 06 - 078.1" type="journal article" year="2008">Cisneros-Heredia, Venegas, Rada, &amp; Schulte 2008</bibRefCitation>
)
</emphasis>
</heading>
</paragraph>
<paragraph id="EE5E36A7FFF6FF90FF7CFF53D275FF3B" blockId="47.[151,1058,152,211]" box="[151,259,186,211]" pageId="47" pageNumber="48">
<figureCitation id="76DA2A22FFF6FF90FF7CFF53D275FF3B" box="[151,259,186,211]" captionStart="FIGURE 27. a, b" captionStartId="46.[151,250,1681,1703]" captionTargetBox="[151,1434,375,1658]" captionTargetId="figure@46.[151,1436,375,1659]" captionTargetPageId="46" captionText="FIGURE 27. a, b) Nymphargus chancas, dorsal and ventral views of CORBIDI 10471, from 5.9 km SE from the type locality of Abra Tangarana, San Martín, 6 ° 19 ' 15.65 &quot; S 76 ° 41 ' 44.16 &quot; W, 973 m; c, d) Teratohyla amelie, dorsal and ventral views of MNCN 45958, from Nuevo Jaen, San Martín 7 ° 39 ' 16.20 &quot; S, 76 ° 39 ' 24.60 &quot; W, 591 m; e) Rulyrana mcdiarmidi, uncollected male from 0.45 km S from the town of Cocachimba, Amazonas, 6 ° 3 ' 44.28 &quot; S, 77 ° 53 ' 36.47 &quot; W, 1811 m; f) Espadarana cf. fernandoi, dorsal view of adult male (CORBIDI 10468) from Quebrada Goca on the Yambrasbamba road, about 1 km N from road entrance, Amazonas, 5 ° 45 ' 50.78 &quot; S, 77 ° 54 ' 46.52 &quot; W, 1711 m." httpUri="https://zenodo.org/record/286951/files/figure.png" pageId="47" pageNumber="48">Figure 27</figureCitation>
</paragraph>
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<subSubSection id="A6FB652CFFF6FF90FF7CFEEBD01EFC1C" pageId="47" pageNumber="48" type="reference_group">
<paragraph id="EE5E36A7FFF6FF90FF7CFEEBD15BFE9C" blockId="47.[151,1436,258,403]" pageId="47" pageNumber="48">
<taxonomicName id="29E14D24FFF6FF90FF7CFEEBD0C4FEF0" ID-CoL="WNS8" authority="Cisneros-Heredia, Venegas, Rada" authorityName="Cisneros-Heredia, Venegas, Rada" authorityYear="2008" box="[151,946,258,280]" class="Amphibia" family="Centrolenidae" genus="Cochranella" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="mcdiarmidi">
<emphasis id="DC95EAB5FFF6FF90FF7CFEEBD2E4FEF0" box="[151,402,258,280]" italics="true" pageId="47" pageNumber="48">Cochranella mcdiarmidi</emphasis>
<bibRefCitation id="8A704B56FFF6FF90FE72FEEBD0C4FEF0" author="Cisneros-Heredia" box="[409,946,258,280]" pageId="47" pageNumber="48" refString="Cisneros-Heredia, D. F., Venegas, P. J., Rada, M. &amp; Schulte, R. (2008) A new species of glassfrog (Anura: Centrolenidae) from the foothill Andean forests of Ecuador and Peru. Herpetologica, 64, 341 - 353. http: // dx. doi. org / 10.1655 / 06 - 078.1" type="journal article" year="2008">Cisneros-Heredia, Venegas, Rada, and Schulte 2008</bibRefCitation>
</taxonomicName>
.
<typeStatus id="315A8805FFF6FF90FC56FEEBD76DFEF0" box="[957,1051,258,280]" pageId="47" pageNumber="48" type="holotype">Holotype</typeStatus>
DFCH-USFQ D132, an adult female from “a small rivulet tributary of the Jambue River, ca.
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S from Zamora (ca.
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,
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,
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), on the western slope of Contrafuerte de Tzunantza, Cordillera Oriental, eastern slopes of the Andes, Provincia de Zamora- Chinchipe,
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</paragraph>
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<emphasis id="DC95EAB5FFF6FF90FF7CFE94D2F0FE7B" box="[151,390,381,403]" italics="true" pageId="47" pageNumber="48">
<taxonomicName id="29E14D24FFF6FF90FF7CFE94D383FE7B" ID-CoL="7B4J" box="[151,245,381,403]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="genus">Rulyrana</taxonomicName>
mcdiarmidi—
</emphasis>
<bibRefCitation id="8A704B56FFF6FF90FE6DFE94D736FE7B" author="Guayasamin" box="[390,1088,381,403]" pageId="47" pageNumber="48" refString="Guayasamin, J. M., Castroviejo-Fisher, S., Trueb, L., Ayarzaguena, J., Rada, M. &amp; Vila, C. (2009) Phylogenetic systematics of Glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1 - 97." type="journal article" year="2009">Guayasamin, Castroviejo-Fisher, Trueb, Ayarzagüena, and Vilá 2009</bibRefCitation>
</paragraph>
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<emphasis id="DC95EAB5FFF6FF90FF7CFE56D2BEFE30" bold="true" box="[151,456,447,472]" pageId="47" pageNumber="48">Background information.</emphasis>
This species was described from a series of five specimens collected from two sites in southern
<collectingCountry id="96F67637FFF6FF90FF14FE0DD22AFE15" box="[255,348,484,509]" name="Ecuador" pageId="47" pageNumber="48">Ecuador</collectingCountry>
and one site in northern
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at elevations between
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. The species was assigned to
<taxonomicName id="29E14D24FFF6FF90FF7CFDEED255FDC8" ID-CoL="3RQB" box="[151,291,519,544]" class="Amphibia" family="Centrolenidae" genus="Cochranella" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="genus">
<emphasis id="DC95EAB5FFF6FF90FF7CFDEED255FDC8" box="[151,291,519,544]" italics="true" pageId="47" pageNumber="48">Cochranella</emphasis>
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on the basis of the humeral spine being absent in males (Cisneros-Heredia
<emphasis id="DC95EAB5FFF6FF90FB83FDE0D7D6FDC8" box="[1128,1184,519,544]" italics="true" pageId="47" pageNumber="48">et al.</emphasis>
2008).
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Guayasamin
<emphasis id="DC95EAB5FFF6FF90FA63FDE0D3C2FDAD" italics="true" pageId="47" pageNumber="48">et al.</emphasis>
(2009)
</bibRefCitation>
assigned this species to the genus
<taxonomicName id="29E14D24FFF6FF90FD7BFDC5D18EFDAD" ID-CoL="7B4J" box="[656,760,556,581]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="genus">
<emphasis id="DC95EAB5FFF6FF90FD7BFDC5D18EFDAD" box="[656,760,556,581]" italics="true" pageId="47" pageNumber="48">Rulyrana</emphasis>
</taxonomicName>
considering morphological similarity to
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<emphasis id="DC95EAB5FFF6FF90FB2CFDC4D6F7FDAD" box="[1223,1409,556,581]" italics="true" pageId="47" pageNumber="48">R. flavopunctata</emphasis>
</taxonomicName>
, a species that had been sequenced in their analysis.
</paragraph>
<paragraph id="EE5E36A7FFF6FF90FF2CFD9DD01EFC1C" blockId="47.[151,1437,447,2021]" pageId="47" pageNumber="48">
<emphasis id="DC95EAB5FFF6FF90FF2CFD9DD232FD65" bold="true" box="[199,324,628,653]" pageId="47" pageNumber="48">New data.</emphasis>
Since 2010 we have encountered this species in a number of sites throughout northern
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, extending the known range of this species roughly
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to the south and east of the Peruvian locality from the original description, and between elevations of
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(
<figureCitation id="76DA2A22FFF6FF90FCADFD55D0EFFD3C" box="[838,921,700,725]" captionStart="FIGURE 14" captionStartId="24.[151,250,1550,1572]" captionTargetBox="[172,1403,193,1497]" captionTargetId="figure@24.[172,1404,193,1498]" captionTargetPageId="24" captionText="FIGURE 14. Type locality of Chimerella corleone new species (blue dot) in San Jose, San Martín, 6 ° 25 ' 2.59 &quot; S, 76 ° 17 ' 21.33 &quot; W, 610 m. Rulyrana mcdiarmidi (purple dots), (1) 1 km S Conchay, Ecuador, approx. 3 ° 6 ' S, 78 ° 25 ' W, 1100 m; (2) type locality, 6 km S from Zamora, Ecuador, approx .. 4 ° 3 ' S 78 ° 56 ' W, 1150 m; (3) La Oliva, Amazonas, 5 ° 18 ' 3.86 &quot; S, 78 ° 23 ' 44.57 &quot; W, 682 m; (4) southwest Cordillera del Condor, Cajamarca, 5 ° 25 ' 16.5 &quot; S, 78 ° 35 ' 23.2 &quot; W, 1210 m; (5) Quebrada Goca, Amazonas, 5 ° 45 ' 50.78 &quot; S, 77 ° 54 ' 46.52 &quot; W, 1711 m; (6) 11.3 km WNW from Pedro Ruiz, Amazonas, 5 ° 54 ' 50.93 &quot; S, 78 ° 4 ' 35.20 &quot; W, 1077 m; (7) 4.7 km NW from Pedro Ruiz, Amazonas, 5 ° 55 ' 5.80 &quot; S, 78 ° 0 ' 38.62 &quot; W, 1238 m; (8) 0.45 km S from the town of Cocachimba, Amazonas, 6 ° 3 ' 44.28 &quot; S, 77 ° 53 ' 36.47 &quot; W, 1811 m. Teratohyla amelie (red dots), (1) type locality, Comunidad de Oglán, Cantón Arajuno, Ecuador, 1 ° 18 ' 65 &quot; S, 77 ° 42 ' 41 &quot; W, 600 m; (2) Huabico, Amazonas, 4 ° 57 ' 16.59 &quot; S, 78 ° 12 ' 39.92 &quot; W, 349 m; (3) Nuevo Jaen, San Martín, 7 ° 39 ' 16.20 &quot; S, 76 ° 39 ' 24.60 &quot; W, 591 m; (4) 10 km from Quincemil towards Pto. Maldonado, Cusco (13 ° 12 ' 3.6 &quot; S, 70 ° 40 ' 28.9 &quot; W, 572 m). Nymphargus chancas (yellow dots), (1) Yurilamas, San Martín, approx. 6 ° 14 ' S, 76 ° 32 ' &quot; W, unknown elevation; (2) type locality, Abra Tangarana, San Martín, 6 ° 16 ' 52.86 &quot; S, 76 ° 43 ' 57.86 &quot; W, 1047 m; (3) 25 km SW from Tarapoto, San Martín, 6 ° 35 ' 38.10 &quot; S, 76 ° 34 ' 17.24 &quot; W, 1131 m; (4) 14 km SE from the town of Shamboyacu, San Martín, 7 ° 4 ' 8.9 '' S, 76 ° 0 ' 55.2 '' W, 1122 m. Asterisks on map indicate type localities for each species." httpUri="https://zenodo.org/record/286937/files/figure.png" pageId="47" pageNumber="48">Fig. 14</figureCitation>
). Here, we provide a list of new localities and associated voucher specimens (when collected): village of Cocachimba, Amazonas,
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, near Gocta waterfall (CORBIDI-HE-2010-6847 and CORBIDI-HE-2010-6848,
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,
<geoCoordinate id="8BD55060FFF6FF90FC2DFCEDD705FCF5" box="[966,1139,772,797]" direction="west" orientation="longitude" pageId="47" pageNumber="48" precision="1" value="-77.89346">77°53'36.47&quot;W</geoCoordinate>
,
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); near the village of La Oliva past the village of Muyo (a larger village roughly
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N from Bagua), Amazonas,
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(MNCN 45957 and MHNC 14015,
<geoCoordinate id="8BD55060FFF6FF90FE20FCA5D125FC8D" box="[459,595,844,869]" direction="south" orientation="latitude" pageId="47" pageNumber="48" precision="1" value="-5.3010726">5°18'3.86&quot;S</geoCoordinate>
,
<geoCoordinate id="8BD55060FFF6FF90FD8AFCA5D079FC8D" box="[609,783,844,869]" direction="west" orientation="longitude" pageId="47" pageNumber="48" precision="1" value="-78.39571">78°23'44.57&quot;W</geoCoordinate>
,
<quantity id="29199B42FFF6FF90FCF7FCA5D01EFC8C" box="[796,872,844,869]" metricMagnitude="2" metricUnit="m" metricValue="6.82" pageId="47" pageNumber="48" unit="m" value="682.0">682 m</quantity>
); south slope of Utcubamba canyon,
<quantity id="29199B42FFF6FF90FAFFFCA5D613FC8D" box="[1300,1381,844,869]" metricMagnitude="3" metricUnit="m" metricValue="4.7" pageId="47" pageNumber="48" unit="km" value="4.7">4.7 km</quantity>
NW from Pedro Ruiz, Amazonas,
<collectingCountry id="96F67637FFF6FF90FE07FC99D156FC6F" box="[492,544,880,903]" name="Peru" pageId="47" pageNumber="48">Peru</collectingCountry>
(uncollected individuals,
<geoCoordinate id="8BD55060FFF6FF90FCA3FC86D0A7FC60" box="[840,977,879,904]" direction="south" orientation="latitude" pageId="47" pageNumber="48" precision="1" value="-5.9182777">5°55'5.80&quot;S</geoCoordinate>
,
<geoCoordinate id="8BD55060FFF6FF90FC34FC86D708FC60" box="[991,1150,879,904]" direction="west" orientation="longitude" pageId="47" pageNumber="48" precision="1" value="-78.01073">78°0'38.62&quot;W</geoCoordinate>
,
<quantity id="29199B42FFF6FF90FB60FC99D790FC60" box="[1163,1254,880,904]" metricMagnitude="3" metricUnit="m" metricValue="1.238" pageId="47" pageNumber="48" unit="m" value="1238.0">1238 m</quantity>
); north slope of Utcubamba canyon,
<quantity id="29199B42FFF6FF90FE7AFC7DD283FC45" box="[401,501,916,941]" metricMagnitude="4" metricUnit="m" metricValue="1.13" pageId="47" pageNumber="48" unit="km" value="11.3">11.3 km</quantity>
WNW from Pedro Ruiz, Amazonas,
<collectingCountry id="96F67637FFF6FF90FC38FC7CD771FC44" box="[979,1031,917,940]" name="Peru" pageId="47" pageNumber="48">Peru</collectingCountry>
(CORBIDI 10470,
<geoCoordinate id="8BD55060FFF6FF90FAEAFC7DD6EEFC45" box="[1281,1432,916,941]" direction="south" orientation="latitude" pageId="47" pageNumber="48" precision="1" value="-5.9141474">5°54'50.93&quot;S</geoCoordinate>
,
<geoCoordinate id="8BD55060FFF6FF90FF7CFC5ED240FC38" box="[151,310,951,976]" direction="west" orientation="longitude" pageId="47" pageNumber="48" precision="1" value="-78.07645">78°4'35.20&quot;W</geoCoordinate>
,
<quantity id="29199B42FFF6FF90FEAFFC51D2E8FC38" box="[324,414,952,976]" metricMagnitude="3" metricUnit="m" metricValue="1.077" pageId="47" pageNumber="48" unit="m" value="1077.0">1077 m</quantity>
); Quebrada Goca on Yambrasbamba road, about
<quantity id="29199B42FFF6FF90FC3FFC51D779FC38" box="[980,1039,951,976]" metricMagnitude="3" metricUnit="m" metricValue="1.0" pageId="47" pageNumber="48" unit="km" value="1.0">1 km</quantity>
N from road entrance, Amazonas,
<collectingCountry id="96F67637FFF6FF90FF7CFC34D3BDFC1C" box="[151,203,989,1012]" name="Peru" pageId="47" pageNumber="48">Peru</collectingCountry>
(CORBIDI 10473,
<geoCoordinate id="8BD55060FFF6FF90FE42FC35D136FC1D" box="[425,576,988,1013]" direction="south" orientation="latitude" pageId="47" pageNumber="48" precision="1" value="-5.7641053">5°45'50.78&quot;S</geoCoordinate>
,
<geoCoordinate id="8BD55060FFF6FF90FDA0FC35D18EFC1D" box="[587,760,988,1013]" direction="west" orientation="longitude" pageId="47" pageNumber="48" precision="1" value="-77.912926">77°54'46.52&quot;W</geoCoordinate>
,
<quantity id="29199B42FFF6FF90FCE8FC35D02DFC1C" box="[771,859,988,1012]" metricMagnitude="3" metricUnit="m" metricValue="1.7109999999999999" pageId="47" pageNumber="48" unit="m" value="1711.0">1711 m</quantity>
).
</paragraph>
</subSubSection>
<subSubSection id="A6FB652CFFF6FF90FF2CFC16D093FA69" pageId="47" pageNumber="48" type="description">
<paragraph id="EE5E36A7FFF6FF90FF2CFC16D0BDFB6D" blockId="47.[151,1437,447,2021]" pageId="47" pageNumber="48">
<emphasis id="DC95EAB5FFF6FF90FF2CFC16D24DFBF0" bold="true" box="[199,315,1023,1048]" pageId="47" pageNumber="48">Variation.</emphasis>
We have not noted variation in this species beyond what is reported in Cisneros-Heredia
<emphasis id="DC95EAB5FFF6FF90FA8BFBE8D6EAFBF0" box="[1376,1436,1023,1048]" italics="true" pageId="47" pageNumber="48">et al.</emphasis>
(2008), except that dorsal coloration ranges from light to dark green and the intensity of dorsal spotting varies among individuals. One juvenile (CORBIDI 10470) had reduced spotting compared to adults photographed in a site nearby. Thus, intensity of spotting may exhibit ontogenetic variation.
</paragraph>
<paragraph id="EE5E36A7FFF6FF90FF2CFB66D093FA69" blockId="47.[151,1437,447,2021]" pageId="47" pageNumber="48">
<emphasis id="DC95EAB5FFF6FF90FF2CFB66D21AFB40" bold="true" box="[199,364,1167,1192]" pageId="47" pageNumber="48">Vocalizations.</emphasis>
We have heard
<taxonomicName id="29E14D24FFF6FF90FDCAFB78D1B4FB40" box="[545,706,1167,1192]" class="Amphibia" family="Centrolenidae" genus="Cochranella" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="mcdiarmidi">
<emphasis id="DC95EAB5FFF6FF90FDCAFB78D1B4FB40" box="[545,706,1167,1192]" italics="true" pageId="47" pageNumber="48">R. mcdiarmidi</emphasis>
</taxonomicName>
calling in two sites (
<quantity id="29199B42FFF6FF90FC45FB79D088FB40" box="[942,1022,1167,1192]" metricMagnitude="3" metricUnit="m" metricValue="4.7" pageId="47" pageNumber="48" unit="km" value="4.7">4.7 km</quantity>
NW from Pedro Ruiz and Quebrada Goca, mentioned above). Unfortunately, we have been unable to obtain a recording of this species. The call sounds similar to that of
<taxonomicName id="29E14D24FFF6FF90FEB1FB30D178FB18" box="[346,526,1239,1264]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="saxiscandens">
<emphasis id="DC95EAB5FFF6FF90FEB1FB30D178FB18" box="[346,526,1239,1264]" italics="true" pageId="47" pageNumber="48">R. saxiscandens</emphasis>
</taxonomicName>
(described below), consisting of a single, piercing “zip” of short duration. In the site near Pedro Ruiz, several males were observed calling from a vertical rock face of a small waterfall. In Quebrada Goca, one male was observed calling (and several others were heard) on a small branch hanging roughly
<quantity id="29199B42FFF6FF90FF7CFAADD3B5FAB4" box="[151,195,1348,1372]" metricMagnitude="0" metricUnit="m" metricValue="1.0" pageId="47" pageNumber="48" unit="m" value="1.0">1 m</quantity>
above the edge of the stream. These frogs were concentrated around fast-flowing sections of the stream, but despite background noise, the call could be heard up to roughly
<quantity id="29199B42FFF6FF90FC8FFA8ED0EBFA68" box="[868,925,1383,1408]" metricMagnitude="1" metricUnit="m" metricValue="5.0" pageId="47" pageNumber="48" unit="m" value="50.0">50 m</quantity>
away.
</paragraph>
</subSubSection>
<subSubSection id="A6FB652CFFF6FF8FFF2CFA65D152FDF0" lastPageId="48" lastPageNumber="49" pageId="47" pageNumber="48" type="distribution">
<paragraph id="EE5E36A7FFF6FF90FF2CFA65D709F9B0" blockId="47.[151,1437,447,2021]" pageId="47" pageNumber="48">
<emphasis id="DC95EAB5FFF6FF90FF2CFA65D282FA4D" bold="true" box="[199,500,1420,1445]" pageId="47" pageNumber="48">Distribution and ecology.</emphasis>
Including the new localities mentioned above,
<taxonomicName id="29E14D24FFF6FF90FBE7FA64D7DBFA4D" box="[1036,1197,1420,1445]" class="Amphibia" family="Centrolenidae" genus="Cochranella" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="mcdiarmidi">
<emphasis id="DC95EAB5FFF6FF90FBE7FA64D7DBFA4D" box="[1036,1197,1420,1445]" italics="true" pageId="47" pageNumber="48">R. mcdiarmidi</emphasis>
</taxonomicName>
is widely distributed in the Utcubamba and central Marañón drainages in Amazonas,
<collectingCountry id="96F67637FFF6FF90FC99FA58D0D3FA20" box="[882,933,1457,1480]" name="Peru" pageId="47" pageNumber="48">Peru</collectingCountry>
and eastern foothills of southern
<collectingCountry id="96F67637FFF6FF90FAC9FA46D609FA20" box="[1314,1407,1455,1480]" name="Ecuador" pageId="47" pageNumber="48">Ecuador</collectingCountry>
at elevations between
<quantity id="29199B42FFF6FF90FE9FFA3DD174FA04" box="[372,514,1492,1517]" metricMagnitude="3" metricUnit="m" metricValue="1.2465" metricValueMax="1.811" metricValueMin="0.682" pageId="47" pageNumber="48" unit="m" value="1246.5" valueMax="1811.0" valueMin="682.0">6821811 m</quantity>
. This species has not been found on the Huallaga-side of Abra Patricia in northern
<collectingCountry id="96F67637FFF6FF90FF7CFA10D3B8F9F8" box="[151,206,1529,1552]" name="Peru" pageId="47" pageNumber="48">Peru</collectingCountry>
.
<taxonomicName id="29E14D24FFF6FF90FF37FA10D2F6F9F8" box="[220,384,1527,1552]" class="Amphibia" family="Centrolenidae" genus="Cochranella" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="mcdiarmidi">
<emphasis id="DC95EAB5FFF6FF90FF37FA10D2F6F9F8" box="[220,384,1527,1552]" italics="true" pageId="47" pageNumber="48">R. mcdiarmidi</emphasis>
</taxonomicName>
occurs in a wide variety of habitats, ranging from pristine rainforest (e.g, near Muyo) to heavily disturbed, scrubby habitats (e.g., near Cocachimba). Adults and juveniles have all been found on rocks or vegetation typically within
<quantity id="29199B42FFF6FF90FE21F9A9D283F9B0" box="[458,501,1600,1624]" metricMagnitude="0" metricUnit="m" metricValue="2.0" pageId="47" pageNumber="48" unit="m" value="2.0">2 m</quantity>
of the water surface. Eggs and tadpoles remain unknown.
</paragraph>
<paragraph id="EE5E36A7FFF6FF90FF2CF98DD27FF900" blockId="47.[151,1437,447,2021]" pageId="47" pageNumber="48">
<taxonomicName id="29E14D24FFF6FF90FF2CF98DD2B7F995" box="[199,449,1636,1661]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="mcdiarmidi">
<emphasis id="DC95EAB5FFF6FF90FF2CF98DD2B7F995" box="[199,449,1636,1661]" italics="true" pageId="47" pageNumber="48">Rulyrana mcdiarmidi</emphasis>
</taxonomicName>
is sympatric with a number of species throughout its known range, including
<taxonomicName id="29E14D24FFF6FF90FF7CF96ED2FDF948" box="[151,395,1671,1696]" class="Amphibia" family="Centrolenidae" genus="Nymphargus" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="posadae">
<emphasis id="DC95EAB5FFF6FF90FF7CF96ED2FDF948" box="[151,395,1671,1696]" italics="true" pageId="47" pageNumber="48">Nymphargus posadae</emphasis>
</taxonomicName>
in the southern Cordillera del
<taxonomicName id="29E14D24FFF6FF90FD05F96ED034F948" box="[750,834,1671,1696]" class="Amphibia" family="Centrolenidae" genus="Centrolene" kingdom="Animalia" order="Anura" pageId="47" pageNumber="58" phylum="Chordata" rank="species" species="condor">Condor</taxonomicName>
in
<collectingCountry id="96F67637FFF6FF90FC82F960D0EBF948" box="[873,925,1673,1696]" name="Peru" pageId="47" pageNumber="48">Peru</collectingCountry>
(Cisneros-Heredia
<emphasis id="DC95EAB5FFF6FF90FB97F960D7C0F948" box="[1148,1206,1671,1696]" italics="true" pageId="47" pageNumber="48">et al.</emphasis>
2008),
<taxonomicName id="29E14D24FFF6FF90FAFBF96ED385F92C" class="Amphibia" family="Centrolenidae" genus="Cochranella" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="erminea">
<emphasis id="DC95EAB5FFF6FF90FAFBF96ED385F92C" italics="true" pageId="47" pageNumber="48">Cochranella erminea</emphasis>
</taxonomicName>
and
<taxonomicName id="29E14D24FFF6FF90FEC3F945D16EF92D" box="[296,536,1708,1733]" class="Amphibia" family="Centrolenidae" genus="Centrolene" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="charapita">
<emphasis id="DC95EAB5FFF6FF90FEC3F945D16EF92D" box="[296,536,1708,1733]" italics="true" pageId="47" pageNumber="48">Centrolene charapita</emphasis>
</taxonomicName>
in the Muyo site in
<collectingCountry id="96F67637FFF6FF90FD13F944D058F92C" box="[760,814,1709,1732]" name="Peru" pageId="47" pageNumber="48">Peru</collectingCountry>
, and
<taxonomicName id="29E14D24FFF6FF90FC83F945D7FFF92D" box="[872,1161,1708,1733]" class="Amphibia" family="Centrolenidae" genus="Espadarana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="fernandoi">
<emphasis id="DC95EAB5FFF6FF90FC83F945D087F92D" box="[872,1009,1708,1733]" italics="true" pageId="47" pageNumber="48">Espadarana</emphasis>
cf.
<emphasis id="DC95EAB5FFF6FF90FBF7F945D7FFF92D" box="[1052,1161,1708,1733]" italics="true" pageId="47" pageNumber="48">fernandoi</emphasis>
</taxonomicName>
in Quebrada Goca,
<collectingCountry id="96F67637FFF6FF90FA82F944D6EAF92C" box="[1385,1436,1709,1732]" name="Peru" pageId="47" pageNumber="48">Peru</collectingCountry>
(
<figureCitation id="76DA2A22FFF6FF90FF74F926D384F900" box="[159,242,1743,1768]" captionStart="FIGURE 27. a, b" captionStartId="46.[151,250,1681,1703]" captionTargetBox="[151,1434,375,1658]" captionTargetId="figure@46.[151,1436,375,1659]" captionTargetPageId="46" captionText="FIGURE 27. a, b) Nymphargus chancas, dorsal and ventral views of CORBIDI 10471, from 5.9 km SE from the type locality of Abra Tangarana, San Martín, 6 ° 19 ' 15.65 &quot; S 76 ° 41 ' 44.16 &quot; W, 973 m; c, d) Teratohyla amelie, dorsal and ventral views of MNCN 45958, from Nuevo Jaen, San Martín 7 ° 39 ' 16.20 &quot; S, 76 ° 39 ' 24.60 &quot; W, 591 m; e) Rulyrana mcdiarmidi, uncollected male from 0.45 km S from the town of Cocachimba, Amazonas, 6 ° 3 ' 44.28 &quot; S, 77 ° 53 ' 36.47 &quot; W, 1811 m; f) Espadarana cf. fernandoi, dorsal view of adult male (CORBIDI 10468) from Quebrada Goca on the Yambrasbamba road, about 1 km N from road entrance, Amazonas, 5 ° 45 ' 50.78 &quot; S, 77 ° 54 ' 46.52 &quot; W, 1711 m." httpUri="https://zenodo.org/record/286951/files/figure.png" pageId="47" pageNumber="48">Fig. 27</figureCitation>
f).
</paragraph>
<paragraph id="EE5E36A7FFF6FF8FFF2CF91DD152FDF0" blockId="47.[151,1437,447,2021]" lastBlockId="48.[151,1437,151,536]" lastPageId="48" lastPageNumber="49" pageId="47" pageNumber="48">
<emphasis id="DC95EAB5FFF6FF90FF2CF91DD2B5F8E5" bold="true" box="[199,451,1780,1805]" pageId="47" pageNumber="48">Taxonomic remarks.</emphasis>
Some of the characters used to differentiate
<taxonomicName id="29E14D24FFF6FF90FC00F91CD7E7F8E5" box="[1003,1169,1780,1805]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="mcdiarmidi">
<emphasis id="DC95EAB5FFF6FF90FC00F91CD7E7F8E5" box="[1003,1169,1780,1805]" italics="true" pageId="47" pageNumber="48">R. mcdiarmidi</emphasis>
</taxonomicName>
from
<taxonomicName id="29E14D24FFF6FF90FB08F91CD6EAF8E5" box="[1251,1436,1780,1805]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="saxiscandens">
<emphasis id="DC95EAB5FFF6FF90FB08F91CD6EAF8E5" box="[1251,1436,1780,1805]" italics="true" pageId="47" pageNumber="48">R. saxiscandens</emphasis>
</taxonomicName>
(following our definition that considers
<taxonomicName id="29E14D24FFF6FF90FD9FF8F0D078F8D8" box="[628,782,1817,1840]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="tangarana">
<emphasis id="DC95EAB5FFF6FF90FD9FF8F0D078F8D8" box="[628,782,1817,1840]" italics="true" pageId="47" pageNumber="48">R. tangarana</emphasis>
</taxonomicName>
as a junior synonym, see below) do not appear to be unambiguously diagnostic. For example, purple coloration in preservative is variable in
<taxonomicName id="29E14D24FFF6FF90FB4AF8D4D62EF8BD" box="[1185,1368,1852,1877]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="saxiscandens">
<emphasis id="DC95EAB5FFF6FF90FB4AF8D4D62EF8BD" box="[1185,1368,1852,1877]" italics="true" pageId="47" pageNumber="48">R. saxiscandens</emphasis>
</taxonomicName>
, with some specimens appearing dull green or grayish lavender (see
<taxonomicName id="29E14D24FFF6FF90FC9BF888D750F890" box="[880,1062,1887,1912]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="saxiscandens">
<emphasis id="DC95EAB5FFF6FF90FC9BF888D750F890" box="[880,1062,1887,1912]" italics="true" pageId="47" pageNumber="48">R. saxiscandens</emphasis>
</taxonomicName>
account below). On average,
<taxonomicName id="29E14D24FFF6FF90FA6FF888D26EF875" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="mcdiarmidi">
<emphasis id="DC95EAB5FFF6FF90FA6FF888D26EF875" italics="true" pageId="47" pageNumber="48">R. mcdiarmidi</emphasis>
</taxonomicName>
likely has a larger SVL, but the two species overlap with respect to this character (two
<taxonomicName id="29E14D24FFF6FF90FB10F86CD6EAF875" box="[1275,1436,1924,1949]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="mcdiarmidi">
<emphasis id="DC95EAB5FFF6FF90FB10F86CD6EAF875" box="[1275,1436,1924,1949]" italics="true" pageId="47" pageNumber="48">R. mcdiarmidi</emphasis>
</taxonomicName>
males
<quantity id="29199B42FFF6FF90FF34F841D2F2F828" box="[223,388,1960,1985]" metricMagnitude="-2" metricUnit="m" metricValue="2.6149999999999998" metricValueMax="2.69" metricValueMin="2.54" pageId="47" pageNumber="48" unit="mm" value="26.15" valueMax="26.9" valueMin="25.4">25.426.9 mm</quantity>
, six
<taxonomicName id="29E14D24FFF6FF90FE53F840D11DF829" box="[440,619,1960,1985]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="saxiscandens">
<emphasis id="DC95EAB5FFF6FF90FE53F840D11DF829" box="[440,619,1960,1985]" italics="true" pageId="47" pageNumber="48">R. saxiscandens</emphasis>
</taxonomicName>
males
<quantity id="29199B42FFF6FF90FD50F841D014F828" box="[699,866,1960,1985]" metricMagnitude="-2" metricUnit="m" metricValue="2.445" metricValueMax="2.66" metricValueMin="2.23" pageId="47" pageNumber="48" unit="mm" value="24.45" valueMax="26.6" valueMin="22.3">22.326.6 mm</quantity>
). Ulnar folds are visible in life in some individuals of
<taxonomicName id="29E14D24FFF6FF90FF52F824D206F80D" box="[185,368,1996,2021]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="saxiscandens">
<emphasis id="DC95EAB5FFF6FF90FF52F824D206F80D" box="[185,368,1996,2021]" italics="true" pageId="47" pageNumber="48">R. saxiscandens</emphasis>
</taxonomicName>
, but generally less-pronounced than in
<taxonomicName id="29E14D24FFF6FF90FCAFF824D09EF80D" box="[836,1000,1996,2021]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="mcdiarmidi">
<emphasis id="DC95EAB5FFF6FF90FCAFF824D09EF80D" box="[836,1000,1996,2021]" italics="true" pageId="47" pageNumber="48">R. mcdiarmidi</emphasis>
</taxonomicName>
. Also,
<taxonomicName id="29E14D24FFF6FF90FBAAF824D793F80D" box="[1089,1253,1996,2021]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="mcdiarmidi">
<emphasis id="DC95EAB5FFF6FF90FBAAF824D793F80D" box="[1089,1253,1996,2021]" italics="true" pageId="47" pageNumber="48">R. mcdiarmidi</emphasis>
</taxonomicName>
is described as having medium-sized eyes (ED/3WD = 1.101.70), even though both male
<typeStatus id="315A8805FFE9FF8FFBFEFF71D7F7FF58" box="[1045,1153,152,176]" pageId="48" pageNumber="49" type="paratype">paratypes</typeStatus>
have ED/3WD of 1.93, again, within the variation we observed in
<taxonomicName id="29E14D24FFE9FF8FFD97FF54D047FF3D" box="[636,817,188,213]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="48" pageNumber="49" phylum="Chordata" rank="species" species="saxiscandens">
<emphasis id="DC95EAB5FFE9FF8FFD97FF54D047FF3D" box="[636,817,188,213]" italics="true" pageId="48" pageNumber="49">R. saxiscandens</emphasis>
</taxonomicName>
(ED/3WD = 1.752.45, N = 6). However, presence of flat warts with low tubercles (which are typically lime green in life) in
<taxonomicName id="29E14D24FFE9FF8FFC51FF08D72DFF10" box="[954,1115,223,248]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="48" pageNumber="49" phylum="Chordata" rank="species" species="mcdiarmidi">
<emphasis id="DC95EAB5FFE9FF8FFC51FF08D72DFF10" box="[954,1115,223,248]" italics="true" pageId="48" pageNumber="49">R. mcdiarmidi</emphasis>
</taxonomicName>
vs. skin uniformly spiculate and shagreened in
<taxonomicName id="29E14D24FFE9FF8FFE86FEECD154FEF5" box="[365,546,260,285]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="48" pageNumber="49" phylum="Chordata" rank="species" species="saxiscandens">
<emphasis id="DC95EAB5FFE9FF8FFE86FEECD154FEF5" box="[365,546,260,285]" italics="true" pageId="48" pageNumber="49">R. saxiscandens</emphasis>
</taxonomicName>
may be a good diagnostic character between these two species. Nevertheless, the results from our phylogenetic analysis do not recover
<taxonomicName id="29E14D24FFE9FF8FFCA2FEC0D098FEA8" box="[841,1006,295,320]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="48" pageNumber="49" phylum="Chordata" rank="species" species="mcdiarmidi">
<emphasis id="DC95EAB5FFE9FF8FFCA2FEC0D098FEA8" box="[841,1006,295,320]" italics="true" pageId="48" pageNumber="49">R. mcdiarmidi</emphasis>
</taxonomicName>
and
<taxonomicName id="29E14D24FFE9FF8FFBC4FEC0D791FEA8" box="[1071,1255,295,320]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="48" pageNumber="49" phylum="Chordata" rank="species" species="saxiscandens">
<emphasis id="DC95EAB5FFE9FF8FFBC4FEC0D791FEA8" box="[1071,1255,295,320]" italics="true" pageId="48" pageNumber="49">R. saxiscandens</emphasis>
</taxonomicName>
as reciprocally monophyletic. Furthermore, nested within this clade is an individual referred to as
<taxonomicName id="29E14D24FFE9FF8FFBC6FEA4D388FE60" authority="Guayasamin et al. 2008" authorityName="Guayasamin et al." authorityYear="2008" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="48" pageNumber="49" phylum="Chordata" rank="species" species="flavopunctata">
<emphasis id="DC95EAB5FFE9FF8FFBC6FEA4D790FE8D" box="[1069,1254,332,357]" italics="true" pageId="48" pageNumber="49">R. flavopunctata</emphasis>
(
<bibRefCitation id="8A704B56FFE9FF8FFB1FFEA5D380FE60" author="Guayasamin" pageId="48" pageNumber="49" refString="Guayasamin, J. M., Castroviejo-Fisher, S., Ayarzaguena, J., Trueb, L. &amp; Vila, C. (2008) Phylogenetic relationships of glassfrogs (Centrolenidae) based on mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution, 48, 574 - 595. http: // dx. doi. org / 10.1016 / j. ympev. 2008.04.012" type="journal article" year="2008">
Guayasamin
<emphasis id="DC95EAB5FFE9FF8FFA63FEA4D3C2FE60" italics="true" pageId="48" pageNumber="49">et al.</emphasis>
2008
</bibRefCitation>
)
</taxonomicName>
from Provincia Morona-Santiago,
<collectingCountry id="96F67637FFE9FF8FFD66FE86D19CFE60" box="[653,746,367,392]" name="Ecuador" pageId="48" pageNumber="49">Ecuador</collectingCountry>
(
<figureCitation id="76DA2A22FFE9FF8FFD10FE86D039FE60" box="[763,847,367,392]" captionStart-0="FIGURE 2 a" captionStart-1="FIGURE 2 c" captionStartId-0="7.[151,250,1894,1916]" captionStartId-1="9.[151,250,1900,1922]" captionTargetBox-0="[151,1436,193,1850]" captionTargetBox-1="[193,1379,190,1860]" captionTargetId-0="figure@7.[151,1436,193,1857]" captionTargetId-1="figure@9.[193,1380,190,1888]" captionTargetPageId-0="7" captionTargetPageId-1="9" captionText-0="FIGURE 2 a. Phylogeny of Centrolenidae, Allophrynidae, and outgroups. Strict consensus tree of 52 most parsimonious trees (cost = 16898 steps). Values above and below branches indicate jackknife and Goodman-Bremer support, respectively. a) outgroups, Allophrynidae, Celsiella, and Hyalinobatrachium; b) Nymphargus and Centrolene; c) Vitreorana, Teratohyla, Chimerella, Espadarana, Cochranella, Sachatamia, and Rulyrana." captionText-1="FIGURE 2 c. Phylogeny of Centrolenidae, Allophrynidae, and outgroups. Strict consensus tree of 52 most parsimonious trees (cost = 16898 steps). Values above and below branches indicate jackknife and Goodman-Bremer support, respectively. a) outgroups, Allophrynidae, Celsiella, and Hyalinobatrachium; b) Nymphargus and Centrolene; c) Vitreorana, Teratohyla, Chimerella, Espadarana, Cochranella, Sachatamia, and Rulyrana." httpUri-0="https://zenodo.org/record/286923/files/figure.png" httpUri-1="https://zenodo.org/record/286925/files/figure.png" pageId="48" pageNumber="49">Fig. 2c</figureCitation>
). Also, the genetic distances among these terminals are low (p-distances = 0.00.6%,
<tableCitation id="A363031CFFE9FF8FFDD1FE7DD1E1FE45" box="[570,663,404,429]" captionStart="TABLE 6" captionStartId="55.[151,239,936,959]" captionTargetBox="[159,1425,1053,1395]" captionTargetPageId="55" captionText="TABLE 6. Percentage of inter- and intraspecific uncorrected genetic distances (diagonal and below the diagonal respectively) within members of Rulyrana (sample size in parentheses) inferred from 537 aligned characters of a fragment of the mitochondrial 16 S gene. In this work we consider R. saxiscandens and R. tangarana as synonyms." httpUri="http://table.plazi.org/id/BA9E662FFFEEFF88FF7CFC41D628FBE8" pageId="48" pageNumber="49" tableUuid="BA9E662FFFEEFF88FF7CFC41D628FBE8">Table 6</tableCitation>
). These results suggest the possibility of a single, widespread, morphologically variable species. However, our very limited sampling of
<taxonomicName id="29E14D24FFE9FF8FFBE1FE50D7BCFE38" box="[1034,1226,439,464]" class="Amphibia" family="Centrolenidae" genus="Rulyrana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="48" pageNumber="49" phylum="Chordata" rank="species" species="flavopunctata">
<emphasis id="DC95EAB5FFE9FF8FFBE1FE50D7BCFE38" box="[1034,1226,439,464]" italics="true" pageId="48" pageNumber="49">R. flavopunctata</emphasis>
</taxonomicName>
(DNA sequences retrieved from GenBank of a specimen identified by others and that we have not examined) preclude us from taking further taxonomic decisions.
</paragraph>
</subSubSection>
</treatment>
</document>