436 lines
40 KiB
XML
436 lines
40 KiB
XML
<document ID-DOI="http://dx.doi.org/10.3897/zookeys.685.12152" ID-GBIF-Dataset="2462b9c5-a860-456d-baba-b57c10667b57" ID-PMC="PMC5646672" ID-Pensoft-Pub="1313-2970-685-65" ID-PubMed="29089838" ID-ZBK="310FD77C029A45F59AE4C59772CA1F83" ModsDocAuthor="" ModsDocDate="2017" ModsDocID="1313-2970-685-65" ModsDocOrigin="ZooKeys 685" ModsDocTitle="Bryophrynephuyuhampatu sp. n., a new species of Cusco Andes frog from the cloud forest of the eastern slopes of the Peruvian Andes (Amphibia, Anura, Craugastoridae)" checkinTime="1502744503733" checkinUser="pensoft" docAuthor="Catenazzi, Alessandro, Ttito, Alex, Diaz, M. Isabel & Shepack, Alexander" docDate="2017" docId="3291630F99790CC9C007AE0B546371A5" docLanguage="en" docName="ZooKeys 685: 65-81" docOrigin="ZooKeys 685" docSource="http://dx.doi.org/10.3897/zookeys.685.12152" docTitle="Bryophryne phuyuhampatu Catenazzi, Ttito, Diaz & Shepack, 2017, sp. n." docType="treatment" docUuid="BB13FD82-3470-4E31-A6EF-87606B0CC356" docUuidSource="ZooBank" docVersion="5" lastPageNumber="71" masterDocId="D504FF8BFFC9FFCC4421AC2E7554F54E" masterDocTitle="Bryophrynephuyuhampatu sp. n., a new species of Cusco Andes frog from the cloud forest of the eastern slopes of the Peruvian Andes (Amphibia, Anura, Craugastoridae)" masterLastPageNumber="81" masterPageNumber="65" pageNumber="66" updateTime="1668164625785" updateUser="ExternalLinkService">
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<mods:titleInfo>
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<mods:title>Bryophrynephuyuhampatu sp. n., a new species of Cusco Andes frog from the cloud forest of the eastern slopes of the Peruvian Andes (Amphibia, Anura, Craugastoridae)</mods:title>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Catenazzi, Alessandro</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Ttito, Alex</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Diaz, M. Isabel</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Shepack, Alexander</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>ZooKeys</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2017</mods:date>
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<mods:detail type="volume">
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<mods:number>685</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>65</mods:start>
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<mods:end>81</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:location>
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<mods:url>http://dx.doi.org/10.3897/zookeys.685.12152</mods:url>
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</mods:location>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.685.12152</mods:identifier>
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<mods:identifier type="Pensoft-Pub">1313-2970-685-65</mods:identifier>
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<mods:identifier type="ZBK">310FD77C029A45F59AE4C59772CA1F83</mods:identifier>
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<mods:identifier type="ZooBank">310FD77C029A45F59AE4C59772CA1F83</mods:identifier>
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</mods:mods>
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<treatment ID-GBIF-Taxon="132648825" LSID="urn:lsid:zoobank.org:act:BB13FD82-3470-4E31-A6EF-87606B0CC356" httpUri="http://treatment.plazi.org/id/3291630F99790CC9C007AE0B546371A5" lastPageId="6" lastPageNumber="71" pageId="1" pageNumber="66">
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<subSubSection pageId="1" pageNumber="66" type="nomenclature">
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<paragraph pageId="1" pageNumber="66">
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<taxonomicName LSID="http://zoobank.org/BB13FD82-3470-4E31-A6EF-87606B0CC356" class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne phuyuhampatu" order="Anura" pageId="1" pageNumber="66" phylum="Chordata" rank="species" species="phuyuhampatu">Bryophryne phuyuhampatu</taxonomicName>
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<taxonomicNameLabel pageId="1" pageNumber="66">sp. n.</taxonomicNameLabel>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="1" pageNumber="66" type="holotype">
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<paragraph pageId="1" pageNumber="66">Holotype.</paragraph>
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<paragraph pageId="1" pageNumber="66">
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(Figs 1-3, Table 2). CORBIDI 18226, an adult male (Figs 2, 3) from
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<geoCoordinate direction="south" orientation="latitude" precision="1" value="-13.370039">13°22'12.14''S</geoCoordinate>
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;
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<geoCoordinate direction="west" orientation="longitude" precision="1" value="-71.11384">71°6'49.82''W</geoCoordinate>
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(WGS84), 2795-2850 m a.s.l., Quispillomayo valley,
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<normalizedToken originalValue="Área">Area</normalizedToken>
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de
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<normalizedToken originalValue="Conservación">Conservacion</normalizedToken>
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Privada (ACP) Ukumari Llaqta, Distrito Paucartambo, Provincia Paucartambo, Departamento de Cusco, Peru, collected by A. Catenazzi, A. Shepack, M. I. Diaz and A. Ttito on 27 May 2016.
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</paragraph>
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<caption pageId="1" pageNumber="66">
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<paragraph pageId="1" pageNumber="66">
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Figure 1. Map of Peru indicating the type localities of species of
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne" order="Anura" pageId="1" pageNumber="66" phylum="Chordata" rank="genus">Bryophryne</taxonomicName>
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.
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</paragraph>
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</caption>
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<caption pageId="1" pageNumber="66">
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<paragraph pageId="1" pageNumber="66">
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Figure 2. Live (left column) and preserved (right column) specimen of the holotype of
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne phuyuhampatu" order="Anura" pageId="1" pageNumber="66" phylum="Chordata" rank="species" species="phuyuhampatu">Bryophryne phuyuhampatu</taxonomicName>
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sp. n., male CORBIDI 18226 (SVL 16.9 mm) in dorsolateral A, B dorsal C, D and ventral E, F views. Photographs by A. Catenazzi.
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</paragraph>
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</caption>
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<caption pageId="1" pageNumber="66">
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<paragraph pageId="1" pageNumber="66">
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Figure 3. Ventral views of hand A and foot B of holotype, CORBIDI 18226 (hand length 4.1 mm, foot length 6.6 mm) of
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne phuyuhampatu" order="Anura" pageId="1" pageNumber="66" phylum="Chordata" rank="species" species="phuyuhampatu">Bryophryne phuyuhampatu</taxonomicName>
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sp. n. Photographs by A. Catenazzi.
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</paragraph>
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</caption>
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</subSubSection>
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<subSubSection pageId="1" pageNumber="66" type="paratopotypes">
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<paragraph pageId="1" pageNumber="66">Paratopotypes.</paragraph>
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<paragraph pageId="1" pageNumber="66">(Fig. 4, Table 2). Four specimens: two females, CORBIDI 18224 and MUBI 14654, and one male, CORBIDI 18225, collected with the holotype on 27 May 2016; and one male, MUBI 14655 collected on 28 May 2016.</paragraph>
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<caption pageId="1" pageNumber="66">
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<paragraph pageId="1" pageNumber="66">
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Figure 4. Dorsolateral and ventral views of four paratypes of
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne phuyuhampatu" order="Anura" pageId="1" pageNumber="66" phylum="Chordata" rank="species" species="phuyuhampatu">Bryophryne phuyuhampatu</taxonomicName>
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sp. n. showing variation in dorsal and ventral coloration. Male CORBIDI 18225, SVL = 14.2 mm A, B Male MUBI 14655, SVL = 15.9 mm C, D Female CORBIDI 18224, SVL = 22.6 mm E, F. Female MUBI 14654, SVL = 22.2 mm G, H Photographs by A. Catenazzi.
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</paragraph>
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</caption>
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<caption pageId="1" pageNumber="66">
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<paragraph pageId="1" pageNumber="66">
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Table 2. Range and average (
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<normalizedToken originalValue="±">+/-</normalizedToken>
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standard deviation) measurements (in mm) of type series of
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne phuyuhampatu" order="Anura" pageId="1" pageNumber="66" phylum="Chordata" rank="species" species="phuyuhampatu">Bryophryne phuyuhampatu</taxonomicName>
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sp. n.
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</paragraph>
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</caption>
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<paragraph pageId="1" pageNumber="66">
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<table pageId="1" pageNumber="66">
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<tr pageId="1" pageNumber="66">
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<th colspan="1" pageId="1" pageNumber="66" rowspan="1">Characters</th>
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<th colspan="1" pageId="1" pageNumber="66" rowspan="1">Females (n = 2)</th>
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<th colspan="1" pageId="1" pageNumber="66" rowspan="1">Males (n = 3)</th>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">SVL</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">TL</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">FL</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">HL</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">HW</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">ED</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">IOD</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">EW</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">IND</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">
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<normalizedToken originalValue="E–N">E-N</normalizedToken>
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</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">TLSVL</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">FLSVL</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">HLSVL</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">HWSVL</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">HWHL</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">
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<normalizedToken originalValue="E–NED">E-NED</normalizedToken>
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</td>
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</tr>
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<tr pageId="1" pageNumber="66">
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<td colspan="1" pageId="1" pageNumber="66" rowspan="1">EWIOD</td>
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</tr>
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</table>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="1" pageNumber="66" type="referred specimens">
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<paragraph pageId="1" pageNumber="66">Referred specimens.</paragraph>
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<paragraph pageId="1" pageNumber="66">Three juveniles, CORBIDI 18227, 18228, and MUBI 14665, collected with the holotype and paratopotypes on 27 May 2016.</paragraph>
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</subSubSection>
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<subSubSection lastPageId="3" lastPageNumber="68" pageId="1" pageNumber="66" type="generic placement">
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<paragraph pageId="1" pageNumber="66">Generic placement.</paragraph>
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<paragraph lastPageId="3" lastPageNumber="68" pageId="1" pageNumber="66">
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A new species of
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne" order="Anura" pageId="1" pageNumber="66" phylum="Chordata" rank="genus">Bryophryne</taxonomicName>
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as defined by
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<bibRefCitation author="Duellman, WE" journalOrPublisher="Natur und Tier Verlag, Muenster" pageId="9" pageNumber="74" title="Terrestrial-breeding frogs (Strabomantidae) in Peru." year="2009">Duellman and Lehr (2009)</bibRefCitation>
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,
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<bibRefCitation author="Hedges, SB" journalOrPublisher="Zootaxa" pageId="9" pageNumber="74" pagination="1 - 182" title="New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and conservation." volume="1737" year="2008">Hedges et al. (2008)</bibRefCitation>
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, and
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<bibRefCitation author="Padial, JM" journalOrPublisher="Zootaxa" pageId="9" pageNumber="74" pagination="1 - 132" title="Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria." url="https://doi.org/10.11646/zootaxa.3825.1.1" volume="3825" year="2014">Padial et al. (2014)</bibRefCitation>
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. Frogs of the genus
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne" order="Anura" pageId="1" pageNumber="66" phylum="Chordata" rank="genus">Bryophryne</taxonomicName>
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are morphologically similar and closely related to
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Barycholos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Barycholos" order="Anura" pageId="1" pageNumber="66" phylum="Chordata" rank="genus">Barycholos</taxonomicName>
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,
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Holoaden" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Holoaden" order="Anura" pageId="1" pageNumber="66" phylum="Chordata" rank="genus">Holoaden</taxonomicName>
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,
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Noblella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Noblella" order="Anura" pageId="2" pageNumber="67" phylum="Chordata" rank="genus">
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<pageBreakToken pageId="2" pageNumber="67" start="start">Noblella</pageBreakToken>
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</taxonomicName>
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and
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Psychrophrynella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Psychrophrynella" order="Anura" pageId="2" pageNumber="67" phylum="Chordata" rank="genus">Psychrophrynella</taxonomicName>
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(
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<bibRefCitation author="Chaparro, JC" journalOrPublisher="Zootaxa" pageId="8" pageNumber="73" pagination="94 - 108" title="A new species of Andean frog of the genus Bryophryne from southern Peru (Anura: Craugastoridae) and its phylogenetic position, with notes on the diversity of the genus." url="https://doi.org/10.11646/zootaxa.3994.1.4" volume="3994" year="2015">Chaparro et al. 2015</bibRefCitation>
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;
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<bibRefCitation author="Hedges, SB" journalOrPublisher="Zootaxa" pageId="9" pageNumber="74" pagination="1 - 182" title="New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and conservation." volume="1737" year="2008">Hedges et al. 2008</bibRefCitation>
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;
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<bibRefCitation author="Heinicke, MP" journalOrPublisher="Proceedings of the National Academy of Sciences of the United States of America" pageId="9" pageNumber="74" pagination="10092 - 10097" title="Major Caribbean and Central American frog faunas originated by ancient oceanic dispersal." url="https://doi.org/10.1073/pnas.0611051104" volume="104" year="2007">Heinicke et al. 2007</bibRefCitation>
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;
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<bibRefCitation author="Padial, JM" journalOrPublisher="Zootaxa" pageId="9" pageNumber="74" pagination="1 - 132" title="Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria." url="https://doi.org/10.11646/zootaxa.3825.1.1" volume="3825" year="2014">Padial et al. 2014</bibRefCitation>
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). Genetic data confirm generic placement of the new species within
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne" order="Anura" pageId="2" pageNumber="67" phylum="Chordata" rank="genus">Bryophryne</taxonomicName>
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(Table 1). We found substantial genetic distances (uncorrected p-distances from 3.7-6.7%; Table 1) between
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<taxonomicName lsidName="B. phuyuhampatu" pageId="2" pageNumber="67" rank="species" species="phuyuhampatu">B. phuyuhampatu</taxonomicName>
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and congeneric species for which mitochondrial sequence data were available (
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<taxonomicName lsidName="B. bakersfield" pageId="2" pageNumber="67" rank="species" species="bakersfield">B. bakersfield</taxonomicName>
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,
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<taxonomicName lsidName="B. bustamantei" pageId="2" pageNumber="67" rank="species" species="bustamantei">B. bustamantei</taxonomicName>
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, and
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<taxonomicName lsidName="B. cophites" pageId="2" pageNumber="67" rank="species" species="cophites">B. cophites</taxonomicName>
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). The most closely related species is
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<taxonomicName lsidName="B. bakersfield" pageId="2" pageNumber="67" rank="species" species="bakersfield">B. bakersfield</taxonomicName>
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(16S uncorrected p-distance: 3.7-4.1%), followed by
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<taxonomicName lsidName="B. cophites" pageId="2" pageNumber="67" rank="species" species="cophites">B. cophites</taxonomicName>
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(5.6-6.2%) and
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<taxonomicName lsidName="B. bustamantei" pageId="2" pageNumber="67" rank="species" species="bustamantei">B. bustamantei</taxonomicName>
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(5.6-6.7%). Regarding species from other genera,
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<taxonomicName lsidName="B. phuyuhampatu" pageId="2" pageNumber="67" rank="species" species="phuyuhampatu">B. phuyuhampatu</taxonomicName>
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had genetic distances ranging from 12.4% (
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Psychrophrynella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Psychrophrynella guillei" order="Anura" pageId="2" pageNumber="67" phylum="Chordata" rank="species" species="guillei">Psychrophrynella guillei</taxonomicName>
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) to 20.8% (
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Barycholos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Barycholos pulcher" order="Anura" pageId="2" pageNumber="67" phylum="Chordata" rank="species" species="pulcher">Barycholos pulcher</taxonomicName>
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). In addition to the molecular data, the new species is assigned to
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne" order="Anura" pageId="2" pageNumber="67" phylum="Chordata" rank="genus">Bryophryne</taxonomicName>
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rather than any of
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<pageBreakToken pageId="3" pageNumber="68" start="start">the</pageBreakToken>
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other genera on the basis of overall morphological resemblance with the type species
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<taxonomicName lsidName="B. cophites" pageId="3" pageNumber="68" rank="species" species="cophites">B. cophites</taxonomicName>
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, including head narrower than body, short limbs, and tympanic membrane and annulus usually absent (absent in most species of
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne" order="Anura" pageId="3" pageNumber="68" phylum="Chordata" rank="genus">Bryophryne</taxonomicName>
|
||
, except for
|
||
<taxonomicName lsidName="B. flammiventris" pageId="3" pageNumber="68" rank="species" species="flammiventris">B. flammiventris</taxonomicName>
|
||
and
|
||
<taxonomicName lsidName="B. gymnotis" pageId="3" pageNumber="68" rank="species" species="gymnotis">B. gymnotis</taxonomicName>
|
||
), and geographic distribution within the Departamento Cusco, where all other species of
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne" order="Anura" pageId="3" pageNumber="68" phylum="Chordata" rank="genus">Bryophryne</taxonomicName>
|
||
occur.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="4" lastPageNumber="69" pageId="3" pageNumber="68" type="diagnosis">
|
||
<paragraph pageId="3" pageNumber="68">Diagnosis.</paragraph>
|
||
<paragraph lastPageId="4" lastPageNumber="69" pageId="3" pageNumber="68">
|
||
A new species of
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne" order="Anura" pageId="3" pageNumber="68" phylum="Chordata" rank="genus">Bryophryne</taxonomicName>
|
||
characterized by: (1) skin on dorsum shagreen; skin on venter areolate, discoidal fold absent, thoracic fold present; dorsolateral folds irregular and discontinuous; (2) tympanic membrane and tympanic annulus absent; (3) snout rounded in dorsal view and in profile; (4) upper eyelid with two small tubercles, narrower than IOD; cranial crests absent; (5) dentigerous process of vomers absent; (6) vocal sac and slits absent; nuptial pads absent; (7) Finger I much shorter than Finger II; tips of digits slightly pointed; (8) fingers lacking lateral fringes; (9) outer edge of forearm bearing small tubercles; (10) heel bearing minute tubercles; inner tarsal fold absent; outer edge of tarsus bearing small tubercles; (11) inner metatarsal tubercle prominent, ovoid, of similar relief and slightly larger than ovoid, outer metatarsal tubercle; supernumerary plantar tubercles indistinct; (12) toes lacking lateral fringes; webbing absent; toes III and V about equal in length; tips of digits slightly pointed; (13) in life, dorsum tan to green and brown with dark brown markings, greenish blue on lower flanks; some specimens with a yellow middorsal line extending from tip of snout to cloaca and to the posterior surface of thighs; interorbital bar present; chest, belly and ventral parts of forearms and legs dark brown with grayish blue
|
||
<pageBreakToken pageId="4" pageNumber="69" start="start">flecks</pageBreakToken>
|
||
; throat brown with flecks turning from gray-blue to copper near tip of mouth; palmar and plantar surfaces brown with lighter fingers and toes; (14) SVL 14.2-16.9 in males (n = 3), 22.2-22.6 in females (n = 2).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="5" lastPageNumber="70" pageId="4" pageNumber="69" type="comparisons">
|
||
<paragraph pageId="4" pageNumber="69">Comparisons.</paragraph>
|
||
<paragraph pageId="4" pageNumber="69">
|
||
The new species differs from other members of the genus by having green coloration on dorsum and blue coloration on flanks and ventral parts. Furthermore,
|
||
<taxonomicName lsidName="B. phuyuhampatu" pageId="4" pageNumber="69" rank="species" species="phuyuhampatu">B. phuyuhampatu</taxonomicName>
|
||
differs from other species by the following combination of characters (condition for comparing species in parenthesis): from
|
||
<taxonomicName lsidName="B. abramalage" pageId="4" pageNumber="69" rank="species" species="abramalage">B. abramalage</taxonomicName>
|
||
by having proportionally longer feet with FL/SVL from 0.41-0.45 (0.37-0.42), narrower head with HW/HL from 0.85-0.86 (0.97-1.07), and inner metatarsal tubercle larger than outer metatarsal (inner half the size of outer metatarsal tubercle);
|
||
<taxonomicName lsidName="B. flammiventris" pageId="4" pageNumber="69" rank="species" species="flammiventris">B. flammiventris</taxonomicName>
|
||
and
|
||
<taxonomicName lsidName="B. gymnotis" pageId="4" pageNumber="69" rank="species" species="gymnotis">B. gymnotis</taxonomicName>
|
||
by lacking a tympanum (present), from
|
||
<taxonomicName lsidName="B. bakersfield" pageId="4" pageNumber="69" rank="species" species="bakersfield">B. bakersfield</taxonomicName>
|
||
and
|
||
<taxonomicName lsidName="B. bustamantei" pageId="4" pageNumber="69" rank="species" species="bustamantei">B. bustamantei</taxonomicName>
|
||
by having discontinuous dorsolateral folds (continuous), from
|
||
<taxonomicName lsidName="B. cophites" pageId="4" pageNumber="69" rank="species" species="cophites">B. cophites</taxonomicName>
|
||
by females being much smaller (22.6 mm vs. 35.8 mm), from
|
||
<taxonomicName lsidName="B. hanssaueri" pageId="4" pageNumber="69" rank="species" species="hanssaueri">B. hanssaueri</taxonomicName>
|
||
by lacking bright orange coloration on throat (present), from
|
||
<taxonomicName lsidName="B. nubilosus" pageId="4" pageNumber="69" rank="species" species="nubilosus">B. nubilosus</taxonomicName>
|
||
by having toes III and V similar in length (toe V> III), and from
|
||
<taxonomicName lsidName="B. zonalis" pageId="4" pageNumber="69" rank="species" species="zonalis">B. zonalis</taxonomicName>
|
||
by having blue-gray mottled coloration on belly (distinctive black mottled coloration, variably confined to lower portion of belly). The new species further differs from
|
||
<taxonomicName lsidName="B. gymnotis" pageId="4" pageNumber="69" rank="species" species="gymnotis">B. gymnotis</taxonomicName>
|
||
by having vomers lacking dentigerous processes (present), from
|
||
<taxonomicName lsidName="B. cophites" pageId="4" pageNumber="69" rank="species" species="cophites">B. cophites</taxonomicName>
|
||
by males lacking nuptial pads (present), and from
|
||
<taxonomicName lsidName="B. bakersfield" pageId="4" pageNumber="69" rank="species" species="bakersfield">B. bakersfield</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="B. bustamantei" pageId="4" pageNumber="69" rank="species" species="bustamantei">B. bustamantei</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="B. flammiventris" pageId="4" pageNumber="69" rank="species" species="flammiventris">B. flammiventris</taxonomicName>
|
||
and
|
||
<taxonomicName lsidName="B. gymnotis" pageId="4" pageNumber="69" rank="species" species="gymnotis">B. gymnotis</taxonomicName>
|
||
for males lacking vocal slits (present).
|
||
</paragraph>
|
||
<paragraph lastPageId="5" lastPageNumber="70" pageId="4" pageNumber="69">
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne phuyuhampatu" order="Anura" pageId="4" pageNumber="69" phylum="Chordata" rank="species" species="phuyuhampatu">Bryophryne phuyuhampatu</taxonomicName>
|
||
(max. SVL 22.6 mm) is much smaller than
|
||
<taxonomicName lsidName="B. bakersfield" pageId="4" pageNumber="69" rank="species" species="bakersfield">B. bakersfield</taxonomicName>
|
||
(31.1 mm),
|
||
<taxonomicName lsidName="B. cophites" pageId="4" pageNumber="69" rank="species" species="cophites">B. cophites</taxonomicName>
|
||
(35.8 mm; pers. obs.),
|
||
<taxonomicName lsidName="B. hanssaueri" pageId="4" pageNumber="69" rank="species" species="hanssaueri">B. hanssaueri</taxonomicName>
|
||
(29.3 mm; pers. obs.), and
|
||
<taxonomicName lsidName="B. zonalis" pageId="5" pageNumber="70" rank="species" species="zonalis">
|
||
<pageBreakToken pageId="5" pageNumber="70" start="start">B</pageBreakToken>
|
||
. zonalis
|
||
</taxonomicName>
|
||
(32.4 mm), smaller than
|
||
<taxonomicName lsidName="B. bustamantei" pageId="5" pageNumber="70" rank="species" species="bustamantei">B. bustamantei</taxonomicName>
|
||
(23.4 mm),
|
||
<taxonomicName lsidName="B. flammiventris" pageId="5" pageNumber="70" rank="species" species="flammiventris">B. flammiventris</taxonomicName>
|
||
(24.1),
|
||
<taxonomicName lsidName="B. hanssaueri" pageId="5" pageNumber="70" rank="species" species="hanssaueri">B. hanssaueri</taxonomicName>
|
||
(24.6 mm), and about the same size of
|
||
<taxonomicName lsidName="B. abramalagae" pageId="5" pageNumber="70" rank="species" species="abramalagae">B. abramalagae</taxonomicName>
|
||
(20.1 mm) and
|
||
<taxonomicName lsidName="B. nubilosus" pageId="5" pageNumber="70" rank="species" species="nubilosus">B. nubilosus</taxonomicName>
|
||
(26.0 mm; pers. obs.). Five other small species of craugastorid frogs of the subfamily
|
||
<taxonomicName lsidName="" pageId="5" pageNumber="70" rank="subfamily" subfamily="Holoadeninae">Holoadeninae</taxonomicName>
|
||
are known to occur in montane forests and high Andean grasslands south of the Apurimac canyon in Peru:
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Noblella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Noblella madreselva" order="Anura" pageId="5" pageNumber="70" phylum="Chordata" rank="species" species="madreselva">Noblella madreselva</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="N. pygmaea" pageId="5" pageNumber="70" rank="species" species="pygmaea">N. pygmaea</taxonomicName>
|
||
,
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Psychrophrynella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Psychrophrynella bagrecito" order="Anura" pageId="5" pageNumber="70" phylum="Chordata" rank="species" species="bagrecito">Psychrophrynella bagrecito</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="P. chirihampatu" pageId="5" pageNumber="70" rank="species" species="chirihampatu">P. chirihampatu</taxonomicName>
|
||
and
|
||
<taxonomicName lsidName="P. usurpator" pageId="5" pageNumber="70" rank="species" species="usurpator">P. usurpator</taxonomicName>
|
||
, which all possess a visible tympanic annulus.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="5" pageNumber="70" type="description">
|
||
<paragraph pageId="5" pageNumber="70">Description of holotype.</paragraph>
|
||
<paragraph pageId="5" pageNumber="70">Adult male (16.9 mm SVL); head narrower than body, its length 33% of SVL; head wider than long, head length 83% of head width; head width 32% of SVL; snout short, rounded in dorsal and lateral views (Fig. 2), eye diameter 35% of head length, its diameter 1.2 times as large as its distance from the nostril; nostrils slightly protuberant, close to snout, directed dorsolaterally; canthus rostralis slightly straight in dorsal view, rounded in profile; loreal region slightly concave; lips rounded; upper eyelids with two small tubercles; upper eyelid width 46% of interorbital distance; interorbital region flat, lacking cranial crests; eye-nostril distance 81% of eye diameter; supratympanic fold short and weak; tympanic membrane and tympanic annulus absent; postrictal tubercles absent. Vocal sac and vocal slits absent. Choanae ovoid, small, positioned far anteriorly and laterally, widely separated from each other; dentigerous processes of vomer and vomerine teeth absent; tongue large, ovoid, about 2.5 times as long as wide, not notched posteriorly.</paragraph>
|
||
<paragraph pageId="5" pageNumber="70">Skin on dorsum shagreen with small, scattered tubercles; dorsolateral folds discontinuous, extending from posterior margin of upper eyelid to sacral region; skin on flanks tuberculate; skin on throat smooth, skin on chest, and belly areolate; thoracic fold present, discoidal fold absent; cloaca slightly protuberant, cloacal sheath short, cloacal region without tubercles. Outer surface of forearm with minute tubercles; palmar tubercle flat and oval, approximately same length but twice the width of elongate, thenar tubercle; few supernumerary tubercles low, ovoid; subarticular tubercles prominent, ovoid in ventral view, rounded in lateral view, largest at base of fingers; fingers lacking lateral fringes; Finger I much shorter than Finger II; relative lengths of fingers 3> 4 = 2> 1 (Fig. 3); tips of digits slightly pointed, lacking circumferential grooves (Fig. 3); forearm lacking tubercles.</paragraph>
|
||
<paragraph pageId="5" pageNumber="70">Hindlimbs short and robust, tibia length 38% of SVL; foot length 39% of SVL; upper surfaces of hindlimbs shagreen with scattered, minute tubercles; posterior surface of thighs tuberculate to areolate, ventral surface areolate; heel with minute tubercles (not visible in preservative); inner edge of tarsus without tubercles, outer edge of tarsus with small tubercles; inner metatarsal tubercle prominent, ovoid, of similar relief and slightly larger than ovoid, outer metatarsal tubercle; supernumerary plantar tubercles indistinct; subarticular tubercles low, ovoid in dorsal view; toes lacking lateral fringes, not webbed; toe tips weakly pointed, not expanded laterally, about as large as those on fingers; relative lengths of toes: 4> 3 = 5> 2> 1 (Fig. 3); foot length 32% of SVL.</paragraph>
|
||
<paragraph pageId="5" pageNumber="70">
|
||
Measurements of holotype (all in mm): SVL 16.9, TL 6.5, FL 6.6, HL 5.5, HW 5.4, ED 1.6, IOD 2.4, EW 1.1, IND 1.5,
|
||
<normalizedToken originalValue="E–N">E-N</normalizedToken>
|
||
1.3.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="5" pageNumber="70" type="coloration of holotype in life">
|
||
<paragraph pageId="5" pageNumber="70">Coloration of holotype in life.</paragraph>
|
||
<paragraph pageId="5" pageNumber="70">(Fig. 2). Dorsum green and brown with a dark brown marking extending from the interorbital bar to a mid-dorsal longitudinal band, a horizontal dark mark near the sacral region, and an oblique dark band on each flank. Dorsal surfaces of arms and legs dark brown, with transverse dark bars on forearms and hind limbs. Lower flanks with greenish blue flecks. Chest, belly and ventral parts of forearms and legs dark brown with grayish blue flecks. Iris grayish blue with a medial copper band. Throat brown with flecks turning from gray-blue to copper near the tip of the mouth. Palmar and plantar surfaces brown; tips of fingers and toes light brown to yellow.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="5" pageNumber="70" type="coloration of holotype in alcohol">
|
||
<paragraph pageId="5" pageNumber="70">Coloration of holotype in alcohol.</paragraph>
|
||
<paragraph pageId="5" pageNumber="70">(Fig. 2). Similar to coloration in life, but dorsal surfaces grayish tan with higher contrast of dorsal markings. Ventral surfaces beige to brown with cream flecks.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="5" pageNumber="70" type="variation">
|
||
<paragraph pageId="5" pageNumber="70">Variation.</paragraph>
|
||
<paragraph pageId="5" pageNumber="70">Coloration in life is based on field notes and photographs taken by A. Catenazzi of the paratopotypes (Fig. 4; photographs available through Calphoto database). The amount of dorsal green coloration varies among specimens. While juvenile MUBI 14665 and male MUBI 14655 are similar to the holotype in having a generally greenish dorsum, all other specimens have dark tan to brown dorsum, with just a few tubercles colored green. Female MUBI 14654, male CORBIDI 18225 and juvenile CORBIDI 18228 have a yellow middorsal line extending from the tip of the snout to the cloaca and to the posterior surface of the thighs.</paragraph>
|
||
<paragraph pageId="5" pageNumber="70">The summary of measurements of all types is reported in Table 2.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="5" pageNumber="70" type="etymology">
|
||
<paragraph pageId="5" pageNumber="70">Etymology.</paragraph>
|
||
<paragraph pageId="5" pageNumber="70">
|
||
The specific name
|
||
<taxonomicName lsidName="phuyuhampatu" pageId="5" pageNumber="70" rank="species" species="phuyuhampatu">phuyuhampatu</taxonomicName>
|
||
is a combination of Quechua words used in apposition meaning
|
||
<normalizedToken originalValue="“toad”">"toad"</normalizedToken>
|
||
(
|
||
<normalizedToken originalValue="“hampa'tu”">"hampa'tu"</normalizedToken>
|
||
) that lives in the
|
||
<normalizedToken originalValue="“fog”">"fog"</normalizedToken>
|
||
(
|
||
<normalizedToken originalValue="“phuyu”">"phuyu"</normalizedToken>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="6" lastPageNumber="71" pageId="5" pageNumber="70" type="distribution">
|
||
<paragraph pageId="5" pageNumber="70">Distribution, natural history, and threats.</paragraph>
|
||
<paragraph pageId="5" pageNumber="70">
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne phuyuhampatu" order="Anura" pageId="5" pageNumber="70" phylum="Chordata" rank="species" species="phuyuhampatu">Bryophryne phuyuhampatu</taxonomicName>
|
||
was discovered during a rapid amphibian survey in the upper Quispillomayo Valley (Fig. 5A) from 22 to 31 May 2016. The Quispillomayo torrent (Fig. 5B) is a tributary of the Nusiniscato River, which reaches the Araza River downstream of Quincemil, in the upper Madre de Dios drainage. During the inventory high-Andean grasslands (puna; 3350-4515 m a.s.l.), a forest patch of tasta (
|
||
<taxonomicName class="Magnoliopsida" family="Escalloniaceae" genus="Escallonia" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Escallonia myrtillioides" order="Escalloniales" pageId="5" pageNumber="70" phylum="Tracheophyta" rank="species" species="myrtillioides">Escallonia myrtillioides</taxonomicName>
|
||
), kishuar (
|
||
<taxonomicName class="Magnoliopsida" family="Scrophulariaceae" genus="Buddleja" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Buddleja incana" order="Lamiales" pageId="5" pageNumber="70" phylum="Tracheophyta" rank="species" species="incana">Buddleja incana</taxonomicName>
|
||
) and
|
||
<normalizedToken originalValue="qeñua">qenua</normalizedToken>
|
||
(
|
||
<taxonomicName class="Magnoliopsida" family="Rosaceae" genus="Polylepis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Polylepis incana" order="Rosales" pageId="5" pageNumber="70" phylum="Tracheophyta" rank="species" species="incana">Polylepis incana</taxonomicName>
|
||
) at 4280 m a.s.l., montane scrub, disturbed areas and other transitional formations along the treeline around 3350 m a.s.l., and the montane cloud forest from 2780-3350 m a.s.l. were sampled. Frogs were searched for under rocks, logs, mosses, and in the leaf litter and the understory in the montane forest. All but one specimens of
|
||
<taxonomicName lsidName="B. phuyuhampatu" pageId="5" pageNumber="70" rank="species" species="phuyuhampatu">B. phuyuhampatu</taxonomicName>
|
||
were found under mosses in the cloud forest around 2850 m a.s.l. (Fig. 5C). Male MUBI 14655 was found ~250 m from this site, under rocks and mosses under the riparian vegetation at the confluence of a small stream at 2795 m a.s.l. Two sympatric frogs,
|
||
<taxonomicName class="Amphibia" family="Hemiphractidae" genus="Gastrotheca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gastrotheca excubitor" order="Anura" pageId="5" pageNumber="70" phylum="Chordata" rank="species" species="excubitor">Gastrotheca cf. excubitor</taxonomicName>
|
||
and
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Psychrophrynella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Psychrophrynella chirihampatu" order="Anura" pageId="5" pageNumber="70" phylum="Chordata" rank="species" species="chirihampatu">Psychrophrynella chirihampatu</taxonomicName>
|
||
, were found under rocks in disturbed habitats (i.e., along streams, landslides) but not in the cloud forest. Two additional amphibian species,
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne" order="Anura" pageId="5" pageNumber="70" phylum="Chordata" rank="genus">Bryophryne</taxonomicName>
|
||
sp. and
|
||
<taxonomicName lsidName="B. zonalis" pageId="5" pageNumber="70" rank="species" species="zonalis">B. cf. zonalis</taxonomicName>
|
||
, were found along with
|
||
<taxonomicName lsidName="G. excubitor" pageId="5" pageNumber="70" rank="species" species="excubitor">G. cf. excubitor</taxonomicName>
|
||
in the grasslands from 3100-3650 m a.s.l.
|
||
</paragraph>
|
||
<caption pageId="5" pageNumber="70">
|
||
<paragraph pageId="5" pageNumber="70">
|
||
Figure 5. Collection localities of
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne phuyuhampatu" order="Anura" pageId="5" pageNumber="70" phylum="Chordata" rank="species" species="phuyuhampatu">Bryophryne phuyuhampatu</taxonomicName>
|
||
sp. n. in the upper Quispillomayo River valley A lookout from 3050 m a.s.l.: frogs were found under mosses, leaves and rocks in the cloud forest along the Quispillomayo River B such as the type locality at 2850 m a.s.l. C and disturbed forest at the confluence with a stream at 2795 m a.s.l. D Photographs by A. Catenazzi.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="5" pageNumber="70">
|
||
Both female paratopotypes had large eggs in their ovaries, indicative of terrestrial breeding and direct development: CORBIDI 18224 contained 15 eggs averaging 1.58
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
0.05 mm in diameter (range 1.20-1.80 mm), while MUBI 16654 contained 16 eggs averaging 2.44
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
0.03 mm in diameter (range 2.30-2.60 mm).
|
||
</paragraph>
|
||
<paragraph pageId="6" pageNumber="71">
|
||
<pageBreakToken pageId="6" pageNumber="71" start="start">The</pageBreakToken>
|
||
type locality (and known distribution range) of the new species lies within the
|
||
<normalizedToken originalValue="Área">Area</normalizedToken>
|
||
de
|
||
<normalizedToken originalValue="Conservación">Conservacion</normalizedToken>
|
||
Privada Ukumari Llaqta (
|
||
<bibRefCitation author="Catenazzi, A" journalOrPublisher="Peerj" pageId="8" pageNumber="73" title="A new species of Psychrophrynella (Amphibia, Anura, Craugastoridae) from the humid montane forests of Cusco, eastern slopes of the Peruvian Andes" volume="4" year="2016">Catenazzi and Ttito 2016</bibRefCitation>
|
||
), a protected area recognized by Peruvian environmental ministerial decree N° 301-2011-MINAM in December 2011. The upper puna and transitional habitats, as well as a narrow elevational band around the treeline are used for agriculture (potato cultivation), livestock (llamas grazing), fishing (exotic trout), and timber extraction. These land use patterns appear sustainable, and the grasslands at Patawasi (3350-3450 m a.s.l.) are in excellent conditions, with large bunchgrasses supporting large populations of
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne" order="Anura" pageId="6" pageNumber="71" phylum="Chordata" rank="genus">Bryophryne</taxonomicName>
|
||
sp. and
|
||
<taxonomicName lsidName="B. zonalis" pageId="6" pageNumber="71" rank="species" species="zonalis">B. cf. zonalis</taxonomicName>
|
||
. There is little indication of human disturbance in the cloud forest, and the main use seems to be limited to trout fishing.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |