459 lines
41 KiB
XML
459 lines
41 KiB
XML
<document ID-DOI="http://dx.doi.org/10.3897/zookeys.713.20776" ID-GBIF-Dataset="e0c76918-0bcc-4425-942f-bee1857e0268" ID-PMC="PMC5704321" ID-Pensoft-Pub="1313-2970-713-131" ID-PubMed="29187793" ID-ZBK="CC5733613D0643C68C4DEEAA14CCA3CD" ModsDocAuthor="" ModsDocDate="2017" ModsDocID="1313-2970-713-131" ModsDocOrigin="ZooKeys 713" ModsDocTitle="A new species of Phrynopus (Amphibia, Anura, Craugastoridae) from upper montane forests and high Andean grasslands of the Pui Pui Protected Forest in central Peru" checkinTime="1509722121965" checkinUser="pensoft" docAuthor="Lehr, Edgar, May, Rudolf von, Moravec, Jiri & Cusi, Juan Carlos" docDate="2017" docId="47B55233901CCAE31F1F788C1BE3C20D" docLanguage="en" docName="ZooKeys 713: 131-157" docOrigin="ZooKeys 713" docSource="http://dx.doi.org/10.3897/zookeys.713.20776" docTitle="Phrynopus inti Lehr, May, Moravec & Cusi, 2017, sp. n." docType="treatment" docUuid="C3E88CD6-7AD2-4CFE-8129-A6DB6D747F70" docUuidSource="ZooBank" docVersion="5" lastPageNumber="145" masterDocId="D05CFFF1BD7DFFD9726D1906987EF831" masterDocTitle="A new species of Phrynopus (Amphibia, Anura, Craugastoridae) from upper montane forests and high Andean grasslands of the Pui Pui Protected Forest in central Peru" masterLastPageNumber="157" masterPageNumber="131" pageNumber="137" updateTime="1668165089873" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>A new species of Phrynopus (Amphibia, Anura, Craugastoridae) from upper montane forests and high Andean grasslands of the Pui Pui Protected Forest in central Peru</mods:title>
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Lehr, Edgar</mods:namePart>
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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>May, Rudolf von</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>Moravec, Jiri</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>Cusi, Juan Carlos</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: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>713</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>131</mods:start>
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<mods:end>157</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.713.20776</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.713.20776</mods:identifier>
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<mods:identifier type="Pensoft-Pub">1313-2970-713-131</mods:identifier>
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<mods:identifier type="ZBK">CC5733613D0643C68C4DEEAA14CCA3CD</mods:identifier>
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<mods:identifier type="ZooBank">CC5733613D0643C68C4DEEAA14CCA3CD</mods:identifier>
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</mods:mods>
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<treatment ID-GBIF-Taxon="137233669" LSID="urn:lsid:zoobank.org:act:C3E88CD6-7AD2-4CFE-8129-A6DB6D747F70" httpUri="http://treatment.plazi.org/id/47B55233901CCAE31F1F788C1BE3C20D" lastPageId="14" lastPageNumber="145" pageId="6" pageNumber="137">
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<subSubSection pageId="6" pageNumber="137" type="nomenclature">
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<paragraph pageId="6" pageNumber="137">
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<taxonomicName LSID="http://zoobank.org/C3E88CD6-7AD2-4CFE-8129-A6DB6D747F70" class="Amphibia" family="Strabomantidae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="6" pageNumber="137" phylum="Chordata" rank="species" species="inti">
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<pageBreakToken pageId="6" pageNumber="137" start="start">Phrynopus</pageBreakToken>
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inti
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</taxonomicName>
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<taxonomicNameLabel pageId="6" pageNumber="137">sp. n.</taxonomicNameLabel>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="6" pageNumber="137" type="reference_group">
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<paragraph pageId="6" pageNumber="137">
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<taxonomicName class="Amphibia" family="Strabomantidae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus" order="Anura" pageId="6" pageNumber="137" phylum="Chordata" rank="genus">Phrynopus</taxonomicName>
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sp. A in Lehr, von May, Moravec, & Cusi (2017)
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</paragraph>
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</subSubSection>
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<subSubSection pageId="6" pageNumber="137" type="common name">
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<paragraph pageId="6" pageNumber="137">Common name.</paragraph>
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<paragraph pageId="6" pageNumber="137">English: Inti Andes Frog. Spanish: Rana Andina Inti.</paragraph>
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</subSubSection>
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<subSubSection pageId="6" pageNumber="137" type="holotype">
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<paragraph pageId="6" pageNumber="137">Holotype</paragraph>
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<paragraph pageId="6" pageNumber="137">
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(Figs 4A,B, 5, 6). MUSM 31183 (IWU 155), adult male from the buffer zone of the Pui Pui Protected Forest, Quebrada Tasta, forest patch near the house of Evaristo Borquez Quintana,
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<geoCoordinate direction="south" orientation="latitude" precision="1" value="-11.446889">11°26'48.8"S</geoCoordinate>
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, 74°54 2.8"W, 3609 m a.s.l. (Figs 2, 10C), Provincia Satipo, Region Junin, Peru, collected on 9 May 2012 by E. Lehr and R. von May.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="6" pageNumber="137" type="paratypes">
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<paragraph pageId="6" pageNumber="137">Paratypes</paragraph>
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<paragraph pageId="6" pageNumber="137">(Figs 7, 8, 9). A total of 15, all from Provincia Satipo, Region Junin, Peru (for detailed information see below): 5 males (MUSM 31976, 31984, 31203, NMP6V 75584, UMMZ 245220), 1 female (MUSM 31968), 9 juveniles (MUSM 31184, 31969, 31974, 31985, NMP6V 75585 87, UMMZ 245218, 245219).</paragraph>
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<paragraph pageId="6" pageNumber="137">
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MUSM 31184, UMMZ 245218, 245219, collected with the holotype on 9 May 2012 by E. Lehr and R. von May; MUSM 31203, near trail from Tasta to Tarhuish (first cumbre),
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<taxonomicName class="Diplopoda" family="Platyrhacidae" genus="Polylepis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Polylepis" order="Polydesmida" pageId="6" pageNumber="137" phylum="Arthropoda" rank="genus">Polylepis</taxonomicName>
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forest patch,
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<geoCoordinate direction="south" orientation="latitude" precision="1" value="-11.435722">11°26'8.6"S</geoCoordinate>
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,
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<geoCoordinate direction="west" orientation="longitude" precision="1" value="-74.899025">74°53'56.5"W</geoCoordinate>
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, 3886 m a.s.l. collected on 20 May 2012 by E. Lehr and R. von May; MUSM 31968, 31969, UMMZ 245220, Toldopampa,
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<geoCoordinate direction="south" orientation="latitude" precision="15" value="-11.504167">11°30'15"S</geoCoordinate>
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,
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<geoCoordinate direction="west" orientation="longitude" precision="15" value="-74.92583">74°55'33"W</geoCoordinate>
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, 3670 m a.s.l., collected on 22 June 2013 by E. Lehr, J. Moravec, and J.C. Cusi; NMP6V 75584, from Sector Carrizal, Satipo-Toldopampa Road at km 134 on left side of road coming from Satipo,
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<geoCoordinate direction="south" orientation="latitude" precision="1" value="-11.484305">11°29'03.5"S</geoCoordinate>
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,
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<geoCoordinate direction="west" orientation="longitude" precision="1" value="-74.890915">74°53'27.3"W</geoCoordinate>
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, 3350 m a.s.l., collected on 23 June 2013 by E. Lehr, J.C. Cusi, and J. Moravec; MUSM 31974, 31976, NMP6V 75585, Antuyo,
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<geoCoordinate direction="south" orientation="latitude" precision="1" value="-11.334361">11°20'03.7"S</geoCoordinate>
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,
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<geoCoordinate direction="west" orientation="longitude" precision="1" value="-74.99697">74°59'49.1"W</geoCoordinate>
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, 3700 m a.s.l., collected on 27 June 2013 by E. Lehr, J.C. Cusi, and J. Moravec; MUSM 31984, 31985, NMP6V 75586, 75587, Laguna Sinchon,
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<geoCoordinate direction="south" orientation="latitude" precision="1" value="-11.282306">11°16'56.3"S</geoCoordinate>
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,
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<geoCoordinate direction="west" orientation="longitude" precision="1" value="-75.05325">75°03'11.7"W</geoCoordinate>
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, 3890 m, collected on 30 June 2013 by E. Lehr, J.C. Cusi, and J. Moravec.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="7" pageNumber="138" type="generic placement">
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<paragraph pageId="7" pageNumber="138">
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<pageBreakToken pageId="7" pageNumber="138" start="start">Generic</pageBreakToken>
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placement.
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</paragraph>
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<paragraph pageId="7" pageNumber="138">
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We assign this species to
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus" order="Anura" pageId="7" pageNumber="138" phylum="Chordata" rank="genus">Phrynopus</taxonomicName>
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based on molecular evidence (Fig. 3).
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</paragraph>
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</subSubSection>
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<subSubSection pageId="7" pageNumber="138" type="diagnosis">
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<paragraph pageId="7" pageNumber="138">Diagnosis.</paragraph>
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<paragraph pageId="7" pageNumber="138">
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A species of
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus" order="Anura" pageId="7" pageNumber="138" phylum="Chordata" rank="genus">Phrynopus</taxonomicName>
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having the following combination of characters: (1) Skin on dorsum and flanks shagreen with scattered, low tubercles, more dense on dorsum; skin on venter smooth; discoidal fold absent, thoracic fold present; prominent supratympanic fold; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus absent; (3) snout rounded in dorsal and lateral views; (4) upper eyelid without enlarged tubercles; width of upper eyelid narrower than IOD; cranial crests absent; (5) dentigerous processes of vomers minute or absent; (6) vocal slits and nuptial pads absent; (7) Finger I shorter than Finger II; tips of digits bulbous, rounded; (8) fingers without lateral fringes; (9) ulnar and tarsal tubercles absent; (10) heel without tubercles; inner tarsal fold absent; (11) inner metatarsal tubercle rounded, about three times as large as ovoid outer metatarsal tubercle; supernumerary plantar tubercles absent; (12) toes without lateral fringes; basal webbing absent; Toe V slightly longer than Toe III; toe tips bulbous, rounded, about as large as those on fingers; (13) in life, dorsum pale grayish brown with or without dark brown blotches or blackish brown with small yellow flecks; throat, chest and venter pale grayish brown with salmon mottling, groin pale grayish brown with salmon colored flecks; iris golden orange with fine dark brown reticulations; (14) SVL 27.2 35.2 mm in males (n = 6), and 40.4 mm in single female.
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="8" lastPageNumber="139" pageId="7" pageNumber="138" type="comparisons">
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<paragraph pageId="7" pageNumber="138">Comparisons.</paragraph>
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<paragraph lastPageId="8" lastPageNumber="139" pageId="7" pageNumber="138">
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="7" pageNumber="138" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
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sp. n. is readily distinguished from its 34 congeners in Peru (
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<bibRefCitation pageId="16" pageNumber="147" url="http://amphibiaweb.org/" year="2017">AmphibiaWeb 2017</bibRefCitation>
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), by its relatively large SVL (except for
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<taxonomicName lsidName="P. juninensis" pageId="7" pageNumber="138" rank="species" species="juninensis">P. juninensis</taxonomicName>
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and
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<taxonomicName lsidName="P. kauneorum" pageId="7" pageNumber="138" rank="species" species="kauneorum">P. kauneorum</taxonomicName>
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) of up to 40.4 mm, by having the groin pale grayish brown with salmon
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<pageBreakToken pageId="8" pageNumber="139" start="start">colored</pageBreakToken>
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flecks, the venter pale grayish brown with salmon mottling and the iris golden orange with fine dark brown reticulations.
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="8" pageNumber="139" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
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sp. n. is most similar to the large central Peruvian species
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<taxonomicName lsidName="P. juninensis" pageId="8" pageNumber="139" rank="species" species="juninensis">P. juninensis</taxonomicName>
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(SVL up to 43.1 mm,
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<bibRefCitation author="Duellman, WE" journalOrPublisher="Natur und Tier-Verlag, Munster" pageId="16" pageNumber="147" title="Terrestrial-breeding Frogs (Strabomantidae) in Peru." year="2009">Duellman and Lehr 2009</bibRefCitation>
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) and
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<taxonomicName lsidName="P. kauneorum" pageId="8" pageNumber="139" rank="species" species="kauneorum">P. kauneorum</taxonomicName>
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(SVL up to 56.4 mm,
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<bibRefCitation author="Lehr, E" journalOrPublisher="Zoologische Abhandlungen Museum fur Tierkunde Dresden" pageId="17" pageNumber="148" pagination="71 - 75" title="Morphological and ecological remarks on Phynopuskauneorum (Amphibia, Anura, Leptodactylidae)." volume="52" year="2002 b">Lehr et al. 2002b</bibRefCitation>
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), Fig. 4. All three species share a gray ground coloration and dark brown canthal and supratympanic stripes, lack dorsolateral folds and males lack vocal slits and nuptial pads, but can be distinguished as follows:
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="8" pageNumber="139" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
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sp. n. has weak postocular folds (absent in both
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<taxonomicName lsidName="P. juninensis" pageId="8" pageNumber="139" rank="species" species="juninensis">P. juninensis</taxonomicName>
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and
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<taxonomicName lsidName="P. kauneorum" pageId="8" pageNumber="139" rank="species" species="kauneorum">P. kauneorum</taxonomicName>
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), has dentigerous processes of vomers (absent in
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<taxonomicName lsidName="P. juninensis" pageId="8" pageNumber="139" rank="species" species="juninensis">P. juninensis</taxonomicName>
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, present in
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<taxonomicName lsidName="P. kauneorum" pageId="8" pageNumber="139" rank="species" species="kauneorum">P. kauneorum</taxonomicName>
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), skin on dorsum shagreen with scattered, low tubercles (smooth to weakly areolate in
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<taxonomicName lsidName="P. juninensis" pageId="8" pageNumber="139" rank="species" species="juninensis">P. juninensis</taxonomicName>
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, smooth in
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<taxonomicName lsidName="P. kauneorum" pageId="8" pageNumber="139" rank="species" species="kauneorum">P. kauneorum</taxonomicName>
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), skin on venter smooth (areolate in
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<taxonomicName lsidName="P. juninensis" pageId="8" pageNumber="139" rank="species" species="juninensis">P. juninensis</taxonomicName>
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, smooth in
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<taxonomicName lsidName="P. kauneorum" pageId="8" pageNumber="139" rank="species" species="kauneorum">P. kauneorum</taxonomicName>
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), dorsum pale grayish brown with or without dark brown blotches or blackish brown with small yellow flecks (dorsum grayish brown with dark brown markings in
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<taxonomicName lsidName="P. juninensis" pageId="8" pageNumber="139" rank="species" species="juninensis">P. juninensis</taxonomicName>
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, dorsum pale brown to tan with dark brown markings in
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<taxonomicName lsidName="P. kauneorum" pageId="8" pageNumber="139" rank="species" species="kauneorum">P. kauneorum</taxonomicName>
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), venter pale grayish brown with salmon mottling (pale brown with gray blotches in
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<taxonomicName lsidName="P. juninensis" pageId="8" pageNumber="139" rank="species" species="juninensis">P. juninensis</taxonomicName>
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, pinkish to grayish tan in
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<taxonomicName lsidName="P. kauneorum" pageId="8" pageNumber="139" rank="species" species="kauneorum">P. kauneorum</taxonomicName>
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), and the iris is golden orange (copper in
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<taxonomicName lsidName="P. juninensis" pageId="8" pageNumber="139" rank="species" species="juninensis">P. juninensis</taxonomicName>
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, dark brown in
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<taxonomicName lsidName="P. kauneorum" pageId="8" pageNumber="139" rank="species" species="kauneorum">P. kauneorum</taxonomicName>
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).
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</paragraph>
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<caption pageId="8" pageNumber="139">
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<paragraph pageId="8" pageNumber="139">
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Figure 4.
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="8" pageNumber="139" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
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sp. n. (A, B holotype, MUSM 31183, male, SVL 32.5 mm),
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<taxonomicName lsidName="P. juninensis" pageId="8" pageNumber="139" rank="species" species="juninensis">P. juninensis</taxonomicName>
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(C, DMUSM 33258, female, SVL 33.0 mm),
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<taxonomicName lsidName="P. kauneorum" pageId="8" pageNumber="139" rank="species" species="kauneorum">P. kauneorum</taxonomicName>
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(E, F holotype, MUSM 20459, female, SVL 29.1 mm) in dorsolateral and ventral views. Photos by E. Lehr and R. von May (C, D).
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</paragraph>
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</caption>
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</subSubSection>
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<subSubSection lastPageId="10" lastPageNumber="141" pageId="8" pageNumber="139" type="description">
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<paragraph pageId="8" pageNumber="139">Description of the holotype.</paragraph>
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<paragraph pageId="8" pageNumber="139">Head as wide as body, wider than long, HW 110% of HL; HW 38% of SVL; HL 35% of SVL; snout short, rounded in dorsal and lateral views (Figs 5A, B), ED larger than E N distance (ED 148% of E N); nostrils protuberant, directed dorsolaterally; canthus rostralis slightly curved in dorsal view, rounded in profile; loreal region slightly concave; lips rounded; upper eyelid without enlarged tubercles; EW slightly narrower than IOD (EW 94% of IOD); postocular folds low, extending from posterior margin of upper eyelid to level of upper arm insertion (Fig. 5B); supratympanic fold broad, extending from posterior corner of eye to level of upper arm insertion; tympanic membrane and tympanic annulus absent, tympanic region without postrictal tubercles. Choanae small, ovoid, close to but not concealed by palatal shelf of maxilla; dentigerous processes of vomers minute, embedded in mucosa of mouth, widely separated; tongue broad, about twice as long as wide, not notched posteriorly, posterior half free; vocal slits absent.</paragraph>
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<caption pageId="8" pageNumber="139">
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<paragraph pageId="8" pageNumber="139">
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Figure 5. Life male holotype (MUSM 31183, SVL 32.5 mm) of
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="8" pageNumber="139" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
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sp. n. in dorsolateral view (A), dorsal view (B), flanks, groin, anterior surfaces of thighs (C), and ventral view (D). Photos by E. Lehr.
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</paragraph>
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</caption>
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<paragraph pageId="8" pageNumber="139">Skin on dorsum shagreen with scattered, low tubercles, more dense on posterior half of body, dorsolateral folds absent (Fig. 5B); skin on flanks shagreen with few scattered, low tubercles; skin on throat, chest and belly smooth (Fig. 5D); discoidal fold absent, thoracic fold present; cloacal sheath not distinct; cloacal region without tubercles. Outer surface of forearm without tubercles; outer palmar tubercle barely visible, low, ovoid, slightly smaller than ovoid inner palmar tubercle; supernumerary tubercles absent; subarticular tubercles low, ovoid, most prominent on base of fingers; fingers without lateral fringes; Finger I shorter than Finger II; tips of digits rounded, bulbous, lacking circumferential grooves; nuptial pads absent (Fig. 6A).</paragraph>
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<caption pageId="8" pageNumber="139">
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<paragraph pageId="8" pageNumber="139">
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Figure 6. Ventral views of right hand (A) and right foot (B) of holotype of
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="8" pageNumber="139" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
|
||
sp. n. (MUSM 31183). Drawings by E. Lehr.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph lastPageId="9" lastPageNumber="140" pageId="8" pageNumber="139">
|
||
Hind limbs long and slender, TL 39% of SVL; FL 43% of SVL; dorsal surface of hind limbs shagreen with few low tubercles; anterior surfaces of thighs shagreen, posterior surfaces of thighs weakly areolate; heel without a conical tubercle; outer surface of tarsus without tubercles; outer metatarsal tubercle rounded, weakly conical, about four times as large as prominent ovoid inner metatarsal tubercle; supernumerary plantar
|
||
<pageBreakToken pageId="9" pageNumber="140" start="start">tubercles</pageBreakToken>
|
||
absent; subarticular tubercles low, ovoid in dorsal view, most distinct on base of toes; toes without lateral fringes; basal webbing absent; toe tips bulbous, rounded, lacking circumferential grooves, about as large as those on fingers; relative lengths of toes: 1 <2 <3 <5 <4; Toe V slightly longer than Toe III (Fig. 6B).
|
||
</paragraph>
|
||
<paragraph pageId="9" pageNumber="140">Measurements of the holotype (in mm).SVL 32.5; tibia length 12.7; foot length 14.0; head length 11.3; head width 12.5; eye diameter 3.4; interorbital distance 3.5; upper eyelid width 3.3; internarial distance 2.9; eye-nostril distance 2.3.</paragraph>
|
||
<paragraph pageId="9" pageNumber="140">Coloration of the holotype in life (Fig. 5). Dorsum pale grayish brown with dark brown blotches, a dark brown X-shaped marking on shoulder region and an irregular shaped dark brown interorbital blotch. Flanks paler than dorsum with few pale brown flecks. Canthal and supratympanic stripes dark brown. Upper lip with few pale brown flecks. Arms and legs dorsally with few pale and dark brown blotches and flecks. Throat, chest and venter pale grayish brown with salmon mottling, denser on posterior half of belly and thighs. Groin, posterior surfaces of thighs, posterior surfaces of tibias and dorsal surfaces of feet vibrant salmon colored. Iris golden orange with fine dark brown reticulations.</paragraph>
|
||
<paragraph pageId="10" pageNumber="141">
|
||
<pageBreakToken pageId="10" pageNumber="141" start="start">Coloration</pageBreakToken>
|
||
of the holotype in preservative. Dorsum tan with dark brown blotches and dark brown X-shaped marking on shoulder region and an irregular shaped dark brown interorbital blotch. Flanks paler than dorsum, with few pale brown flecks. Canthal and supratympanic stripes dark brown. Upper lip with few pale brown flecks. Arms and legs dorsally tan with few pale and dark brown blotches and flecks. Groin creamy white. Throat, chest and venter creamy white and pale gray mottled. Ventral surfaces of hand and feet creamy white. Iris pale gray.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="12" lastPageNumber="143" pageId="10" pageNumber="141" type="variation">
|
||
<paragraph pageId="10" pageNumber="141">Variation.</paragraph>
|
||
<paragraph lastPageId="12" lastPageNumber="143" pageId="10" pageNumber="141">
|
||
All paratypes (Figs 7 9) are similar to the holotype regarding morphology and proportions (Tables 3, 4). Besides differences in SVL (Tables 3, 4), coloration variation in life is notable. Three males (MUSM 31203, UMMZ 245220 (Fig. 7A C),
|
||
<pageBreakToken pageId="11" pageNumber="142" start="start">285</pageBreakToken>
|
||
) are similar to the holotype in coloration except for having much less salmon coloration. One male (MUSM 31976, Fig. 7D F) has the dorsum uniformly grayish brown without dark brown blotches. One male (MUSM 31984, Fig. 7G I) has the dorsum blackish brown with small yellow flecks. The single female (MSUM 31968, Fig. 8) is similar in coloration to the holotype except for only having few small flecks of salmon
|
||
<pageBreakToken pageId="12" pageNumber="143" start="start">in</pageBreakToken>
|
||
groin, and ventrally on thighs and shanks. The dorsal coloration of the juveniles (Fig. 9) is similar to the adults (dorsum pale grayish brown with dark brown blotches in MUSM 31969 [Fig. 9A, B], 31974, NMP6V 75585, blackish brown with small yellow flecks in NMP6V 75586, 75587 [Fig. 9D, E], uniformly blackish brown in MUSM 31985). The ventral coloration is different in juveniles. One juvenile (MSUM 31969, Fig. 9C) has the venter reddish brown, three juveniles (MUSM 31974, NMP6V 75585, 75587 (Fig. 9F)) have the venter reddish brown and tan mottled.
|
||
</paragraph>
|
||
<caption pageId="12" pageNumber="143">
|
||
<paragraph pageId="12" pageNumber="143">
|
||
Table 3. Measurements (in mm) of adult type specimens of
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="12" pageNumber="143" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
|
||
sp. n. M = male, F = female. For other abbreviations see materials and methods.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="12" pageNumber="143">
|
||
<table pageId="12" pageNumber="143">
|
||
<tr pageId="12" pageNumber="143">
|
||
<th colspan="1" pageId="12" pageNumber="143" rowspan="2">Characters</th>
|
||
<th colspan="1" pageId="12" pageNumber="143" rowspan="1">
|
||
<abbrev content-type="institution" id="ABBRID0EBOCK" pageId="12" pageNumber="143">MUSM</abbrev>
|
||
</th>
|
||
<th colspan="1" pageId="12" pageNumber="143" rowspan="1">
|
||
<named-content content-type="dwc:institutional_code" pageId="12" pageNumber="143">UMMZ</named-content>
|
||
</th>
|
||
<th colspan="1" pageId="12" pageNumber="143" rowspan="1">
|
||
<abbrev content-type="institution" id="ABBRID0EROCK" pageId="12" pageNumber="143">MUSM</abbrev>
|
||
</th>
|
||
<th colspan="1" pageId="12" pageNumber="143" rowspan="1">
|
||
<abbrev content-type="institution" id="ABBRID0EZOCK" pageId="12" pageNumber="143">NMP6V</abbrev>
|
||
</th>
|
||
<th colspan="1" pageId="12" pageNumber="143" rowspan="1">
|
||
<abbrev content-type="institution" id="ABBRID0EBPCK" pageId="12" pageNumber="143">MUSM</abbrev>
|
||
</th>
|
||
<th colspan="1" pageId="12" pageNumber="143" rowspan="1">
|
||
<abbrev content-type="institution" id="ABBRID0EJPCK" pageId="12" pageNumber="143">MUSM</abbrev>
|
||
</th>
|
||
<th colspan="1" pageId="12" pageNumber="143" rowspan="1">
|
||
<abbrev content-type="institution" id="ABBRID0ERPCK" pageId="12" pageNumber="143">MUSM</abbrev>
|
||
</th>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<th colspan="1" pageId="12" pageNumber="143" rowspan="1">31203</th>
|
||
<th colspan="1" pageId="12" pageNumber="143" rowspan="1">245220</th>
|
||
<th colspan="1" pageId="12" pageNumber="143" rowspan="1">31183</th>
|
||
<th colspan="1" pageId="12" pageNumber="143" rowspan="1">75584</th>
|
||
<th colspan="1" pageId="12" pageNumber="143" rowspan="1">31984</th>
|
||
<th colspan="1" pageId="12" pageNumber="143" rowspan="1">31976</th>
|
||
<th colspan="1" pageId="12" pageNumber="143" rowspan="1">31968</th>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">SVL</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">TL</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">FL</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">HL</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">HW</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">ED</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">IOD</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">EW</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">IND</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<caption ID-Table-UUID="D85C471B78A90A01DD40046C5AE35560" httpUri="http://table.plazi.org/id/D85C471B78A90A01DD40046C5AE35560" pageId="12" pageNumber="143">
|
||
<paragraph pageId="12" pageNumber="143">
|
||
Table 4. Measurements (in mm) and proportions of male type specimens of
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="12" pageNumber="143" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
|
||
sp. n.; ranges followed by means and one standard deviation in parentheses. For abbreviations see materials and methods.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="12" pageNumber="143">
|
||
<table pageId="12" pageNumber="143">
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="12" pageNumber="143" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
|
||
</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">SVL</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">TL</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">FL</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">HL</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">HW</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">ED</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">IOD</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">EW</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">IND</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">TLSVL</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">FLSVL</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">HLSVL</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">HWSVL</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">HWHL</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">ED</td>
|
||
</tr>
|
||
<tr pageId="12" pageNumber="143">
|
||
<td colspan="1" pageId="12" pageNumber="143" rowspan="1">EWIOD</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<caption pageId="12" pageNumber="143">
|
||
<paragraph pageId="12" pageNumber="143">
|
||
Figure 7. Variation of male paratypes of
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="12" pageNumber="143" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
|
||
sp. n. in dorsolateral, dorsal, and ventral views. A C (UMMZ 245220, SVL 27.4 mm), D F (MUSM 31976, SVL 35.2 mm), G I (MSUM 31984, SVL 35.1 mm). Photos by E. Lehr, and by J.C. Cusi (E).
|
||
</paragraph>
|
||
</caption>
|
||
<caption pageId="12" pageNumber="143">
|
||
<paragraph pageId="12" pageNumber="143">
|
||
Figure 8. Female paratype of
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="12" pageNumber="143" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
|
||
sp. n. (MUSM 31968, SVL 40.4 mm) in dorsolateral (A), dorsal (B), and ventral views (C). Photos by E. Lehr and J. Moravec (A).
|
||
</paragraph>
|
||
</caption>
|
||
<caption pageId="12" pageNumber="143">
|
||
<paragraph pageId="12" pageNumber="143">
|
||
Figure 9. Variation of juvenile paratypes of
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="12" pageNumber="143" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
|
||
sp. n. in dorsolateral, dorsal, and ventral views. A C (MUSM 31969, SVL 16.0 mm), D F (NMP6V 75587, SVL 20.3 mm). Photos by E. Lehr.
|
||
</paragraph>
|
||
</caption>
|
||
</subSubSection>
|
||
<subSubSection pageId="12" pageNumber="143" type="etymology">
|
||
<paragraph pageId="12" pageNumber="143">Etymology.</paragraph>
|
||
<paragraph pageId="12" pageNumber="143">
|
||
The species epithet
|
||
<taxonomicName lsidName="inti" pageId="12" pageNumber="143" rank="species" species="inti">inti</taxonomicName>
|
||
is derived from the Quechuan noun Inti, the Incan sun god. The golden-orange iris reminds us of the sun.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="14" lastPageNumber="145" pageId="12" pageNumber="143" type="distribution">
|
||
<paragraph pageId="12" pageNumber="143">Distribution, natural history, and threat status.</paragraph>
|
||
<paragraph pageId="12" pageNumber="143">
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="12" pageNumber="143" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
|
||
sp. n. is known from four localities outside and two localities inside the Pui Pui Protected Forest between 3350 and 3890 m a.s.l., covering an estimated area of 101.3 km2 (Figs 1, 2).
|
||
</paragraph>
|
||
<paragraph lastPageId="14" lastPageNumber="145" pageId="12" pageNumber="143">
|
||
The type locality, Quebrada Tasta (Fig. 2), is outside the PPPF. The holotype and three paratypes (MUSM 31184, UMMZ 245218, 245219) were found in the afternoon in a forest patch under rocks near the house of Evaristo Borquez Quintana, on 9 May
|
||
<pageBreakToken pageId="13" pageNumber="144" start="start">2012</pageBreakToken>
|
||
at 3609 m a.s.l. (Fig. 10C). The vegetation at the type locality consists of
|
||
<taxonomicName class="Diplopoda" family="Platyrhacidae" genus="Polylepis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Polylepis" order="Polydesmida" pageId="13" pageNumber="144" phylum="Arthropoda" rank="genus">Polylepis</taxonomicName>
|
||
trees, small bushes, ferns, moss, and Peruvian feather grass. No sympatric anurans were recorded. One specimen (MUSM 31203) was found in the afternoon under moss in a
|
||
<taxonomicName class="Diplopoda" family="Platyrhacidae" genus="Polylepis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Polylepis" order="Polydesmida" pageId="13" pageNumber="144" phylum="Arthropoda" rank="genus">Polylepis</taxonomicName>
|
||
forest patch near the trail from Tasta to Tarhuish at 3886 m a.s.l. Three specimens (MUSM 31968, 31969, UMMZ 245220) were collected in the morning under rocks and in moss in the mountain slopes of the Toldopampa valley close to Toldopampa
|
||
<pageBreakToken pageId="14" pageNumber="145" start="start">at</pageBreakToken>
|
||
3670 m a.s.l. (Fig. 10B). Specimens were found under rocks and in moss. Sympatric anurans include
|
||
<taxonomicName class="Amphibia" family="Hemiphractidae" genus="Gastrotheca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gastrotheca griswoldi" order="Anura" pageId="14" pageNumber="145" phylum="Chordata" rank="species" species="griswoldi">Gastrotheca griswoldi</taxonomicName>
|
||
Shreve, 1941. One specimen (NMP6V 75584) was collected under moss in the early afternoon at the Satipo-Toldopampa Road at km 134 on the left side of the road coming from Satipo at 3350 m a.s.l. (Fig. 10A). Sympatric anurans here include
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis bounides" order="Anura" pageId="14" pageNumber="145" phylum="Chordata" rank="species" species="bounides">Pristimantis bounides</taxonomicName>
|
||
(MUSM 31970, 31971) and
|
||
<taxonomicName class="Amphibia" family="Hemiphractidae" genus="Gastrotheca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gastrotheca griswoldi" order="Anura" pageId="14" pageNumber="145" phylum="Chordata" rank="species" species="griswoldi">Gastrotheca griswoldi</taxonomicName>
|
||
(MUSM 31972). Three specimens (MUSM 31974, 31976, NMP6V 75585) were found under rocks and in moss in Antuyo at 3700 m a.s.l. (Fig. 10D). Sympatric anurans here include
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="14" pageNumber="145" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
|
||
(MUSM 31975) and
|
||
<taxonomicName class="Amphibia" family="Hemiphractidae" genus="Gastrotheca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gastrotheca griswoldi" order="Anura" pageId="14" pageNumber="145" phylum="Chordata" rank="species" species="griswoldi">Gastrotheca griswoldi</taxonomicName>
|
||
(IWU 290). Four specimens (MUSM 31984, 31985, NMP6V 75586, 75587) were found in the puna in the afternoon in moss close to the Laguna Sinchon at 3890 m a.s.l. (Fig. 10E). Sympatric anurans here include
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis puipui" order="Anura" pageId="14" pageNumber="145" phylum="Chordata" rank="species" species="puipui">Pristimantis puipui</taxonomicName>
|
||
(MSUM 31981 83).
|
||
</paragraph>
|
||
<caption pageId="14" pageNumber="145">
|
||
<paragraph pageId="14" pageNumber="145">
|
||
Figure 10. Type locality and habitats of
|
||
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="14" pageNumber="145" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
|
||
sp. n. Satipo-Toldopampa Road at km 134 on left side of street coming from Satipo, 3350 m a.s.l., 23 June 2013 (A); Quebrada Toldopampa, 3670 m a.s.l., 22 June 2013 (B); Type locality, Quebrada Tasta, 3609 m a.s.l., 20 May 2012 (C); Antuyo, PPPF, 3700 m a.s.l., 27 June 2013 (D); Laguna Sinchon, PPPF, 3890 m a.s.l., 29 June 2013 (E). Photos by E. Lehr.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="14" pageNumber="145">
|
||
One male specimen (MUSM 31203) had as ectoparasites five trombiculid mites on the right side in the area of the upper arm insertion. Such parasites are not uncommon in Andean frogs (e.g.,
|
||
<bibRefCitation author="Quinzio, S" journalOrPublisher="Salamandra" pageId="18" pageNumber="149" pagination="263 - 268" title="Intradermal infections by chigger mites (Hannemania spp.) in the Andean frog Telmatobiusatacamensis (Anura, Telmatobiidae)." volume="51" year="2015">Quinzio and Goldberg 2015</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Lehr, E" journalOrPublisher="Zootaxa" pageId="17" pageNumber="148" pagination="301 - 336" title="Three new species of Pristimantis (Amphibia, Anura, Craugastoridae) from upper montane forests and high Andean grasslands of the Pui Pui Protected Forest in central Peru." url="https://doi.org/10.11646/zootaxa.4299.3.1" volume="4299" year="2017">Lehr et al. 2017</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph pageId="14" pageNumber="145">
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The IUCN Red List criteria (
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<bibRefCitation author="Heinicke, MP" journalOrPublisher="Proceedings of the National Academy of Sciences, USA" pageId="17" pageNumber="148" url="http://www.iucnredlist" year="2001">IUCN 2001</bibRefCitation>
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) consider that if a species occurs in fewer than 10 threat-defined locations and the extent of occurrence (EOO) is <20,000 km2, it should be classified as Vulnerable or Endangered.
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="14" pageNumber="145" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
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sp. n. is known from six localities distributed in the PPPF and its buffer zone (Fig. 10), with an estimated EOO of 101.3 km2. As such, this new species might be classified as Vulnerable if we take into account these criteria. However, given that the PPPF may host a greater number of locations (two of them are inside the protected area), we propose that
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="14" pageNumber="145" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
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sp. n. should likely be categorized as Near Threatened (NT). Despite that two locations of the known distribution of
|
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<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus inti" order="Anura" pageId="14" pageNumber="145" phylum="Chordata" rank="species" species="inti">Phrynopus inti</taxonomicName>
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sp. n. are within the PPPF (Fig. 10) and formally protected, other factors such as fungal infections, climate change, pollution, and man-made fires (used to expand grazing areas for livestock) continue to be threats for many Andean amphibians even inside protected areas (
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<bibRefCitation author="Catenazzi, A" editor="Heatwole, H" journalOrPublisher="Herpetological Monographs" pageId="16" pageNumber="147" pagination="1 - 23" title="Conservation status of amphibians in Peru." url="https://doi.org/10.1655/HERPMONOGRAPHS-D-13-00003" volume="28" volumeTitle="Status of Conservation and Decline of Amphibians: Western Hemisphere (Volume 9) Amphibian Biology." year="2014">Catenazzi and von May 2014</bibRefCitation>
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). Agriculture and cattle raising are more acute in the Toldopampa valley than in the Tasta valley.
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</paragraph>
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</subSubSection>
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</treatment>
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</document> |