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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.660.11394" ID-GBIF-Dataset="a69b75d2-80b6-4cf9-98a8-87a53aa74b35" ID-PMC="PMC5549528" ID-Pensoft-Pub="1313-2970-660-17" ID-PubMed="28794672" ID-ZBK="CB98BAD25B8D43D9AAE86077314E6E94" ModsDocAuthor="" ModsDocDate="2017" ModsDocID="1313-2970-660-17" ModsDocOrigin="ZooKeys 660" ModsDocTitle="A new species of terrestrial-breeding frog (Amphibia, Craugastoridae, Pristimantis) from high elevations of the Pui Pui Protected Forest in central Peru" checkinTime="1488970275490" checkinUser="pensoft" docAuthor="Lehr, Edgar &amp; May, Rudolf von" docDate="2017" docId="1E6994BAA61313A700E882C5628D0E18" docLanguage="en" docName="ZooKeys 660: 17-42" docOrigin="ZooKeys 660" docSource="http://dx.doi.org/10.3897/zookeys.660.11394" docTitle="Pristimantis attenboroughi Lehr &amp; May, 2017, sp. n." docType="treatment" docUuid="DCE88D49-0EB1-4DA4-A672-5341763B3236" docUuidSource="ZooBank" docVersion="5" lastPageNumber="31" masterDocId="FF93ED493419FFF4FF9A9042FF87FFDB" masterDocTitle="A new species of terrestrial-breeding frog (Amphibia, Craugastoridae, Pristimantis) from high elevations of the Pui Pui Protected Forest in central Peru" masterLastPageNumber="42" masterPageNumber="17" pageNumber="21" updateTime="1668164281347" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>A new species of terrestrial-breeding frog (Amphibia, Craugastoridae, Pristimantis) from high elevations of the Pui Pui Protected Forest in central Peru</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Lehr, Edgar</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>May, Rudolf von</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:title>ZooKeys</mods:title>
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<mods:date>2017</mods:date>
<mods:detail type="volume">
<mods:number>660</mods:number>
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<mods:start>17</mods:start>
<mods:end>42</mods:end>
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<mods:url>http://dx.doi.org/10.3897/zookeys.660.11394</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.660.11394</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-660-17</mods:identifier>
<mods:identifier type="ZBK">CB98BAD25B8D43D9AAE86077314E6E94</mods:identifier>
<mods:identifier type="ZooBank">CB98BAD25B8D43D9AAE86077314E6E94</mods:identifier>
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<treatment ID-GBIF-Taxon="127886945" LSID="urn:lsid:zoobank.org:act:DCE88D49-0EB1-4DA4-A672-5341763B3236" httpUri="http://treatment.plazi.org/id/1E6994BAA61313A700E882C5628D0E18" lastPageId="14" lastPageNumber="31" pageId="4" pageNumber="21">
<subSubSection pageId="4" pageNumber="21" type="nomenclature">
<paragraph pageId="4" pageNumber="21">
<taxonomicName LSID="http://zoobank.org/DCE88D49-0EB1-4DA4-A672-5341763B3236" class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="4" pageNumber="21" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
<taxonomicNameLabel pageId="4" pageNumber="21">sp. n.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection lastPageId="5" lastPageNumber="22" pageId="4" pageNumber="21" type="common name">
<paragraph pageId="4" pageNumber="21">Common name.</paragraph>
<paragraph pageId="4" pageNumber="21">
English:
<normalizedToken originalValue="Attenboroughs">Attenborough's</normalizedToken>
Rubber Frog. Spanish: Rana
<normalizedToken originalValue="cutín">cutin</normalizedToken>
Attenborough.
</paragraph>
<paragraph pageId="4" pageNumber="21">
Holotype. MUSM 31196 (IWU 178, Figs 3, 4), adult male from the Pui Pui Protected Forest, Provincia Satipo,
<normalizedToken originalValue="Región">Region</normalizedToken>
<normalizedToken originalValue="Junín">Junin</normalizedToken>
, Peru, Upper part of Quebrada Tarhuish, &quot;Laguna Udrecocha&quot;, Puna, open area on east side of Laguna Udrecocha,
<geoCoordinate direction="south" orientation="latitude" precision="1" value="-11.390028">11°23'24.1&quot;S</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-74.97569">74°58'32.5&quot;W</geoCoordinate>
, 3936 m a.s.l. (Fig. 8A), collected on 17 May 2012 by E. Lehr and R. von May.
</paragraph>
<caption pageId="4" pageNumber="21">
<paragraph pageId="4" pageNumber="21">
Figure 3. Life male holotype (MUSM 31196, SVL 18.9 mm) of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="4" pageNumber="21" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
sp. n. in dorsolateral view (A), dorsal view (B), flanks, groin, anterior surfaces of thighs (C), posterior surfaces of thighs (D), and ventral view (E). Photos by E. Lehr.
</paragraph>
</caption>
<caption pageId="4" pageNumber="21">
<paragraph pageId="4" pageNumber="21">
Figure 4. Ventral views of right hand (A) and right foot (B) of holotype of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="4" pageNumber="21" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
sp. n. (MUSM 31196). Drawings by E. Lehr.
</paragraph>
</caption>
<paragraph pageId="4" pageNumber="21">
Paratypes. A total of 33 (Figs 5-7, 8C), all from inside the PPPF (except for: MUSM 31199-31202, NMP6V 75526-29), Provincia Satipo,
<normalizedToken originalValue="Región">Region</normalizedToken>
<normalizedToken originalValue="Junín">Junin</normalizedToken>
: 10 adult females (MUSM 31977, 31980, 31987, 31201, NMP6V 75076, 75522 [GenBank accession numbers KY594753, KY594761], 75523, 75528 [GenBank accession numbers KY594756, KY594764], 75529 [GenBank accession number KY594757], 75534), 20 adult males (MUSM 31186 [GenBank accession number KY594752], 31195, 31199, 31202, 31975, 31979, 31988, 31989, 31992, 31993, NMP6V 75077-75079, 75524 [GenBank accession numbers KY594754, KY594762], 75525 [GenBank accession numbers KY594755, KY594763], 75526, 75527, 75533, UMMZ 244726, 244727), 3 juveniles (MUSM 31187, 31990, 31200).
</paragraph>
<caption pageId="4" pageNumber="21">
<paragraph pageId="4" pageNumber="21">
Figure 5. Variation of male paratypes of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="4" pageNumber="21" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
sp. n. in dorsolateral, dorsal, and ventral views.
<normalizedToken originalValue="AC">A-C</normalizedToken>
(MUSM 31186, SVL 18.6 mm)
<normalizedToken originalValue="DF">D-F</normalizedToken>
(MUSM 31195, SVL 16.9 mm)
<normalizedToken originalValue="GI">G-I</normalizedToken>
(MUSM 31992, SVL 15.9 mm). Photos by E. Lehr.
</paragraph>
</caption>
<caption pageId="4" pageNumber="21">
<paragraph pageId="4" pageNumber="21">
Figure 6. Variation of female paratypes of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="4" pageNumber="21" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
sp. n. in dorsolateral, dorsal, and ventral views.
<normalizedToken originalValue="AC">A-C</normalizedToken>
(NMP6V 75522, SVL 19.2 mm)
<normalizedToken originalValue="DF">D-F</normalizedToken>
(MUSM 31987, SVL 23.0 mm)
<normalizedToken originalValue="GI">G-I</normalizedToken>
(MUSM 31977, SVL 21.9 mm). Photos by E. Lehr.
</paragraph>
</caption>
<caption pageId="4" pageNumber="21">
<paragraph pageId="4" pageNumber="21">
Figure 7. Variation of juvenile paratypes of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="4" pageNumber="21" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
sp. n. in dorsolateral, dorsal, and ventral views.
<normalizedToken originalValue="AC">A-C</normalizedToken>
(MUSM 31990, SVL 14.0 mm)
<normalizedToken originalValue="DF">D-F</normalizedToken>
(MUSM 31187, SVL 12.5 mm)
<normalizedToken originalValue="GI">G-I</normalizedToken>
(MUSM 31200, SVL 14.0 mm). Photos by E. Lehr.
</paragraph>
</caption>
<paragraph lastPageId="5" lastPageNumber="22" pageId="4" pageNumber="21">
MUSM 31186, MUSM 31187, NMP6V 75522, 75523: Quebrada Tarhuish, left bank of Antuyo River,
<normalizedToken originalValue="“Shiusha”">&quot;Shiusha&quot;</normalizedToken>
, upper montane forest,
<geoCoordinate direction="south" orientation="latitude" precision="1" value="-11.36775">11°22'3.9&quot;S</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-74.93686">74°56'12.7&quot;W</geoCoordinate>
, 3414 m a.s.l. collected on 12 May 2012 by E. Lehr and R. von May. MUSM 31195, NMP6V 75524, 75524: collected at the type locality along with the holotype. MUSM 31199, 31200, MUSM 31201, 31202, NMP6V
<pageBreakToken pageId="5" pageNumber="22" start="start">75526</pageBreakToken>
, 75527: Upper part of Quebrada Tasta, &quot;Laguna Luichococha&quot;, Puna,
<geoCoordinate direction="south" orientation="latitude" precision="1" value="-11.456583">11°27'23.7&quot;S</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-74.91961">74°55'10.6&quot;W</geoCoordinate>
, 3708 m a.s.l. collected on 20 May 2012 by E. Lehr and R. von May. NMP6V 75528, 75529: near trail from Tasta to Tarhuish (first mountain peak),
<taxonomicName class="Diplopoda" family="Platyrhacidae" genus="Polylepis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Polylepis" order="Polydesmida" pageId="5" pageNumber="22" phylum="Arthropoda" rank="genus">Polylepis</taxonomicName>
forest patch,
<geoCoordinate direction="south" orientation="latitude" precision="1" value="-11.435722">11°26'8.6&quot;S</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-74.899025">74°53'56.5&quot;W</geoCoordinate>
, 3886 m a.s.l. collected on 20 May 2012 by E. Lehr and R. von May. MUSM 31975: Antuyo,
<geoCoordinate direction="south" orientation="latitude" precision="1" value="-11.334361">11°20'03.7&quot;S</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-74.99697">74°59'49.1&quot;W</geoCoordinate>
, 3700 m a.s.l. collected on 27 June 2013 by E. Lehr, J. Moravec, and J.C. Cusi. MUSM 31977, 31979, MUSM 31980, NMP6V 75076, UMMZ 244726: Hatunpata,
<geoCoordinate direction="south" orientation="latitude" precision="1" value="-11.302195">11°18'07.9&quot;S</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-75.02639">75°01'35.0&quot;W</geoCoordinate>
, 3710 m a.s.l. collected on 28 June 2013 by by E. Lehr, J. Moravec, and J.C. Cusi. MUSM 31987-31990, NMP6V 75077, 75078, 75533, UMMZ 244727: Trancapampa,
<geoCoordinate direction="south" orientation="latitude" precision="1" value="-11.297001">11°17'49.2&quot;S</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-75.01286">75°00'46.3&quot;W</geoCoordinate>
, 3550 m a.s.l. collected on 2 July 2013 by E. Lehr, J. Moravec, and J.C. Cusi. MUSM 31992, 31993, NMP6V 75079, 75534: Antuyo Bajo,
<geoCoordinate direction="south" orientation="latitude" precision="1" value="-11.314834">11°18'53.4&quot;S</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="1" value="-74.993">74°59'34.8&quot;W</geoCoordinate>
, 3400 m a.s.l. collected on 4 July 2013 by E. Lehr, J. Moravec, and J.C. Cusi.
</paragraph>
</subSubSection>
<subSubSection lastPageId="9" lastPageNumber="26" pageId="6" pageNumber="23" type="generic placement">
<paragraph pageId="6" pageNumber="23">
<pageBreakToken pageId="6" pageNumber="23" start="start">Generic</pageBreakToken>
placement.
</paragraph>
<paragraph pageId="6" pageNumber="23">
We assign this species to
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis" order="Anura" pageId="6" pageNumber="23" phylum="Chordata" rank="genus">Pristimantis</taxonomicName>
based on our molecular data (Fig. 2).
</paragraph>
<paragraph lastPageId="8" lastPageNumber="25" pageId="6" pageNumber="23">
Diagnosis. A new species of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis" order="Anura" pageId="6" pageNumber="23" phylum="Chordata" rank="genus">Pristimantis</taxonomicName>
assigned to the
<taxonomicName lsidName="danae" pageId="6" pageNumber="23" rank="species" species="danae">danae</taxonomicName>
species Group having the following combination of characters: (1) Skin on dorsum shagreen with low scattered tubercles, skin on flanks tuberculate, skin on venter areolate; discoidal fold absent, thoracic fold present; irregularly shaped, discontinuous dorsolateral folds present; (2) tympanic membrane and tympanic annulus absent; (3) snout short, rounded in dorsal and in lateral views; (4) upper eyelid without enlarged conical tubercles; EW shorter
<pageBreakToken pageId="7" pageNumber="24" start="start">than</pageBreakToken>
IOD; cranial crests absent; (5) dentigerous processes of vomers present; (6) males without vocal slits, nuptial pads absent; (7) Finger I shorter than Finger II; tips of digits narrow, rounded, lacking circumferential grooves; (8) fingers without lateral fringes; (9) small conical ulnar and tarsal tubercles present; (10) heel with a small conical tubercle; inner tarsal fold usually absent; (11) inner metatarsal tubercle ovoid, 1.5 times as large as outer; outer metatarsal tubercle small, rounded; vie low supernumerary plantar tubercles; (12) toes without lateral fringes; basal toe webbing absent; Toe V longer than Toe III; tips of digits narrow, rounded, lacking circumferential grooves, toe tips slightly smaller than those on fingers; (13) in life, dorsal ground coloration pale or dark gray, reddish brown or brownish olive with dark gray scattered flecks, some with X-shaped mark on scapular and ill-defined diagonal bars on flanks; dark grayish-brown canthal and supratympanic stripes usually present; groin dark gray or pale reddish brown with a pale
<pageBreakToken pageId="8" pageNumber="25" start="start">red</pageBreakToken>
to pink tint in some; venter dark gray, pale gray, grayish brown or pale grayish green and in some dark gray mottled; iris pale grayish green with fine black vermiculation and brownish-orange horizontal streak across pupil and lower half of iris; (14) SVL in adult males 14.6-19.2 mm (n = 21), in adult females 19.2-23.0 mm (n = 10).
</paragraph>
<paragraph lastPageId="9" lastPageNumber="26" pageId="8" pageNumber="25">
Comparisons.
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
is readily distinguished from its congeners in Ecuador (176 species,
<bibRefCitation pageId="15" pageNumber="32" title="AmphibiaWeb. Information on amphibian biology and conservation" url="http://amphibiaweb.org/" year="2016">AmphibiaWeb 2016</bibRefCitation>
), Peru (128 species,
<bibRefCitation pageId="15" pageNumber="32" title="AmphibiaWeb. Information on amphibian biology and conservation" url="http://amphibiaweb.org/" year="2016">AmphibiaWeb 2016</bibRefCitation>
), and Bolivia (17 species,
<bibRefCitation pageId="15" pageNumber="32" title="AmphibiaWeb. Information on amphibian biology and conservation" url="http://amphibiaweb.org/" year="2016">AmphibiaWeb 2016</bibRefCitation>
) by having narrow digits without circumferential grooves, by lacking a tympanic annulus and tympanic membrane, and by having irregularly shaped, discontinuous dorsolateral folds. In Peru 18 species of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="genus">Pristimantis</taxonomicName>
lack a tympanum; these are
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis academicus" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="academicus">Pristimantis academicus</taxonomicName>
Lehr, Moravec, and Gagliardi Urrutia, 2010,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis altamazonicus" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="altamazonicus">Pristimantis altamazonicus</taxonomicName>
(Barbour and Dunn, 1921),
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis ashaninka" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="ashaninka">Pristimantis ashaninka</taxonomicName>
Lehr and Moravec, 2017,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis colodactylus" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="colodactylus">Pristimantis colodactylus</taxonomicName>
(Lynch, 1979),
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis coronatus" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="coronatus">Pristimantis coronatus</taxonomicName>
Lehr and Duellman, 2007a,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis croceoinguinis" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="croceoinguinis">Pristimantis croceoinguinis</taxonomicName>
(Lynch, 1968),
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis cruciocularis" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="cruciocularis">Pristimantis cruciocularis</taxonomicName>
(Lehr, Lundberg, Aguilar, and von May, 2006),
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis flavobracatus" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="flavobracatus">Pristimantis flavobracatus</taxonomicName>
(Lehr, Lundberg, Aguilar, and von May, 2006),
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis imitatrix" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="imitatrix">Pristimantis imitatrix</taxonomicName>
(Duellman, 1978b),
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis lirellus" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="lirellus">Pristimantis lirellus</taxonomicName>
(Dwyer, 1995),
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis leucorrhinus" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="leucorrhinus">Pristimantis leucorrhinus</taxonomicName>
Boano, Mazzotti, and Sindaco, 2008,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis martiae" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="martiae">Pristimantis martiae</taxonomicName>
(Lynch, 1974),
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis minutulus" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="minutulus">Pristimantis minutulus</taxonomicName>
Duellman and Hedges, 2007,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis rhabdocnemus" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="rhabdocnemus">Pristimantis rhabdocnemus</taxonomicName>
(Duellman and Hedges, 2005),
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis simonsii" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="simonsii">Pristimantis simonsii</taxonomicName>
(Boulenger, 1900),
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis tantanti" order="Anura" pageId="8" pageNumber="25" phylum="Chordata" rank="species" species="tantanti">Pristimantis tantanti</taxonomicName>
(Lehr, Torres-Gastello,
<pageBreakToken pageId="9" pageNumber="26" start="start">and</pageBreakToken>
<normalizedToken originalValue="Suárez-Segovia">Suarez-Segovia</normalizedToken>
, 2007),
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis ventrimarmoratus" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="ventrimarmoratus">Pristimantis ventrimarmoratus</taxonomicName>
(Boulenger, 1912), and
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis vilcabambae" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="vilcabambae">Pristimantis vilcabambae</taxonomicName>
Lehr, 2007. Of these, only
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis simonsii" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="simonsii">Pristimantis simonsii</taxonomicName>
from northern Peru has narrow digits without circumferential grooves.
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
and
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis simonsii" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="simonsii">Pristimantis simonsii</taxonomicName>
lack circumferential grooves and a tympanum, and both have dorsolateral folds, but
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
is smaller than
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis simonsii" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="simonsii">Pristimantis simonsii</taxonomicName>
(female SVL 26.2-33.3 mm in
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis simonsii" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="simonsii">Pristimantis simonsii</taxonomicName>
), and male
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
lack nuptial pads which are present in
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis simonsii" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="simonsii">Pristimantis simonsii</taxonomicName>
.
</paragraph>
<paragraph pageId="9" pageNumber="26">
Members of the
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis orestes" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="orestes">Pristimantis orestes</taxonomicName>
species Group are terrestrial and inhabit high elevations in southern Ecuador and in Peru (Duellman and Lehr, 2009) and have narrow digits, and only one of the 17 species (
<bibRefCitation author="Guayasamin, JM" journalOrPublisher="Zootaxa" pageId="17" pageNumber="34" pagination="345 - 356" title="A new species of the Pristimantisorestes group (Amphibia: Strabomantidae) from the high Andes of Ecuador, Reserva Mazar." url="https://doi.org/10.11646/zootaxa.3616.4.3" volume="3616" year="2013">Guayasamin and Artega 2013</bibRefCitation>
) lacks circumferential grooves (
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis simonsii" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="simonsii">Pristimantis simonsii</taxonomicName>
), and only two (
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="genus">Pristimantis</taxonomicName>
seorsus,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis simonsii" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="simonsii">Pristimantis simonsii</taxonomicName>
) lack a tympanum. Furthermore
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
is phylogenetically distant from members of this group which is considered to be not monophyletic (
<bibRefCitation author="Duellman, WE" journalOrPublisher="Natur und Tier-Verlag, Muenster" pageId="16" pageNumber="33" title="Terrestrial-breeding Frogs (Strabomantidae) in Peru." year="2009">Duellman and Lehr 2009</bibRefCitation>
, Fig. 2).
</paragraph>
<paragraph pageId="9" pageNumber="26">
Among the three other new species of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="genus">Pristimantis</taxonomicName>
from the upper montane forests and Puna of the PPPF, only
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="genus">Pristimantis</taxonomicName>
sp. n. E lacks circumferential grooves and a tympanum. However,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
and
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="genus">Pristimantis</taxonomicName>
sp. n. E both differ regarding other morphological traits, coloration, and genetically.
</paragraph>
<paragraph pageId="9" pageNumber="26">
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
shares with
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis stipa" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="stipa">Pristimantis stipa</taxonomicName>
Venegas and Duellman, 2012 from the Puna of northern Peru (
<bibRefCitation author="Venegas, PJ" journalOrPublisher="Zootaxa" pageId="19" pageNumber="36" pagination="47 - 59" title="Two syntopic new species of the Pristimantisorestes Group (Anura: Strabomantidae) from northwestern Peru." volume="3249" year="2012">Venegas and Duellman 2012</bibRefCitation>
) narrow digits without circumferential grooves and dorsolateral folds. However,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
is smaller (female SVL 19.2-23.0 mm [n = 10] vs. 35.1 mm [n = 1]), lacks a tympanum (present in
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis stipa" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="stipa">Pristimantis stipa</taxonomicName>
), and has ulnar tubercles not coalesced into fold (coalesced into low fold in
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis stipa" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="stipa">Pristimantis stipa</taxonomicName>
),
<bibRefCitation author="Venegas, PJ" journalOrPublisher="Zootaxa" pageId="19" pageNumber="36" pagination="47 - 59" title="Two syntopic new species of the Pristimantisorestes Group (Anura: Strabomantidae) from northwestern Peru." volume="3249" year="2012">Venegas and Duellman (2012)</bibRefCitation>
.
</paragraph>
<paragraph pageId="9" pageNumber="26">
The new species shares narrow digits without circumferential grooves and the absence of a tympanic annulus and tympanic membrane with the Andean genera
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="genus">Phrynopus</taxonomicName>
Peters, 1873 (except for
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus auriculatus" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="auriculatus">Phrynopus auriculatus</taxonomicName>
Duellman and Hedges, 2008, and
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus peruanus" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="peruanus">Phrynopus peruanus</taxonomicName>
Peters, 1873), 28 species from elevations between 2200 and 4400 m a.s.l. in central and northern Peru, Duellman and Lehr, 2009) and
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Bryophryne" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bryophryne" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="genus">Bryophryne</taxonomicName>
Hedges, Duellman, and Heinicke, 2008 (8 species from elevations between 2900 and 4120 m a.s.l. in southern Peru,
<bibRefCitation author="Duellman, WE" journalOrPublisher="Natur und Tier-Verlag, Muenster" pageId="16" pageNumber="33" title="Terrestrial-breeding Frogs (Strabomantidae) in Peru." year="2009">Duellman and Lehr 2009</bibRefCitation>
),
<bibRefCitation pageId="15" pageNumber="32" title="AmphibiaWeb. Information on amphibian biology and conservation" url="http://amphibiaweb.org/" year="2016">AmphibiaWeb (2016)</bibRefCitation>
.
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
is most similar with
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus chaparroi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="chaparroi">Phrynopus chaparroi</taxonomicName>
Mamani and Malqui, 2014 which was described based on morphological characters and found at elevations between 4205 and 4490 m a.s.l. in southern
<normalizedToken originalValue="Región">Region</normalizedToken>
<normalizedToken originalValue="Junín">Junin</normalizedToken>
(
<bibRefCitation author="Mamani, L" journalOrPublisher="Zootaxa" pageId="18" pageNumber="35" pagination="207 - 214" title="A new species of Phrynopus (Anura: Craugastoridae) from the central Peruvian Andes." url="https://doi.org/10.11646/zootaxa.3838.2.5" volume="3838" year="2014">Mamani and Malqui 2014</bibRefCitation>
). Both
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
and
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus chaparroi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="chaparroi">Phrynopus chaparroi</taxonomicName>
lack a tympanum and have narrow digits without circumferential grooves. However,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
is smaller than
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus chaparroi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="chaparroi">Phrynopus chaparroi</taxonomicName>
(female SVL 19.2-23.0 mm [n = 10] vs. 30.0-32.2 [n = 4]), lacks protuberant subconical postrictal tubercles (present in
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus chaparroi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="chaparroi">Phrynopus chaparroi</taxonomicName>
), has dorsolateral folds (absent in
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus chaparroi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="chaparroi">Phrynopus chaparroi</taxonomicName>
), dentigerous processes of vomers present (absent in
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus chaparroi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="chaparroi">Phrynopus chaparroi</taxonomicName>
), and males lack nuptial pads (present in
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus chaparroi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="chaparroi">Phrynopus chaparroi</taxonomicName>
).
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus chaparroi" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="species" species="chaparroi">Phrynopus chaparroi</taxonomicName>
might belong to
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis" order="Anura" pageId="9" pageNumber="26" phylum="Chordata" rank="genus">Pristimantis</taxonomicName>
, but molecular characters need to be applied to confirm our suspicion.
</paragraph>
</subSubSection>
<subSubSection lastPageId="10" lastPageNumber="27" pageId="9" pageNumber="26" type="description">
<paragraph pageId="9" pageNumber="26">Description of the holotype.</paragraph>
<paragraph lastPageId="10" lastPageNumber="27" pageId="9" pageNumber="26">
Head about as long as wide; head length 39.7% of SVL; head width 38.6% of SVL; cranial crests absent; snout short, rounded in dorsal view, rounded in lateral view (Fig. 3A, B); eye-nostril distance 70% of eye diameter; nostrils slightly protuberant, directed dorsolaterally; canthus rostralis short, rounded
<pageBreakToken pageId="10" pageNumber="27" start="start">in</pageBreakToken>
lateral view, weakly concave in dorsal view; loreal region concave; lips rounded; outer margin of upper eyelid each with few slightly enlarged conical tubercles; upper eyelid width 51.9% of IOD (see photo in life Fig. 3); supratympanic fold short and broad, extending from posterior margin of upper eyelid slightly curved to insertion of arm; tympanic membrane and annulus absent; distinct conical postrictal tubercles present bilaterally. Choanae small, ovoid, not concealed by palatal shelf of maxilla; dentigerous processes of vomers positioned posterior to level of choanae, oblique, narrowly separated; tongue long, oval, about three times as long as wide, not notched posteriorly, posterior half free.
</paragraph>
<paragraph pageId="10" pageNumber="27">Skin on dorsum shagreen with low scattered tubercles, skin on flanks tuberculate, irregularly shaped, discontinuous dorsolateral folds present extending from posterior level of tympanic area to level of hind limb insertion; skin on throat, chest, and belly areolate; discoidal fold absent, thoracic fold present; cloacal sheath short.</paragraph>
<paragraph pageId="10" pageNumber="27">Outer ulnar surface each with a row of four minute low tubercles; palmar tubercle bifid; thenar tubercle ovoid; subarticular tubercles well defined, most prominent on base of fingers, round in ventral view, subconical in lateral view; supernumerary tubercles indistinct; fingers short and stout lacking lateral fringes, Finger I shorter than Finger II; tips of digits of fingers narrow, round, lacking circumferential grooves (Fig. 4A).</paragraph>
<paragraph pageId="10" pageNumber="27">Hind limbs short, slender, tibia length 40.2% of SVL; foot length 41.3% of SVL; dorsal surfaces of hind limbs tuberculate; inner surface of thighs smooth, posterior surfaces of thighs tuberculate, ventral surfaces of thighs areolate; heels each with a small conical tubercle; outer surface of tarsus with few scattered minute low tubercles; inner tarsal fold absent, but small tubercle proximal to metatarsal tubercle; inner metatarsal tubercle ovoid, one and a half times the size of round outer metatarsal tubercle; subarticular tubercles well defined, round in ventral view, subconical in lateral view; few plantar supernumerary tubercles, about one third the size of subarticular tubercles; toes without lateral fringes; basal webbing absent; tips of digits narrow, round, less expanded than those on fingers, lacking circumferential grooves; relative length of toes: 1&lt;2&lt;5&lt;3&lt;4; Toe V slightly longer than Toe III (tip of digit of Toe III and Toe V not reaching distal subarticular tubercle on Toe IV; Fig. 4B).</paragraph>
</subSubSection>
<subSubSection pageId="10" pageNumber="27" type="description">
<paragraph pageId="10" pageNumber="27">Measurements (in mm) of the holotype.</paragraph>
<paragraph pageId="10" pageNumber="27">
SVL 18.9; tibia length 7.6; foot length 7.8; head length 7.5; head width 7.3; eye diameter 2.0; inter orbital distance 2.7; upper eyelid width 1.4; internarial distance 1.9;
<normalizedToken originalValue="eyenostril">eye-nostril</normalizedToken>
distance 1.4.
</paragraph>
</subSubSection>
<subSubSection pageId="10" pageNumber="27" type="coloration of the holotype in life">
<paragraph pageId="10" pageNumber="27">Coloration of the holotype in life</paragraph>
<paragraph pageId="10" pageNumber="27">(Fig. 3). The dorsal ground coloration is pale reddish brown with few dark brown flecks; narrow dark brown canthal and supratympanic stripes; flanks pale reddish brown with dark brown flecks forming irregularly shaped diagonal bars; groin and anterior surfaces of thighs reddish brown with dark brown flecks and pale reddish tint; chest, belly, and ventral surfaces of thighs dark grayish brown, throat pale reddish brown and pale gray mottled; palmar and plantar surfaces, and fingers and toes dark grayish brown; iris pale grayish green with fine black vermiculation and brownish-orange horizontal streak across pupil and lower half of iris.</paragraph>
<caption pageId="10" pageNumber="27">
<paragraph pageId="10" pageNumber="27">
Table 3. Measurements (in mm) of selected adult type specimens of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="10" pageNumber="27" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
sp. n. M = male, F = female. For other abbreviations see methods.
</paragraph>
</caption>
<paragraph pageId="10" pageNumber="27">
<table pageId="10" pageNumber="27">
<tr pageId="10" pageNumber="27">
<th colspan="1" pageId="10" pageNumber="27" rowspan="1">Characters</th>
<th colspan="1" pageId="10" pageNumber="27" rowspan="1">
<named-content content-type="dwc:institutional_code" pageId="10" pageNumber="27">MUSM</named-content>
31988
</th>
<th colspan="1" pageId="10" pageNumber="27" rowspan="1">
<named-content content-type="dwc:institutional_code" pageId="10" pageNumber="27">MUSM</named-content>
31992
</th>
<th colspan="1" pageId="10" pageNumber="27" rowspan="1">
<named-content content-type="dwc:institutional_code" pageId="10" pageNumber="27">MUSM</named-content>
31186
</th>
<th colspan="1" pageId="10" pageNumber="27" rowspan="1">
<named-content content-type="dwc:institutional_code" pageId="10" pageNumber="27">UMMZ</named-content>
244727
</th>
<th colspan="1" pageId="10" pageNumber="27" rowspan="1">
<named-content content-type="dwc:institutional_code" pageId="10" pageNumber="27">NMP6V</named-content>
75523
</th>
<th colspan="1" pageId="10" pageNumber="27" rowspan="1">
<named-content content-type="dwc:institutional_code" pageId="10" pageNumber="27">MUSM</named-content>
31980
</th>
<th colspan="1" pageId="10" pageNumber="27" rowspan="1">
<named-content content-type="dwc:institutional_code" pageId="10" pageNumber="27">MUSM</named-content>
31977
</th>
<th colspan="1" pageId="10" pageNumber="27" rowspan="1">
<named-content content-type="dwc:institutional_code" pageId="10" pageNumber="27">NMP6V</named-content>
75076
</th>
<th colspan="1" pageId="10" pageNumber="27" rowspan="1">
<named-content content-type="dwc:institutional_code" pageId="10" pageNumber="27">MUSM</named-content>
31987
</th>
</tr>
<tr pageId="10" pageNumber="27">
<td colspan="1" pageId="10" pageNumber="27" rowspan="1">SVL</td>
</tr>
<tr pageId="10" pageNumber="27">
<td colspan="1" pageId="10" pageNumber="27" rowspan="1">TL</td>
</tr>
<tr pageId="10" pageNumber="27">
<td colspan="1" pageId="10" pageNumber="27" rowspan="1">FL</td>
</tr>
<tr pageId="10" pageNumber="27">
<td colspan="1" pageId="10" pageNumber="27" rowspan="1">HL</td>
</tr>
<tr pageId="10" pageNumber="27">
<td colspan="1" pageId="10" pageNumber="27" rowspan="1">HW</td>
</tr>
<tr pageId="10" pageNumber="27">
<td colspan="1" pageId="10" pageNumber="27" rowspan="1">ED</td>
</tr>
<tr pageId="10" pageNumber="27">
<td colspan="1" pageId="10" pageNumber="27" rowspan="1">IOD</td>
</tr>
<tr pageId="10" pageNumber="27">
<td colspan="1" pageId="10" pageNumber="27" rowspan="1">EW</td>
</tr>
<tr pageId="10" pageNumber="27">
<td colspan="1" pageId="10" pageNumber="27" rowspan="1">IND</td>
</tr>
<tr pageId="10" pageNumber="27">
<td colspan="1" pageId="10" pageNumber="27" rowspan="1">E-N</td>
</tr>
</table>
</paragraph>
</subSubSection>
<subSubSection lastPageId="12" lastPageNumber="29" pageId="10" pageNumber="27" type="coloration of the holotype in preservative">
<paragraph pageId="10" pageNumber="27">Coloration of the holotype in preservative.</paragraph>
<paragraph lastPageId="11" lastPageNumber="28" pageId="10" pageNumber="27">
The dorsal ground coloration is pale brown with few dark brown flecks; narrow dark brown canthal and supratympanic
<pageBreakToken pageId="11" pageNumber="28" start="start">stripes</pageBreakToken>
; flanks pale brown with many dark brown flecks forming irregularly shaped diagonal bars; groin and anterior surfaces of thighs brown with dark brown flecks; chest, belly, and ventral surfaces of thighs dark brown, throat pale brown and pale gray mottled; palmar and plantar surfaces, and fingers and toes dark brown; iris pale gray.
</paragraph>
<caption pageId="11" pageNumber="28">
<paragraph pageId="11" pageNumber="28">
Table 4. Measurements (in mm) and proportions of adult male and adult female type specimens of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="11" pageNumber="28" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
sp. n.; ranges followed by means and one standard deviation in parentheses. For abbreviations see methods.
</paragraph>
</caption>
<paragraph pageId="11" pageNumber="28">
<table pageId="11" pageNumber="28">
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">Characters</td>
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">Males (n = 21)</td>
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">Females (n = 10)</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">SVL</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">TL</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">FL</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">HL</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">HW</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">ED</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">IOD</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">EW</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">IND</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">E-N</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">TLSVL</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">FLSVL</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">HLSVL</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">HWSVL</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">HWHL</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">E-NED</td>
</tr>
<tr pageId="11" pageNumber="28">
<td colspan="1" pageId="11" pageNumber="28" rowspan="1">EWIOD</td>
</tr>
</table>
</paragraph>
<paragraph pageId="11" pageNumber="28">
Variation. All paratypes (Figs 5-7) are similar to the holotype regarding morphology and proportions (Tables 3, 4). Besides differences in SVL, notable morphological variation includes prominence of dorsolateral folds (e.g., prominent dorsolateral folds in MUSM 31192, 31195, Fig. 5
<normalizedToken originalValue="DF">D-F</normalizedToken>
,
<normalizedToken originalValue="GI">G-I</normalizedToken>
; weak dorsolateral folds in MUSM 31186, 31975, 31977, NMP6V 75522, 75528, 75529, Fig. 6
<normalizedToken originalValue="GI">G-I</normalizedToken>
), and coarseness of tuberculate skin texture on flanks and hind limbs (skin coarsely tuberculate in MUSM 31186, 31192, 31195, NMP6V 75525, Fig. 5; skin weakly tubercular MUSM 31987, 31997, NMP6V 75528, 75529). Two specimens (NMP6V 75529, 75534) have a tubercle-like inner tarsal fold present.
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="11" pageNumber="28" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
demonstrates a remarkable polymorphism in coloration (Figs 5-7).
</paragraph>
<paragraph lastPageId="12" lastPageNumber="29" pageId="11" pageNumber="28">
The dorsal coloration ranges from pale gray (MUSM 31987, NMP6V 75533, Fig. 6
<normalizedToken originalValue="DF">D-F</normalizedToken>
), dark gray (MSUM 31186, 3199, NMP6V 75522, 75523, 75528, 75529,
<pageBreakToken pageId="12" pageNumber="29" start="start">Fig</pageBreakToken>
. 6
<normalizedToken originalValue="AC">A-C</normalizedToken>
), reddish brown (MUSM 31195, 31975, NMP6V 75525, Figs 5
<normalizedToken originalValue="DF">D-F</normalizedToken>
) to brownish olive (MUSM 31992, 31997, Figs 5
<normalizedToken originalValue="GI">G-I</normalizedToken>
, 6
<normalizedToken originalValue="GI">G-I</normalizedToken>
) with dark gray scattered flecks. Some have an X-shaped mark on scapular (MUSM 31200, 31975, 31990), some ill-defined diagonal bars on the flanks (MUSM 31195). Dark grayish-brown canthal and supratympanic stripes are usually present except for dark gray specimens (MSUM 31186, 3199, NMP6V 75522, 75523, 75528, 75529). The groin is dark gray (MSUM 31186, 3199, NMP6V 75522, 75523, 75528, 75529) or pale reddish brown with a pale red to pink tint in some specimens (MUSM 31195, 31196). The venter is dark gray (NMP6V 75522, 75523, 75528, 75529, Fig. 6C), pale gray (MUSM 31987, Fig. 6F), grayish brown (MUSM 31186, 31195, NMP6V 75525, Fig. 5C, F) or pale grayish green and gray mottled (MUSM 31197, Fig. 6I) or dark gray and pale gray mottled (MUSM 31199, 31975, 31992, NMP6V 75533, Fig. 5I).
</paragraph>
<paragraph pageId="12" pageNumber="29">Juveniles (MUSM 31187, 31990, 31200, Fig. 7) have a paler coloration (yellowish to reddish brown) with contrasting dark brown flecks and distinct canthal and supratympanic stripes. All have the iris pale grayish green with fine black vermiculation and brownish-orange horizontal streak across pupil and lower half of iris, and usually a narrow vertical dark gray streak from pupil through middle of lower iris.</paragraph>
</subSubSection>
<subSubSection pageId="12" pageNumber="29" type="etymology">
<paragraph pageId="12" pageNumber="29">Etymology.</paragraph>
<paragraph pageId="12" pageNumber="29">We dedicate this species to Sir David Frederick Attenborough in honor for his educational documentaries on wildlife, especially on amphibians (e.g., Life in Cold Blood, Fabulous Frogs), and for raising awareness about the importance of wildlife conservation. The specific epithet is used as noun in apposition.</paragraph>
</subSubSection>
<subSubSection lastPageId="14" lastPageNumber="31" pageId="12" pageNumber="29" type="distribution">
<paragraph pageId="12" pageNumber="29">Distribution, natural history, and conservation status.</paragraph>
<paragraph lastPageId="14" lastPageNumber="31" pageId="12" pageNumber="29">
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="12" pageNumber="29" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
is known from six localities inside the PPPF (Puna of Quebrada Tarhuish at
<pageBreakToken pageId="13" pageNumber="30" start="start">Laguna</pageBreakToken>
Udrecocha, Fig. 8A; upper montane forest of Quebrada Tarhuish on the left bank
<normalizedToken originalValue="“Shiusha”">&quot;Shiusha&quot;</normalizedToken>
of Antuyo River; Antuyo; Antuyo Bajo; Hatunpata, and Trancapampa, Figs 8B, 9) and from two outside the PPPF (upper part of Quebrada Tasta close to Laguna Luichococha; in
<taxonomicName class="Diplopoda" family="Platyrhacidae" genus="Polylepis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Polylepis" order="Polydesmida" pageId="13" pageNumber="30" phylum="Arthropoda" rank="genus">Polylepis</taxonomicName>
forest of first mountain peak next to trail from Tasta to Tarhuish), and is distributed at elevations between 3400 and 3936 m a.s.l., Fig. 9. The type locality (Figs 8A, 9), upper part of Quebrada Tarhuish, on the east side of Laguna Udrecocha at 3936 m a.s.l., belongs to the Puna ecoregion (
<bibRefCitation pageId="13" pageNumber="30">Brack 1986</bibRefCitation>
). The vegetation consists of Peruvian feather grass (
<taxonomicName class="Liliopsida" family="Poaceae" genus="Stipa" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Stipa ichu" order="Poales" pageId="13" pageNumber="30" phylum="Tracheophyta" rank="species" species="ichu">Stipa ichu</taxonomicName>
), mosses, and small bushes. The holotype was found inside moss in the afternoon on 17 May 2012. No sympatric anurans were found at the type locality. At the upper montane forest of Quebrada Tarhuish on the left bank
<normalizedToken originalValue="“Shiusha”">&quot;Shiusha&quot;</normalizedToken>
of Antuyo River,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="13" pageNumber="30" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
was found deep inside large moss layers. Sympatric anurans are
<taxonomicName class="Amphibia" family="Hemiphractidae" genus="Gastrotheca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gastrotheca griswoldi" order="Anura" pageId="13" pageNumber="30" phylum="Chordata" rank="species" species="griswoldi">Gastrotheca griswoldi</taxonomicName>
(MUSM 31193),
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis" order="Anura" pageId="14" pageNumber="31" phylum="Chordata" rank="genus">
<pageBreakToken pageId="14" pageNumber="31" start="start">Pristimantis</pageBreakToken>
</taxonomicName>
sp. n. C (MUSM 31190-92),
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis" order="Anura" pageId="14" pageNumber="31" phylum="Chordata" rank="genus">Pristimantis</taxonomicName>
sp. n. D (MUSM 31197-98), and
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Phrynopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynopus" order="Anura" pageId="14" pageNumber="31" phylum="Chordata" rank="genus">Phrynopus</taxonomicName>
sp. n. A (MUSM 31203).
</paragraph>
<caption pageId="14" pageNumber="31">
<paragraph pageId="14" pageNumber="31">
Figure 8. Habitats of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="14" pageNumber="31" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
sp. n. in the PPPF: A type locality in the upper Tarhuish valley at Laguna Udrecocha, Puna at 3936 m a.s.l., 17 May 2012 B upper montane forest at 3550 m a.s.l. where
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="14" pageNumber="31" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
sp. n. was found in moss pads C female
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="14" pageNumber="31" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
sp. n. (MUSM 31980, SVL 21.5 mm) guarding a clutch in a moss pad. Photos by E. Lehr.
</paragraph>
</caption>
<paragraph pageId="14" pageNumber="31">
A female
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="14" pageNumber="31" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
(MUSM 31980, Fig. 8C) guarding 20 eggs was found at Hatunpata inside moss, 3710 m a.s.l., on 28 June 2013. The eggs were pale cream colored and had an average diameter of 3.5
<normalizedToken originalValue="±">+/-</normalizedToken>
0.1 mm (3.3-3.6 mm, n = 20).
</paragraph>
<paragraph pageId="14" pageNumber="31">
The IUCN Red List criteria (
<bibRefCitation author="Heinicke, MP" journalOrPublisher="IUCN Species Survival Commission, IUCN, Gland, Switzerland and Cambridge, UK" pageId="17" pageNumber="34" title="IUCN Red List Categories and Criteria. Version 3.1" url="http://www.iucnredlist.org/technical-documents/categories-and-criteria" year="2001">IUCN 2001</bibRefCitation>
) consider that if a species occurs in fewer than 10 threat-defined locations and the extent of occurrence (EOO) is &lt;20,000 km2, it should be classified as Vulnerable or Endangered.
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="14" pageNumber="31" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
is known from seven localities distributed in the PPPF and its buffer zone (Fig. 9), with an estimated EOO of 66.54 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 and most of them are inside the protected area, we propose that
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="14" pageNumber="31" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
should likely be categorized as Near Threatened (NT).
</paragraph>
<caption pageId="14" pageNumber="31">
<paragraph pageId="14" pageNumber="31">
Figure 9. Distribution of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="14" pageNumber="31" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
sp. n. in the PPPF and its surroundings: type locality: Laguna Udrecocha, 3936 m a.s.l.; 1 Upper part of Quebrada Tasta, &quot;Laguna Luichococha&quot;, 3708 m a.s.l. 2 near trail from Tasta to Tarhuish (first mountain peak),
<taxonomicName class="Diplopoda" family="Platyrhacidae" genus="Polylepis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Polylepis" order="Polydesmida" pageId="14" pageNumber="31" phylum="Arthropoda" rank="genus">Polylepis</taxonomicName>
forest patch, 3886 m a.s.l. 3 Quebrada Tarhuish, left bank of Antuyo River,
<normalizedToken originalValue="“Shiusha”">&quot;Shiusha&quot;</normalizedToken>
, 3414 m a.s.l. 4 Antuyo, 3700 m a.s.l. 5 Antuyo Bajo, 3400 m a.s.l. 6 Hatunpata, 3710 m a.s.l. 7 Trancapampa, 3550 m a.s.l.
</paragraph>
</caption>
<paragraph pageId="14" pageNumber="31">
Given that the known distribution of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis attenboroughi" order="Anura" pageId="14" pageNumber="31" phylum="Chordata" rank="species" species="attenboroughi">Pristimantis attenboroughi</taxonomicName>
overlaps with the PPPF, a substantial portion of the habitat of this species is formally protected. However, 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 (
<bibRefCitation author="Catenazzi, A" journalOrPublisher="Herpetological Monographs" pageId="16" pageNumber="33" pagination="1 - 23" title="Conservation status of amphibians in Peru." url="https://doi.org/10.1655/HERPMONOGRAPHS-D-13-00003" volume="28" year="2014">Catenazzi and von May 2014</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>