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<document ID-CLB-Dataset="81241" ID-DOI="http://dx.doi.org/10.3897/zse.98.84963" ID-GBIF-Dataset="1b01b657-a168-4fc5-a13e-6caee8e94f87" ID-Pensoft-Pub="1860-0743-2-213" ID-Pensoft-UUID="86D36A45524C5B3CBF9BB1209ADE1633" ID-ZooBank="959396406B7F41E9825CA53024113F54" ModsDocID="1860-0743-98-2-213" checkinTime="1656145962600" checkinUser="pensoft" docAuthor="Koehler, Joern, Castillo-Urbina, Ernesto, Aguilar-Puntriano, Cesar, Vences, Miguel &amp; Glaw, Frank" docDate="2022" docId="0617C8BA682B5AA08B7C998290664158" docLanguage="en" docName="ZoosystEvol 98(2): 213-232" docOrigin="Zoosystematics and Evolution 98 (2)" docPubDate="2022-06-24" docSource="http://dx.doi.org/10.3897/zse.98.84963" docTitle="Pristimantis nebulosus" docType="treatment" docVersion="6" id="86D36A45524C5B3CBF9BB1209ADE1633" lastPageNumber="213" masterDocId="86D36A45524C5B3CBF9BB1209ADE1633" masterDocTitle="Rediscovery, redescription and identity of Pristimantis nebulosus (Henle, 1992), and description of a new terrestrial-breeding frog from montane rainforests of central Peru (Anura, Strabomantidae)" masterLastPageNumber="232" masterPageNumber="213" pageNumber="213" updateTime="1732938932683" updateUser="ExternalLinkService">
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<mods:title id="F293E1C1B1DA3F828E4DC048F28A93C6">Rediscovery, redescription and identity of Pristimantis nebulosus (Henle, 1992), and description of a new terrestrial-breeding frog from montane rainforests of central Peru (Anura, Strabomantidae)</mods:title>
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<mods:namePart id="9181F42B61C563351635A9D0D02DF29B">Koehler, Joern</mods:namePart>
<mods:affiliation id="38A941D0EA95ADD015D59E3EDF11B885">Hessisches Landesmuseum Darmstadt, Friedensplatz 1, 64283 Darmstadt, Germany</mods:affiliation>
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<mods:namePart id="47E445CEF06FB11EB7F41B80DF3D0FDE">Castillo-Urbina, Ernesto</mods:namePart>
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<mods:affiliation id="090B878FFCE19156B809C4A7040B1950">Universidad Nacional Mayor de San Marcos, Museo de Historia Natural, Departamento de Herpetologia, Av. Arenales 1256, Lima 11, Peru</mods:affiliation>
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<mods:namePart id="E4EB7BD62CFD5795A87EFC26D8B006A2">Aguilar-Puntriano, Cesar</mods:namePart>
<mods:nameIdentifier id="EF1E3DF1373AE96CCB37596E451DB5C1" type="ORCID">https://orcid.org/0000-0001-6372-7926</mods:nameIdentifier>
<mods:affiliation id="9BC76DBED5CEA967FF8678ECFB05AB44">Universidad Nacional Mayor de San Marcos, Museo de Historia Natural, Departamento de Herpetologia, Av. Arenales 1256, Lima 11, Peru</mods:affiliation>
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<mods:namePart id="D685D0E78523ED577098E3D1C61FBB3C">Vences, Miguel</mods:namePart>
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<mods:affiliation id="974FE2F1F903555E12A3AB325090851E">Zoological Institute, Technische Universitaet Braunschweig, Mendelssohnstr. 4, 38106 Braunschweig, Germany</mods:affiliation>
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<mods:namePart id="1B6B3F9BF0F63A1504A2C0A38AB38FC0">Glaw, Frank</mods:namePart>
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<mods:affiliation id="02F2BAC4194DB49B3F7279938E7E2567">Zoologische Staatssammlung Muenchen (ZSM-SNSB), Muenchhausenstr. 21, 81247 Muenchen, Germany</mods:affiliation>
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<mods:date id="2E75650E8E6319F1BCDF4C8604768C8A">2022</mods:date>
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<treatment id="0617C8BA682B5AA08B7C998290664158" ID-GBIF-Taxon="196283140" LSID="urn:lsid:plazi:treatment:0617C8BA682B5AA08B7C998290664158" httpUri="http://treatment.plazi.org/id/0617C8BA682B5AA08B7C998290664158" lastPageNumber="213" pageId="0" pageNumber="213" scope_family="Strabomantidae" scope_order="Anura">
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<taxonomicName id="A5D36030048E85E7071290FE6263A220" LSID="0617C8BA-682B-5AA0-8B7C-998290664158" authority="(Henle, 1992)" baseAuthorityName="Henle" baseAuthorityYear="1992" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis nebulosus" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="species" species="nebulosus">Pristimantis nebulosus (Henle, 1992)</taxonomicName>
</paragraph>
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<paragraph id="1F9618437334BB8DFFF185B08CA08D86" pageId="0" pageNumber="213">Relationships.</paragraph>
<paragraph id="4D30CF113E8251DA58D94F99596B7954" pageId="0" pageNumber="213">
The Maximum Likelihood tree, based on an alignment of 584 bp of the 16S gene, is insufficient to establish a reliable hypothesis of phylogenetic relationships among the
<taxonomicName id="6FF64D823135D09F673FB02A0533C58A" authorityName="Jimenez de la Espada" authorityYear="1870" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="genus">
<emphasis id="DA8477F1B8B2E4FCEF489245D2A09C28" italics="true" pageId="0" pageNumber="213">Pristimantis</emphasis>
</taxonomicName>
samples included (Fig.
<figureCitation id="7DEE8D7FF3184F778132D2174988562F" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Maximum Likelihood tree inferred from a 584 bp fragment of the mitochondrial 16 S gene of Pristimantis species. Values at nodes are bootstrap proportions in percent (values below 49 % not shown). Newly obtained sequences for this study are in red font. For other samples used, species names are followed by GenBank accession number and locality. Yunganastes pluvicanorus was used as outgroup (not shown for graphic reasons only). Inset photos depict Pristimantis bipunctatus (MUSM 31120), P. nebulosus (MUSM 40299, FGZC 6254) and P. symptosus sp. nov. (MUSM 40256, FGZC 6207) in life." figureDoi="10.3897/zse.98.84963.figure4" httpUri="https://binary.pensoft.net/fig/705931" pageId="0" pageNumber="213">4</figureCitation>
). The
<taxonomicName id="6C4FB0F811F0E999D22B5E735244FDF5" lsidName="P. conspicillatus" pageId="0" pageNumber="213" rank="species" species="conspicillatus">
<emphasis id="E55B5C2AEA6830899E1D04CB2AFC0F89" italics="true" pageId="0" pageNumber="213">P. conspicillatus</emphasis>
</taxonomicName>
species group is recovered paraphyletic with respect to the sample of
<taxonomicName id="012F7862DCFBCEBBB736B22E4C82860C" lsidName="P. gaigeae" pageId="0" pageNumber="213" rank="species" species="gaigeae">
<emphasis id="440C9BEC60B2F31153CBEC09FDD151BE" italics="true" pageId="0" pageNumber="213">P. gaigeae</emphasis>
</taxonomicName>
from Panama, a member of the
<taxonomicName id="9D2B4771AD5AD6EA7DED57ED1C9D9EA2" lsidName="P. conspicillatus" pageId="0" pageNumber="213" rank="species" species="conspicillatus">
<emphasis id="7D3BA74A74A6604854BCC55B46E33C96" italics="true" pageId="0" pageNumber="213">P. conspicillatus</emphasis>
</taxonomicName>
group sensu
<bibRefCitation id="B9F9483FB2AD056FB327C00B00B67BB7" DOI="https://doi.org/10.11646/zootaxa.1737.1.1" author="Hedges, SB" journalOrPublisher="Zootaxa" pageId="0" pageNumber="213" pagination="1 - 182" refId="B18" refString="Hedges, SB, Duellman, WE, Heinicke, MP, 2008. New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa 1737 (1): 1 - 182, DOI: https://doi.org/10.11646/zootaxa.1737.1.1" title="New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and conservation." url="https://doi.org/10.11646/zootaxa.1737.1.1" volume="1737" year="2008">Hedges et al. (2008)</bibRefCitation>
and
<bibRefCitation id="502DBB23C70714A189234D834575C294" DOI="https://doi.org/10.11646/zootaxa.3825.1.1" author="Padial, JM" journalOrPublisher="Zootaxa" pageId="0" pageNumber="213" pagination="1 - 132" refId="B44" refString="Padial, JM, Grant, T, Frost, DR, 2014. Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria. Zootaxa 3825: 1 - 132, DOI: https://doi.org/10.11646/zootaxa.3825.1.1" title="Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria." url="https://doi.org/10.11646/zootaxa.3825.1.1" volume="3825" year="2014">Padial et al. (2014)</bibRefCitation>
. Together with species representing the
<taxonomicName id="224763DF317973769624E9233A56D392" lsidName="P. lacrimosus" pageId="0" pageNumber="213" rank="species" species="lacrimosus">
<emphasis id="2022905F8893AB42D93FDC35CD04E0EC" italics="true" pageId="0" pageNumber="213">P. lacrimosus</emphasis>
</taxonomicName>
,
<taxonomicName id="17B4DE7C085A7200C79C226C44581B62" lsidName="P. ridens" pageId="0" pageNumber="213" rank="species" species="ridens">
<emphasis id="9F9D65F039AA0A7C1FC6568C33C794CC" italics="true" pageId="0" pageNumber="213">P. ridens</emphasis>
</taxonomicName>
and
<taxonomicName id="C5D610B6962478992A6937DF6E0B21DF" lsidName="P. danae" pageId="0" pageNumber="213" rank="species" species="danae">
<emphasis id="BA49A1F891B49BD99E6348DCAC0152CF" italics="true" pageId="0" pageNumber="213">P. danae</emphasis>
</taxonomicName>
species groups,
<taxonomicName id="8084779F6FEF88E2C788F2F9418EE64D" lsidName="P. gaigeae" pageId="0" pageNumber="213" rank="species" species="gaigeae">
<emphasis id="551F734793FFDB835F1B35B10984BBB8" italics="true" pageId="0" pageNumber="213">P. gaigeae</emphasis>
</taxonomicName>
is placed sister to the remaining species of the
<taxonomicName id="7B087039C60A6BBF09E8FA6A587E9B73" lsidName="P. conspicillatus" pageId="0" pageNumber="213" rank="species" species="conspicillatus">
<emphasis id="8F7ACD99864DB63AB84747DD5F139CFE" italics="true" pageId="0" pageNumber="213">P. conspicillatus</emphasis>
</taxonomicName>
group. Within the
<taxonomicName id="C001CEB3E3B1FC1F126862EEE4CF950C" lsidName="P. conspicillatus" pageId="0" pageNumber="213" rank="species" species="conspicillatus">
<emphasis id="68B67B8E3449ACE46A5F4A0EC6FB82F7" italics="true" pageId="0" pageNumber="213">P. conspicillatus</emphasis>
</taxonomicName>
group, most relationships are not resolved and basal nodes did not receive significant support. However, the three specimens of the focal
<taxonomicName id="C2429A9F6FBA9ECC34DEB5290A97322B" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="DED8B5A4D3F48512C6B3736BDCA1A7FF" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
are recovered with high support (bootstrap value 98%) as members of a clade containing nominal
<taxonomicName id="06B97031EE4C672F1E14E87D8AE644AA" lsidName="P. bipunctatus" pageId="0" pageNumber="213" rank="species" species="bipunctatus">
<emphasis id="062BBE330B18068D5F33892C6713C0A8" italics="true" pageId="0" pageNumber="213">P. bipunctatus</emphasis>
</taxonomicName>
and an undescribed candidate species, with
<taxonomicName id="674B0675038E691E22CDD2E93F927590" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="99E3026595C045DDBF2ED537B49D24DC" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
placed sister to nominal
<taxonomicName id="6EBC5A8AF75AB7A337C98DE0E80F267D" lsidName="P. bipunctatus" pageId="0" pageNumber="213" rank="species" species="bipunctatus">
<emphasis id="85C240310DA1FBD8AD1571B82758D60F" italics="true" pageId="0" pageNumber="213">P. bipunctatus</emphasis>
</taxonomicName>
(without significant support; 36%). The clade containing
<taxonomicName id="4D8CFFDDF9ED90C912C05022C73A5015" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="E8B4315552916A27753D69B872BFE263" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
,
<taxonomicName id="E7DC1562A99E3662F46B3C9A9233DD68" lsidName="P. bipunctatus" pageId="0" pageNumber="213" rank="species" species="bipunctatus">
<emphasis id="10793B5113FA4CD1BDA02EBD0A40A4D3" italics="true" pageId="0" pageNumber="213">P. bipunctatus</emphasis>
</taxonomicName>
, and the candidate species are recovered sister (76%) to a clade containing two samples of
<taxonomicName id="7C505FCE598AF3D79B85A6343028045B" lsidName="P. skydmainos" pageId="0" pageNumber="213" rank="species" species="skydmainos">
<emphasis id="EE3FA45CFB4858C962A2E36140CA6262" italics="true" pageId="0" pageNumber="213">P. skydmainos</emphasis>
</taxonomicName>
(a close relationship of
<taxonomicName id="63EE08D2994BD0EBE0C748B9CB2E1B16" lsidName="P. bipunctatus" pageId="0" pageNumber="213" rank="species" species="bipunctatus">
<emphasis id="844ACD67F760C773DCD0FAEB9967C43E" italics="true" pageId="0" pageNumber="213">P. bipunctatus</emphasis>
</taxonomicName>
and
<taxonomicName id="6B0583557DA5718A7BF2F6A8B20A6A6B" lsidName="P. skydmainos" pageId="0" pageNumber="213" rank="species" species="skydmainos">
<emphasis id="CC92F10E290D6F2FAFA1ADEAC8C7AEEE" italics="true" pageId="0" pageNumber="213">P. skydmainos</emphasis>
</taxonomicName>
has already been revealed by
<bibRefCitation id="9867E798F074E8E6613CC3D0E7D5F67C" DOI="https://doi.org/10.11646/zootaxa.3825.1.1" author="Padial, JM" journalOrPublisher="Zootaxa" pageId="0" pageNumber="213" pagination="1 - 132" refId="B44" refString="Padial, JM, Grant, T, Frost, DR, 2014. Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria. Zootaxa 3825: 1 - 132, DOI: https://doi.org/10.11646/zootaxa.3825.1.1" title="Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria." url="https://doi.org/10.11646/zootaxa.3825.1.1" volume="3825" year="2014">Padial et al. 2014</bibRefCitation>
). Our
<taxonomicName id="4336D0CF1EFDCBF57A6DAB19453EE798" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="430814AB2A3EA29371CC3C128B0B7D40" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
specimens had uncorrected p-distances in the studied 16S gene fragment of 5.4-6.2% to
<taxonomicName id="A8AB02C393A53851279053F71942F272" lsidName="P. bipunctatus" pageId="0" pageNumber="213" rank="species" species="bipunctatus">
<emphasis id="8D7A82D31C4A139EAB24767157D96CFA" italics="true" pageId="0" pageNumber="213">P. bipunctatus</emphasis>
</taxonomicName>
(with the greatest distance of 6.2% referring to the
<taxonomicName id="D10386C2F5136523F23544F6C543681C" lsidName="P. bipunctatus" pageId="0" pageNumber="213" rank="species" species="bipunctatus">
<emphasis id="FFC467AFB640BDC834D2E185CA46AAE8" italics="true" pageId="0" pageNumber="213">P. bipunctatus</emphasis>
</taxonomicName>
holotype KU291638) and 4.1% to the mentioned candidate species (Table
<tableCitation id="2021A93F8C8F7781A8BC192B6A22C6AB" captionStart="Table 2" captionStartId="T2" captionText="Table 2. Estimates of uncorrected p-distances in a 491 bp fragment of the mitochondrial 16 S gene among selected samples of the Pristimantis conspicillatus species group, calculated with MEGA 7 (all ambiguous positions were removed for each sequence pair)." httpUri="http://table.plazi.org/id/65FB5CBE73548D2CDB3D7308B54A4C0C" pageId="0" pageNumber="213" tableUuid="65FB5CBE73548D2CDB3D7308B54A4C0C">2</tableCitation>
). P-distances to other species in the
<taxonomicName id="82B13E7AFA5CD68E241808DC8F5247DF" lsidName="P. conspicillatus" pageId="0" pageNumber="213" rank="species" species="conspicillatus">
<emphasis id="B64DDB620A7C7E786DEB676B35A82B63" italics="true" pageId="0" pageNumber="213">P. conspicillatus</emphasis>
</taxonomicName>
species group range from 5.8-17.8%.
</paragraph>
<caption id="65FB5CBE73548D2CDB3D7308B54A4C0C" ID-Table-UUID="65FB5CBE73548D2CDB3D7308B54A4C0C" httpUri="http://table.plazi.org/id/65FB5CBE73548D2CDB3D7308B54A4C0C" pageId="0" pageNumber="213" start="Table 2" startId="T2">
<paragraph id="69A874A7374E24B6133722A26500F032" pageId="0" pageNumber="213">
<emphasis id="1C885058C366D3BA4D6346FC74483ED2" bold="true" pageId="0" pageNumber="213">Table 2.</emphasis>
Estimates of uncorrected p-distances in a 491 bp fragment of the mitochondrial 16S gene among selected samples of the
<taxonomicName id="BC664B7B2E25099C18D928FA0C1B5F3C" baseAuthorityName="Gunther" baseAuthorityYear="1858" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis conspicillatus" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="species" species="conspicillatus">
<emphasis id="344F2B674CDA779D0D237C5B1941FF49" italics="true" pageId="0" pageNumber="213">Pristimantis conspicillatus</emphasis>
</taxonomicName>
species group, calculated with MEGA7 (all ambiguous positions were removed for each sequence pair).
</paragraph>
</caption>
<paragraph id="4A8DF49999B96583E1BBB6E32877A2BD" pageId="0" pageNumber="213">
<table id="222EB22C04BEDA541609D9FBC31856AB" pageId="0" pageNumber="213">
<tr id="354FBACBA8F29E416DF075017CD4EF55" pageId="0" pageNumber="213">
<th id="FC7B1F2AC91A149059C912B067A3A90B" colspan="1" isEmpty="true" pageId="0" pageNumber="213" rowspan="1">-</th>
<th id="4C1DEDE328E4F678EDBEB9741374C7C1" colspan="1" pageId="0" pageNumber="213" rowspan="1">FGZC 6253</th>
<th id="0A81C6568E383A19733467046A7092F6" colspan="1" pageId="0" pageNumber="213" rowspan="1">FGZC 6254</th>
<th id="1CC4B1ACC2CAF3996CD980945DFF2A52" colspan="1" pageId="0" pageNumber="213" rowspan="1">FGZC 6281</th>
<th id="E09728124A9F7D96928DA6FC56D470A7" colspan="1" pageId="0" pageNumber="213" rowspan="1">FGZC 6207</th>
<th id="FCE9151FB667742182ED7B91FB24B18C" colspan="1" pageId="0" pageNumber="213" rowspan="1">KU 291638</th>
<th id="CA9122CB9C8597501BDE5AC9D3351D5E" colspan="1" pageId="0" pageNumber="213" rowspan="1">MUSM 31179</th>
<th id="D86B47B5F00A81C09182DB0C847F4277" colspan="1" pageId="0" pageNumber="213" rowspan="1">MUSM 31120</th>
</tr>
<tr id="587CB2C1610C9E7209618F415403DDD4" pageId="0" pageNumber="213">
<td id="9603B32FAE0DF0E9B5CC25F760A1B888" colspan="1" pageId="0" pageNumber="213" rowspan="1">
<taxonomicName id="2ACF647CC9DE378AF389BCC46CE487EA" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="EE9C29BD5265118D2BB57F488E82CAB5" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
FGZC 6253 (Abra La Divisoria)
</td>
<td id="FDF9E9F1238A2707C70C6CFC69860AAB" colspan="1" isEmpty="true" pageId="0" pageNumber="213" rowspan="1">-</td>
<td id="7BC842FA3EAFBD3FA4918099C643F21F" colspan="1" isEmpty="true" pageId="0" pageNumber="213" rowspan="1">-</td>
<td id="A1CFA3F6AE7A1E0FE02890CF43AEFC72" colspan="1" isEmpty="true" pageId="0" pageNumber="213" rowspan="2">-</td>
<td id="7E5600F0C84D852B56443B3518E9C43A" colspan="1" isEmpty="true" pageId="0" pageNumber="213" rowspan="3">-</td>
<td id="BFAF1A16F9E5DC5B7B314816A74570A7" colspan="1" isEmpty="true" pageId="0" pageNumber="213" rowspan="4">-</td>
<td id="641F718E2B9B362614046982F0C03008" colspan="1" isEmpty="true" pageId="0" pageNumber="213" rowspan="5">-</td>
<td id="F5E39FC800E737DE305A1AD6A009C12D" colspan="1" isEmpty="true" pageId="0" pageNumber="213" rowspan="6">-</td>
</tr>
<tr id="BE469498A1BE0CD55BDF98A005E4381B" pageId="0" pageNumber="213">
<td id="14D7207E9BD42BD41CF349E157ED8DB3" colspan="1" pageId="0" pageNumber="213" rowspan="1">
<taxonomicName id="B7D0F76C0B78FE2E7C1F8D05ACC2A5DF" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="B22D56EC96E48F117BD33D292D251538" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
FGZC 6254 (Abra La Divisoria)
</td>
<td id="677EBC220B60D9E2D6383387FD72E411" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.000</td>
<td id="86EEAB196C02A9ECCD6854BCE1ABD50B" colspan="1" isEmpty="true" pageId="0" pageNumber="213" rowspan="1">-</td>
</tr>
<tr id="2244CC8A0F7AFBC1611CA7EDA4E6BB13" pageId="0" pageNumber="213">
<td id="0D7D28F5964A65897814BB33575E6BC1" colspan="1" pageId="0" pageNumber="213" rowspan="1">
<taxonomicName id="4BFFEFD958A7F4CB254DFBB6006CFA7E" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="2F77DB8994F1361BDC7ABB7C366FF4EE" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
FGZC 6281 (Abra La Divisoria)
</td>
<td id="CEF274C5DDF0C6618EAE946C42413A4B" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.000</td>
<td id="3B0808C73988760E7A091ADA575362BA" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.000</td>
<td id="CBAF9F44660B516733466F9E9AFB2B87" colspan="1" isEmpty="true" pageId="0" pageNumber="213" rowspan="1">-</td>
</tr>
<tr id="4B8D6F5E6912B01E73259F45958BD97B" pageId="0" pageNumber="213">
<td id="0CE265BDEDA4330FC479BCD513F721F2" colspan="1" pageId="0" pageNumber="213" rowspan="1">
<taxonomicName id="08617FB3097026F5E0C3FBF58D9C03DB" lsidName="P. symptosus" pageId="0" pageNumber="213" rank="species" species="symptosus">
<emphasis id="6E3D454803762C58DE01C21ED529175F" italics="true" pageId="0" pageNumber="213">P. symptosus</emphasis>
</taxonomicName>
FGZC 6207 (Cordillera de Carpish)
</td>
<td id="99DBA442D3A5B46C4FC5D55A6E007897" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.041</td>
<td id="AF100DE555FDF72674C966774659AA3E" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.041</td>
<td id="D43D908765CB9CF1A95344470F296824" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.041</td>
<td id="4D42969F2C8DC9B0F2BF9C360E58C9B9" colspan="1" isEmpty="true" pageId="0" pageNumber="213" rowspan="1">-</td>
</tr>
<tr id="590F9C5B4C5EAD01B30FDE245107D37E" pageId="0" pageNumber="213">
<td id="22C5507CE80B3FF859458C1B4CC03B1F" colspan="1" pageId="0" pageNumber="213" rowspan="1">
<taxonomicName id="B3CBD06FBAEE2DA36525952307E3CE62" lsidName="P. bipunctatus" pageId="0" pageNumber="213" rank="species" species="bipunctatus">
<emphasis id="2024E25242F321A18B73D5CA497B749C" italics="true" pageId="0" pageNumber="213">P. bipunctatus</emphasis>
</taxonomicName>
KU 291638 (E of Oxapampa)
</td>
<td id="CA95C1434FD44C09B681037E2A2C32DA" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.062</td>
<td id="A1442AB8B241C210AF7B11A527374BBA" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.062</td>
<td id="7DDC3D87CD2605E84C125197D3E5BD00" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.062</td>
<td id="1C79462761C371489A0A5FB1454FB482" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.041</td>
<td id="DF7E3523AE868A9A4CA19242463DBDB7" colspan="1" isEmpty="true" pageId="0" pageNumber="213" rowspan="1">-</td>
</tr>
<tr id="139F40521617B8DA959265CE848F007D" pageId="0" pageNumber="213">
<td id="AE653AA18370A56CC232D2D14C0DA64A" colspan="1" pageId="0" pageNumber="213" rowspan="1">
<taxonomicName id="2BAF75A91285855477881C8C90074BD4" lsidName="P. bipunctatus" pageId="0" pageNumber="213" rank="species" species="bipunctatus">
<emphasis id="6C0522AE3FAE27AFADA796ECDC55F445" italics="true" pageId="0" pageNumber="213">P. bipunctatus</emphasis>
</taxonomicName>
MUSM 31179 (Pui Pui Forest)
</td>
<td id="B8F61474787C339C2DE7150F5DE48B5F" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.054</td>
<td id="EA41C92AB1E1BC164A04E2C6E9962881" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.054</td>
<td id="A4E0567C81FEB03AF19B358BF58A67C6" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.054</td>
<td id="703D4A6C0D526111C834D28AA5DA2293" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.033</td>
<td id="419C6827E0C5A83770BFF209E0B6EFA0" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.037</td>
<td id="CAF64858B77411E58A78D0A847F069A7" colspan="1" isEmpty="true" pageId="0" pageNumber="213" rowspan="1">-</td>
</tr>
<tr id="05B01343F2F4D55F14C4C343E5A995D6" pageId="0" pageNumber="213">
<td id="56E7BEB18168D74D8FBF9D0F4CA6941E" colspan="1" pageId="0" pageNumber="213" rowspan="1">
<taxonomicName id="D19B06FDB610C536E6603793E772E2B4" lsidName="P. bipunctatus" pageId="0" pageNumber="213" rank="species" species="bipunctatus">
<emphasis id="FD21BEE2292F18CA6582D0DFAD2D1EE7" italics="true" pageId="0" pageNumber="213">P. bipunctatus</emphasis>
</taxonomicName>
MUSM 31120 (Yanachaga
<normalizedToken id="F599F9C01BD5A012C901D8229F65C1EF" originalValue="Chemillén">Chemillen</normalizedToken>
)
</td>
<td id="F1AE46AA866D86D3EB27E5CE326223D2" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.058</td>
<td id="8E6A21B29CB355390ED6BDE7111BD482" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.058</td>
<td id="42FA73240873613E86259DC15FEED630" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.058</td>
<td id="EE9D5405DF2D842421BD51819F979920" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.039</td>
<td id="4387B7EA3A291D076F8C115BEC36375F" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.006</td>
<td id="2F6C309BABBAD4233E0E0360F0DFDA45" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.031</td>
<td id="F8097DB198CC75666826DD9D059E2027" colspan="1" isEmpty="true" pageId="0" pageNumber="213" rowspan="1">-</td>
</tr>
<tr id="5454614A952044F8DA3D773DCB7C4376" pageId="0" pageNumber="213">
<td id="7D3597E97CF7376057B29077BA67FFA0" colspan="1" pageId="0" pageNumber="213" rowspan="1">
<taxonomicName id="36272B98D63C733B6823916CA696CBC2" lsidName="P. bipunctatus" pageId="0" pageNumber="213" rank="species" species="bipunctatus">
<emphasis id="EADB9F1FEB102674A5DE7A73FCFC936F" italics="true" pageId="0" pageNumber="213">P. bipunctatus</emphasis>
</taxonomicName>
NMP6V/75062 (Pui Pui Forest)
</td>
<td id="428568A5A164AF4A473CF69F6E4A8AA3" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.054</td>
<td id="A02EA802598FCF20C4074FD555D1DE96" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.054</td>
<td id="D883A6C33F8A768944BAEAEF2D1E5606" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.054</td>
<td id="11372B738764A9FCAB9F7EB1278AAAC4" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.033</td>
<td id="A02CBA4C4C55E9A0B2C36FD2498AD994" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.037</td>
<td id="34EA27078BB97BB42CE5580D9A2E4ED2" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.000</td>
<td id="A5D820E49B91B4D3804B4CD400F69FE6" colspan="1" pageId="0" pageNumber="213" rowspan="1">0.031</td>
</tr>
</table>
</paragraph>
<caption id="CAEAF9A599AD6EA132307AB979BF09FA" doi="10.3897/zse.98.84963.figure4" httpUri="https://binary.pensoft.net/fig/705931" pageId="0" pageNumber="213" start="Figure 4" startId="F4">
<paragraph id="270F062C1A482EDF5D808C99B6E2C50C" pageId="0" pageNumber="213">
<emphasis id="4EC049A7D349AF5EDEC0FB66C10A5765" bold="true" pageId="0" pageNumber="213">Figure 4.</emphasis>
Maximum Likelihood tree inferred from a 584 bp fragment of the mitochondrial 16S gene of
<taxonomicName id="BB0405F44DADBC404EDC9ED800894B28" authorityName="Jimenez de la Espada" authorityYear="1870" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="genus">
<emphasis id="6057B5E108956AF2C34152280BFA0A8C" italics="true" pageId="0" pageNumber="213">Pristimantis</emphasis>
</taxonomicName>
species. Values at nodes are bootstrap proportions in percent (values below 49% not shown). Newly obtained sequences for this study are in red font. For other samples used, species names are followed by GenBank accession number and locality.
<taxonomicName id="7CE37884A503B08F4B95975AFC13F9D4" baseAuthorityName="De la Riva &amp; Lynch" baseAuthorityYear="1997" class="Amphibia" family="Craugastoridae" genus="Yunganastes" kingdom="Animalia" lsidName="Yunganastes pluvicanorus" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="species" species="pluvicanorus">
<emphasis id="1C4E06C662DA36D6C14CDBF1E1CC292B" italics="true" pageId="0" pageNumber="213">Yunganastes pluvicanorus</emphasis>
</taxonomicName>
was used as outgroup (not shown for graphic reasons only). Inset photos depict
<taxonomicName id="908C06BBFACD90170B8E825AD99226C1" baseAuthorityName="Duellman &amp; Hedges" baseAuthorityYear="2005" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis bipunctatus" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="species" species="bipunctatus">
<emphasis id="120F99242884D4AE11AE8014127B7918" italics="true" pageId="0" pageNumber="213">Pristimantis bipunctatus</emphasis>
</taxonomicName>
(MUSM 31120),
<taxonomicName id="2D72A1CE6C173A8432BC647A4D0705EB" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="D4FAF2585E502DDD153DBB254D4E6B5D" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
(MUSM 40299, FGZC 6254) and
<taxonomicName id="D8A2EB9FE0EDE82D600191423613AAE5" lsidName="P. symptosus" pageId="0" pageNumber="213" rank="species" species="symptosus">
<emphasis id="1E2732CA3BABC34B55486B05873E5B97" italics="true" pageId="0" pageNumber="213">P. symptosus</emphasis>
</taxonomicName>
sp. nov. (MUSM 40256, FGZC 6207) in life.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="176EA6AD47D47856E3D60AB8779B01F7" pageId="0" pageNumber="213" type="diagnosis">
<paragraph id="98FEC8E135F4E68CFC8EEA4E1334A8AB" pageId="0" pageNumber="213">Diagnosis.</paragraph>
<paragraph id="0A6C11C5603912EC8016531B9F776DC5" pageId="0" pageNumber="213">
Based on the holotype and the newly collected material, we provide the following updated diagnosis for
<taxonomicName id="B10DC7F9CD50266029679A019097756C" baseAuthorityName="Henle" baseAuthorityYear="1992" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis nebulosus" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="species" species="nebulosus">
<emphasis id="4740DA5D8D76EE986CF57D26E48966BB" italics="true" pageId="0" pageNumber="213">Pristimantis nebulosus</emphasis>
</taxonomicName>
(Henle, 1992): a medium-sized species of the
<taxonomicName id="7045C563EE5FBD5ABBB759FAA8E18ACE" baseAuthorityName="Gunther" baseAuthorityYear="1858" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis conspicillatus" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="species" species="conspicillatus">
<emphasis id="5D6B058452C80428D38A9727ED2C26FF" italics="true" pageId="0" pageNumber="213">Pristimantis conspicillatus</emphasis>
</taxonomicName>
species group (based on molecular relationships and morphological similarity), with 27.7-29.1 mm SVL in adult males (n = 3), characterised by: (1) skin on dorsum shagreen, with few scattered small tubercles; flanks shagreen; throat, chest and venter weakly areolate, smooth only in the middle and anterior part; posterior surfaces of limbs smooth; discoidal fold conspicuous; dorsolateral folds present, but low, partly interrupted; dorsal folds absent; two prominent postrictal conical tubercles present; upper eyelid lacking tubercles and granules; (2) tympanic membrane and annulus distinct, nearly round, their length slightly less than half of eye diameter; supratympanic fold long, prominent, curved, surrounding upper tympanum; (3) head slightly longer than wide; snout subacuminate in dorsal view, rounded in lateral profile; canthus rostralis straight in dorsal view, sharp in profile; (4) cranial crests absent; (5) dentigerous process of vomers large, triangular in outline, oblique, situated posteromedial to choanae; (6) males with vocal slits, single subgular vocal sac, and nuptial asperities on dorsal surface of thenar tubercle; (7) hands with long and slender fingers, first finger only very slightly longer than second; subarticular tubercles subconical, prominent; supernumerary tubercles round, smaller than subarticular tubercles; palmar tubercle bifid; thenar tubercle elongated; terminal discs of inner two fingers enlarged and round, those of external fingers enlarged, ovate to triangular, about twice the width of digit proximal to disc; circumferential grooves conspicuous, ungual flap slightly indented; lateral fringes and keels on fingers absent; basal webbing between fingers II and III; (8) ulnar tubercles absent; (9) tubercles on heel and tarsus absent, tarsal fold present, narrow and oblique; (10) inner metatarsal tubercle ovate, prominent; outer metatarsal tubercle round, flat, inconspicuous; supernumerary tubercles absent; (11) toes long and slender (FootL 54-56% SVL); narrow lateral fringes weakly expressed, basal toe webbing present; toe V reaching mid-level of penultimate subarticular tubercle of toe IV; toe V slightly longer than toe III; tips of toes rounded to slightly truncate, enlarged; circumferential grooves evident, ungual flap not indented; (12) in life, dorsal colouration overall brown of different shades (sometimes with orange to reddish tint), with dark brown chevrons or irregular flecks on dorsum, outlined with cream; dark brown bars on dorsal surfaces of arms and legs, outlined with cream; brown interorbital bar, at least anteriorly outlined with cream; a pair of black occipital spots: broad black supratympanic stripe; greyish-black canthal stripe; lips dark brown, barred with cream; tip of snout with triangular cream-coloured fleck; belly yellowish-cream; throat yellow or yellowish-cream, mottled with grey spots; ventral surfaces of thighs and shanks orange brown to reddish-brown; posterior surface of thighs reddish-brown with small light spots; iris bronze, with some dark brown reticulation and a median reddish-brown streak; posterior iris periphery turquoise; bones white; (13) advertisement call consisting of a single pulsed note of 206-382 ms duration with a pulse rate of 156-174 pulses/second, emitted in short series (see below).
</paragraph>
</subSubSection>
<subSubSection id="59AC94280779E0A27434B00743F4E606" pageId="0" pageNumber="213" type="comparisons">
<paragraph id="37010E5CDA697149BD80DEB0D6301106" pageId="0" pageNumber="213">Comparison.</paragraph>
<paragraph id="ADB949479A29174957A3671403850041" pageId="0" pageNumber="213">
Among the now 41 recognised species in the
<taxonomicName id="4A68AF90FD6E08C393DA3026017D77A8" lsidName="P. conspicillatus" pageId="0" pageNumber="213" rank="species" species="conspicillatus">
<emphasis id="5E7CAC164820D0710F624355A6E73EB6" italics="true" pageId="0" pageNumber="213">P. conspicillatus</emphasis>
</taxonomicName>
species group (
<bibRefCitation id="F67821CBA88E4571F7CD797AF1458E5E" DOI="https://doi.org/10.11646/zootaxa.3825.1.1" author="Padial, JM" journalOrPublisher="Zootaxa" pageId="0" pageNumber="213" pagination="1 - 132" refId="B44" refString="Padial, JM, Grant, T, Frost, DR, 2014. Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria. Zootaxa 3825: 1 - 132, DOI: https://doi.org/10.11646/zootaxa.3825.1.1" title="Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria." url="https://doi.org/10.11646/zootaxa.3825.1.1" volume="3825" year="2014">Padial et al. 2014</bibRefCitation>
,
<bibRefCitation id="D02CAFBCA9A6C9585EA1B32ED2D103DB" DOI="https://doi.org/10.2992/007.083.0302" author="Padial, JM" journalOrPublisher="Annals of Carnegie Museum" pageId="0" pageNumber="213" pagination="207 - 218" refId="B45" refString="Padial, JM, Gagliardi-Urrutia, G, Chaparro, JC, Gutierrez, RC, 2016. A new species of the Pristimantis conspicillatus group from the Peruvian Amazon (Anura: Craugastoridae). Annals of Carnegie Museum 83 (3): 207 - 218, DOI: https://doi.org/10.2992/007.083.0302" title="A new species of the Pristimantis conspicillatus group from the Peruvian Amazon (Anura: Craugastoridae)." url="https://doi.org/10.2992/007.083.0302" volume="83" year="2016">2016</bibRefCitation>
;
<bibRefCitation id="AEBDEF2D3A90C0CF58E7E1DB785B01F7" DOI="https://doi.org/10.3897/zookeys.687.13221" author="de Oliveira, EA" journalOrPublisher="ZooKeys" pageId="0" pageNumber="213" pagination="101 - 129" refId="B6" refString="de Oliveira, EA, Rodrigues, LR, Kaefer, IL, Pinto, KC, Hernandez-Ruz, EJ, 2017. A new species of Pristimantis from eastern Brazilian Amazonia (Anura, Craugastoridae). ZooKeys 687: 101 - 129, DOI: https://doi.org/10.3897/zookeys.687.13221" title="A new species of Pristimantis from eastern Brazilian Amazonia (Anura, Craugastoridae)." url="https://doi.org/10.3897/zookeys.687.13221" volume="687" year="2017">de Oliveira et al. 2017</bibRefCitation>
,
<bibRefCitation id="DDC3FAF0DBD7ABB3B112C025FA87BDF5" DOI="https://doi.org/10.1371/journal.pone.0229971" author="de Oliveira, EA" journalOrPublisher="ZooKeys" pageId="0" pageNumber="213" refId="B7" refString="de Oliveira, EA, da Silva, LA, Silva, EAP, Guimaraes, KLA, Penhacek, M, Martinez, JG, Rodrigues, LR, Santana, DJ, Hernandez-Ruz, EJ, 2020. Four new species of Pristimantis Jimenez de la Espada, 1870 (Anura: Craugastoridae) in the eastern Amazon. PLoS ONE 15(11): e0243182. https://doi.org/10.1371/journal.pone.0229971" title="Four new species of Pristimantis Jimenez de la Espada, 1870 (Anura: Craugastoridae) in the eastern Amazon. PLoS ONE 15 (11): e 0243182." url="https://doi.org/10.1371/journal.pone.0229971" year="2020">2020</bibRefCitation>
;
<bibRefCitation id="B08B3E038418ED01E578B35F84C512B5" DOI="https://doi.org/10.1670/19-114" author="Taucce, PPG" journalOrPublisher="Journal of Herpetology" pageId="0" pageNumber="213" pagination="245 - 257" refId="B52" refString="Taucce, PPG, Nascimento, JS, Trevisan, CC, Leite, FSF, Santana, DJ, Haddad, CFB, Napoli, MF, 2020. A new rupicolous species of the Pristimantis conspicillatus group (Anura: Brachycephaloidea: Craugastoridae) from central Bahia, Brazil. Journal of Herpetology 54 (2): 245 - 257, DOI: https://doi.org/10.1670/19-114" title="A new rupicolous species of the Pristimantis conspicillatus group (Anura: Brachycephaloidea: Craugastoridae) from central Bahia, Brazil." url="https://doi.org/10.1670/19-114" volume="54" year="2020">Taucce et al. 2020</bibRefCitation>
), 12 species, including
<taxonomicName id="958ADB49FE5368C0D4C06B8FB0FE2057" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="E616791AA86CEC879467476ACEBF86A7" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
, exhibit dorsolateral folds (
<bibRefCitation id="83303F43C287DC1C9E48621E5C2D8D08" author="Duellman, WE" journalOrPublisher="Natur und Tier Verlag, Muenster" pageId="0" pageNumber="213" refId="B10" refString="Duellman, WE, Lehr, E, 2009. Terrestrial-breeding frogs (Strabomantidae) in Peru. Natur und Tier Verlag, Muenster" title="Terrestrial-breeding frogs (Strabomantidae) in Peru." year="2009">Duellman and Lehr 2009</bibRefCitation>
;
<bibRefCitation id="4DB6F6717E42FC6C030D46AE92559A32" DOI="https://doi.org/10.2992/007.083.0302" author="Padial, JM" journalOrPublisher="Annals of Carnegie Museum" pageId="0" pageNumber="213" pagination="207 - 218" refId="B45" refString="Padial, JM, Gagliardi-Urrutia, G, Chaparro, JC, Gutierrez, RC, 2016. A new species of the Pristimantis conspicillatus group from the Peruvian Amazon (Anura: Craugastoridae). Annals of Carnegie Museum 83 (3): 207 - 218, DOI: https://doi.org/10.2992/007.083.0302" title="A new species of the Pristimantis conspicillatus group from the Peruvian Amazon (Anura: Craugastoridae)." url="https://doi.org/10.2992/007.083.0302" volume="83" year="2016">Padial et al. 2016</bibRefCitation>
). Among these,
<taxonomicName id="BC59E458D1C29C4FB0BFA5132C503FB1" lsidName="P. avicuporum" pageId="0" pageNumber="213" rank="species" species="avicuporum">
<emphasis id="0441A0EE7B0FC05A23AD72BB61163715" italics="true" pageId="0" pageNumber="213">P. avicuporum</emphasis>
</taxonomicName>
mainly differs by the presence of an interocular dermal fold, fingers bearing lateral fringes, and an upper eyelid with numerous tubercles (
<bibRefCitation id="C769ABB149D41070038CBF7D8F155F3A" DOI="https://doi.org/10.5962/bhl.title.16169" author="Duellman, WE" journalOrPublisher="Scientific Papers of the Natural History Museum, the University of Kansas" pageId="0" pageNumber="213" pagination="1 - 78" refId="B11" refString="Duellman, WE, Pramuk, JB, 1999. Frogs of the genus Eleutherodactylus (Anura: Leptodactylidae) in the Andes of northern Peru. Scientific Papers of the Natural History Museum, the University of Kansas 13: 1 - 78, DOI: https://doi.org/10.5962/bhl.title.16169" title="Frogs of the genus Eleutherodactylus (Anura: Leptodactylidae) in the Andes of northern Peru." url="https://doi.org/10.5962/bhl.title.16169" volume="13" year="1999">Duellman and Pramuk 1999</bibRefCitation>
;
<bibRefCitation id="B0D88A557C932E0CE7E7FA639373B655" author="Duellman, WE" journalOrPublisher="Natur und Tier Verlag, Muenster" pageId="0" pageNumber="213" refId="B10" refString="Duellman, WE, Lehr, E, 2009. Terrestrial-breeding frogs (Strabomantidae) in Peru. Natur und Tier Verlag, Muenster" title="Terrestrial-breeding frogs (Strabomantidae) in Peru." year="2009">Duellman and Lehr 2009</bibRefCitation>
);
<taxonomicName id="645A023D0D2137866BC40DF5FC81DF50" lsidName="P. buccinator" pageId="0" pageNumber="213" rank="species" species="buccinator">
<emphasis id="D21C6F897082AE978109B010347945D8" italics="true" pageId="0" pageNumber="213">P. buccinator</emphasis>
</taxonomicName>
mainly differs by continuous and prominent dorsolateral folds, an X-shaped or V-shaped dorsal fold and presence of an interorbital dermal ridge (
<bibRefCitation id="E8222B35B12AADE4F9E64DEDE471E152" author="Rodriguez, LO" journalOrPublisher="Alytes" pageId="0" pageNumber="213" pagination="49 - 63" refId="B48" refString="Rodriguez, LO, 1994. A new species of the Eleutherodactylus conspicillatus group (Leptodactylidae) from Peru, with comments on its call. Alytes 12: 49 - 63" title="A new species of the Eleutherodactylus conspicillatus group (Leptodactylidae) from Peru, with comments on its call." volume="12" year="1994">
<normalizedToken id="18A092E230CA88A985FAD9BD7C3E168B" originalValue="Rodríguez">Rodriguez</normalizedToken>
1994
</bibRefCitation>
);
<taxonomicName id="D063CD3C6FE76A0DC145B6B1A0F51F37" lsidName="P. condor" pageId="0" pageNumber="213" rank="species" species="condor">
<emphasis id="9D2BA1FD6CDA8F9654A932FA86405B5D" italics="true" pageId="0" pageNumber="213">P. condor</emphasis>
</taxonomicName>
mainly differs by larger male size (SVL 32.1-39.5 mm), finger I distinctly longer than finger II, lack of basal webbing between fingers II and III, and lack of basal webbing between toes (
<bibRefCitation id="825331F2843452B7137A395D4EABFFCC" DOI="https://doi.org/10.5962/bhl.title.16222" author="Lynch, JD" journalOrPublisher="University of Kansas, Museum of Natural History, Miscellaneous Publication" pageId="0" pageNumber="213" pagination="1 - 86" refId="B34" refString="Lynch, JD, Duellman, WE, 1980. The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). University of Kansas, Museum of Natural History, Miscellaneous Publication 69: 1 - 86, DOI: https://doi.org/10.5962/bhl.title.16222" title="The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae)." url="https://doi.org/10.5962/bhl.title.16222" volume="69" year="1980">Lynch and Duellman 1980</bibRefCitation>
);
<taxonomicName id="1CD62B37DFB626FA2256DE07F8847E1D" lsidName="P. conspicillatus" pageId="0" pageNumber="213" rank="species" species="conspicillatus">
<emphasis id="BD0537FE5D2840EA180C27EAB68D748A" italics="true" pageId="0" pageNumber="213">P. conspicillatus</emphasis>
</taxonomicName>
mainly differs by finger I being distinctly longer than finger II, fingers with lateral keels, lack of basal webbing between fingers II and III, and posterior surfaces of thighs with orange spots in life (
<bibRefCitation id="95BA3DCF82F5072F74B855521A62EF6A" author="Duellman, WE" journalOrPublisher="Natur und Tier Verlag, Muenster" pageId="0" pageNumber="213" refId="B10" refString="Duellman, WE, Lehr, E, 2009. Terrestrial-breeding frogs (Strabomantidae) in Peru. Natur und Tier Verlag, Muenster" title="Terrestrial-breeding frogs (Strabomantidae) in Peru." year="2009">Duellman and Lehr 2009</bibRefCitation>
);
<taxonomicName id="CD76D63C5A2D6C39340F9003EFFBBC6D" lsidName="P. malkini" pageId="0" pageNumber="213" rank="species" species="malkini">
<emphasis id="890624EBF433193006B3594A29011800" italics="true" pageId="0" pageNumber="213">P. malkini</emphasis>
</taxonomicName>
mainly differs by slightly larger male size (SVL 30.4-37.2 mm), finger I distinctly longer than finger II, and more extensive webbing between toes (
<bibRefCitation id="B9539F4D46689841C254B6088865E60D" author="Lynch, JD" journalOrPublisher="American Museum Novitates" pageId="0" pageNumber="213" pagination="1 - 24" publicationUrl="https://digitallibrary.amnh.org/handle/2246/5429" refId="B32" refString="Lynch, JD, 1980. A taxonomic and distributional synopsis of the Amazonian frogs of the genus Eleutherodactylus. American Museum Novitates 2696: 1 - 24, https://digitallibrary.amnh.org/handle/2246/5429" title="A taxonomic and distributional synopsis of the Amazonian frogs of the genus Eleutherodactylus." url="https://digitallibrary.amnh.org/handle/2246/5429" volume="2696" year="1980">Lynch 1980</bibRefCitation>
;
<bibRefCitation id="CD002722D3D5BD7F34840E119D7D66E9" author="Duellman, WE" journalOrPublisher="Natur und Tier Verlag, Muenster" pageId="0" pageNumber="213" refId="B10" refString="Duellman, WE, Lehr, E, 2009. Terrestrial-breeding frogs (Strabomantidae) in Peru. Natur und Tier Verlag, Muenster" title="Terrestrial-breeding frogs (Strabomantidae) in Peru." year="2009">Duellman and Lehr 2009</bibRefCitation>
);
<taxonomicName id="7E19C560463CB574751B1DDA8C6FCF84" lsidName="P. meridionalis" pageId="0" pageNumber="213" rank="species" species="meridionalis">
<emphasis id="3131A247FCE9DA5977A76769ADC6CF78" italics="true" pageId="0" pageNumber="213">P. meridionalis</emphasis>
</taxonomicName>
mainly differs by finger I being shorter than finger II, dentigerous processes of vomers small and round, and fingers bearing lateral fringes (
<bibRefCitation id="E8871C9D9F863525E6365B2C94B2D8B9" author="Duellman, WE" journalOrPublisher="Natur und Tier Verlag, Muenster" pageId="0" pageNumber="213" refId="B10" refString="Duellman, WE, Lehr, E, 2009. Terrestrial-breeding frogs (Strabomantidae) in Peru. Natur und Tier Verlag, Muenster" title="Terrestrial-breeding frogs (Strabomantidae) in Peru." year="2009">Duellman and Lehr 2009</bibRefCitation>
);
<taxonomicName id="C7A90D5DA8196A9A7B61D7B69CE39E78" lsidName="P. peruvianus" pageId="0" pageNumber="213" rank="species" species="peruvianus">
<emphasis id="FCF3AF3674C0B3F46B1136E62EB67E7E" italics="true" pageId="0" pageNumber="213">P. peruvianus</emphasis>
</taxonomicName>
mainly differs by prominent and continuous dorsolateral folds, finger I distinctly longer than finger II, fingers with lateral fringes, and lack of basal webbing between toes (
<bibRefCitation id="EB0743BA3A475D72B86E4AAC65FD4A5D" author="Koehler, J" journalOrPublisher="Bonner zoologische Monographien" pageId="0" pageNumber="213" pagination="1 - 243" publicationUrl="https://www.zobodat.at/pdf/Bonner-Zoologische-Monographien_48_0001-0243.pdf" refId="B24" refString="Koehler, J, 2000. Amphibian diversity in Bolivia: a study with special reference to montane forest regions. Bonner zoologische Monographien 48: 1 - 243, https://www.zobodat.at/pdf/Bonner-Zoologische-Monographien_48_0001-0243.pdf" title="Amphibian diversity in Bolivia: a study with special reference to montane forest regions." url="https://www.zobodat.at/pdf/Bonner-Zoologische-Monographien_48_0001-0243.pdf" volume="48" year="2000">
<normalizedToken id="078DF23A9FFFF516F120BB8952AC1F1C" originalValue="Köhler">Koehler</normalizedToken>
2000
</bibRefCitation>
;
<bibRefCitation id="152036EA889BD5419765EEB7066F12E9" DOI="https://doi.org/10.1111/j.1096-3642.2008.00424.x" author="Padial, JM" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="0" pageNumber="213" pagination="97 - 122" refId="B42" refString="Padial, JM, De la Riva, I, 2009. Integrative taxonomy reveals cryptic Amazonian species of Pristimantis (Anura: Strabomantidae). Zoological Journal of the Linnean Society 155 (1): 97 - 122, DOI: https://doi.org/10.1111/j.1096-3642.2008.00424.x" title="Integrative taxonomy reveals cryptic Amazonian species of Pristimantis (Anura: Strabomantidae)." url="https://doi.org/10.1111/j.1096-3642.2008.00424.x" volume="155" year="2009">Padial and De la Riva 2009</bibRefCitation>
);
<taxonomicName id="527746165907EABD0F6F9695096F59D2" lsidName="P. skydmainos" pageId="0" pageNumber="213" rank="species" species="skydmainos">
<emphasis id="39C732A4A48D1C979CF952D6C9D6170D" italics="true" pageId="0" pageNumber="213">P. skydmainos</emphasis>
</taxonomicName>
mainly differs by more prominent and continuous dorsolateral folds, finger I being shorter than finger II, and a black mid-dorsal tubercle (
<bibRefCitation id="5746C4846F06270FBC1E28C69920510C" DOI="https://doi.org/10.2307/1447759" author="Flores, G" journalOrPublisher="Copeia" pageId="0" pageNumber="213" pagination="388 - 394" refId="B14" refString="Flores, G, Rodriguez, LO, 1997. Two new species of the Eleutherodactylus conspicillatus group (Anura: Leptodactylidae) from Peru. Copeia 1997 (2): 388 - 394, DOI: https://doi.org/10.2307/1447759" title="Two new species of the Eleutherodactylus conspicillatus group (Anura: Leptodactylidae) from Peru." url="https://doi.org/10.2307/1447759" volume="1997" year="1997">
Flores and
<normalizedToken id="8B462CDF2350017D74F7F60281A9F2FD" originalValue="Rodríguez">Rodriguez</normalizedToken>
1997
</bibRefCitation>
;
<bibRefCitation id="D54667C21DB8627AC9D39C1959331C76" DOI="https://doi.org/10.1163/156853805774806359" author="Padial, JM" journalOrPublisher="Amphibia-Reptilia" pageId="0" pageNumber="213" pagination="553 - 556" refId="B41" refString="Padial, JM, De la Riva, I, 2005. The taxonomic status of Eleutherodactylus skydmainos Flores &amp; Rodriguez, 1997, and E. karcharias Flores &amp; Rodriguez, 1997 (Anura: Leptodactylidae). Amphibia-Reptilia 26 (4): 553 - 556, DOI: https://doi.org/10.1163/156853805774806359" title="The taxonomic status of Eleutherodactylus skydmainos Flores &amp; Rodriguez, 1997, and E. karcharias Flores &amp; Rodriguez, 1997 (Anura: Leptodactylidae)." url="https://doi.org/10.1163/156853805774806359" volume="26" year="2005">Padial and De la Riva 2005</bibRefCitation>
). Superficially,
<taxonomicName id="4658C1F116490D827CF31B6692C5BD64" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="8533A7BB632CEBF7A3B5F956DDFFB33C" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
is similar to
<taxonomicName id="B128F143984C014A25FE9EB692537071" lsidName="P. iiap" pageId="0" pageNumber="213" rank="species" species="iiap">
<emphasis id="B51DEAA818E919EA3D70F489FF626383" italics="true" pageId="0" pageNumber="213">P. iiap</emphasis>
</taxonomicName>
, with which it shares similar adult male size, inconspicuous and partly interrupted dorsolateral folds, finger I only very slightly longer or equal in length to finger II, presence of postrictal tubercles, and similar colour pattern (
<bibRefCitation id="C8AA26984ECD7B471B3A95CB1DB8A310" DOI="https://doi.org/10.2992/007.083.0302" author="Padial, JM" journalOrPublisher="Annals of Carnegie Museum" pageId="0" pageNumber="213" pagination="207 - 218" refId="B45" refString="Padial, JM, Gagliardi-Urrutia, G, Chaparro, JC, Gutierrez, RC, 2016. A new species of the Pristimantis conspicillatus group from the Peruvian Amazon (Anura: Craugastoridae). Annals of Carnegie Museum 83 (3): 207 - 218, DOI: https://doi.org/10.2992/007.083.0302" title="A new species of the Pristimantis conspicillatus group from the Peruvian Amazon (Anura: Craugastoridae)." url="https://doi.org/10.2992/007.083.0302" volume="83" year="2016">Padial et al. 2016</bibRefCitation>
). However,
<taxonomicName id="0DBE4DE21B4C4E7DAE7043EAAC8ACCDC" lsidName="P. iiap" pageId="0" pageNumber="213" rank="species" species="iiap">
<emphasis id="B9E75981DC3EDA977216504679C3DBCD" italics="true" pageId="0" pageNumber="213">P. iiap</emphasis>
</taxonomicName>
mainly differs by the lack of basal webbing between fingers II and III (present in
<taxonomicName id="2AAC8F51A118B4BA57F8E494EE9F8B0A" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="4C71218D5AD735867CAF0053643F2019" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
), lack of basal toe webbing (present), upper eyelid with small granules (absent), a copper-coloured iris in life (bronze), and advertisement call (
<bibRefCitation id="E701F9E9C80A08A96863DEA64BF7919D" DOI="https://doi.org/10.2992/007.083.0302" author="Padial, JM" journalOrPublisher="Annals of Carnegie Museum" pageId="0" pageNumber="213" pagination="207 - 218" refId="B45" refString="Padial, JM, Gagliardi-Urrutia, G, Chaparro, JC, Gutierrez, RC, 2016. A new species of the Pristimantis conspicillatus group from the Peruvian Amazon (Anura: Craugastoridae). Annals of Carnegie Museum 83 (3): 207 - 218, DOI: https://doi.org/10.2992/007.083.0302" title="A new species of the Pristimantis conspicillatus group from the Peruvian Amazon (Anura: Craugastoridae)." url="https://doi.org/10.2992/007.083.0302" volume="83" year="2016">Padial et al. 2016</bibRefCitation>
; see below). The closely related species
<taxonomicName id="8AA8468442B0B11111C374536463F72D" lsidName="P. bipunctatus" pageId="0" pageNumber="213" rank="species" species="bipunctatus">
<emphasis id="02EBFA7724F1F352A257689D459C5E24" italics="true" pageId="0" pageNumber="213">P. bipunctatus</emphasis>
</taxonomicName>
also shares many characters with
<taxonomicName id="5148DF1CDFF7FDA27B46F0DACBB1D8FC" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="9CFC32E5C7DAA134FF6FC2CCE9DEC075" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
, such as similar male size, dorsolateral folds present but inconspicuous, finger I about equal in length to finger II, upper eyelid lacking tubercles or granules, and similar colour pattern. However, it differs from
<taxonomicName id="8E03100B954A977076F94004CF8045C2" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="4730536CDFDA2D2A30826234F09EC4B3" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
by the lack of basal webbing between fingers II and III (present), bluntly rounded snout in dorsal view (subacuminate), truncate in profile (rounded), basal webbing between toes IV and V (basal webbing between all toes) (
<bibRefCitation id="540E2FD5102B172026B89593753F59EB" DOI="https://doi.org/10.1643/CH-05-019R" author="Duellman, WE" journalOrPublisher="Copeia" pageId="0" pageNumber="213" pagination="526 - 538" refId="B8" refString="Duellman, WE, Hedges, SB, 2005. Eleutherodactyline frogs (Anura: Leptodactylidae) from the Cordillera Yanachaga in Central Peru. Copeia 2005 (3): 526 - 538, DOI: https://doi.org/10.1643/CH-05-019R" title="Eleutherodactyline frogs (Anura: Leptodactylidae) from the Cordillera Yanachaga in Central Peru." url="https://doi.org/10.1643/CH-05-019R" volume="2005" year="2005">Duellman and Hedges 2005</bibRefCitation>
;
<bibRefCitation id="CC7BEEE3DBC7B55BEB8F0FA310E2ED52" author="Duellman, WE" journalOrPublisher="Natur und Tier Verlag, Muenster" pageId="0" pageNumber="213" refId="B10" refString="Duellman, WE, Lehr, E, 2009. Terrestrial-breeding frogs (Strabomantidae) in Peru. Natur und Tier Verlag, Muenster" title="Terrestrial-breeding frogs (Strabomantidae) in Peru." year="2009">Duellman and Lehr 2009</bibRefCitation>
), and considerable divergence in the mitochondrial 16S gene. According to
<bibRefCitation id="476DFFA7C32E40E5809499FCD1B4F101" author="Duellman, WE" journalOrPublisher="Natur und Tier Verlag, Muenster" pageId="0" pageNumber="213" refId="B10" refString="Duellman, WE, Lehr, E, 2009. Terrestrial-breeding frogs (Strabomantidae) in Peru. Natur und Tier Verlag, Muenster" title="Terrestrial-breeding frogs (Strabomantidae) in Peru." year="2009">Duellman and Lehr (2009)</bibRefCitation>
,
<taxonomicName id="AFB5FB9D626C966F4C9CA3ACD3A3CC69" lsidName="P. adiastolus" pageId="0" pageNumber="213" rank="species" species="adiastolus">
<emphasis id="846DB0F1BF91D7C55A9EC479416FA22F" italics="true" pageId="0" pageNumber="213">P. adiastolus</emphasis>
</taxonomicName>
is morphologically cryptic to
<taxonomicName id="CE3BD5AAE0D62D26B8A0189D737B03A1" lsidName="P. bipunctatus" pageId="0" pageNumber="213" rank="species" species="bipunctatus">
<emphasis id="A0C5096369FCA010E6DEE66989893008" italics="true" pageId="0" pageNumber="213">P. bipunctatus</emphasis>
</taxonomicName>
. However,
<taxonomicName id="7313B90B3168A853C8A2C83DA49BFD1F" lsidName="P. adiastolus" pageId="0" pageNumber="213" rank="species" species="adiastolus">
<emphasis id="19B74860C953E43D7E426F9EE9C6C481" italics="true" pageId="0" pageNumber="213">P. adiastolus</emphasis>
</taxonomicName>
mainly differs from
<taxonomicName id="D16E52AEB2210D52A66A05740C4B0468" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="49CABF3F7E914D4E549D87963395CFFF" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
by more prominent and continuous dorsolateral folds, lack of basal webbing between fingers, and finger I slightly shorter than finger II (
<bibRefCitation id="9AF8E8C9517A8B10E2F9822A2BFB17DA" DOI="https://doi.org/10.11606/issn.2316-9079.v6i2p119-135" author="Duellman, WE" journalOrPublisher="Phyllomedusa" pageId="0" pageNumber="213" pagination="119 - 135" refId="B9" refString="Duellman, WE, Hedges, SB, 2007. Three new species of Pristimantis (Lissamphibia, Anura) from montane forests of the Cordillera Yanachaga in central Peru. Phyllomedusa 6 (2): 119 - 135, DOI: https://doi.org/10.11606/issn.2316-9079.v6i2p119-135" title="Three new species of Pristimantis (Lissamphibia, Anura) from montane forests of the Cordillera Yanachaga in central Peru." url="https://doi.org/10.11606/issn.2316-9079.v6i2p119-135" volume="6" year="2007">Duellman and Hedges 2007</bibRefCitation>
). Individuals of the recently described species
<taxonomicName id="0A6B629E6F35E30C36F63F363FBAB8CC" lsidName="P. pictus" pageId="0" pageNumber="213" rank="species" species="pictus">
<emphasis id="85993C1D962BF40F352C8E96239927EB" italics="true" pageId="0" pageNumber="213">P. pictus</emphasis>
</taxonomicName>
,
<taxonomicName id="F30DCF378EF2DD1A4C7C779683B5677B" lsidName="P. pluvian" pageId="0" pageNumber="213" rank="species" species="pluvian">
<emphasis id="E7651B338411CE57E80EAEBD46AA6AEA" italics="true" pageId="0" pageNumber="213">P. pluvian</emphasis>
</taxonomicName>
and
<taxonomicName id="A3DE69E14A70044BE33337A70F132730" lsidName="P. moa" pageId="0" pageNumber="213" rank="species" species="moa">
<emphasis id="358426DD34EE9C2101A51422D50F3460" italics="true" pageId="0" pageNumber="213">P. moa</emphasis>
</taxonomicName>
from Amazonian Brazil may sometimes exhibit weakly developed dorsolateral folds, but all differ from
<taxonomicName id="D0F53CCC461716CB8875C92364B3B068" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="19F090CF94A48CD3BC7CDB77E3127AEE" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
by smooth mid-venter (versus areolate) and advertisement call (
<bibRefCitation id="0747543EE3070D91DC905042FC0F2DC8" DOI="https://doi.org/10.1371/journal.pone.0229971" author="de Oliveira, EA" journalOrPublisher="ZooKeys" pageId="0" pageNumber="213" refId="B7" refString="de Oliveira, EA, da Silva, LA, Silva, EAP, Guimaraes, KLA, Penhacek, M, Martinez, JG, Rodrigues, LR, Santana, DJ, Hernandez-Ruz, EJ, 2020. Four new species of Pristimantis Jimenez de la Espada, 1870 (Anura: Craugastoridae) in the eastern Amazon. PLoS ONE 15(11): e0243182. https://doi.org/10.1371/journal.pone.0229971" title="Four new species of Pristimantis Jimenez de la Espada, 1870 (Anura: Craugastoridae) in the eastern Amazon. PLoS ONE 15 (11): e 0243182." url="https://doi.org/10.1371/journal.pone.0229971" year="2020">de Oliveira et al. 2020</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="0F506222F1B8981FDED74BFCA8BD0AF4" pageId="0" pageNumber="213" type="natural history">
<paragraph id="C67948977556569DDA369ED72CFC5A2B" pageId="0" pageNumber="213">Natural history.</paragraph>
<paragraph id="3B421C04C120433C5FBB47686B988A74" pageId="0" pageNumber="213">
<taxonomicName id="3577B5007F012BAE26F13CEB4C34C721" baseAuthorityName="Henle" baseAuthorityYear="1992" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis nebulosus" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="species" species="nebulosus">
<emphasis id="EF31E99D5615C48729D9E23975BE5A97" italics="true" pageId="0" pageNumber="213">Pristimantis nebulosus</emphasis>
</taxonomicName>
occurs in an area of evergreen montane rainforest of moderate height. Forest grows at moderate to steeply sloped terrain (Fig.
<figureCitation id="D1C35F54BD26F92F53C6CF94D48B906B" captionStart="Figure 5" captionStartId="F6" captionText="Figure 5. View of the forest at the type locality of Pristimatis nebulosus at Abra La Divisoria, southern Cordillera Azul, Departamento Huanuco, Peru, at 1650 m a. s. l. Photo taken on 8 November 2019." figureDoi="10.3897/zse.98.84963.figure5" httpUri="https://binary.pensoft.net/fig/705932" pageId="0" pageNumber="213">5</figureCitation>
). See also
<bibRefCitation id="B51D876FF8ECFC4851195112E98D13DF" author="Myers, CW" journalOrPublisher="American Museum Novitates" pageId="0" pageNumber="213" pagination="1 - 24" publicationUrl="https://digitallibrary.amnh.org/handle/2246/5326" refId="B38" refString="Myers, CW, Daly, JW, 1979. A name for the poison frog of Cordillera Azul, eastern Peru, with notes on its biology and skin toxins (Dendrobatidae). American Museum Novitates 2674: 1 - 24, https://digitallibrary.amnh.org/handle/2246/5326" title="A name for the poison frog of Cordillera Azul, eastern Peru, with notes on its biology and skin toxins (Dendrobatidae)." url="https://digitallibrary.amnh.org/handle/2246/5326" volume="2674" year="1979">Myers and Daly (1979)</bibRefCitation>
for a detailed description of the area. The area has been heavily altered by different kinds of plantations (e.g. tea). Individuals were found in disturbed habitat along a narrow dirt road. Males were calling from a bushy area during light rain at night, one individual called from near the ground, another one was sitting on a leaf about 30 cm above the ground. The species was found in syntopy with
<taxonomicName id="9B81501D22FFA75BF874B269C9ECF822" authorityName="Jimenez de la Espada" authorityYear="1870" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="genus">
<emphasis id="E28F8242485D59A9C654C9853ADAB9B7" italics="true" pageId="0" pageNumber="213">Pristimantis</emphasis>
</taxonomicName>
sp. (
<taxonomicName id="6DC5DD5EFC35B0B2182E2693D23B31F3" authorityName="Lehr &amp; von May" authorityYear="2009" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis divnae" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="species" species="divnae">Pristimantis aff. divnae</taxonomicName>
),
<taxonomicName id="10CE5AB5631FD0A34F96AEBB10F441A0" genus="P." lsidName="P." pageId="0" pageNumber="213" rank="genus">
<emphasis id="7860ACA190CC6D31A5504A29CD44C419" italics="true" pageId="0" pageNumber="213">P.</emphasis>
</taxonomicName>
sp. (
<taxonomicName id="6BEFA1E1A44262133CA911C486D37C32" baseAuthorityName="Jimenez de la Espada" baseAuthorityYear="1875" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis lacrimosus" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="species" species="lacrimosus">
<emphasis id="B17A6F573F3041E88D8609B04F0A5CA3" italics="true" pageId="0" pageNumber="213">Pristimantis lacrimosus</emphasis>
</taxonomicName>
group),
<taxonomicName id="9B29EAA0A5406738732ECC2130C9A56D" authorityName="Fitzinger" authorityYear="1826" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="genus">
<emphasis id="B2332735128FEC8D2CA1D7DA83A646E9" italics="true" pageId="0" pageNumber="213">Rhinella</emphasis>
</taxonomicName>
sp. (
<emphasis id="5EF9ECA7CD3587F1EE77432FD132CF8D" italics="true" pageId="0" pageNumber="213">margaritifera</emphasis>
group),
<taxonomicName id="CB1B842EBFF6249D6D5FC566D4500B72" baseAuthorityName="Cope" baseAuthorityYear="1871" class="Amphibia" family="Hylidae" genus="Boana" kingdom="Animalia" lsidName="Boana lanciformis" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="species" species="lanciformis">
<emphasis id="76A2D05578BDFE1F76701AD24D20CE53" italics="true" pageId="0" pageNumber="213">Boana lanciformis</emphasis>
</taxonomicName>
,
<taxonomicName id="E2C78FA23B9147E7A709A0BBD46BBE8B" baseAuthorityName="Duellman" baseAuthorityYear="1982" class="Amphibia" family="Hylidae" genus="Dendropsophus" kingdom="Animalia" lsidName="Dendropsophus aperomeus" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="species" species="aperomeus">
<emphasis id="90681C267448EC0F28E779643ACAFF7B" italics="true" pageId="0" pageNumber="213">Dendropsophus aperomeus</emphasis>
</taxonomicName>
,
<taxonomicName id="13CB2AD978A746ECC40986E3218D34CE" baseAuthorityName="Miranda-Ribeiro" baseAuthorityYear="1926" class="Amphibia" family="Hylidae" genus="Scinax" kingdom="Animalia" lsidName="Scinax garbei" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="species" species="garbei">
<emphasis id="473F0B27337393239DFEDD57A4E7C8BF" italics="true" pageId="0" pageNumber="213">Scinax garbei</emphasis>
</taxonomicName>
,
<taxonomicName id="FC004E6910AE2F3AEA0FA9D99E244846" genus="S." lsidName="S." pageId="0" pageNumber="213" rank="genus">
<emphasis id="F05B2D9ABB6A07C0F739C5CB98F94536" italics="true" pageId="0" pageNumber="213">S.</emphasis>
</taxonomicName>
sp. (
<taxonomicName id="35FB89236DCD4AAA84C958A4B3078E26" baseAuthorityName="Laurenti" baseAuthorityYear="1768" class="Amphibia" family="Hylidae" genus="Scinax" kingdom="Animalia" lsidName="Scinax ruber" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="species" species="ruber">Scinax aff. ruber</taxonomicName>
) and
<taxonomicName id="ACEB642177339E12A5DDEC8BE044100B" authorityName="Steindachner" authorityYear="1867" class="Amphibia" family="Leptodactylidae" genus="Adenomera" kingdom="Animalia" lsidName="Adenomera" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="genus">
<emphasis id="E3E4F5837D792F0E6B1DB77F523F628C" italics="true" pageId="0" pageNumber="213">Adenomera</emphasis>
</taxonomicName>
sp.
</paragraph>
<caption id="2A2D5E09AE2B2D3FFD631C87AB61623D" doi="10.3897/zse.98.84963.figure5" httpUri="https://binary.pensoft.net/fig/705932" pageId="0" pageNumber="213" start="Figure 5" startId="F6">
<paragraph id="E460C1EFFC4003845D1FC71603F837B9" pageId="0" pageNumber="213">
<emphasis id="9EA23425FAB34751698EDEF615B16C4F" bold="true" pageId="0" pageNumber="213">Figure 5.</emphasis>
View of the forest at the type locality of
<taxonomicName id="7D0F8505B59361AFADBF571D952C541F" genus="Pristimatis" lsidName="Pristimatis nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="800433B87E663DCFFF57151ED2E49748" italics="true" pageId="0" pageNumber="213">Pristimatis nebulosus</emphasis>
</taxonomicName>
at Abra La Divisoria, southern Cordillera Azul, Departamento
<normalizedToken id="7E0A5432E89BE250333FC22F12CD4236" originalValue="Huánuco">Huanuco</normalizedToken>
, Peru, at 1650 m a.s.l. Photo taken on 8 November 2019.
</paragraph>
</caption>
<caption id="DE0E168D8C7FA1A2BD57FADC32B50D94" doi="10.3897/zse.98.84963.figure6" httpUri="https://binary.pensoft.net/fig/705933" pageId="0" pageNumber="213" start="Figure 6" startId="F5">
<paragraph id="F543B68F76BF6563243C6D5645808B09" pageId="0" pageNumber="213">
<emphasis id="D2AE70C357A674453237B974E7AD32A4" bold="true" pageId="0" pageNumber="213">Figure 6.</emphasis>
On top, audiospectrogram and corresponding oscillogram of one advertisement call of a topotypic
<taxonomicName id="3D9424F770F28BE66566C276A79E6768" baseAuthorityName="Henle" baseAuthorityYear="1992" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis nebulosus" order="Anura" pageId="0" pageNumber="213" phylum="Chordata" rank="species" species="nebulosus">
<emphasis id="CA199C8CB505483FC3DD4D481E2E0999" italics="true" pageId="0" pageNumber="213">Pristimantis nebulosus</emphasis>
</taxonomicName>
from Abra la Divisoria, Cordillera Azul, 1650 m, Peru, at 1000 ms time scale. Below, an oscillogram at 8000 ms time scale showing one complete call series consisting of 7 calls of the same individual. Recording band-pass filtered at 450-9600 Hz.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="AF568423A045360AA1AF1DDA1CA3A468" pageId="0" pageNumber="213" type="vocalisation">
<paragraph id="BA7FE57628052B9A485BE1607F9E8FA7" pageId="0" pageNumber="213">Vocalisation.</paragraph>
<paragraph id="45DDD11A5F352F0EF28D1DEE23E1A60D" pageId="0" pageNumber="213">
Calls were recorded on 8 November 2019, around 20:00 h, at Abra la Divisoria, Departamento
<normalizedToken id="EEA2F5E9B6017DBF477742DC05861DD7" originalValue="Huánuco">Huanuco</normalizedToken>
, 1650 m a.s.l. (air temperature estimated 18 °C). The recorded individual could not be observed calling, but searching at the spot of sound emission revealed an individual of
<taxonomicName id="02EA61BDF88B580289E7C1E953CB326A" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="A26AE4AFF66E92C24BC294AB7F51C662" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
, leaving little doubt that recorded calls actually correspond to this species. The advertisement call consists of a single regularly pulsed note, repeated at a rather regular succession in short call series (Fig.
<figureCitation id="B29361AF74E49150A842AE6C1F696822" captionStart="Figure 6" captionStartId="F5" captionText="Figure 6. On top, audiospectrogram and corresponding oscillogram of one advertisement call of a topotypic Pristimantis nebulosus from Abra la Divisoria, Cordillera Azul, 1650 m, Peru, at 1000 ms time scale. Below, an oscillogram at 8000 ms time scale showing one complete call series consisting of 7 calls of the same individual. Recording band-pass filtered at 450 - 9600 Hz." figureDoi="10.3897/zse.98.84963.figure6" httpUri="https://binary.pensoft.net/fig/705933" pageId="0" pageNumber="213">6</figureCitation>
). Call series consist of seven calls (n = 2), with the initial call being the longest in duration and separated from the remaining calls of the series by a longer inter-call interval with about two times the duration of other inter-call intervals. Pulses within calls (= notes) are broadly fused, but fairly countable in the oscillogram. Apart from the pulse structure, there is amplitude modulation within each call, with the greatest amplitude occurring at the beginning of the call, then decreasing to about half the initial amplitude at about one third of the
<normalizedToken id="35EA6A739B18DC080EC5ACFE4FFBF086" originalValue="call’s">call's</normalizedToken>
duration and rapidly decreasing to zero at the
<normalizedToken id="AC716BA7BA7A3A351B1E4EDF2617617B" originalValue="call’s">call's</normalizedToken>
end, giving the call a bell-shaped appearance in the oscillogram. Numerical parameters of 14 analysed calls (two call series) of one male are as follows: call duration (= note duration) 206-382 ms (254.1
<normalizedToken id="0001F2FCF368B210AD9CEED9D799374D" originalValue="±">+/-</normalizedToken>
40.3 ms); inter-call interval within call series 507-1730 ms (796.1
<normalizedToken id="833A4BB7B766395A52517BD41E33D346" originalValue="±">+/-</normalizedToken>
348.8 ms); pulses per call 33-37 (35.4
<normalizedToken id="339F9AE5241131857E1672CB050F8910" originalValue="±">+/-</normalizedToken>
1.5); pulses per second 156-174 (162.8
<normalizedToken id="9CB0DBF78D9FE092DD11ADF9AB50EA03" originalValue="±">+/-</normalizedToken>
7.0); dominant frequency 3186-3359 Hz (3230
<normalizedToken id="3101765602ACF2BE56FE8B9003561663" originalValue="±">+/-</normalizedToken>
57 Hz); prevalent bandwidth 1500-6600 Hz, but bands of very low call energy recognisable up to 9600 Hz. Apart from the dominant frequency band, additional frequency bands, apparently induced by the pulse rate (not harmonics; see
<bibRefCitation id="87AAB5F90559E1F9309BCFFCAEC05538" DOI="https://doi.org/10.11646/zootaxa.4251.1.1" author="Koehler, J" journalOrPublisher="Zootaxa" pageId="0" pageNumber="213" pagination="1 - 124" refId="B26" refString="Koehler, J, Jansen, M, Rodriguez, A, Kok, PJR, Toledo, LF, Emmrich, M, Glaw, F, Haddad, CFB, Roedel, MO, Vences, M, 2017. The use of bioacoustics in anuran taxonomy: theory, terminology, methods and recommendations for best practice. Zootaxa 4251 (1): 1 - 124, DOI: https://doi.org/10.11646/zootaxa.4251.1.1" title="The use of bioacoustics in anuran taxonomy: theory, terminology, methods and recommendations for best practice." url="https://doi.org/10.11646/zootaxa.4251.1.1" volume="4251" year="2017">
<normalizedToken id="2C671D64F65E941EAC787B0758DE4A4B" originalValue="Köhler">Koehler</normalizedToken>
et al. 2017
</bibRefCitation>
), occur at approximately 1400-1900, 4400-5000, 6000-6600, and 7900-8400 Hz, respectively. Within regular call series, calls were repeated at a rate of 54-72 calls/minute. Duration of call series was 7.13 and 6.05 seconds, respectively.
</paragraph>
</subSubSection>
<subSubSection id="A43957AE553C69B9661883C5DD44C68C" pageId="0" pageNumber="213" type="distribution">
<paragraph id="44491C03FE497BB6532AAA4306C35E97" pageId="0" pageNumber="213">Distribution.</paragraph>
<paragraph id="17DBFC8B38E57D7B8471C35F32EA7AF0" pageId="0" pageNumber="213">
So far,
<taxonomicName id="D5F2CE3CF522DA58F691CC8718E220EC" lsidName="P. nebulosus" pageId="0" pageNumber="213" rank="species" species="nebulosus">
<emphasis id="C55A18C596DF3AFC8DFBAA1699C4B65B" italics="true" pageId="0" pageNumber="213">P. nebulosus</emphasis>
</taxonomicName>
is only known from its type locality in the southern Cordillera Azul, at the border of the Departamentos
<normalizedToken id="949BA18E29FAC874F6C40964D8552724" originalValue="Huánuco">Huanuco</normalizedToken>
and Ucayali. However, it is very likely that the species occupies a wider range, at least along the same elevational corridor within the Cordillera Azul. Possibly, there are additional records of this species represented by unidentified or misidentified specimens in scientific collections.
</paragraph>
</subSubSection>
</treatment>
</document>
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